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Triacylglycerol and coronary heart disease.

Based on meta-analysis of prospective studies from the epidemiological literature, TAG is a risk factor for CVD, independent of HDL-cholesterol. The RR values were 1.3 and 1.8 for a 1 mmol/l increase in TAG among men and women respectively. Adjustment for HDL-cholesterol and other risk factors attenuated these estimates, but they remained statistically significant. Recent prospective findings from the Stanford Five City Project (Gardner et al. 1996) and the Physicians' Health Study (Stampfer et al. 1996) further demonstrate that TAG and LDL size are highly inter-related risk factors for CHD. Quantitative genetic analysis from large-scale family studies show that these correlations reflect common genetic influences that may be important for understanding genetic susceptibility to CHD.

Coronary Disease↗

Lactational traits of importance in dairy cows and applications for emerging biotechnologies.

New molecular and quantitative genetic technologies are the latest in a long list of technologies that have been introduced to dairy industries over many decades to improve the performance of cows. The catalysts for future advances will be sequencing of the bovine genome and development of high-throughput technologies to identify and exploit relevant variation in DNA sequences. The new technologies will allow the selection of animals based on specific genotypes that suit specific applications such as once-daily (OAD) milking or extended lactation. These technologies will also allow exploitation of between-cow variation in milk composition, which is currently hidden by bulking of milk on and between farms. Thus, there are opportunities to produce milk from herds of cows selected for specific milk composition that might be, for example, more suitable for cheese-making or have milkfat with specific properties to enhance human health. Identification of genes and gene polymorphisms associated with improved resistance to specific diseases in cows and other farm species also represents a real opportunity to improve animal health over the coming decade. New levels of genetic regulation have been identified, e.g. microRNA and epigenetics, the impacts of which on the performance of cows and humans are only just beginning to be understood. There is a potential role for veterinarians to provide or be the interface for provision of genetic advice to farmers in much the same way that nutritional advice is currently given.

Animals↗

Heritability of plasma von Willebrand factor antigen concentration in German Wirehaired pointers.

We applied quantitative genetic analyses to a population of German Wirehaired pointer dogs affected with type 2 von Willebrand disease. Plasma von Willebrand factor (vWF) protein concentration measured as vWF antigen (vWF:Ag), clinical history, and pedigree data were compiled for 331 dogs over a 5-year test period. Eight dogs had histories of abnormal bleeding and had markedly decreased plasma vWF:Ag concentrations (<1%). Four per cent of the dogs were inbred, with an average inbreeding of 2.52%. The estimated heritability of plasma vWF concentration was 0.52. We found a major gene effect on vWF concentration. Using a single gene locus model and two different prediction methods, the upper threshold value for the aa genotype was less than 1% vWF:Ag, and the optimal threshold value for discrimination between the AA and Aa genotypes was between 68% and 72% vWF:Ag. Our analyses indicate that phenotype, assigned on the basis of a single vWF:Ag determination, is heritable and can be applied for selective breeding in a von Willebrand disease test programme.

Animals↗

Variation among families for characteristics of the adhesive plaque in the barnacle Balanus amphitrite.

A quantitative genetics approach was used to examine variation in the characteristics of the adhesive plaque of the barnacle Balanus amphitrite Darwin attached to two silicone substrata. Barnacles settled on silicone polymer films occasionally form thick, soft adhesive plaques, in contrast to the thin, hard plaques characteristic of attachment to other surfaces. The proportion of barnacles producing a thick adhesive plaque was 0.31 for Veridian, a commercially available silicone fouling-release coating, and 0.18 for Silastic T-2, a silicone rubber used for mold-making. For both materials, significant variation among maternal families in the proportion of barnacles producing a thick adhesive plaque was observed, which suggests the presence of genetic variation, or maternal environmental effects, for this plaque characteristic. For the Veridian coating, barnacles expressing the thick adhesive plaque also exhibited significantly reduced tenacity. This represents the first reported case for potential genetic control of intraspecific phenotypic variation in the physical characteristics and tenacity of the adhesive of a fouling invertebrate.

Animals↗

A general theory for the evolutionary dynamics of virulence.

Most theory on the evolution of virulence is based on a game-theoretic approach. One potential shortcoming of this approach is that it does not allow the prediction of the evolutionary dynamics of virulence. Such dynamics are of interest for several reasons: for experimental tests of theory, for the development of useful virulence management protocols, and for understanding virulence evolution in situations where the epidemiological dynamics never reach equilibrium and/or when evolutionary change occurs on a timescale comparable to that of the epidemiological dynamics. Here we present a general theory similar to that of quantitative genetics in evolutionary biology that allows for the easy construction of models that include both within-host mutation as well as superinfection and that is capable of predicting both the short- and long-term evolution of virulence. We illustrate the generality and intuitive appeal of the theory through a series of examples showing how it can lead to transparent interpretations of the selective forces governing virulence evolution. It also leads to novel predictions that are not possible using the game-theoretic approach. The general theory can be used to model the evolution of other pathogen traits as well.

Animals↗

Selection, inheritance, and the evolution of parent-offspring interactions.

Very few studies have examined parent-offspring interactions from a quantitative genetic perspective. We used a cross-fostering design and measured genetic correlations and components of social selection arising from two parental and two offspring behaviors in the burying beetle Nicrophorus vespilloides. Genetic correlations were assessed by examining behavior of relatives independent of common social influences. We found positive genetic correlations between all pairs of behaviors, including between parent and offspring behaviors. Patterns of selection were assessed by standardized performance and selection gradients. Parental provisioning had positive effects on offspring performance and fitness, while remaining near the larvae without feeding them had negative effects. Begging had positive effects on offspring performance and fitness, while increased competition among siblings had negative effects. Coadaptations between parenting and offspring behavior appear to be maintained by genetic correlations and functional trade-offs; parents that feed their offspring more also spend more time in the area where they can forage for themselves. Families with high levels of begging have high levels of sibling competition. Integrating information from genetics and selection thus provides a general explanation for why variation persists in seemingly beneficial traits expressed in parent-offspring interactions and illustrates why it is important to measure functionally related suites of behaviors.

Animals↗

Temporal variation can facilitate niche evolution in harsh sink environments.

We examine the impact of temporal variation on adaptive evolution in "sink" environments, where a species encounters conditions outside its niche. Sink populations persist because of recurrent immigration from sources. Prior studies have highlighted the importance of demographic constraints on adaptive evolution in sinks and revealed that adaptation is less likely in harsher sinks. We examine two complementary models of population and evolutionary dynamics in sinks: a continuous-state quantitative-genetics model and an individual-based model. In the former, genetic variance is fixed; in the latter, genetic variance varies because of mutation, drift, and sampling. In both models, a population in a constant harsh sink environment can exist in alternative states: local maladaptation (phenotype comparable to immigrants from the source) or adaptation (phenotype near the local optimum). Temporal variation permits transitions between these states. We show that moderate amounts of temporal variation can facilitate adaptive evolution in sinks, permitting niche evolution, particularly for slow or autocorrelated variation. Such patterns of temporal variation may particularly pertain to sinks caused by biotic interactions (e.g., predation). Our results are relevant to the evolutionary dynamics of species' ranges, the fate of exotic invasive species, and the evolutionary emergence of infectious diseases into novel hosts.

Adaptation, Biological↗

The evolution of infidelity in socially monogamous passerines: the strength of direct and indirect selection on extrapair copulation behavior in females.

Many studies have been aimed at understanding the maintenance of female infidelity in socially monogamous birds. Because engaging in extrapair copulations (EPCs) is believed to be costly for females, it has been argued that EPC behavior must bring indirect benefits to females by elevating offspring fitness. We use empirical data from the literature to assess the relative strength of indirect and direct selection on female EPC behavior, using quantitative genetic approximations of selection. This analysis confirmed that there is generally negative direct selection on EPC behavior caused by depressed paternal investment by social males. In contrast, there was no significant positive indirect selection on EPC behavior in females. A comparison between the two types of selection suggests that the force of direct negative selection is generally much stronger than that of indirect positive selection. Indirect selection is thus unlikely to maintain EPC behavior in the face of direct selection against it. We suggest that EPCs may instead be the result of antagonistic selection on loci influencing the outcome of male-female encounters and that EPC behavior per se may not be adaptive for females but may reflect sexual conflict due to strong selection in males to achieve extrapair copulations.

Animals↗

Escalation, retreat, and female indifference as alternative outcomes of sexually antagonistic coevolution.

Verbal and quantitative genetic models of sexually antagonistic coevolution suggest that coevolutionary arms races should be common. Sexual selection favors exaggeration of male persistence traits that are costly to females, and females, in turn, are selected to resist these traits. The heightened resistance by females is thought to then favor further exaggeration in the male trait, leading to an escalating coevolutionary arms race between persistence and resistance traits. Much of this theory, however, is based on an (implicit) assumption that there are tight constraints on how female resistance can evolve. We develop a theory that identifies and relaxes these constraints, allowing female resistance to evolve in a fashion that better reflects known empirical patterns in the evolution of female preference functions (the resistance trait). Our results suggest that evolutionary arms races that lead to the exaggeration of persistence and resistance will be much less common than formerly predicted. Females sometimes evolve indifference to male traits rather than resistance and can even evolve to discriminate against these traits. These alternative outcomes depend on the existence of genetic variance in the components of the female sensory system underlying female resistance and on the strength of natural selection acting on these components. Female indifference tends to evolve when natural selection on the sensory system is weak, and under these conditions, sexually antagonistic coevolution tends not to reduce female fitness significantly at equilibrium. When natural selection on the female sensory system is strong, however, then arms races are more likely, and female fitness is then sometimes significantly depressed at equilibrium. Sexually antagonistic coevolution is thus likely to have strong deleterious effects on population fitness only when female sensory traits are under strong natural selection to perform functions in addition to those involved with mating. Together, these results suggest that identifying the nature of genetic variation in and the strength of natural selection on female resistance should be a central goal of future studies of sexual conflict.

Animal Communication↗

Identification of selective sources: partitioning selection based on interactions.

Interspecific interactions are an inescapable reality in nature. The evolution of a species is largely determined by the environment, abiotic or biotic, in which selection occurs. Quantifying the magnitude of selection is crucial to understanding which aspects of the environment are important to the evolution of a species. Such knowledge is particularly important to fields such as conservation biology, which attempts to maintain a suitable environment for the prosperity of a species, or coevolution, where dynamics are determined by the strength of reciprocal selection between species. I present a general method by which selection due to interspecific interactions may be quantified. This technique is based on past quantitative genetic models of selection and can be used with other methodologies that build on these standard models. The approach may be expanded to account for n-species interactions (e.g., a plant with two pollinators). Simulation studies conducted using this method indicate that the magnitude of selection between two species is strongly correlated with the presence of nonrandom interactions.

Biological Evolution↗

Variation in continuous reaction norms: quantifying directions of biological interest.

Thermal performance curves are an example of continuous reaction norm curves of common shape. Three modes of variation in these curves--vertical shift, horizontal shift, and generalist-specialist trade-offs--are of special interest to evolutionary biologists. Since two of these modes are nonlinear, traditional methods such as principal components analysis fail to decompose the variation into biological modes and to quantify the variation associated with each mode. Here we present the results of a new method, template mode of variation (TMV), that decomposes the variation into predetermined modes of variation for a particular set of thermal performance curves. We illustrate the method using data on thermal sensitivity of growth rate in Pieris rapae caterpillars. The TMV model explains 67% of the variation in thermal performance curves among families; generalist-specialist trade-offs account for 38% of the total between-family variation. The TMV method implemented here is applicable to both differences in mean and patterns of variation, and it can be used with either phenotypic or quantitative genetic data for thermal performance curves or other continuous reaction norms that have a template shape with a single maximum. The TMV approach may also apply to growth trajectories, age-specific life-history traits, and other function-valued traits.

Animals↗

The coadaptation of parental supply and offspring demand.

The evolution of parent-offspring interactions for the provisioning of care is usually explained as the phenotypic outcome of resolved conflicting selection pressures. However, parental care and offspring solicitation are expected to have complex patterns of inheritance. Here we present a quantitative genetic model of parent-offspring interactions that allows us to investigate the evolutionary maintenance of a state of resolved conflict. We show that offspring solicitation and parental provisioning are expected to become genetically correlated through coadaptation and that their genetic architecture is dictated by an interaction between patterns of selection and the proximate mechanisms regulating supply and demand. When selection is predominately on offspring solicitation, our model suggests that the genetic correlations between provisioning and solicitation are usually positive if provisioning reduces solicitation. Conversely, when selection is predominately on parental provisioning, the correlations are mostly negative as long as parents show a positive response to offspring demand. Empirical estimates of the genetic architecture of traits involved in family interactions fit these predictions. Our model demonstrates how the evolutionary maintenance of parent-offspring interactions can result in variable patterns of coadaptation, and it provides an explanation for the diversity of family interactions within and among species.

Adaptation, Physiological↗

Evolutionary response to selection on clutch size in a long-term study of the mute swan.

Life-history traits in wild populations are often regarded as being subject to directional selection, and the existence of substantial variation and microevolutionary stasis of these characters is therefore a problem in need of explanation. Avian clutch size is an archetypal life-history trait in this context, and many studies have sought to test explanations for stasis in clutch size. Surprisingly, there are many fewer studies that used long-term data to ask how selection acts on clutch size, particularly in a multivariate framework. In this article, we report selection, inheritance, and evolution of clutch size over 25 years in a colony of mute swans using a multivariate quantitative genetic framework to control for correlations with breeding time. We show that clutch size is influenced by both additive genetic and permanent environmental effects and that selection acts on clutch size in combination with breeding time. Natural selection on clutch size is strongly directional, favoring larger clutches, and we observe an increase in clutch size of 0.35 standard deviations, consistent with the expected response based on selection and inheritance of clutch size. We hypothesize that these changes result from recent relaxation of food constraints and predation risks experienced by this colony.

Animals↗

The evolution of infidelity in socially monogamous passerines: neglected components of direct and indirect selection.

A recent study by Goran Arnqvist and Mark Kirkpatrick in the American Naturalist (165:S26-S37) suggested that female polyandry in birds is not driven by females because quantitative genetic approximations of selection demonstrated that indirect selection for female infidelity is weaker than natural selection against it. Instead, it was argued that extrapair copulations are the result of antagonistic selection on male behavior driving female coercion. While the approach and framework of the study were very good, the conclusions of the study were premature because a number of potential adaptive components of polyandry were unaccounted for, and several critical assumptions are unsupported by the current empirical data. Our understanding of extrapair paternity in birds, and perhaps polyandry in general, will be improved by a better empirical understanding of the direct benefits of fertility assurance and postcopulatory cryptic female choice and the relationship between polyandry and male investment. In addition, we need to develop a greater awareness of the limitations of trying to study behavior by proxy in the molecular laboratory. Together, these challenges and the framework recently presented should improve our understanding of the true function of extrapair paternity in birds.

Agonistic Behavior↗

Using clusters of computers for large QU-GENE simulation experiments.

The QU-GENE Computing Cluster (QCC) is a hardware and software solution to the automation and speedup of large QU-GENE (QUantitative GENEtics) simulation experiments that are designed to examine the properties of genetic models, particularly those that involve factorial combinations of treatment levels. QCC automates the management of the distribution of components of the simulation experiments among the networked single-processor computers to achieve the speedup.

Computer Simulation↗

Computer simulation of selection in a hypothetical crop species.

A computer simulation model demonstrating the effect of directional selection upon a simplified quantitative genetic character, is presented. Complementary homozygous parents were crossed to generate a fully heterozygous F1 population. Phenotypic truncation selection was effected upon the F2 and subsequent generations, and genotypic and phenotypic response to selection, as well as the change in population heterozygosity, was monitored to generation F9. The model simulates selection for alternative breeding regimes, modes of allelic interaction, selection pressure, population size and levels of environmental variation. Three alternative methods for the simulation of the environmental component of phenotypic variance are incorporated, and mechanisms and relative merits of the rival methods are discussed. Data generated by the model conformed closely to theoretical expectation, lending support to the use of the model as a platform for development of more comprehensive simulation models. Potential improvements to the model are discussed.

Algorithms↗

Variation in amylase haplotypes among congenic lines of the house mouse.

Pancreatic amylase in the mouse displays considerable quantitative genetic variation. Agar gel electrophoresis reveals that homozygous animals have either one form of the enzyme, type A, or two forms, type AB. Only few animals have been found that contradict this statement, namely among Mus musculus castaneous from Thailand, which has a single-banded B type. Double-banded homozygous specimens of various origins have different relative proportions of the two isoenzymes. By measuring the A:B ratios in such animals, a number of distinct haplotypes or amylase complexes, determining ratios ranging from 61% A:39%, B to 12% A:88% B, have been recognized. These complexes differ also with respect to the total amount of amylase produced. If the reference stock C3H/As is given the value 1, then other haplotypes have values ranging from 1.0 to 0.27. Nineteen amylase haplotypes have been established in congenic lines on a C3H/As background. Some of these lines contain at least four active pancreatic amylase structural genes and breeding experiments have demonstrated that the genetic elements regulating total amylase production and relative proportions of the isoenzymes are located within the amylase complex, cis-acting, and very closely linked to the structural genes.

Amylases↗

Multivariate mutation-selection balance with constrained pleiotropic effects.

A multivariate quantitative genetic model is analyzed that is based on the assumption that the genetic variation at a locus j primarily influences an underlying physiological variable yj, while influence on the genotypic values is determined by a kind of "developmental function" which is not changed by mutations at this locus. Assuming additivity among loci the developmental function becomes a linear transformation of the underlying variables y onto the genotypic values x, x = By. In this way the pleiotropic effects become constrained by the structure of the B-matrix. The equilibrium variance under mutation-stabilizing selection balance in infinite and finite populations is derived by using the house of cards approximation. The results are compared to the predictions given by M. Turelli in 1985 for pleiotropic two-character models. It is shown that the B-matrix model gives the same results as Turelli's five-allele model, suggesting that the crucial factor determining the equilibrium variance in multivariate models with pleiotropy is the assumption about constraints on the pleiotropic effects, and not the number of alleles as proposed by Turelli. Finally it is shown that under Gaussian stabilizing selection the structure of the B-matrix has effectively no influence on the mean equilibrium fitness of an infinite population. Hence the B-matrix and consequently to some extent also the structure of the genetic correlation matrix is an almost neutral character. The consequences for the evolution of genetic covariance matrices are discussed.

Genetic Variation↗