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Life history trade-offs are one of the central tenets of evolutionary demography. Trade-offs, depicting negative covariances between individuals' life history traits, can arise from genetic constraints, or from a finite amount of resources that each individual has to allocate in a zero-sum game between somatic and reproductive functions. While theory predicts that trade-offs are ubiquitous, empirical studies have often failed to detect such negative covariances in wild populations. One way to improve the detection of trade-offs is by accounting for the environmental context, as trade-off expression may depend on environmental conditions. However, current methodologies usually search for fixed covariances between traits, thereby ignoring their context dependence. Here, we present a hierarchical multivariate 'covariance reaction norm' model, adapted from Martin (2023), to help detect context dependence in the expression of life-history trade-offs using demographic data. The method allows continuous variation in the phenotypic correlation between traits. We validate the model on simulated data for both intraindividual and intergenerational trade-offs. We then apply it to empirical datasets of yellow-bellied marmots (Marmota flaviventer) and Soay sheep (Ovis aries) as a proof-of-concept showing that new insights can be gained by applying our methodology, such as detecting trade-offs only in specific environments. We discuss its potential for application to many of the existing long-term demographic datasets and how it could improve our understanding of trade-off expression in particular, and life history theory in general.

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The negative effects of heat stress on swine reproduction have been well documented and the recent global warming trend caused by climate change is leading to more days with high temperatures every year. This has caused a reduction in litter trait performance of Landrace sows in Taiwan, a country extending across tropical and subtropical oceanic zones. Therefore, this study developed a modified model to determine which stages of pregnancy, before, early, middle, and late, had the largest impacts of heat stress on litter traits. A reaction norm model (RNM) was used to identify sows with high resilience to heat stress for litter traits followed by analysis of the modified model. Data from Landrace sows were collected from 2 farms in Taiwan between 2008 and 2021. A total of 11,059 records were collected for total number born (TNB), number born alive (NBA), and stillborn rate (STBR). The results showed that the heritabilities of TNB, NBA, and STBR were 0.170, 0.115, and 0.077, respectively. These results were similar between the conventional model and the modified model. In the modified model, the before and early stages of sow pregnancy were the significant periods for TNB and NBA (P < 0.05), while the early and middle stages were significant for STBR (P < 0.05). According to the RNM results, the heritability estimates for TNB, NBA, and STBR were 0.23 to 0.11, 0.18 to 0.08, and 0.10 to 0.04, respectively, showing a decrease from low temperature-humidity index (THI) to high THI. The minimum genetic correlations between the highest and the lowest THI for TNB, NBA, and STBR were 0.85, 0.64, and 0.80, respectively. The results of the RNM for breeding value showed re-ranking across THI values. In conclusion, similar results were obtained for heritability when the model was modified for heat stress estimation. Yet re-ranking of breeding values across THI could help farmers to select not only for improved litter trait performance but also for heat stress resilience of Landrace sows in Taiwan.

Heat stress caused by climate change is a challenge for the pig industry, especially in countries located in tropical and subtropical zones, such as Taiwan. It can adversely affect litter traits, leading to less pork production and higher economic losses to farms. Therefore, identifying sows with the potential to tolerate high heat with high humid condition is an important task for the pig industry. This article proposes a reaction norm model to determine the trend in breeding value across temperature and humidity index values and its implications for litter traits of sows. Our results indicate that litter traits can be used to select sows with high heat tolerance.

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With global warming, there are growing challenges for raising taurine and composite beef cattle populations in tropical regions, including elevated temperatures, limited forage availability, parasite infestation, and infectious diseases. These environmental factors can trigger specific physiological responses in the developing fetus, which may have long-term implications on its performance. Therefore, the main objective of this study was to assess the influence of naturally induced thermal stress during the gestation period on the subsequent performance of tropical composite beef cattle progeny. Furthermore, we aimed to investigate the impact of genotype-by-gestational thermal environment interaction (G×Eg) on traits under selection pressure in the breeding population. A total of 157,414 animals from 58 farms located in various Brazilian states were recorded for birth weight (BW), preweaning weight gain (PWG), yearling weight (YW), hip height (HH), scrotal circumference (SC), and days to first calving (DFC). We first applied a linear regression model to the BW data, which revealed that the last 40 d of gestation were suitable for calculating the mean temperature humidity index (THIg). Subsequent regression analyses revealed that for every 10-unit increase in THIg, detrimental effects of approximately 1.13% to 16.34% are expected for all traits evaluated. Genetic parameters were estimated through a reaction norm model using THIg as the environmental descriptor. The posterior means of heritability estimates (SD) were 0.35 (0.07), 0.25 (0.03), 0.31 (0.03), 0.37 (0.01), 0.29 (0.07), and 0.20 (0.09) for the direct effect of BW, PWG, YW, HH, SC, and DFC, respectively. These estimates varied along the range of THIg values, suggesting a variable response to selection depending on the thermal environment during gestation. Genetic correlation estimates between more divergent THIg values were low or negative for YW, PWG, and DFC, indicating that the best-performing individuals at low THIg values may not perform as well at high THIg values and vice versa. Overall, thermal stress during gestation impacts the future performance of beef cattle offspring. Our results indicate the need for developing effective breeding strategies that take into account G×Eg effects and the re-ranking of breeding animals along the THIg scale, particularly for traits such as DFC that are highly sensitive to thermal stress.

With global warming posing increasing challenges in tropical regions, this study aimed to assess the impact of thermal stress during gestation on the performance of composite beef cattle offspring. Environmental factors such as high temperatures, humidity, limited forage availability, and parasite infestation can elicit physiological responses in the developing fetus, affecting its long-term performance and welfare. Using the temperature humidity index (THIg) of the late gestation as a measure of thermal environment, a reaction norm model was applied to analyze the birth weight, preweaning weight gain, yearling weight, hip height, scrotal circumference, and days to first calving (DFC). Results revealed that increasing THIg values were associated with a detrimental effect in these traits. Genotype-by-environment interaction was found to significantly influence trait variability, with DFC showing the strongest effect. Negative genetic correlations were observed between divergent THIg values, suggesting that individuals performing well in mild thermal environments may not excel in high thermal stress conditions. The heritability estimates varied along the THIg scale, indicating that selection response may vary depending on the thermal environment during gestation. These findings emphasize the need for breeding strategies that account for genotype-by-environment effects and consider the impact of thermal stress on cattle performance.

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Introgression of non-native conspecifics changes the genetic composition of wild populations, potentially leading to loss of local adaptations and fitness declines. However, long-term data from wild populations are still relatively few. Here, we studied the effects of introgression in a Danish brown trout (Salmo trutta, L.) population, subjected to intensive stocking with domesticated hatchery fish of non-native origin. We used wild-caught genetically wild and admixed trout as well as fish from the partly domesticated hatchery strain used for stocking the river up until ~15 years prior to this study, to produce 22 families varying in hatchery/wild admixture. Following a replicated common-garden experiment conducted in fish tanks from first feeding through 23 weeks at 7, 12, and 16°C, we observed a significant positive relationship between family admixture and fish size upon termination, an effect observed through all levels of admixture. Furthermore, the admixture effect was most distinct at the higher rearing temperatures. Although the hatchery strain used for stocking had been in culture for ~7 generations, it had not been deliberately selected for increased growth. These data thus demonstrate: (i) that growth had increased in the hatchery strain even in the absence of deliberate directional selection for this trait, (ii) that the increasing effect of admixture by temperature could represent inadvertent selection for performance in the hatchery strain at higher temperatures, and most significantly, (iii) that despite undergoing up to five generations of natural selection in the admixed wild population, the genetically increased growth potential was still detectable and thus persistent. Our findings suggest that altered growth patterns and potentially their cascading effects are of importance to the severity of hatchery/wild introgression, especially under changing-climate scenarios and are of general significance to conservation practitioners seeking to evaluate long-term effects of intra-specific hybridization including under recovery.

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Ongoing climate change poses an increasing threat to biodiversity. To avoid decline or extinction, species need to either adjust or adapt to new environmental conditions or track their climatic niches across space. In sessile organisms such as plants, phenotypic plasticity can help maintain fitness in variable and even novel environmental conditions and is therefore likely to play an important role in allowing them to survive climate change, particularly in the short term. Understanding a species' response to rising temperature is crucial for planning well-targeted and cost-effective conservation measures. We sampled seeds of three Hypericum species (H. maculatum, H. montanum, and H. perforatum), from a total of 23 populations originating from different parts of their native distribution areas in Europe. We grew them under four different temperature regimes in a greenhouse to simulate current and predicted future climatic conditions in the distribution areas. We measured flowering start, flower count, and subsequent seed weight, allowing us to study variations in the thermal plasticity of flowering phenology and its relation to fitness. Our results show that individuals flowered earlier with increasing temperature, while the degree of phenological plasticity varied among species. More specifically, the plasticity of H. maculatum varied depending on population origin, with individuals from the leading range edge being less plastic. Importantly, we show a positive relationship between higher plasticity and increased flower production, indicating adaptive phenological plasticity. The observed connection between plasticity and fitness supports the idea that plasticity may be adaptive. This study underlines the need for information on plasticity for predicting species' potential to thrive under global change and the need for studies on whether higher phenotypic plasticity is currently being selected as natural populations experience a rapidly changing climate.

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AbstractIn ectotherms, temperature has a strong effect on metabolic rate (MR), yet the extent to which the thermal sensitivity of MR varies among versus within individuals is largely unknown. This is of interest because significant among-individual variation is a prerequisite for the evolution of metabolic thermal sensitivity. Here, we estimated the repeatability (R) of the thermal sensitivity of MR in individual virgin, adult male Drosophila melanogaster (N=316) by taking repeated overnight measures of their MRs at two temperatures (~24°C and ~27°C). At the population level, thermal sensitivity decreased with locomotor activity, and older individuals showed a higher thermal sensitivity of MR than younger individuals. Taking these effects (and body mass) into account, we detected significant repeatability in both the centered intercept (Rint=0.52±0.04) and the slope (Rslp=0.21±0.07) of the metabolic thermal reaction norms, which respectively represent average MR and thermal sensitivity of MR. Furthermore, individuals with a higher overall MR also displayed greater increases in MR as temperature increased from ~24°C to ~27°C (rind=0.32±0.14). Average MR and thermal sensitivity of MR were also positively correlated within individuals (re=0.15±0.07). Our study represents a point of departure for future larger studies, in which more complex protocols (e.g., wider temperature range, breeding design) can be applied to quantify the causal components of variation in thermal sensitivity that are needed to make accurate predictions of adaptive responses to global warming.

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We performed a plateau-linear reaction norm model (RNM) analysis of number born alive (NBA) in purebred Landrace pigs, where breeding value changes according to maximum temperature at mating day, using public meteorological observation data in Japan. We analysed 52,668 NBA records obtained from 10,320 Landrace sows. Pedigree data contained 99,201 animals. Off-farm daily temperature data at the nearest weather station from each of the farms were downloaded from the Japan Meteorological Agency website. A plateau-linear RNM analysis based on daily maximum temperature on mating day (threshold temperature of 16.6°C) was performed. The percentage of the records with daily maximum temperatures at mating days of ≤16.6, ≥25.0, ≥30.0 and ≥35.0°C were 34.3%, 33.6%, 14.0% and 0.8%, respectively. The value of Akaike's information criterion for the plateau-linear RNM was lower than that for a simple repeatability model (RM). With the plateau-linear RNM, estimated value of heritability ranged from 0.14 to 0.15, while that from the RM analysis was 0.15. Additive genetic correlation between intercept and slope terms was estimated to be -0.52 from the plateau-linear RNM analysis. Estimated additive genetic correlations were >0.9 between NBA at different temperatures ranging from 16.6 to 37.6°C. For the 10,320 sows, average values of prediction reliability of the intercept and slope terms for breeding values in the plateau-linear RNM were 0.47 and 0.16, respectively. Increasing weight for slope term in linear selection index could bring positive genetic gain in the slope part, but prediction accuracy would decrease. Our results imply that genetically improving heat tolerance in sows reared in Japan focusing on NBA using RNM is possible, while RNM is more complex to implement and interpret. Therefore, further study should be encouraged to make genetic improvement for heat tolerance in sows more efficient.

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Phenotypic plasticity helps animals to buffer the effects of increasing thermal and nutritional stress created by climate change. Plastic responses to single and combined stressors can vary among genetically diverged populations. However, less is known about how plasticity in response to combined stress varies among individuals within a population or whether such variation changes across life-history traits. This is important because individual variation within populations shapes population-level responses to environmental change. Here, we used isogenic lines of Drosophila melanogaster to assess the plasticity of egg-to-adult viability and sex-specific body size for combinations of 2 temperatures (25 °C or 28 °C) and 3 diets (standard diet, low caloric diet, or low protein:carbohydrate ratio diet). Our results reveal substantial within-population genetic variation in plasticity for egg-to-adult viability and wing size in response to combined thermal-nutritional stress. This genetic variation in plasticity was a result of cross-environment genetic correlations that were often < 1 for both traits, as well as changes in the expression of genetic variation across environments for egg-to-adult viability. Cross-sex genetic correlations for body size were weaker when the sexes were reared in different conditions, suggesting that the genetic basis of traits may change with the environment. Furthermore, our results suggest that plasticity in egg-to-adult viability is genetically independent from plasticity in body size. Importantly, plasticity in response to diet and temperature individually differed from plastic shifts in response to diet and temperature in combination. By quantifying plasticity and the expression of genetic variance in response to combined stress across traits, our study reveals the complexity of animal responses to environmental change, and the need for a more nuanced understanding of the potential for populations to adapt to ongoing climate change.

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Understanding the evolution of traits subject to trade-offs is challenging because phenotypes can (co)vary at both the among- and within-individual levels. Among-individual covariation indicates consistent, possibly genetic, differences in how individuals resolve the trade-off, while within-individual covariation indicates trait plasticity. There is also the potential for consistent among-individual differences in behavioral plasticity, although this has rarely been investigated. We studied the sources of (co)variance in two characteristics of an acoustic advertisement signal that trade-off with one another and are under sexual selection in the gray treefrog, Hyla chrysoscelis: call duration and call rate. We recorded males on multiple nights calling spontaneously and in response to playbacks simulating different competition levels. Call duration, call rate, and their product, call effort, were all repeatable both within and across social contexts. Call duration and call rate covaried negatively, and the largest covariance was at the among-individual level. There was extensive plasticity in calling with changes in social competition, and we found some evidence for among-individual variance in call rate plasticity. The significant negative among-individual covariance in trait values is perpendicular to the primary direction of sexual selection in this species, indicating potential limits on the response to selection.

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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.

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Phenotypic plasticity is often posited as an avenue for adaptation to environmental change, whereby environmental influences on phenotypes could shift trait expression toward new optimal values. Conversely, plastic trait expression may inhibit adaptation to environmental change by reducing selective pressure on ill-adapted traits. While plastic responses are often assumed to be linear, nonlinear phenotype-environment relationships are common, especially in thermally sensitive traits. Here we examine nonlinear plasticity in a trait with great ecological and evolutionary significance: sexual phenotype in species with environmental sex determination (ESD). In species with ESD, development switches between male and female at an environmental threshold (the inflection point). The inflection point is a key trait for adaptive responses to changing environments and should evolve toward the new optimum in order to maintain evolutionarily stable sex ratios. We used an individual-based theoretical model to investigate how two forms of plasticity in the ESD reaction norm-the nonlinear slope of the reaction norm and a linear shift in the inflection point-influence the evolution of the inflection point under climate warming. We found that steeper reaction norm slopes (high nonlinear plasticity) promoted evolution toward new optimal phenotypes (higher inflection points). In contrast, increased linear plasticity in the inflection point (shift) hindered adaptive evolution. Additionally, populations in moderate warming scenarios showed greater adaptive evolution of the inflection point compared with populations in extreme warming scenarios, suggesting that the proximity of existing phenotypes to new optimal phenotypes influences evolutionary outcomes. Unexpectedly, we found greater population persistence under high climate variability, due to the increased production of rare-sex individuals in unusually cold years. Our results demonstrate that different forms of phenotypic plasticity have crucially different effects on adaptive evolution. Plasticity that prevented sex ratio bias hindered the evolution of the inflection point, while plasticity that exacerbated sex ratio bias promoted adaptation to environmental change.

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AbstractWhen organisms respond behaviorally to a stimulus, they exhibit plasticity, but some individuals respond to the same stimulus consistently differently than others, thereby also exhibiting personality differences. Parent house sparrows express individual differences in how often they feed offspring and how that feeding rate changes with nestling age. Mean feeding rate and its slope with respect to nestling age were positively correlated at median nestling ages but not at hatching, indicating that individuality is primarily in plasticity. Individual differences could arise because of (1) interactions between environmental variables, (2) differences in underlying state or "quality," or (3) differences in the ability to update cues of changing nestling demand. Individual slopes were modestly repeatable across breeding attempts, hinting at the likely action of additional environmental variables, but only brood size was important. I also found few correlates suggesting quality differences. I used short-term brood size manipulations at two nestling ages to test divergent predictions between the three hypotheses. The pattern of correlations between response to the manipulation and individual slope did not fit any single hypothesis. Patterns of sparrow parental care reveal that personality and plasticity are not cleanly separable, and their biology is likely intertwined. New thinking may be needed about the factors parents use in decisions about care and the relevant fitness consequences.

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Conspecifics of the same age and size differ consistently in the pace with which they expend energy. This among-individual variation in metabolic rate is thought to influence behavioural variation, since differences in energy requirements should motivate behaviours that facilitate energy acquisition, such as being bold or active in foraging. While there is evidence for links between metabolic rate and behaviour in constant environments, we know little about whether metabolic rate and behaviour change together when the environment changes-that is, if metabolic and behavioural plasticity co-vary. We investigated this using a fish that becomes dormant in winter and strongly reduces its activity when the environment cools, the cunner (Tautogolabrus adspersus). We found strong and predictable among-individual variation in thermal plasticity of metabolic rates, from resting to maximum levels, but no evidence for among-individual variation in thermal plasticity of movement activity, meaning that these key physiological and behavioural traits change independently when the environment changes. The strong among-individual variation in metabolic rate plasticity resulted in much higher repeatability (among-individual consistency) of metabolic rates at warm than cold temperatures, indicating that the potential for metabolic rate to evolve under selection is temperature-dependent, as repeatability can set the upper limit to heritability. This article is part of the theme issue 'The evolutionary significance of variation in metabolic rates'.

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This study provides a regional picture of long-term changes in Atlantic salmon growth at the southern edge of their distribution, using a multi-population approach spanning 49 years and five populations. We provide empirical evidence of salmon life history being influenced by a combination of common signals in the marine environment and population-specific signals. We identified an abrupt decline in growth from 1976 and a more recent decline after 2005. As these declines have also been recorded in northern European populations, our study significantly expands a pattern of declining marine growth to include southern European populations, thereby revealing a large-scale synchrony in marine growth patterns for almost five decades. Growth increments during their sea sojourn were characterized by distinct temporal dynamics. At a coarse temporal resolution, growth during the first winter at sea seemed to gradually improve over the study period. However, the analysis of finer seasonal growth patterns revealed ecological bottlenecks of salmon life histories at sea in time and space. Our study reinforces existing evidence of an impact of early marine growth on maturation decision, with small-sized individuals at the end of the first summer at sea being more likely to delay maturation. However, each population was characterized by a specific probabilistic maturation reaction norm, and a local component of growth at sea in which some populations have better growth in some years might further amplify differences in maturation rate. Differences between populations were smaller than those between sexes, suggesting that the sex-specific growth threshold for maturation is a well-conserved evolutionary phenomenon in salmon. Finally, our results illustrate that although most of the gain in length occurs during the first summer at sea, the temporal variability in body length at return is buffered against the decrease in post-smolt growth conditions. The intricate combination of growth over successive seasons, and its interplay with the maturation decision, could be regulating body length by maintaining diversity in early growth trajectories, life histories, and the composition of salmon populations.

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Phenotypic plasticity can allow organisms to cope with environmental changes. Although reaction norms are commonly used to quantify plasticity along gradients of environmental conditions, they often miss the temporal dynamics of phenotypic change, especially the speed at which it occurs. Here, we argue that studying the rate of phenotypic plasticity is a crucial step to quantify and understand its adaptiveness. Iteratively measuring plastic traits allows us to describe the actual dynamics of phenotypic changes and avoid quantifying reaction norms at times that do not truly reflect the organism's capacity for plasticity. Integrating the temporal component in how we describe, quantify, and conceptualise phenotypic plasticity can change our understanding of its diversity, evolution, and consequences.

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Phenotypic plasticity is an important topic in biology and evolution. However, how to generate broadly applicable insights from individual studies remains a challenge. Here, with flowering time observed from a large geographical region for sorghum and rice genetic populations, we examine the consistency of parameter estimation for reaction norms of genotypes across different subsets of environments and searched for potential strategies to inform the study design. Both sample size and environmental mean range of the subset affected the consistency. The subset with either a large range of environmental mean or a large sample size resulted in genetic parameters consistent with the overall pattern. Furthermore, high accuracy through genomic prediction was obtained for reaction norm parameters of untested genotypes using models built from tested genotypes under the subsets of environments with either a large range or a large sample size. With 1428 and 1674 simulated settings, our analyses suggested that the distribution of environmental index values of a site should be considered in designing experiments. Overall, we showed that environmental context was critical, and considerations should be given to better cover the intended range of the environmental variable. Our findings have implications for the genetic architecture of complex traits, plant-environment interaction, and climate adaptation.

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In ectotherms, body temperature is a crucial determinant of performance and fitness, as captured by thermal performance curves (TPCs). Since survival in variable environments is often facilitated by phenotypic plasticity, to reliably assess an organism's ability to cope with thermal changes, it is necessary to characterize not only TPCs but also their reaction norms. While previous studies have investigated plasticity in TPCs, these studies focus only on selected parameters and a few developmental temperatures. They may, therefore, overlook the complexity of developmental plasticity in TPCs. Here, we examined the full extent of thermal developmental plasticity in TPCs for fecundity and hatchability in Drosophila melanogaster. By employing a factorial design with ten developmental and twelve adult temperatures, our study enabled a comprehensive characterization of reaction norms of all key TPC parameters. We found that developmental temperature had a significant impact on the egg production rate, with minor effects on other TPC parameters. Nonoptimal developmental temperatures negatively affected most TPC parameters, whereas development at 22-26 °C maximized reproductive fitness. We also revealed that developmental plasticity for the maximum reproductive performance might be predominantly caused by developmentally-induced changes in ovariole number. Our results in conjunction with previous studies on intraspecific variation suggest that any changes in TPC for reproduction primarily involve the egg production rate, indicating that adaptive evolution and phenotypic plasticity followed the same pathway. Overall, our findings underscore the limitations of developmental plasticity in enhancing reproductive fitness, suggesting that while certain traits, such as egg production rate, may be relatively plastic, these changes may not be sufficient to enable effective adjustment to environmental shifts.

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The presence or absence of awns-whether wheat heads are 'bearded' or 'smooth' - is the most visible phenotype distinguishing wheat cultivars. Previous studies suggest that awns may improve yields in heat or water-stressed environments, but the exact contribution of awns to yield differences remains unclear. Here we leverage historical phenotypic, genotypic, and climate data for wheat (Triticum aestivum) to estimate the yield effects of awns under different environmental conditions over a 12-year period in the southeastern USA. Lines were classified as awned or awnless based on sequence data, and observed heading dates were used to associate grain fill periods of each line in each environment with climatic data and grain yield. In most environments, awn suppression was associated with higher yields, but awns were associated with better performance in heat-stressed environments more common at southern locations. Wheat breeders in environments where awns are only beneficial in some years may consider selection for awned lines to reduce year-to-year yield variability, and with an eye towards future climates.

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Fish often spawn eggs with ovarian fluids that have been hypothesized to support the sperm of some males over others (cryptic female choice). Alternatively, sperm reactions to ovarian fluids could reveal male strategies. We used wild-caught lake char (Salvelinus umbla) to experimentally test whether sperm react differently to the presence of ovarian fluid, and whether any differential sperm reaction could be predicted by male breeding coloration, male inbreeding coefficients (based of 4150 SNPs) or the kinship coefficients between males and females. Male coloration was positively linked to body size and current health (based on lymphocytosis and thrombocytosis) but was a poor predictor of inbreeding or kinship coefficients. We found that sperm of more colourful males were faster in diluted ovarian fluids than in water only, while sperm of paler males were faster in water than in ovarian fluids. We then let equal numbers of sperm compete for fertilizations in the presence or absence of ovarian fluids and genetically assigned 1464 embryos (from 70 experimental trials) to their fathers. The presence of ovarian fluids significantly increased the success of the more colourful competitors. Sperm of less inbred competitors were more successful when tested in water only than in diluted ovarian fluids. The kinship coefficients had no significant effects on sperm traits or fertilization success in the presence of ovarian fluids, although parallel stress tests on embryos had revealed that females would profit more from mating with least related males rather than most coloured ones. We conclude that sperm of more colourful males are best adapted to ovarian fluids, and that the observed reaction norms suggest male strategies rather than cryptic female choice.

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For theoretical studies, reaction norm evolution in a changing environment can be modeled by means of the multivariate breeder's equation, with the reaction norm parameters treated as traits in their own right. This is, however, not a feasible approach for the use of field data, where the intercept and slope values are not available. An alternative approach is to use infinite-dimensional characters and smooth covariance function estimates found by, e.g., random regression. This is difficult because of the need to find, for example, polynomial basis functions that fit the data reasonably well over time, and because reaction norms in multivariate cases are correlated, such that they cannot be modeled independently. Here, I present an alternative approach based on a multivariate linear mixed model of any order, with dynamical incidence and residual covariance matrices that reflect the changing environment. From such a mixed model follows a dynamical BLUP model for the estimation of the individual reaction norm parameter values at any given parent generation, and for updating of the mean reaction norm parameter values from generation to generation by means of Robertson's secondary theorem of natural selection. This will, for example, make it possible to disentangle the microevolutionary and plasticity components in climate change responses. The BLUP model incorporates the additive genetic relationship matrix in the usual way, and overlapping generations can easily be accommodated. Additive genetic and environmental model parameters are assumed to be known and constant, but it is discussed how they can be estimated by means of a prediction error method. The identifiability by the use of field or laboratory data containing environmental, phenotypic, fitness, and additive genetic relationship data is an important feature of the proposed model.