Plasticity in probabilistic reaction norms for maturation in a salmonid fish

The relationship between body size and the probability of maturing, often referred to as the probabilistic maturation reaction norm (PMRN), has been increasingly used to infer genetic variation in maturation schedule. Despite this trend, few studies have directly evaluated plasticity in the PMRN. A transplant experiment using white-spotted charr demonstrated that the PMRN for precocious males exhibited plasticity. A smaller threshold size at maturity occurred in charr inhabiting narrow streams where more refuges are probably available for small charr, which in turn might enhance the reproductive success of sneaker precocious males. Our findings suggested that plastic effects should clearly be included in investigations of variation in PMRNs.     

Functional mapping of reaction norms to multiple environmental signals

Whether there are different genes involved in response to different environmental signals and how these genes interact to determine the final expression of the trait are of fundamental importance in agricultural and biological research. We present a statistical framework for mapping environment-induced genes (or quantitative trait loci, QTLs) of major effects on the expression of a trait that respond to changing environments. This framework is constructed with a maximum-likelihood-based mixture model, in which the mean and covariance structure of environment-induced responses is modelled. The means for responses to continuous environmental states, referred to as reaction norms, are approximated for different QTL genotypes by mathematical equations that were derived from fundamental biological principles or based on statistical goodness-of-fit to observational data. The residual covariance between different environmental states was modelled by autoregressive processes. Such an approach to studying the genetic control of reaction norms can be expected to be advantageous over traditional mapping approaches in which no biological principles and statistical structures are considered. We demonstrate the analytical procedure and power of this approach by modelling the photosynthetic rate process as a function of temperature and light irradiance. Our approach allows for testing how a QTL affects the reaction norm of photosynthetic rate to a specific environment and whether there exist different QTLs to mediate photosynthetic responses to temperature and light irradiance, respectively.     

The selective advantage of reaction norms for environmental tolerance

A tolerance curve defines the dependence of a genotype's fitness on the state of an environmental gradient. It can be characterized by a mode (the genotype's optimal environment) and a width (the breadth of adaptation). It seems possible that one or both of these characters can be modified in an adaptive manner, at least partially, during development. Thus, we extend the theory of environmental tolerance to include reaction norms for the mode and the width of the tolerance curve. We demonstrate that the selective value of such reaction norms increases with increasing spatial heterogeneity and between-generation temporal variation in the environment and with decreasing within-generation temporal variation. Assuming that the maintenance of a high breadth of adaptation is costly, reaction, norms are shown to induce correlated selection for a reduction in this character. Nevertheless, regardless of the magnitude of the reaction norm, there is a nearly one to one relationship between the optimal breadth of adaptation and the within-generation temporal variation perceived by the organism. This suggests that empirical estimates of the breadth of adaptation may provide a useful index of this type of environmental variation from the organism's point of view.     

Relating environmental variation to selection on reaction norms: an experimental test

Theoretical models predict that selection on reaction norms should depend on the relative frequency of environmental states experienced by a population. We report a laboratory experimental test of this prediction for thermal performance curves of larval growth rate in Pieris rapae in relation to their thermal environment. We measured short-term relative growth rate (RGR) for each individual at a series of five temperatures, and then we assigned individuals randomly to warm or cool selection treatments, which differ in the frequency distributions of environmental temperatures. Selection gradient analyses of two independent experiments demonstrated significant positive selection for increasing RGR, primarily through its effects on survival to adulthood and on development rate. In both the warm and cool selection treatments, the magnitude of directional selection on RGR was consistently greater at lower (suboptimal) temperatures than at higher temperatures; differences in selection between the treatments did not match model predictions. The temporal order and duration of environmental conditions may affect patterns of selection on thermal performance curves and other continuous reaction norms, complicating the connections between variation in environment, phenotype, and fitness.     

Convergent lifespan reaction norms in the yeast cultures exposed to different environmental stresses

Lifespan extension under mild stress is frequently observed although difficult to quantify and generalize as previous studies differed substantially in specific experimental arrangements. We cultured the budding yeast in several environments defined by different temperature, source of energy, saline concentration or combinations of these factors. Cells obtained under different growth regimes were transferred to identical and generally nonstressful conditions except for an absence of organic carbon. Chronological lifespan (CL) of the starving cells showed an approximately common norm of reaction when plotted against the growth rate which served as a measure of stress intensity. CL increased roughly 50% in cultures raised at moderately slower pace, regardless of what particular single or multiple stress signals were present, and then decreased gradually with a deepening growth deceleration. We suggest that the strongly nonlinear relation between the metabolic rate and longevity can be a potent constraint controlling norms of phenotypic reaction in a variety of environmental gradients.     

Latitudinal and voltinism compensation shape thermal reaction norms for growth rate

Latitudinal variation in thermal reaction norms of key fitness traits may inform about the response of populations to climate warming, yet their adaptive nature and evolutionary potential are poorly known. We assessed the contribution of quantitative genetic, neutral genetic and environmental effects to thermal reaction norms of growth rate for populations of the damselfly Ischnura elegans. Among populations, reaction norms differed primarily in elevation, suggesting that time constraints associated with shorter growth seasons in univoltine, high-latitude as well as multivoltine, low-latitude populations selected for faster growth rates. Phenotypic divergence among populations is consistent with selection rather than drift as Q(ST) was greater than F(ST) in all cases. Q(ST) estimates increased with experimental temperature and were influenced by genotype by environment interactions. Substantial additive genetic variation for growth rate in all populations suggests that evolution of trait means in different environments is not constrained. Heritability of growth rates was higher at high temperature, driven by increased genetic rather than environmental variance. While environment-specific nonadditive effects also may contribute to heritability differences among temperatures, maternal effects did not play a significant role (where these could be accounted for). Genotype by environment interactions strongly influenced the adaptive potential of populations, and our results suggest the potential for microevolution of thermal reaction norms in each of the studied populations. In summary, the observed latitudinal pattern in growth rates is adaptive and results from a combination of latitudinal and voltinism compensation. Combined with the evolutionary potential of thermal reaction norms, this may affect populations' ability to respond to future climate warming.     

Translating environmental gradients into discontinuous reaction norms via hormone signalling in a polyphenic butterfly

Polyphenisms—the expression of discrete phenotypic morphs in response to environmental variation—are examples of phenotypic plasticity that may potentially be adaptive in the face of predictable environmental heterogeneity. In the butterfly Bicyclus anynana, we examine the hormonal regulation of phenotypic plasticity that involves divergent developmental trajectories into distinct adult morphs for a suite of traits as an adaptation to contrasting seasonal environments. This polyphenism is induced by temperature during development and mediated by ecdysteroid hormones. We reared larvae at separate temperatures spanning the natural range of seasonal environments and measured reaction norms for ecdysteroids, juvenile hormones (JHs) and adult fitness traits. Timing of peak ecdysteroid, but not JH titres, showed a binary response to the linear temperature gradient. Several adult traits (e.g. relative abdomen mass) responded in a similar, dimorphic manner, while others (e.g. wing pattern) showed a linear response. This study demonstrates that hormone dynamics can translate a linear environmental gradient into a discrete signal and, thus, that polyphenic differences between adult morphs can already be programmed at the stage of hormone signalling during development. The range of phenotypic responses observed within the suite of traits indicates both shared regulation and independent, trait-specific sensitivity to the hormone signal.     

Plasticity of size and growth in fluctuating thermal environments: comparing reaction norms and performance curves

Ectothermic animals exhibit two distinct kinds of plasticityin response to temperature: Thermal performance curves (TPCs),in which an individual's performance (e.g., growth rate) variesin response to current temperature; and developmental reactionnorms (DRNs), in which the trait value (e.g., adult body sizeor development time) of a genotype varies in response to developmentaltemperatures experienced over some time period during development.Here we explore patterns of genetic variation and selectionon TPCs and DRNs for insects in fluctuating thermal environments.First, we describe two statistical methods for partitioningtotal genetic variation into variation for overall size or performanceand variation in plasticity, and apply these methods to availabledatasets on DRNs and TPCs for insect growth and size. Our resultsindicate that for the datasets we considered, genetic variationin plasticity represents a larger proportion of the total geneticvariation in TPCs compared to DRNs, for the available datasets.Simulations suggest that estimates of the genetic variationin plasticity are strongly affected by the number and rangeof temperatures considered, and by the degree of nonlinearityin the TPC or DRN. Second, we review a recent analysis of fieldselection studies which indicates that directional selectionfavoring increased overall size is common in many systems—thatbigger is frequently fitter. Third, we use a recent theoreticalmodel to examine how selection on thermal performance curvesrelates to environmental temperatures during selection. Themodel predicts that if selection acts primarily on adult sizeor development time, then selection on thermal performance curvesfor larval growth or development rates is directly related tothe frequency distribution of temperatures experienced duringlarval development. Using data on caterpillar temperatures inthe field, we show that the strength of directional selectionon growth rate is predicted to be greater at the modal (mostfrequent) temperatures, not at the mean temperature or at temperaturesat which growth rate is maximized. Our results illustrate someof the differences in genetic architecture and patterns of selectionbetween thermal performance curves and developmental reactionnorms.     

Dimorphic growth patterns and sex‐specific reaction norms in the butterfly Lycaena hippothoe sumadiensis

Directly developing larvae of the butterfly Lycaena hippothoe sumadiensis exhibited two growth strategies with one cohort passing four larval instars at high growth rates, and the other five instars at lower growth rates. The 4‐instar‐cohort displayed decreased development times, in combination with slightly reduced pupal and adult weights. In addition to adjustment of growth rate, omitting a larval instar may comprise a further mechanism to decrease development time when needed. Using the 4‐instar‐cohort, sex‐related differences in reaction norms were investigated over a temperature gradient. At high temperatures, protandrous males showed early emergence at a reduced size, whereas weight of females remained similar throughout. These differences suggest that large size is more important for female than for male fitness. The pattern is similar to that previously reported for alpine L. tityrus, indicating that sex‐specific reaction norms might be widespread in species living under severe time constraints.     

Maternal effects of egg size in brown trout (Salmo trutta): norms of reaction to environmental quality

The magnitude of fitness variation caused by maternal effects and, thus, the adaptive significance of maternal traits may depend on environmental quality, generating crossing reaction norms among offspring phenotypes that shape life-history evolution. By manipulating intraclutch variation in egg size and comparing siblings we examined the maternal effects of egg size on offspring performance and tested for the existence of reaction norms to environmental quality using the brown trout Salmo trutta. When sibling groups of small and large eggs were reared separately in a hatchery environment initial size differences disappeared rapidly. However, in semi-natural environments and under direct competition, juveniles from large eggs experienced growth and survival advantages over siblings from small eggs. Moreover, distinct reaction norms existed, with the differences in performance of juveniles from small and large eggs being most pronounced in the poorer growth environments. Our results provide the first direct evidence, to our knowledge, for a causal relationship between egg size and fitness-related traits in fishes, independent of potentially confounding genetic effects. Moreover, they indicate that previous studies have been biased by experimental conditions that excluded competitive asymmetries and environmental variability. The existence of reaction norms indicates a shift in optimal egg size across gradients of environmental quality that probably shapes the evolution of this trait.     

Impact of environmental disturbances on estimated genetic parameters and breeding values for growth traits in pigs

Due to the diversification of farming systems and climate change, farm animals are exposed to environmental disturbances to which they respond differently depending on their robustness. Disturbances such as heat stress or sanitary challenges (not always recorded, especially when they are of short duration and low intensity) have a transitory impact on animals, resulting in changes in phenotypes of production (feed intake, BW, etc.). The aim of this study was to evaluate the impact of such unknown disturbances on the estimated genetic parameters and breeding values (BV) for production traits. A population of 6 120 individuals over five generations divided into eight batches of 10 pens was generated, each individual underwent an ?100-day test period. A longitudinal phenotype mimicking piglet weight during the fattening period was simulated for each individual in two situations: disturbed and non-disturbed. The disturbed phenotype was modified according to the robustness of the animal and the intensity and duration of the disturbance that the animal was subjected to. Various sets of simulations (1 000 replicates per set) were considered depending on the type of disturbance (at the level of the batch, pen, or individual), the genetic correlation (negative, neutral, or positive) between the two components of the robustness (resistance and resilience), the genetic correlation (negative, neutral, or positive) between growth and the components of robustness, and the heritability of the components of robustness (weak or moderate). An animal model was used to estimate the genetic parameters and BV for two production traits: the BW at 100 days of age (BW₁₀₀) and average daily gain (ADG). The estimated heritability of the production traits was lower in the disturbed situation compared to the non-disturbed one (reduction of 0.08 and 0.05 points respectively for BW₁₀₀ and ADG). The correlations between estimated breeding values of the observed phenotypes (EBV) and BV for production traits in absence of disturbance were lower in the disturbed situation (reduction of 0.04 and 0.06 points for BW₁₀₀ and ADG respectively) while the partial correlation between EBV and BV for robustness was not significantly different from 0 in the two situations. These results suggest that selection in a well-controlled environment with random disturbances of low intensities does not allow to improve animal robustness while it is less effective for improving production traits than selection under no environmental disturbances.     

A quantitative genetic model for growth,shape, reaction norms,and other infinite-dimensional characters

Infinite-dimensional characters are those in which the phenotype of an individual is described by a function, rather than by a finite set of measurements. Examples include growth trajectories, morphological shapes, and norms of reaction. Methods are presented here that allow individual phenotypes, population means, and patterns of variance and covariance to be quantified for infinite-dimensional characters. A quantitative-genetic model is developed, and the recursion equation for the evolution of the population mean phenotype of an infinite-dimensional character is derived. The infinite-dimensional method offers three advantages over conventional finite-dimensional methods when applied to this kind of trait: (1) it describes the trait at all points rather than at a finite number of landmarks, (2) it eliminates errors in predicting the evolutionary response to selection made by conventional methods because they neglect the effects of selection on some parts of the trait, and (3) it estimates parameters of interest more efficiently.     

Diet-dependent body morphology and ontogenetic reaction norms in Eurasian perch

Individual morphology and performance are directly or indirectly under the influence of variation in resource levels. To study the effects of different resource conditions and their effects on morphology and ontogenetic reaction norms in young‐of‐the‐year (YOY) perch (Perca fluviatilis), we used three different approaches. First, we examined the morphological trajectories over early ontogeny in relation to lake‐specific resources in a field study. Second, one lake that lacked perch recruitment was stocked with perch eggs from a control lake in a whole‐lake experiment to study ontogenetic reaction norms. Third, we compared the development of YOY perch in the three lakes that mainly inhabited the littoral zone with YOY perch experimentally confined to enclosures in the pelagic zone of the lakes.
Overall body morphology of the YOY perch changed both as a function of size and as a function of diet. As perch increased in size they developed a deeper body morphology corresponding to an increased proportion of benthic macroinvertebrates in their diet. In pelagic enclosures where perch were constrained to feed mainly on zooplankton they had a more fusiform body morphology than perch in the lakes that fed on a mixture of zooplankton and macroinvertebrates. Similarly, the ontogenetic reaction norm of perch was related to the diet and lake‐specific zooplankton levels in the whole‐lake experiment.
In the pelagic enclosures, perch with high growth rates had a more fusiform body morphology than slow‐growing perch, whereas the opposite was found in the lakes, where perch included more macroinvertebrates in their diets. Perch in lakes with a higher proportion of macroinvertebrates in the diet also had deeper body morphology. The opposite morphology – growth rate relationship found between perch in the pelagic versus those using the whole lake suggest a morphological trade‐off between foraging on zooplankton and foraging on macroinvertebrates. Our results suggest that YOY perch show different ontogenetic reaction norms as a function of lake‐specific resource levels, which may allow YOY recruitment to later stages. Our results further suggest that diet‐related changes in morphology are a rapid process.     

The characterization of complex continuous norms of reaction

Recent focus on the array of phenotypes expressed under differing environmental conditions, or phenotypic plasticity, has led to increased understanding of its genetic basis as well as its adaptive significance. However, the quantification of plasticity has proven difficult, hampered by both the limited number of environments over which plasticity may typically be assessed and by the need to assume, a priori, the general form of reaction norms under study. Our understanding of the shapes of continuous norms of reaction and, consequently, the subtle differences that may exist in shapes among genotypes or populations is rudimentary. Here, we propose the use of the loess smoothing function to analyze complex norms of reaction and to quantify total plasticity over many environments. A thermogradient incubator offers an ideal means to provide many environments for a demonstration of the use of the loess method. We test seed germination in three populations of two monocarpic plant species for population differentiation in plasticity to temperature. First, we test for differentiation in norms of reaction to 30 temperature environments among three populations of the monocarpic perennial, Lobelia inflata . The second demonstration assesses plasticity to eight temperature environments of three populations of the arctic-alpine annual, Koenigia islandica . Our demonstration shows that the loess technique can detect significant genetic differentiation among populations in complex norms of reaction for both species studied, and suggests that the use of this procedure should be considered where the form of norms of reaction might be complex. The general applicability of the approach is discussed.     

Effect of electrochemical redox reaction on growth and metabolism of Saccharomyces cerevisiae as an environmental factor

The effect of an electrochemically generated oxidation-reduction potential and electric pulse on ethanol production and growth of Saccharomyces cerevisiae ATCC 26603 was experimented and compared with effects of electron mediators (neutral red, benzyl viologen, and thionine), chemical oxidants (hydrogen peroxide and hypochlorite), chemical reductants (sulfite and nitrite), oxygen, and hydrogen. The oxidation (anodic) and reduction (cathodic) potential and electric pulse activated ethanol production and growth, and changed the total soluble protein pattern of the test strain. Neutral red electrochemically reduced activated ethanol production and growth of the test strain, but benzyl viologen and thionine did not. Nitrite inhibited ethanol production but did not influence growth of the test strain. Hydrogen peroxide, hypochlorite, and sulfite did not influence ethanol production and growth of the test strain. Hydrogen and oxygen also did not influence the growth and ethanol production. It shows that the test strain may perceive electrochemically generated oxidation-reduction potential and electric pulse as an environmental factor.     

Impact of heat waves on mortality in Croatia

The aim of this work was to determine the criteria for heat loads associated with an increase in mortality in different climatic regions of Croatia. The relationship between heat stress and mortality was analysed for the period 1983–2008. The input series is excess mortality defined as the deviations of mortality from expected values determined by means of a Gaussian filter of 183 days. The assessment of the thermal environment was performed by means of physiologically equivalent temperature (PET). The curve depicting the relationship between mortality and temperature has a U shape, with increased mortality in both the cold and warm parts of the scale but more pronounced in the warm part. The threshold temperature for increased mortality was determined using a scatter plot and fitting data by means of moving average of mortality; the latter is defined as the temperature at which excess mortality becomes significant. The values are higher in the continental part of Croatia than at the coast due to the refreshing influence of the sea during the day. The same analysis on a monthly basis shows that at the beginning of the warm season increased mortality occurs at a lower temperature compared with later on in the summer, and the difference is up to 15 °C between August and April. The increase in mortality is highest during the first 3–5 days and after that it decreases and falls below the expected value. Long-lasting heat waves present an increased risk, but in very long heat waves the increase in mortality is reduced due to mortality displacement.     

Evolution of oxygen evolving reaction center-II of thylakoids

A model for evolution of oxygen evolving reaction center II of higher plant initiating from a chlorophyll--quinone complex is proposed. The reaction center gradually incorporates pheophytin, Fe and Mn to finally achieve oxidation of water to oxygen. The structural and functional pattern during evolution is proposed to descend from higher order of symmetry to lower one.     

Host plant adaptation and the evolution of thermal reaction norms

For most ectotherms, increasing the rearing temperature reduces the final (adult) body size, producing a negative slope for the thermal reaction norm. Recent studies show that this relationship may be reversed under conditions of low resource quality, producing a positive slope for the thermal reaction norm. If populations or species differ in the degree of evolutionary adaptation to a resource, how does this differential adaptation alter their thermal reaction norms? We used a common garden experiment with the tobacco hornworm, Manduca sexta, to address this question. We examined the thermal reaction norms for body size of two populations of M. sexta that differ in their evolutionary exposures to an atypical, low-quality resource (devil's claw; Proboscidea louisianica), but have comparable exposures to a typical, high-quality resource (tobacco; Nicotiana tabacum). Both populations had increased mean larval mortalities and development times when reared on devil's claw compared with tobacco, but the magnitudes of these increases differed between populations. Both populations had similar, negatively sloped thermal reaction norms on the typical, high-quality resource (tobacco), but had divergent, non-negative thermal reaction norms on the atypical, low-quality resource (devil's claw): the population with the longer evolutionary history of exposure to the atypical resource exhibited a flat (rather than positive) reaction norm. These results suggest that population differences in host plant adaptation can predictably influence the slopes of thermal reaction norms.