The incubation environment does not explain significant variation in heart rate plasticity among avian embryos.

The conditions an organism experiences during development can modify how they plastically respond to short-term changes in their environment later in life. This can be adaptive because the optimal average trait value and the optimal plastic change in trait value in response to the environment may differ across different environments. For example, early developmental temperatures can adaptively modify how reptiles, fish and invertebrates metabolically respond to temperature. However, whether individuals within populations respond differently (a prerequisite to adaptive evolution), and whether this occurs in birds, which are only ectothermic for part of their life cycle, is not known. We experimentally tested these possibilities by artificially incubating the embryos of Pekin ducks (Anas platyrhynchos domesticus) at constant or variable temperatures. We measured their consequent heart rate reaction norms to short-term changes in egg temperature and tracked their growth. Contrary to expectations, the early thermal environment did not modify heart rate reaction norms, but regardless, these reaction norms differed among individuals. Embryos with higher average heart rates were smaller upon hatching, but heart rate reaction norms did not predict subsequent growth. Our data also suggests that the thermal environment may affect both the variance in heart rate reaction norms and their covariance with growth. Thus, individual avian embryos can vary in their plasticity to temperature, and in contrast to fully ectothermic taxa, the early thermal environment does not explain this variance. Because among-individual variation is one precondition to adaptive evolution, the factors that do contribute to such variability may be important.

Early-life telomeres are influenced by environments acting at multiple temporal and spatial scales.

An individual's telomere length early in life may reflect or contribute to key life-history processes sensitive to environmental variation. Yet, the relative importance of genetic and environmental factors in shaping early-life telomere length is not well understood as it requires samples collected from multiple generations with known developmental histories. We used a confirmed pedigree and conducted an animal model analysis of telomere lengths obtained from nestling house sparrows (Passer domesticus) sampled over a span of 22 years. We found significant additive genetic variation for early-life telomere length, but it comprised a small proportion (9%) of the total biological variation. Three sources of environmental variation were important: among cohorts, among-breeding attempts within years, and among nestmates. The magnitude of variation among breeding attempts and among nestmates also differed by cohort, suggesting that interactive effects of environmental factors across time or spatial scales were important, yet we were unable to identify the specific causes of these interactions. The mean amount of precipitation during the breeding season positively predicted telomere length, but neither weather during a given breeding attempt nor date in the breeding season contributed to an offspring's telomere length. At the level of individual nestlings, offspring sex, size and mass at 10 days of age also did not predict telomere length. Environmental effects appear especially important in shaping early-life telomere length in some species, and more focus on how environmental factors that interact across scales may help to explain some of the variation observed among studies.

Temperature-mediated plasticity in incubation schedules is unlikely to evolve to buffer embryos from climatic challenges in a seasonal songbird.

Phenotypic plasticity is hypothesized to facilitate adaptive responses to challenging conditions, such as those resulting from climate change. However, tests of the key predictions of this 'rescue hypothesis', that variation in plasticity exists and can evolve to buffer unfavourable conditions, remain rare. Here, we investigate among-female variation in temperature-mediated plasticity of incubation schedules and consequences for egg temperatures using the chestnut-crowned babbler (Pomatostomus ruficeps) from temperate regions of inland south-eastern Australia. Given recent phenological advances in this seasonal breeder and thermal requirements of developing embryos (>~25°C, optimally ~38°C), support for evolutionary rescue-perhaps paradoxically-requires that plasticity serves to buffer embryos more from sub-optimally low temperatures. We found significant variation in the duration of incubation bouts (mean ± SD = 27 ± 22 min) and foraging bouts (mean ± SD = 17 ± 11 min) in this maternal-only incubator. However, variation in each arose because of variation in the extent to which mothers increased on- and off-bout durations when temperatures (0-36°C) were more favourable rather than unfavourable as required under rescue. In addition, there was a strong positive intercept-slope correlation in on-bout durations, indicating that those with stronger plastic responses incubated more at average temperatures (~19°C). Combined, these effects reduced the functional significance of plastic responses: an individual's plasticity was neither associated with daily contributions to incubation (i.e. attentiveness) nor average egg temperatures. Our results highlight that despite significant among-individual variation in environmental-sensitivity, plasticity in parental care traits need not evolve to facilitate buffering against unfavourable conditions.

Variation in Embryonic Metabolic Reaction Norms and the Role of the Environment.

AbstractEarly developmental environments can shape how organisms respond to later environments, but despite the potential for this phenomenon to alter the evolution of phenotypes and their underlying mechanisms in variable environments, details of this process are not understood. For example, both temperature and parental age can alter offspring metabolic plasticity and growth within species, yet the extent of such effects is unknown. We measured the reaction norms of embryonic heart rate in response to egg temperature and the change in egg mass over the incubation period in wild house sparrows. Using Bayesian linear mixed models, we estimated covariation in the intercepts and slopes of these reaction norms among clutches and eggs. We found that heart rate intercepts, not slopes, varied among clutches and that neither intercepts nor slopes varied among eggs within clutches. In contrast, egg mass intercepts and slopes varied among clutches and eggs. Ambient temperature did not explain variance in reaction norms. Instead, individuals from older mothers were more metabolically sensitive to egg temperature and lost less mass over the incubation period than individuals from younger mothers. Nevertheless, heart rate reaction norms and egg mass reaction norms did not covary. Our results suggest that early environments influenced by parents may contribute to variation in embryonic reaction norms. The structure of variation in embryonic reaction norms that exists both among clutches and among eggs demonstrates a complexity in plastic phenotypes that should be explored in future work. Furthermore, the potential for the embryonic environment to shape the reaction norms of other traits has implications for the evolution of plasticity more broadly.

Quantitative Evaluation of Tactile Foraging Behavior in Pekin and Muscovy Ducks.

Ducks have developed a variety of foraging strategies that utilize touch sensitive bills to match their ecological niche within wetlands. These techniques include diving, sieving, dabbling, and grazing. Ducks exhibiting tactile specialization in foraging outperform visual and non-tactile foraging ducks in behavioral experiments and have a higher percentage of light-touch mechanoreceptor neurons expressing Piezo2 in the trigeminal ganglia. Belonging to two different tribes of Anseriformes, the well-studied tactile specialist Pekin (Tribe Anatini: Anas platyrhynchos domestica) and lesser studied Muscovy (Tribe Cairinini: Cairina moschata domestica) ducks were tested on a series of experiments to assess these birds' functional tactile acuity. Both species of duck were able to separate out and consume edible items from increasing amounts of inedible plastiline clay distractors. They could also both be trained to associate a food reward with plastiline stimuli of differing size and shape using touch alone. However, only females of each species could learn to associate food reward with otherwise identical stimuli differing only in hardness. Pekin females performed significantly better than Muscovy females suggesting the anatomical specializations present in many Anatini may contribute to this type of tactile acuity. These findings have potential relevance in understanding the evolution of tactile ability and feeding ecology.

Advancing breeding phenology does not affect incubation schedules in chestnut-crowned babblers: Opposing effects of temperature and wind.

Projecting population responses to climate change requires an understanding of climatic impacts on key components of reproduction. Here, we investigate the associations among breeding phenology, climate and incubation schedules in the chestnut-crowned babbler (Pomatostomus ruficeps), a 50 g passerine with female-only, intermittent incubation that typically breeds from late winter (July) to early summer (November). During daylight hours, breeding females spent an average of 33 min on the nest incubating (hereafter on-bouts) followed by 24-min foraging (hereafter off-bouts), leading to an average daytime nest attentiveness of 60%. Nest attentiveness was 25% shorter than expected from allometric calculations, largely because off-bout durations were double the expected value for a species with 16 g clutches (4 eggs × 4 g/egg). On-bout durations and daily attentiveness were both negatively related to ambient temperature, presumably because increasing temperatures allowed more time to be allocated to foraging with reduced detriment to egg cooling. By contrast, on-bout durations were positively associated with wind speed, in this case because increasing wind speed exacerbated egg cooling during off-bouts. Despite an average temperature change of 12°C across the breeding season, breeding phenology had no effect on incubation schedules. This surprising result arose because of a positive relationship between temperature and wind speed across the breeding season: Any benefit of increasing temperatures was canceled by apparently detrimental consequences of increasing wind speed on egg cooling. Our results indicate that a greater appreciation for the associations among climatic variables and their independent effects on reproductive investment are necessary to understand the effects of changing climates on breeding phenology.