Change In Mean Phenotype Research Articles

Is evolution predictable? Quantitative genetics under complex genotype-phenotype maps.

A fundamental aim of post-genomic 21st century biology is to understand the genotype-phenotype map (GPM) or how specific genetic variation relates to specific phenotypic variation. Quantitative genetics approximates such maps using linear models, and has developed methods to predict the response to selection in a population. The other major field of research concerned with the GPM, developmental evolutionary biology, or evo-devo, has found the GPM to be highly nonlinear and complex. Here, we quantify how the predictions of quantitative genetics are affected by a complex, nonlinear map based on the development of a multicellular organ. We compared the predicted change in mean phenotype for a single generation using the multivariate breeder's equation, with the change observed from the model of development. We found that there are frequent disagreements between predicted and observed responses to selection due to the nonlinear nature of the genotype-phenotype map. Our results are a step toward integrating the fields studying the GPM.

Temperature has a causal and plastic effect on timing of breeding in a small songbird.

Phenotypic plasticity is an important mechanism by which an individual can adapt its seasonal timing to predictable, short-term environmental changes by using predictive cues. Identification of these cues is crucial to forecast the response of species to long-term environmental change and to study their potential to adapt. Individual great tits (Parus major) start reproduction early under warmer conditions in the wild, but whether this effect is causal is not well known. We housed 36 pairs of great tits in climate-controlled aviaries and 40 pairs in outdoor aviaries, where they bred under artificial contrasting temperature treatments or in semi-natural conditions, respectively, for two consecutive years, using birds from lines selected for early and late egg laying. We thus obtained laying dates in two different thermal environments for each female. Females bred earlier under warmer conditions in climate-controlled aviaries, but not in outdoor aviaries. The latter was inconsistent with laying dates from our wild population. Further, early selection line females initiated egg laying consistently ∼9 days earlier than late selection line females in outdoor aviaries, but we found no difference in the degree of plasticity (i.e. the sensitivity to temperature) in laying date between selection lines. Because we found that temperature causally affects laying date, climate change will lead to earlier laying. This advancement is, however, unlikely to be sufficient, thereby leading to selection for earlier laying. Our results suggest that natural selection may lead to a change in mean phenotype, but not to a change in the sensitivity of laying dates to temperature.

Evolutionary consequences of nonselective harvesting in density-dependent populations.

There is now considerable empirical evidence that evolutionary changes in many phenotypic characters, such as body mass, age at maturation, and timing of breeding, often occur in populations subject to intense harvesting over longer periods. Here, we analyze the evolutionary component of the selection due to nonselective harvesting, which will operate even under selective harvesting and may generate a large evolutionary response. If phenotype affects susceptibility to density dependence-for example, through resource limitation-then nonselective harvesting can induce evolutionary change through its effect on population density. We provide a model for evolution of a quantitative character in such a fluctuating density-dependent population, using the diffusion approximation to describe jointly the temporal changes in mean phenotype and log population size. We show how nonselective harvesting in particular generates r-selection governed by genetic variation in the strength of density regulation and the magnitude of population fluctuations. We show that r-selection caused by nonselective harvesting is proportional to the mean fraction of the population harvested. We then compare the short-term as well as the long-term evolutionary impact of nonselective harvesting for different harvesting strategies by using the mean harvest fraction for different strategies. This comparison is performed for three different harvesting strategies: constant, proportional, and threshold harvesting. The more ecologically sustainable strategies also produce smaller evolutionary changes.

AN EXACT FORM OF THE BREEDER'S EQUATION FOR THE EVOLUTION OF A QUANTITATIVE TRAIT UNDER NATURAL SELECTION

Starting with the Price equation, I show that the total evolutionary change in mean phenotype that occurs in the presence of fitness variation can be partitioned exactly into five components representing logically distinct processes. One component is the linear response to selection, as represented by the breeder's equation of quantitative genetics, but with heritability defined as the linear regression coefficient of mean offspring phenotype on parent phenotype. The other components are identified as constitutive transmission bias, two types of induced transmission bias, and a spurious response to selection caused by a covariance between parental fitness and offspring phenotype that cannot be predicted from parental phenotypes. The partitioning can be accomplished in two ways, one with heritability measured before (in the absence of) selection, and the other with heritability measured after (in the presence of) selection. Measuring heritability after selection, though unconventional, yields a representation for the linear response to selection that is most consistent with Darwinian evolution by natural selection because the response to selection is determined by the reproductive features of the selected group, not of the parent population as a whole. The analysis of an explicitly Mendelian model shows that the relative contributions of the five terms to the total evolutionary change depends on the level of organization (gene, individual, or mated pair) at which the parent population is divided into phenotypes, with each frame of reference providing unique insight. It is shown that all five components of phenotypic evolution will generally have nonzero values as a result of various combinations of the normal features of Mendelian populations, including biparental sex, allelic dominance, inbreeding, epistasis, linkage disequilibrium, and environmental covariances between traits. Additive genetic variance can be a poor predictor of the adaptive response to selection in these models. The narrow-sense heritability sigma2A/sigma2P should be viewed as an approximation to the offspring-parent linear regression rather than the other way around.

AN EXACT FORM OF THE BREEDER'S EQUATION FOR THE EVOLUTION OF A QUANTITATIVE TRAIT UNDER NATURAL SELECTION

Starting with the Price equation, I show that the total evolutionary change in mean phenotype that occurs in the presence of fitness variation can be partitioned exactly into five components representing logically distinct processes. One component is the linear response to selection, as represented by the breeder's equation of quantitative genetics, but with heritability defined as the linear regression coefficient of mean offspring phenotype on parent phenotype. The other components are identified as constitutive transmission bias, two types of induced transmission bias, and a spurious response to selection caused by a covariance between parental fitness and offspring phenotype that cannot be predicted from parental phenotypes. The partitioning can be accomplished in two ways, one with heritability measured before (in the absence of) selection, and the other with heritability measured after (in the presence of) selection. Measuring heritability after selection, though unconventional, yields a representation for the linear response to selection that is most consistent with Darwinian evolution by natural selection because the response to selection is determined by the reproductive features of the selected group, not of the parent population as a whole. The analysis of an explicitly Mendelian model shows that the relative contributions of the five terms to the total evolutionary change depends on the level of organization (gene, individual, or mated pair) at which the parent population is divided into phenotypes, with each frame of reference providing unique insight. It is shown that all five components of phenotypic evolution will generally have nonzero values as a result of various combinations of the normal features of Mendelian populations, including biparental sex, allelic dominance, inbreeding, epistasis, linkage disequilibrium, and environmental covariances between traits. Additive genetic variance can be a poor predictor of the adaptive response to selection in these models. The narrow-sense heritability σA2/σP2 should be viewed as an approximation to the offspring-parent linear regression rather than the other way around.

Floral trait variation in Spergularia marina (Caryophyllaceae): ontogenetic, maternal family, and population effects

Traits expressed by modular organisms present difficulties when estimating the genetic component to their variation if their phenotype changes as an individual ages, confounding ontogenetic and genetic sources of phenotypic variation. For such traits, it is necessary to control for ontogenetic effects in order to estimate accurately the degree of genetic variation in a trait. To measure the magnitude of ontogenetic change in floral traits and to determine whether it may obscure underlying genetic sources of floral trait variation in the autogamous annual, Spergularia marina (Caryophyllaceae), we monitored the expression of floral traits over a five-week period in greenhouse-raised plants from four California wild populations. From 130 individuals representing 8–10 maternal families per population, we sampled one flower per week to record the number of ovules, normal anthers (the number of which is positively correlated with pollen production per flower), abnormal anthers (those that grade phenotypically into petals), and petals; and the areas of a single petal and the entire corolla. All traits except the number of abnormal anthers exhibited strong temporal changes in phenotype, although the direction and magnitude of the change differed among traits. To determine whether populations appear to be evolving independently, we examined differences among them in floral trait means, in the magnitude of among-family variation, and in the degree to which they show temporal change in floral trait means. There were significant differences among population means for all traits except the number of ovules and petals per flower. In addition, populations differ in those traits that exhibit significant differences among maternal family means. For most traits, populations also differ in the magnitude of week-to-week changes in mean phenotype, suggesting the presence of genetic variation among populations in the expression of temporal change in floral traits. Finally, for most traits, the magnitude and significance of differences among populations and among maternal family means changed over time. Consequently, broad-sense heritability estimates, predictions of the response to selection, and measures of interpopulation divergence for floral traits are sensitive to the time of flower sampling in S. marina. The role of natural selection in moulding ontogenetic variation in floral traits has not been extensively studied in any species, but the presence of genetic variation among populations observed here suggests that this character is open to evolutionary change.

Genetic analysis of shell-shape variation in Littorina saxatilis on an environmental cline

Three shell-shape parameters of Littorina saxatilis were measured and found to vary in a regular pattern with distance up an estuary. The translation rate of the shell increased, the rate of whorl expansion decreased and the circularity of the aperture decreased proceeding from the exposed shore to the protected shore. The genetic variance of these traits was estimated from the full-sib covariance and the motheroffspring covariance. The genetic variance of the translation rate and the circularity of the aperture was low in all populations, but the genetic variance of the rate of whorl expansion was high on the exposed shore and low on the protected shore. It is argued that the change in mean phenotype of these traits is the result of natural selection produced by varying degrees of wave action and desiccation. The observed genetic variance is consistent with the theory that a trait under selection will show little additive genetic variance.