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Quantitative genetics of floral traits in a gynodioecious wild strawberry Fragaria virginiana: implications for the independent evolution of female and hermaphrodite floral phenotypes.

The independent evolution of floral phenotype is an important part of the process of gender specialization during the evolution of dioecy from hermaphroditism. However, we have little information on the genetic variation of floral traits in species with separate genders. Gynodioecious species (co-occurrence of females and hermaphrodites) have a breeding system intermediate between hermaphroditism and complete separation of the sexes (dioecy) and thus can provide insight into the genetic architecture underlying floral phenotype with respect to both primary (stamens and carpels) and secondary (petals) sexual traits. I used a nested breeding design to examine the potential for response to selection on floral traits and to examine whether this response would be similar in the two sex morphs of gynodioecious Fragaria virginiana. There was significant genetic variation underlying all floral traits, although narrow-sense heritabilities (ranging from -0.25 to 0.44) were, in most cases, much lower than broad-sense ones (ranging from 0.28 to 1. 53). Moreover, the sex morphs differed significantly in their heritabilities for shared traits, such as stamen length, and showed a tendency towards differing significantly in others, like carpel number and petal length. In addition, correlations between the sex morphs for these traits (ranging from 0.41 to 0.58) were significantly greater than 0, but less than 1. These results indicate that greater sexual dimorphism could evolve in this population of F. virginiana, even if selection on these traits is not divergent. However, strong developmental integration of floral traits (e.g. stamen length and petal length) and high levels of nonadditive genetic variance may represent barriers to the evolution of complete sexual dimorphism.

Biological Evolution↗

Quantitative genetics of female choice in an ultrasonic pyralid moth, Achroia grisella: variation and evolvability of preference along multiple dimensions of the male advertisement signal.

The mating system of Achroia grisella (Lepidoptera: Pyralidae) is characterized by male ultrasonic advertisement signalling to which females orientate. Although males provide no direct, somatic benefits to their mates, females prefer males whose signal characters are more exaggerated than the population means. Previous studies showed that the signal characters influencing mate attraction are highly repeatable and heritable. We measured the phenotypic and additive genetic variances (heritability) of female preference in A. grisella, as this additive genetic variance is one of the genetic assumptions of indirect models of sexual selection. We determined the preference index of female A. grisella by repeated phonotaxis trials in which a choice of simulated male signals was presented. These playback experiments showed that female preference indices varied but were repeatable within individuals. Specifically, females differ in the relative importance of the several signal characters during mate assessment. A subsequent half-sib breeding design revealed an amount of additive genetic variance for the female preference index (h(s)2 = 0.212, SE = 0.1347, P = 0.0611; CVA = 0.1826). Our study highlights the importance of careful preparation of test signals and experimental design for quantifying individual variation in (female) preference along multiple signal dimensions.

Animals↗

Quantitative genetic models for the balance between migration and stabilizing selection.

The evolution of a quantitative trait subject to stabilizing selection and immigration, with the immigrants deviating from the local optimum, is considered under a number of different models of the underlying genetic basis of the trait. By comparing exact predictions under the infinitesimal model obtained using numerical methods with predictions of a simplified approximate model based on ignoring linkage disequilibrium, the increase in the expressed genetic variance as a result of linkage disequilibrium generated by migration is shown to be relatively small and negligible, provided that the genetic variance relative to the squared deviation of immigrants from the local optimum is sufficiently large or selection and migration is sufficiently weak. Deviation from normality is shown to be less important by comparing predictions of the infinitesimal model with a model presupposing normality. For a more realistic symmetric model, involving a finite number of loci only, no linkage and equal effects and frequencies across loci, additional changes in the genetic variance arise as a result of changes in underlying allele frequencies. Again, provided that the genetic variance relative to the squared deviation of the immigrants from the local optimum is small, the difference between the predictions of infinitesimal and the symmetric model are small unless the number of loci is very small. However, if the genetic variance relative to the squared deviation of the immigrants from the local optimum is large, or if selection and migration are strong, both linkage disequilibrium and changes in the genetic variance as a result of changes in underlying allele frequencies become important.

Base Sequence↗

Quantitative genetic analysis of ventral midbrain and liver iron in BXD recombinant inbred mice.

Male and female mice from 15 of the BXD/Ty recombinant inbred strain panel were examined for regional brain and liver iron content. Brain regions included medial prefrontal cortex, nucleus accumbens, caudate-putamen and ventral midbrain. Our focal tissue was the ventral midbrain, containing the ventral tegmentum and substantia nigra. This area contains the perikarya of the dopamine neurons that project to nucleus accumbens and caudate-putamen. Genetic correlations between ventral midbrain and liver iron content were not statistically significant, suggesting that peripheral and central iron regulatory systems are largely independent. Correlations between ventral midbrain iron and iron in the caudate-putamen and nucleus accumbens, but not the prefrontal cortex were moderately high and significant. Ventral midbrain and liver iron contents were subjected to quantitative trait loci analysis to identify associated chromosomal locations. This analysis revealed several suggestive loci for iron content in ventral midbrain but fewer loci for liver. Genetic correlations between ventral midbrain iron and published dopamine functional indices were significant, suggesting a link between ventral midbrain iron status and central dopamine neurobiology. This work shows the value of quantitative genetic analysis in the neurobiology of iron and in showing the close association between ventral midbrain iron and nigrostriatal/mesolimbic dopamine function.

Animals↗

The nature of quantitative genetic variation revisited: lessons from Drosophila bristles.

Most characters that distinguish one individual from another, like height or weight, vary continuously in populations. Continuous variation of these 'quantitative' traits is due to the simultaneous segregation of multiple quantitative trait loci (QTLs) as well as environmental influences. A major challenge in human medicine, animal and plant breeding and evolutionary genetics is to identify QTLs and determine their genetic properties. Studies of the classic quantitative traits, abdominal and sternopleural bristle numbers of Drosophila, have shown that: (1) many loci have small effects on bristle number, but a few have large effects and cause most of the genetic variation; (2) 'candidate' loci involved in bristle development often have large quantitative effects on bristle number; and (3) alleles at QTLs affecting bristle number have variable degrees of dominance, interact with each other, and affect other quantitative traits, including fitness. Lessons learned from this model system will be applicable to studies of the genetic basis of quantitative variation in other species.

Animals↗

Quantitative genetic variation: a post-modern view.

It has become commonplace to map individual quantitative trait loci (QTL) in experimental organisms; the means (line-crosses and dense maps of markers) and motivation (the close relationship between continuous physiological traits and common, complex diseases) are self-evident. Progress in mapping human QTL has been more gradual, an inevitable consequence of genetic mapping in a natural population setting. The common objective of these studies has been to understand the molecular mechanisms underlying individual QTL. Recent theoretical and practical advances shift this focus to a more comprehensive or genomic perspective on quantitative variation. Fisher's infinitesimal model of adaptive evolution, which satisfied quantitative geneticists for over 50 years, has been modified in the light of data from QTL mapping experiments in plants and animals. The resulting exponential model provides a pleasing empirical fit to the distribution of QTL effect sizes, predicts that a large amount of quantitative variation will be explained by a limited number of genes and suggests a new mathematical framework for linkage mapping. Molecular analysis of QTL suggests that coding variants (e.g. allozymes) underlie a fraction of quantitative variation and that variants that affect gene expression (expression QTL, eQTL) have a substantial role. This is supported by genomic experiments that combine expression profiling with classical genetic mapping approaches to reveal a remarkable wealth of quantitative heritable variation in the transcriptome and that cis-and trans-acting regulatory factors are organized in networks reflecting pleiotropy. It is hoped that these advances will enhance our understanding of the genetic basis of complex inherited diseases.

Animals↗

Quantitative genetic variation of leaf size and shape in a mixed diploid and triploid population of Populus.

In the interspecific cross of Populus trichocarpa x P. deltoides, unexpected simultaneous occurrence of diploid hybrids and triploid hybrids (with two alleles from the female parent and one from the male parent at each locus) led us to examine the evolutionary genetic significance of this phenomenon. As expected, leaf size and shape of the triploid progeny are closer to the female P. trichocarpa than male P. deltoides parent. Although the pure triploid progeny population did not have higher genetic variance in leaf traits than the pure diploid population, the former appears to hide much non-additive genetic variance and display strong genetic control over the phenotypic plasticity of leaf traits. It is suggested that the cryptic non-additive variance, especially epistasis, can be released when a population is disturbed by changes in the environment. A mixed diploid and triploid progeny population combines phenotypic and genetic characteristics of both pure hybrids and is considered to be of adaptive significance for populars to survive and evolve in a fluctuating environment. The significant effect due to general and specific combining ability differences at the population level suggests that the population divergence of these two species is under additive and non-additive genetic control.

Diploidy↗

A test of quantitative genetic theory using Drosophila- effects of inbreeding and rate of inbreeding on heritabilities and variance components.

Inbreeding is expected to decrease the heritability within populations. However, results from empirical studies are inconclusive. In this study, we investigated the effects of three breeding treatments (fast and slow rate of inbreeding - inbred to the same absolute level - and a control) on heritability, phenotypic, genetic and environmental variances of sternopleural bristle number in Drosophila melanogaster. Heritability, and phenotypic, genetic and environmental variances were estimated in 10 replicate lines within each of the three treatments. Standard least squares regression models and Bayesian methods were used to analyse the data. Heritability and additive genetic variance within lines were higher in the control compared with both inbreeding treatments. Heritabilities and additive genetic variances within lines were higher in slow compared with fast inbred lines, indicating that slow inbred lines retain more evolutionary potential despite the same expected absolute level of inbreeding. The between line variance was larger with inbreeding and more than twice as large in the fast than in the slow inbred lines. The different pattern of redistribution of genetic variance within and between lines in the two inbred treatments cannot be explained invoking the standard model based on selective neutrality and additive gene action. Environmental variances were higher with inbreeding, and more so with fast inbreeding, indicating that inbreeding and the rate of inbreeding affect environmental sensitivity. The phenotypic variance decreased with inbreeding, but was not affected by the rate of inbreeding. No inbreeding depression for mean sternopleural bristle number was observed in this study. Considerable variance between lines in additive genetic variance within lines was observed, illustrating between line variation in evolutionary potential.

Analysis of Variance↗

Genetic polymorphism and trade-offs in the early life-history strategy of the Pacific oyster, Crassostrea gigas (Thunberg, 1795): a quantitative genetic study.

We investigated genetic variability and genetic correlations in early life-history traits of Crassostrea gigas. Larval survival, larval development rate, size at settlement and metamorphosis success were found to be substantially heritable, whereas larval growth rate and juvenile traits were not. We identified a strong positive genetic correlation between larval development rate and size at settlement, and argue that selection could optimize both age and size at settlement. However, trade-offs, resulting in costs of metamorphosing early and large, were suggested by negative genetic correlations or covariances between larval development rate/size at settlement and both metamorphosis success and juvenile survival. Moreover, size advantage at settlement disappeared with time during the juvenile stage. Finally, we observed no genetic correlations between larval and juvenile stages, implying genetic independence of life-history traits between life-stages. We suggest two possible scenarios for the maintenance of genetic polymorphism in the early life-history strategy of C. gigas.

Animals↗

Influence of dominance, leptokurtosis and pleiotropy of deleterious mutations on quantitative genetic variation at mutation-selection balance.

In models of maintenance of genetic variance (V (G)) it has often been assumed that mutant alleles act additively. However, experimental data show that the dominance coefficient varies among mutant alleles and those of large effect tend to be recessive. On the basis of empirical knowledge of mutations, a joint-effect model of pleiotropic and real stabilizing selection that includes dominance is constructed and analyzed. It is shown that dominance can dramatically alter the prediction of equilibrium V (G). Analysis indicates that for the situations where mutations are more recessive for fitness than for a quantitative trait, as supported by the available data, the joint-effect model predicts a significantly higher V (G) than does an additive model. Importantly, for what seem to be realistic distributions of mutational effects (i.e., many mutants may not affect the quantitative trait substantially but are likely to affect fitness), the observed high levels of genetic variation in the quantitative trait under strong apparent stabilizing selection can be generated. This investigation supports the hypothesis that most V (G) comes from the alleles nearly neutral for fitness in heterozygotes while apparent stabilizing selection is contributed mainly by the alleles of large effect on the quantitative trait. Thus considerations of dominance coefficients of mutations lend further support to our previous conclusion that mutation-selection balance is a plausible mechanism of the maintenance of the genetic variance in natural populations.

Genes, Dominant↗

Quantitative genetic aspects of coat color in horses.

The aim of this study was to estimate genetic parameters for coat color in horses. Besides defining coat color classes (gray, chestnut, bay, and black), the phenotypes were also measured quantitatively according to standardized international procedures (Commission Internationale de l'Eclairage L*, a*, b*), where L* describes lightness, a* describes color saturation from red to green, and b* describes color saturation from yellow to blue. The total color saturation was derived from a* and b* and referred to as Chroma. A total of 294 horses from the breeds Lipizzan, Nonius, Arabian Pure Bred, Shagya Arabian, and Gidran were measured at neck, shoulder, and belly. Heritabilities (within and between breeds or color classes) and repeatabilities were estimated using REML from univariate animal models defined separately for gray and nongray horses. For gray horses, the estimated within-breed heritabilities for L* ranged from 0.45 to 0.49 and for a*, b*, and Chroma from 0.09 to 0.52, indicating moderate polygenic effect. For nongray horses, between-color class heritabilities were high (0.70 to 0.85) and within-color class heritabilities were negligible (except for L* measured on neck and belly, 0.21 and 0.34, respectively). Additionally, the importance of L* was described by the relation with the total melanin content of horse coat hair; for gray and nongray horses, a strong negative linear relationship was detected (P < 0.01). The spectrometric measures and the results of this study demonstrate a possible approach to the estimation of the polygenic component involved in coat color inheritance.

Animals↗

Quantitative genetics of secondary hip joint osteoarthritis in a Labrador Retriever-Greyhound pedigree.

OBJECTIVE: To evaluate the quantitative inheritance of secondary hip joint osteoarthritis in a canine pedigree. ANIMALS: 137 Labrador Retrievers, Greyhounds, and mixed-breed dogs. PROCEDURES: Necropsy scores ranging from 0 to 4 were obtained for each hip joint. Seven unaffected Greyhounds with normal hip joint conformation were also used for genetic modeling, but were not euthanized. Sixty-six male and 71 female dogs were allocated to 2 groups (< or = 12 months of age and > 12 months of age). Statistical models were developed to establish the inheritance pattern of hip joint osteoarthritis that developed secondary to hip dysplasia. RESULTS: 62 dogs had evidence of osteoarthritis in a hip joint, and 75 had no evidence of osteoarthritis. After sex was adjusted for, the necropsy score was found to be inherited additively but without dominance. Each Labrador Retriever allele increased the necropsy score by 0.7 to 0.9 points, compared with the Greyhound allele, and male sex increased the necropsy score 0.74 over female sex. Approximately 10% of the variation in necropsy score was attributable to the litter of puppies' origin. CONCLUSIONS AND CLINICAL RELEVANCE: Because secondary hip joint osteoarthritis is inherited additively, selection pressure could be applied to reduce its incidence. Similar statistical models can be used in linkage and association mapping to detect the genes in the underlying quantitative trait loci that contribute to hip joint osteoarthritis.

Age Factors↗

Quantitative genetic analysis of skin reflectance: a multivariate approach.

Skin color is a polygenically determined quantitative trait. Although it has been used extensively in studies of between-population variation, there have been relatively few studies of the inheritance of skin color. In this article we use measurements on 359 members of the Jirel population of eastern Nepal to assess the heritabilities and additive genetic correlations of three skin reflectance measures. Skin color was measured at the upper inner arm site at three wavelengths. A maximum likelihood approach was used to estimate sex and age effects on skin reflectance, heritabilities, and phenotypic variances at each wavelength and both additive genetic and environmental correlations between wavelengths. This technique incorporated information from 36 pedigrees with 2-25 members and 173 independent individuals. Likelihood ratio tests were used to assess the significance of specific variance/covariance components. The results indicate that skin reflectances are moderately heritable at all three wavelengths. The pairwise phenotypic correlations ranged from 0.76 to 0.88. The observed additive genetic correlations were not significantly different from 1.00, suggesting that the same loci influence variation at each wavelength. This evidence for relatively complete pleiotropy implies that measurements at multiple wavelengths yield little additional genetic information, although they may be useful for reducing measurement error. Based on estimates of the genetic and phenotypic covariance matrices, we determined that skin reflectance measurements are expected to provide only as much information for assessing local between-population genetic variation as a single two-allele polymorphic marker. Therefore microevolutionary studies based on skin color variation should be viewed with caution.

Adolescent↗

Quantitative genetics of signal evolution: a comparison of the pheromonal signal in two populations of the cabbage looper, Trichoplusia ni.

Pheromones are important in reproductive isolation among populations of moths, but the genetics associated with diversification of pheromonal signals is poorly understood. To gain insight into processes that may lead to diversification we examined the genetic architecture underlying the production of the sex pheromone of the cabbage looper moth, Trichoplusia ni. We compared genetic parameters of two populations; one with a wild-type pheromone phenotype (N) and one where a single-gene mutation affecting the pheromone blend produced by females had been established (M). Using a half-sib breeding design we estimated heritabilities, coefficients of additive genetic variation, and phenotypic, genetic, and environmental correlations of the pheromone components. In both populations, narrow sense heritabilities were generally moderate and genetic correlations were mostly positive. Comparisons between the two populations showed that, while the pattern of phenotypic correlations showed significant agreement between populations, the patterns of genetic (co)variation (i.e. the shapes of the within population matrix) were dissimilar between the two populations. The presence of additive genetic variation in both populations indicates that there is the potential for further evolution of individual pheromone components. However, because of the differences between the populations in the pattern of genetic variation and covariation, the populations will evolve along different evolutionary trajectories even under identical selection pressures. These results suggest that single gene mutations, once established, can be associated with further alterations in the genetic architecture and this has implications for the evolution of pheromone communication.

Animals↗

Quantitative genetics of ultrasonic advertisement signalling in the lesser waxmoth Achroia grisella (Lepidoptera: pyralidae).

Males of the lesser waxmoth Achroia grisella (Lepidoptera: Pyralidae) produce ultrasonic advertisement signals attractive to females within several metres. Previous studies showed that females prefer male signals that are louder, delivered at a faster rate, and have a greater asynchrony between pulses produced by the right and left wings. These three signal characters vary considerably within populations but are repeatable within individuals. Breeding experiments employing half-sib designs were conducted on both collectively and individually reared moths to determine genetic variance within and covariance among these signal characters. Heritabilities of all signal characters were significant among collectively reared moths. Heritabilities for signal rate and right-left wing asynchrony interval were not significant, however, among individually reared moths, suggesting the presence of significant nonadditive genetic variance or common environmental variation. Development time was also significantly heritable, but only under individual rearing. The only significant genetic correlation was between signal rate and length of the right-left wing asynchrony and this was negative. Our findings on heritability of signal characters are consistent with a coevolutionary sexual selection mechanism, but the absence of signal x development genetic correlation fails to support specifically a good-genes mechanism. The variation in heritability among conditions suggests that environmental variance may be high, and may render selection on signal characters by female choice ineffective. Thus, additive genetic variance for these characters may be maintained in the presence of directional female choice.

Animals↗

Using the quantitative genetic threshold model for inferences between and within species.

Sewall Wright's threshold model has been used in modelling discrete traits that may have a continuous trait underlying them, but it has proven difficult to make efficient statistical inferences with it. The availability of Markov chain Monte Carlo (MCMC) methods makes possible likelihood and Bayesian inference using this model. This paper discusses prospects for the use of the threshold model in morphological systematics to model the evolution of discrete all-or-none traits. There the threshold model has the advantage over 0/1 Markov process models in that it not only accommodates polymorphism within species, but can also allow for correlated evolution of traits with far fewer parameters that need to be inferred. The MCMC importance sampling methods needed to evaluate likelihood ratios for the threshold model are introduced and described in some detail.

Bayes Theorem↗