Heritability of corticosterone response and changes in life history traits during selection in the zebra finch

Vertebrates respond to environmental stressors through the neuro-endocrine stress response, which involves the production of glucocorticoids. We have selected independent, duplicate divergent lines of zebra finches for high, low and control corticosterone responses to a mild stressor. This experiment has shown that over the first four generations, the high lines have demonstrated a significant realized heritability of about 20%. However, the low lines have apparently not changed significantly from controls. This asymmetry in response is potentially because of the fact that all birds appear to be showing increased adaptation to the environment in which they are housed, with significant declines in corticosterone response in control lines as well as low lines. Despite the existence of two- to threefold difference in mean corticosterone titre between high and low lines, there were no observed differences in testosterone titre in adult male birds from the different groups. In addition, there were no consistent, significant differences between the lines in any of the life history variables measured--number of eggs laid per clutch, number of clutches or broods produced per pair, number of fledglings produced per breeding attempt, nor in any of egg, nestling and fledgling mortality. These results highlight the fact that the mechanisms that underlie variation in the avian physiological system can be modified to respond to differences between environments through selection. This adds an additional level of flexibility to the avian physiological system, which will allow it to respond to environmental circumstances.     

Genetic and environmental influences on natal dispersal distance in a resident bird species

We analyzed more than 1,600 dispersal events from two populations of a North American cooperatively breeding woodpecker species to determine what factors influence natal dispersal distance and whether distance traveled affects reproduction later in life. We found significant heritability of natal dispersal distance, in both males and females, indicating substantial additive genetic variance for this behavioral trait. Natal dispersal distance additionally was affected by social and ecological factors: individuals dispersing in their first year of life moved longer distances than those staying on their natal site as helpers for a prolonged time prior to dispersal, and increasing territory isolation led to longer dispersal distances. Successful dispersers incurred fitness costs, with lifetime fledgling production (in both sexes) and lifetime production of recruits to the breeding population (in females only) decreasing with increasing natal dispersal distance. We conclude that natal dispersal distance has a genetic basis but is modulated by environmental and social factors and that natal dispersal distance in this species is (currently) under selection.     

Small‐scale spatial variation in evolvability for life‐history traits in the storm petrel

The evolutionary potential in the timing of recruitment and reproduction may be crucial for the ability of populations to buffer against environmental changes, allowing them to avoid unfavourable breeding conditions. The evolution of a trait in a local population is determined by its heritability and selection. In the present study, we performed pedigree‐based quantitative genetic analyses for two life‐history traits (recruiting age and laying date) using population data of the storm petrel over an 18‐year period in two adjacent breeding colonies (only 150 m apart) that share the same environmental conditions. In both traits, natal colony effect was the main source of the phenotypic variation among individuals, and cohort variance for recruitment age and additive genetic variance for laying date were natal colony‐specific. We found significant heritability only in laying date and, more specifically, only in birds born in one of the colonies. The difference in genetic variance between the colonies was statistically significant. Interestingly, selection on earlier breeding birds was detected only in the colony in which heritable variation in laying date was found. Therefore, local evolvability for a life‐history trait may vary within a unexpectedly small spatial scale, through the diversifying natural selection and insulating gene flow. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 439–446.     

Intraseasonal variation in immune defence,body mass and hematocrit in adult house martins Delichon urbica

The intensity of parasite infections often increases during the reproductive season of the host as a result of parasite reproduction, increased parasite transmission and increased host susceptibility. We report within‐individual variation in immune parameters, hematocrit and body mass in adult house martins Delichon urbica rearing nestlings in nests experimentally infested with house martin bugs Oeciacus hirundinis and birds rearing nestlings in initially parasite‐free nests. From first to second broods body mass and hematocrit of breeding adult house martins decreased. In contrast leucocytes and immunoglobulins became more abundant. When their nests were infested with ectoparasites adults lost more weight compared with birds raising nestlings in nests treated with pyrethrin, whereas the decrease in hematocrit was more pronounced during infection with blood parasites. Neither experimental infestation with house martin bugs nor blood parasites had a significant effect on the amount of immune defences.     

Genetic and environmental effects on morphology and fluctuating asymmetry in nestling barn swallows

A barn swallow Hirundo rustica partial cross‐fostering experiment with simultaneous brood size manipulation was conducted in two years with contrasting weather conditions, to estimate heritable variation in tarsus, tail and wing size and fluctuating asymmetry. Environmental stress had contrasting effects depending on trait type. Significant heritabilities for tarsus, tail and wing size were found only in enlarged broods irrespective of year effects, while tarsus asymmetry was significantly heritable in the year with benign weather conditions irrespective of brood size manipulation effects. Tail, wing and composite (multicharacter) asymmetry were never significantly heritable. The environment with the higher heritability generally had higher additive genetic variance and lower environmental variance, irrespective of trait type. Heritability was larger for trait size than for trait asymmetry. Patterns of genetic variation in nestlings do not necessarily translate to the juvenile or adult stage, as indicated by lack of correlation between nestling and fledgling traits.     

Gender and viability selection on morphology in fledgling pied flycatchers

Until recently, analyses of gender-dependent differences in viability selection and the ontogeny of sexual size dimorphism have been plagued by difficulties in determining the sex of nestling birds on the basis of morphology. Recently, this problem was overcome using molecular sex identification to report for the first time body-size-mediated antagonistic selection on the viability of male and female collared flycatchers. We used molecular sex identification to analyse natural selection on fledgling viability, sexual size dimorphism and effects of parasites in relation to gender in a Mediterranean population of the related pied flycatcher Ficedula hypoleuca. There was directional positive selection on fledgling weight but no selection on tarsus length. Fledgling weight was the most important determinant of fledgling survival, with heavier fledglings having increased viability. Although selective trends were of the same sign for both sexes, only among female fledglings were selection differentials and gradients statistically significant. Therefore, similar trends in selection were revealed in analyses of a data set where sex was ignored and in separate analyses using same-sex sibship trait means. Mite nest ectoparasites negatively affected fledgling weight, and the effects were stronger in female than male fledglings. There was no effect of parasitism on the tarsus length in males, as previously reported in retrospective analyses performed without knowledge of sex until recruitment. Overall, selection on fledgling viability on the basis of morphological traits and hatching date was not confounded by an individual's gender.     

Phenotypic selection on a heritable size trait revisited

One of evolutionary biology's most persistent puzzles is the fact that apparent directional selection on a heritable trait in a natural population often does not produce a selection response. We tested three possible explanations for this problem using data on body size of more than 23,000 individuals, measured over 18 yr, in a collared flycatcher (Ficedula albicollis) population. Using a restricted maximum likelihood "animal model," we found a narrow-sense heritability for fledgling tarsus length of [Formula: see text] SE and substantial common environment effects ([Formula: see text] SE). For survival to adulthood, the selection differential on tarsus length was [Formula: see text] SE. There was, however, no response to this selection over the study period. One explanation for the lack of response might have been that selection was associated with only the environmental (nonheritable) component of phenotype, but we found significant selection on breeding values (the heritable component). There was also no evidence of fluctuating selection pressures or of antagonistic selection between the sexes in selection pressures. Thus, in contrast to earlier investigations in this same population, none of the potential explanations for the absence of a selection response was supported; we discuss alternative hypotheses yet to be investigated.     

Comparing food limitation among three stages of nesting: supplementation experiments with the burrowing owl

Food availability is an important limiting factor for avian reproduction. In altricial birds, food limitation is assumed to be more severe during the nestling stage than during laying or incubation, but this has yet to be adequately tested. Using food‐supplementation experiments over a 5‐year period, we determined the degree and timing of food limitation for burrowing owls (Athene cunicularia) breeding in Canada. Burrowing owls are an endangered species and food limitation during the nestling stage could influence reproductive performance of this species at the northern extent of their range. Supplemented pairs fledged on average 47% more owlets than unfed pairs, except during a year when natural food was not limiting (i.e., a prey irruption year). The difference in fledgling production resulted from high nestling mortality in unfed broods, with 96% of all nestling deaths being attributed to food shortage. Supplemental feeding during the nestling period also increased fledgling structural size. Pairs fed from the start of laying produced the same number of hatchlings as pairs that received no supplemental food before hatch. Furthermore, pairs supplemented from egg laying to fledging and pairs supplemented during the nestling period alone had the same patterns of nestling survival, equal numbers of fledglings, and similar fledgling mass and structural size. Our results provide empirical support for the hypothesis that the nestling period is the most food‐limited phase of the breeding cycle. The experimental design we introduce here could be used with other altricial species to examine how the timing of food limitation differs among birds with a variety of life‐history strategies. For burrowing owls, and other species with similar life histories, long‐term, large‐scale, and appropriately timed habitat management increasing prey abundance or availability is critical for conservation.     

Effects of common origin and common environment on nestling plumage coloration in the great tit (Parus major)

Abstract.— Carotenoids cannot be synthesized by birds and thus have to be ingested with food, suggesting that ca-rotenoid-based plumage coloration is environmentally determined. However signaling functions ascribed to plumage imply that plumage coloration is the outcome of an evolutionary process based on genetic variation. By means of a cross-fostering design we show significant effects of both a common rearing environment and the brood from which a nestling originally came from (common origin) on the plumage coloration of nestling great tits ( Parus major ). This demonstration of origin-related variation in carotenoid-based plumage coloration suggests that the observed variation of the trait has a partial genetic basis. Consistent with environmental determination of this trait, we also found a significant positive correlation between the color saturation of nestlings and their foster-father's plumage. There was no significant correlation between nestling plumage coloration and the food quantity provided to the nestlings by the male, the female, or both parents. This suggests that the nestling-foster father correlation arises by the carotenoid quantity ingested rather than the food quantity per se. No significant nestling-true father correlation was found, which suggests that nestling plumage coloration did not indirectly evolve due to sexual selection. Consistent with this result there was no significant correlation between the nestling's plumage color and its coloration as a breeding adult the following year, suggesting that nestling plumage color is a different trait than the first year plumage.     

Age-related fledgling production in great cormorants Phalacrocorax carbo: influence of individual competence and disappearance of phenotypes

Age-related reproductive performance of great cormorants Phalacrocorax carbo sinensis was studied in a tree nesting colony in Denmark in relation to age-related improvements of competence and progressive disappearance of phenotypes. Within-individual changes in fledgling production were measured, and cross-sectional analyses were applied. The within-individual analyses showed that competence improved with age and/or that individuals showed restraint to optimize their reproductive effort. The within-individual improvements were three to six times higher among individuals that survived and returned to breed beyond the fourth breeding attempt than among individuals disappearing from the breeding population before the fourth breeding attempt. Taking this into account the within-individual improvements explained 70–90% of the age-effect observed in the population over the youngest ages. Effects of breeding experience were significant for females, but only within the group of individuals that were present in the breeding population beyond the age of five years. In males, improvements arose because of unknown factors related to age. Individual great cormorants that bred beyond the age of five years had higher reproductive success, on average, than birds disappearing from the breeding population earlier in life. This supports the differential survival hypothesis. However, the effect on the population mean was partly counterbalanced by late recruitment of other inferior breeders. It is concluded that the enhancement in fledgling production with increasing age was primarily an effect of age-related improvements of competence and secondly an effect of progressive disappearance of phenotypes.     

Variation in heritability of immune function in the tree swallow

Heritability is an important component of the ability of a trait to respond to natural selection; variation in heritability can lead to differences in how a trait responds to selection pressures. Here we test whether an important physiological trait, immune function, varies by comparing heritability estimates through cross-fostering brood manipulation at three wide-spread sites in the tree swallow (Tachycineta bicolor): Alaska, New York and Tennessee. In two of three sites, there was no additive genetic component to nestling immune response to the mitogen phytohaemagglutinin, while immune response had a heritable component in Tennessee. Bootstrapping revealed significant differences in estimated heritability. This conclusion was supported by mother–offspring regressions; in Tennessee breeding females mounting strong immune responses tended to have offspring with strong immune responses, while in New York and Alaska, there was no relationship between the immune responses of mothers and offspring. These results suggest that studies investigating the roles of common origin and rearing environment should consider yearly or spatial variation within a species.     

Breeding performance and survival in the peregrine falcon Falco peregrinus support an age‐related competence improvement hypothesis mediated via an age threshold

We studied the effects of age on breeding performance and survival probability in a peregrine falcon population, using data from a long term monitoring programme (carried out over 16 yr), in which we were able to identify individual birds. We compared the breeding performance and survival of yearling breeders, first‐time adult breeders and adult breeders. We found significant differences in breeding performance but not in survival. Yearling breeders had lower breeding success than older individuals but the breeding performance of inexperienced adults did not differ from that of experienced adults. We did not find changes in terminal breeding success since peregrines in their last year of life sustained the performance levels shown in previous years although with increasing variability. We found no evidence that attempting to breed affected survival probability in any age group. We argue that differences in breeding performance are related to age, not to breeding experience, and that there is an age threshold, coincident with the development of adult plumage, after which breeding performance is not affected either by age or experience. Peregrines that start breeding as yearlings are likely to have greater lifetime reproductive success than birds entering the breeding pool as adults. Consequently, such birds may represent a set of high quality individuals. Our results support the age‐related competence improvement hypothesis as being the relevant explanation for the increase in breeding performance with age.     

Climate change leads to differential shifts in the timing of annual cycle stages in a migratory bird

Shifts in reproductive phenology due to climate change have been well documented in many species but how, within the same species, other annual cycle stages (e.g. moult, migration) shift relative to the timing of breeding has rarely been studied. When stages shift at different rates, the interval between stages may change resulting in overlaps, and as each stage is energetically demanding, these overlaps may have negative fitness consequences. We used long‐term data of a population of European pied flycatchers (Ficedula hypoleuca) to investigate phenological shifts in three annual cycle stages: spring migration (arrival dates), breeding (egg‐laying and hatching dates) and the onset of postbreeding moult. We found different advancements in the timing of breeding compared with moult (moult advances faster) and no advancement in arrival dates. To understand these differential shifts, we explored which temperatures best explain the year‐to‐year variation in the timing of these stages, and show that they respond differently to temperature increases in the Netherlands, causing the intervals between arrival and breeding and between breeding and moult to decrease. Next, we tested the fitness consequences of these shortened intervals. We found no effect on clutch size, but the probability of a fledged chick to recruit increased with a shorter arrival‐breeding interval (earlier breeding). Finally, mark–recapture analyses did not detect an effect of shortened intervals on adult survival. Our results suggest that the advancement of breeding allows more time for fledgling development, increasing their probability to recruit. This may incur costs to other parts of the annual cycle, but, despite the shorter intervals, there was no effect on adult survival. Our results show that to fully understand the consequences of climate change, it is necessary to look carefully at different annual cycle stages, especially for organisms with complex cycles, such as migratory birds.     

Density dependence in an island population of silvereyes

The population of silvereyes Zosterops lateralis chlorocephalus , on Heron Island, Great Barrier Reef has been monitored accurately since 1965. Between 1979 and 1993, the breeding success of all birds was determined by monitoring nests. The population fluctuated between 225 and 483 individuals. Four cyclones led to substantial mortality. As this data set is long-term, has little observation error, and is from an effectively closed population, it provides an unusual opportunity to examine density dependence in reproduction or mortality. Using a stochastic logistic model, we found clear evidence of density dependence in adult population size. Logistic regression suggested that fledgling survival decreased with the numbers of birds attempting to breed. There was also some suggestion that adult survival might be density dependent. The fitted stochastic logistic model predicts negligible risks of extinction for this population, in contrast to the predictions of a published population viability analysis. Whilst our statistical model including density dependence may provide better predictions of the "usual" behaviour of a population than a population viability analysis, we suggest that caution should be exercised when statistically fitted models are used to predict the behaviour of the population at extremes, such as near extinction.     

Persistence of accuracy of genomic estimated breeding values over generations in layer chickens

Background

The predictive ability of genomic estimated breeding values (GEBV) originates both from associations between high-density markers and QTL (Quantitative Trait Loci) and from pedigree information. Thus, GEBV are expected to provide more persistent accuracy over successive generations than breeding values estimated using pedigree-based methods. The objective of this study was to evaluate the accuracy of GEBV in a closed population of layer chickens and to quantify their persistence over five successive generations using marker or pedigree information.

Methods

The training data consisted of 16 traits and 777 genotyped animals from two generations of a brown-egg layer breeding line, 295 of which had individual phenotype records, while others had phenotypes on 2,738 non-genotyped relatives, or similar data accumulated over up to five generations. Validation data included phenotyped and genotyped birds from five subsequent generations (on average 306 birds/generation). Birds were genotyped for 23,356 segregating SNP. Animal models using genomic or pedigree relationship matrices and Bayesian model averaging methods were used for training analyses. Accuracy was evaluated as the correlation between EBV and phenotype in validation divided by the square root of trait heritability.

Results

Pedigree relationships in outbred populations are reduced by 50% at each meiosis, therefore accuracy is expected to decrease by the square root of 0.5 every generation, as observed for pedigree-based EBV (Estimated Breeding Values). In contrast the GEBV accuracy was more persistent, although the drop in accuracy was substantial in the first generation. Traits that were considered to be influenced by fewer QTL and to have a higher heritability maintained a higher GEBV accuracy over generations. In conclusion, GEBV capture information beyond pedigree relationships, but retraining every generation is recommended for genomic selection in closed breeding populations.     

Qualitative population divergence in proximate determination of a sexually selected trait in the collared flycatcher

We examined proximate determination of sexually selected forehead patch size in a Central‐European population of Ficedula albicollis, the collared flycatcher, using a 9‐year database, and compared our results with those obtained in other populations of the same and the sister species. Between‐individual variation of forehead patch size was large, its repeatability larger than, and heritability similar to the Swedish population. Unlike in the other populations, the trait proved unaffected by body condition, and only very slightly influenced by age. There was no relationship between forehead patch size and breeding lifespan, and a marginal negative association with survivorship in adult males. Our results suggest that additive genetic variance of the trait in this population is large, but genes act independently of body condition, and there is no viability indicator value of the trait. This is the first report of a qualitative intraspecific difference in proximate determination of a sexually selected trait.     

Effects of inbreeding and other genetic components on equine fertility

The Finnish mating records of Standardbred trotters (SB; n = 33 679) and Finnhorses (FH; n = 32 731) were analysed to study the effect of the level of inbreeding on foaling rates and to estimate the heritability of foaling rate. A linear mixed model was assumed, with the outcome of the foaling (foal or no foal) as the trait of the study. A restricted maximum likelihood-based method was used to calculate the estimates of the variance components. Predictions of breeding values and estimates of fixed effects were also calculated. The average level of inbreeding was 9.9% in the SB and 3.6% in the FH. The average foaling rates were better in the SB (72.6%) than in the FH (66.3%), but within each breed intense inbreeding had a statistically significant negative effect on foaling rate (P < 0.05). Also, the mating type, the age and breeding type of the mare, and the age of the stallion had statistically significant effects on foaling rate (P < 0.001). The heritability of foaling rate was between 3.4% and 3.7% in SBs and between 5.5% and 9.8% in FHs, when the outcome of the foaling was considered to be a trait of the expected foal. With the same model, the estimates of maternal genetic effect were 4.7% for SBs and 3.2% for FHs, and the estimates of the permanent environmental effects of the stallion were between 1.3% and 1.7%. Avoiding matings with very high inbreeding coefficients would improve foaling rates. It would also be possible to devise a breeding program for better equine fertility, but because the heritability is low, improvement of environmental factors deserves special attention.     

Adult survival and numbers in a coastal breeding population of Redshank Tringa totanus in northwest England

The results of a long-term capture-mark-recapture ringing programme carried out on a coastal population of breeding Redshanks Tringa totanus between 1974 and 1988 are presented.
Both sexes were equally likely to be recaptured in subsequent years, as were birds captured for the first time compared with those that had been captured previously. Older birds were more frequently recaptured than were young birds. There was no significant difference in male and female adult survival rates, with a mean of 72% of females and 75% of males surviving each year.
The breeding population fluctuated annually with estimated breeding densities of 122–285 pairs per km². Variation in breeding numbers (males), but not survival, was partially attributable to winter air temperature.     

Effects of geolocator attachments on breeding parameters of Lesser Kestrels

ABSTRACT Light level geolocators, also known as GLS loggers, are electronic devices intended for tracking the location of wide‐ranging animals using ambient light to estimate latitude and longitude. Miniaturized geolocators that can be used on relatively small migratory birds have recently become available, but little is known about the potential harmful effects of geolocators on birds. We examined the possible effects of 1.5‐g geolocators (dimensions: 21 × 6.5 × 9 mm) on the breeding success and survival of migratory Lesser Kestrels (Falco naumanni). During the 2007 breeding season, kestrels were fitted with geolocators using two attachment methods (Teflon wing harnesses and darvic bands), and geolocators were removed in 2008 after the birds returned to the breeding grounds. We found no differences in the breeding success of control and tagged pairs during the 2007 breeding season, but tagged pairs had greater fledgling mortality in the following breeding season. Furthermore, nestlings of tagged individuals had higher triglyceride and uric acid concentrations in their blood than control nestlings during the breeding season following tagging. As for return rates, 75% of tagged birds came back to the colony after the nonbreeding period, a proportion similar to that reported in previous studies. Although back‐mounts are slightly heavier and require more skill to attach, we recommend their use on small migratory raptors because most leg‐mounted geolocators in this study were damaged or rendered useless by dirt obscuring the light sensor.     

The timing of breeding and offspring size in Great Tits Parus major

The relationship between the timing of breeding, offspring weight and offspring recruitment was investigated in two populations of Great Tits Parus major. In two of six seasons a significant negative correlation was found between laying date and fledgling weight, in three seasons the opposite was true and in one season no significant correlation existed between these variables. Offspring recruitment was quantified in five seasons, and late-nesting birds were found to produce significantly fewer recruiting offspring in two of these, the same two seasons when late fledglings had significantly lower weights. No significant correlation existed between the number of recruits and laying date in the three seasons when late fledglings were as heavy as or heavier than early fledglings. The potential detriment to reproductive success of lateness thus was offset when late-nesting parents managed to produce heavy young. Between-year variations in the seasonal pattern of reproductive success has previously been suggested to contribute to the maintenance of variability in the timing of breeding in Great Tits. In this paper we suggest that producing heavy offspring late in the season might be a proximate factor influencing the seasonal patterns of reproductive success. Producing heavy offspring late in the season is probably not a "strategy" but rather an effect of chance differences between years in the seasonal availability of food or the seasonality of other factors important for the nesting birds.