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Genetic Variance (genetic + variance)
Kinds of Genetic Variance Selected AbstractsMULTIVARIATE QUANTITATIVE GENETICS AND THE LEK PARADOX: GENETIC VARIANCE IN MALE SEXUALLY SELECTED TRAITS OF DROSOPHILA SERRATA UNDER FIELD CONDITIONSEVOLUTION, Issue 12 2004Emma Hine Abstract Single male sexually selected traits have been found to exhibit substantial genetic variance, even though natural and sexual selection are predicted to deplete genetic variance in these traits. We tested whether genetic variance in multiple male display traits of Drosophila serrata was maintained under field conditions. A breeding design involving 300 field-reared males and their laboratory-reared offspring allowed the estimation of the genetic variance-covariance matrix for six male cuticular hydrocarbons (CHCs) under field conditions. Despite individual CHCs displaying substantial genetic variance under field conditions, the vast majority of genetic variance in CHCs was not closely associated with the direction of sexual selection measured on field phenotypes. Relative concentrations of three CHCs correlated positively with body size in the field, but not under laboratory conditions, suggesting condition-dependent expression of CHCs under field conditions. Therefore condition dependence may not maintain genetic variance in preferred combinations of male CHCs under field conditions, suggesting that the large mutational target supplied by the evolution of condition dependence may not provide a solution to the lek paradox in this species. Sustained sexual selection may be adequate to deplete genetic variance in the direction of selection, perhaps as a consequence of the low rate of favorable mutations expected in multiple trait systems. [source] THE ADDITIVE GENETIC VARIANCE AFTER BOTTLENECKS IS AFFECTED BY THE NUMBER OF LOCI INVOLVED IN EPISTATIC INTERACTIONSEVOLUTION, Issue 4 2003Yamama Naciri-Graven Abstract We investigated the role of the number of loci coding for a neutral trait on the release of additive variance for this trait after population bottlenecks. Different bottleneck sizes and durations were tested for various matrices of genotypic values, with initial conditions covering the allele frequency space. We used three different types of matrices. First, we extended Cheverud and Routman's model by defining matrices of "pure" epistasis for three and four independent loci; second, we used genotypic values drawn randomly from uniform, normal, and exponential distributions; and third we used two models of simple metabolic pathways leading to physiological epistasis. For all these matrices of genotypic values except the dominant metabolic pathway, we find that, as the number of loci increases from two to three and four, an increase in the release of additive variance is occurring. The amount of additive variance released for a given set of genotypic values is a function of the inbreeding coefficient, independently of the size and duration of the bottleneck. The level of inbreeding necessary to achieve maximum release in additive variance increases with the number of loci. We find that additive-by-additive epistasis is the type of epistasis most easily converted into additive variance. For a wide range of models, our results show that epistasis, rather than dominance, plays a significant role in the increase of additive variance following bottlenecks. [source] PECTIVE: HERE'S TO FISHER, ADDITIVE GENETIC VARIANCE, AND THE FUNDAMENTALTHEOREM OF NATURAL SELECTIONEVOLUTION, Issue 7 2002James F. Crow Abstract Fisher's fundamental theorem of natural selection, that the rate of change of fitness is given by the additive genetic variance of fitness, has generated much discussion since its appearance in 1930. Fisher tried to capture in the formula the change in population fitness attributable to changes of allele frequencies, when all else is not included. Lessar's formulation comes closest to Fisher's intention, as well as this can be judged. Additional terms can be added to account for other changes. The "theorem" as stated by Fisher is not exact, and therefore not a theorem, but it does encapsulate a great deal of evolutionary meaning in a simple statement. I also discuss the effectiveness of reproductive-value weighting and the theorem in integrated form. Finally, an optimum principle, analogous to least action and Hamilton's principle in physics, is discussed. [source] Female Mate Choice, Calling Song and Genetic Variance in the Cricket, Gryllodes sigillatusETHOLOGY, Issue 3 2008Jocelyn Champagnon Female preferences for song patterns of males of Gryllodes sigillatus and genetic variance of morphological traits correlated with them were analyzed. Females preferred short pulses associated with large males. The males' thorax width, wing length and femur III length showed stronger relationship with the song pulse duration, whereas the relationship between pulse duration and wing width was not significant. Interestingly, this last trait was the only one that showed significant levels of genetic variance. Perhaps these results could be explained by the evolutionary response to sexual selection. Sexual selection could deplete the genetic variance in the male traits related to male-mating success. [source] The Moderating Effect of Religiosity on the Genetic Variance of Problem Alcohol UseALCOHOLISM, Issue 9 2010Tanya M. M. Button Background:, Previous studies have demonstrated that the heritability of alcohol-related phenotypes depends upon the social background in which it is measured (e.g., urbanicity, marital status, and religiosity). The aim of the current study was to identify whether religiosity moderated the genetic variance of problem alcohol use in men and women at two time points: adolescence and early adulthood. Method:, Participants were 312 male MZ pairs, 379 female MZ pairs, 231 male DZ pairs, 235 female DZ pairs, and 275 opposite sex DZ pairs participating in the University of Colorado Center on Antisocial Drug Dependence. Religiosity was measured using the Value on Religion Scale (Jessor and Jessor, 1977), and problem alcohol use was measured using the Composite International Diagnostic Interview,Substance Abuse Module (Cottler et al., 1989). Data were analyzed using a model-fitting approach to the twin data. Results:, In adolescence, genetic variance of problem alcohol use decreased significantly with increasing levels of religiosity in both men and women, whereas in early adulthood, religiosity did not moderate the genetic variance of problem alcohol use in either men or women. Conclusion:, Religiosity appears to moderate the genetic effects on problem alcohol use during adolescence, but not during early adulthood. The reduced genetic variance for problem alcohol use in adolescence may be the consequence of greater social control in adolescence than in young adulthood. [source] Hierarchical Spatial Modeling of Additive and Dominance Genetic Variance for Large Spatial Trial DatasetsBIOMETRICS, Issue 2 2009Andrew O. Finley Summary This article expands upon recent interest in Bayesian hierarchical models in quantitative genetics by developing spatial process models for inference on additive and dominance genetic variance within the context of large spatially referenced trial datasets. Direct application of such models to large spatial datasets are, however, computationally infeasible because of cubic-order matrix algorithms involved in estimation. The situation is even worse in Markov chain Monte Carlo (MCMC) contexts where such computations are performed for several iterations. Here, we discuss approaches that help obviate these hurdles without sacrificing the richness in modeling. For genetic effects, we demonstrate how an initial spectral decomposition of the relationship matrices negate the expensive matrix inversions required in previously proposed MCMC methods. For spatial effects, we outline two approaches for circumventing the prohibitively expensive matrix decompositions: the first leverages analytical results from Ornstein,Uhlenbeck processes that yield computationally efficient tridiagonal structures, whereas the second derives a modified predictive process model from the original model by projecting its realizations to a lower-dimensional subspace, thereby reducing the computational burden. We illustrate the proposed methods using a synthetic dataset with additive, dominance, genetic effects and anisotropic spatial residuals, and a large dataset from a Scots pine (Pinus sylvestris L.) progeny study conducted in northern Sweden. Our approaches enable us to provide a comprehensive analysis of this large trial, which amply demonstrates that, in addition to violating basic assumptions of the linear model, ignoring spatial effects can result in downwardly biased measures of heritability. [source] Genetic variances due to imprinted genes in cattleJOURNAL OF ANIMAL BREEDING AND GENETICS, Issue 3 2002TH. ENGELLANDT The effect of paternally expressed, i.e. maternally imprinted, genes on slaughter records from 2744 German Gelbvieh finishing bulls were estimated. Significant effects of paternal gametes were found for two fatness traits and an estimate of meat content. Paternally expressed genes explained 14 and 16% of the phenotypic variances for pelvic fat and kidney fat, respectively. Ignoring paternal gametic effects resulted in inflated estimates of the additive genetic variances. The heritabilities of pelvic and kidney fat dropped from 0.31 to 0.16 and from 0.59 to 0.28, respectively, when paternal gametes were fitted. A 15% influence of paternally expressed genes and a reduction in heritability of 20% were also found for estimated meat content. Simulation studies demonstrated that the uncorrelated random effect of the sire is a useful indicator for the presence of paternal gametic effects in variance component estimations. The presented results correspond well with findings in swine, where a paternally expressed QTL at the Igf2 gene influences similar trait complexes. A viable speculation could therefore be that an imprinted bovine Igf2 gene caused the effects described here. Genetische Variation auf Grund von paternal exprimierten Genen beim Rind In der vorliegenden Studie wurde der Effekt von paternal exprimierten Genen auf Merkmale der Schlachtleistung und des Schlachtkörperwertes von 2744 Mastbullen der Rasse Gelbvieh geschätzt. Signifikante Effekte von paternal exprimierten Genen wurden für zwei Fettabschnitte des Schlachtkörpers und für den geschätzten Fleischanteil gefunden, sie erklärten 14% und 16% der phänotypischen Varianzen des Becken- und Nierenfettes. Das Nichtbeachten dieser Effekte im linearen Schätzmodell führte zu einer Überschätzung der additiv genetischen Varianzen in den jeweiligen Merkmalen. Wurden die Effekte der paternalen Gameten im Schätzmodell berücksichtigt, fielen die geschätzten Heritabilitäten des Becken- und Nierenfettes von h2=0,31 auf h2=0,16 bzw. von h2=0,59 auf h2=0,28. Der Einfluß der paternalen Gameten auf die phänotypische Varianz des geschätzten Fleischanteils des Schlachkörpers wurde auf 15% geschätzt. Bei Berücksichtigung dieser Effekte im linearen Schätzmodell fiel die Heritabilität des Merkmals Fleischanteil um 20%. In Simulationsstudien konnte gezeigt werden, daß der unkorrelierte, zufällige Effekt des Vaters ein nützlicher Indikator für eventuell vorhandene paternal gametische Effekte bei der Schätzung von Varianzkomponenten sein kann. Die gefundenen Ergebnisse weisen Parallelen zu Untersuchungen beim Schwein auf, wo ein paternal exprimierter QTL am Igf2 Locus ähnliche Merkmalskomplexe beeinflußt. Eine mögliche Spekulation könnte daher sein, daß die hier beschriebenen Effekte beim Rind durch einen Igf2 Locus verursacht werden, der dem Phänomen des Imprinting unterliegt. [source] Genetic analysis of residual feed intakes and other performance test traits of Japanese Black cattle from revised protocolANIMAL SCIENCE JOURNAL, Issue 3 2008Takeshi OKANISHI ABSTRACT The performance test protocol for Japanese Black cattle was revised in April 2002. This resulted in restriction of access to concentrate (based on body weight) and modification of the concentrate's ingredients. Genetic parameters of growth and feed utilization traits of the performance test were estimated using 1304 records using the revised protocol. Residual feed intakes (RFIs) as alternative indicators for feed utilization efficiency were included. (Co)variance components were estimated by EM-REML. Heritabilities for growth traits were between 0.26 and 0.47. Heritabilities for feed intakes and conversions ranged from 0.25 to 0.37 and from 0.03 to 0.29, respectively. Genetic variances and heritabilities were lower for the revised protocol. Highly positive genetic correlations of daily gain (DG) with feed intakes indicated selection on DG is expected to increase feed intake. Selection on feed conversion may lead to higher DG. The heritability estimates for RFIs ranged from 0.10 to 0.33 and were generally higher than corresponding estimates for feed conversion ratios. RFI of TDN showed positive genetic correlations with all feed intakes. The reduction of feed intakes could be expected through selection on the RFI without changing body size. RFIs were considered to be alternative indicators to improve feed utilization efficiency under the new performance test. [source] Letter to the editor: Chronicle for an orphan trait: Comment on Hofer, Shair, Masmela, & Brunelli, "Developmental effects of selective breeding for an infantile trait: The rat pup ultrasonic isolation call"DEVELOPMENTAL PSYCHOBIOLOGY, Issue 4 2001Pierre L. Roubertoux Abstract Hofer, Brunelli, Shair, and Masmela (2001) examined several behavioral and physiological measures in low, high, and unselected lines obtained from a divergent selection for ultrasound production (USP) in young rats. Although the response to selection was clear-cut, few correlated responses appeared. This surprising result could be explained by two reasons. USP has polygenic correlates in this population, and most of the chromosomal regions that are linked with these measures only contribute to a small part of the genetic variance. Therefore, correlated responses to selection might exist, but the common genetic variance between the trait under selection and the indirectly selected trait is too small to be detected by a selective breeding strategy. © 2001 John Wiley & Sons, Inc. Dev Psychobiol 39: 251,254, 2001.. [source] Female Mate Choice, Calling Song and Genetic Variance in the Cricket, Gryllodes sigillatusETHOLOGY, Issue 3 2008Jocelyn Champagnon Female preferences for song patterns of males of Gryllodes sigillatus and genetic variance of morphological traits correlated with them were analyzed. Females preferred short pulses associated with large males. The males' thorax width, wing length and femur III length showed stronger relationship with the song pulse duration, whereas the relationship between pulse duration and wing width was not significant. Interestingly, this last trait was the only one that showed significant levels of genetic variance. Perhaps these results could be explained by the evolutionary response to sexual selection. Sexual selection could deplete the genetic variance in the male traits related to male-mating success. [source] Variation and Repeatability of Female Choice in a Chorusing Katydid, Ephippiger ephippiger: an Experimental Exploration of the Precedence EffectETHOLOGY, Issue 4 2004Michael D. Greenfield Female choice in various species of acoustic insects and anurans entails a psychoacoustic preference for male calls that lead their neighbors by a brief time interval. This discrimination, which can be termed a precedence effect, may select for various mechanisms with which males adjust call rhythm and thus reduce their incidence of ineffective following calls. At a collective level, alternating and synchronous choruses may emerge from these call timing mechanisms. Using playback experiments, we characterized the precedence effect in females of the katydid Ephippiger ephippiger, an alternating choruser in which males use a rhythm adjustment mechanism that prevents calling during brief intervals following their neighbors' calls. E. ephippiger females oriented toward leading male calls in >75% of trials when relatively young (<40 d old) and when playbacks were timed so that following calls began within 100,250 ms of the leading ones. However, this preference declined to below 60% as females aged and the interval separating leading and following call onsets increased. The strength of this precedence effect varied greatly between females, but within broad age classes the effect in a given female was statistically repeatable. Such repeatability indicates the possibility that additive genetic variance could be a significant component of variation in the precedence effect. We discuss the implications of our findings and inference on genetic variance for evolution of the precedence effect and for chorusing. [source] REVIEW: Consilient research approaches in studying gene × environment interactions in alcohol researchADDICTION BIOLOGY, Issue 2 2010Kenneth J. Sher ABSTRACT This review article discusses the importance of identifying gene-environment interactions for understanding the etiology and course of alcohol use disorders and related conditions. A number of critical challenges are discussed, including the fact that there is no organizing typology for classifying different types of environmental exposures, many key human environmental risk factors for alcohol dependence have no clear equivalents in other species, much of the genetic variance of alcohol dependence in human is not ,alcohol specific', and the potential range of gene-environment interactions that could be considered is so vast that maintaining statistical control of Type 1 errors is a daunting task. Despite these and other challenges, there appears to be a number of promising approaches that could be taken in order to achieve consilience and ecologically valid translation between human alcohol dependence and animal models. Foremost among these is to distinguish environmental exposures that are thought to have enduring effects on alcohol use motivation (and self-regulation) from situational environmental exposures that facilitate the expression of such motivations but do not, by themselves, have enduring effects. In order to enhance consilience, various domains of human approach motivation should be considered so that relevant environmental exposures can be sampled, as well as the appropriate species to study them in (i.e. where such motivations are ecologically relevant). Foremost among these are social environments, which are central to the initiation and escalation of human alcohol consumption. The value of twin studies, human laboratory studies and pharmacogenetic studies is also highlighted. [source] The evolutionary genetics of personality,EUROPEAN JOURNAL OF PERSONALITY, Issue 5 2007Lars Penke Abstract Genetic influences on personality differences are ubiquitous, but their nature is not well understood. A theoretical framework might help, and can be provided by evolutionary genetics. We assess three evolutionary genetic mechanisms that could explain genetic variance in personality differences: selective neutrality, mutation-selection balance, and balancing selection. Based on evolutionary genetic theory and empirical results from behaviour genetics and personality psychology, we conclude that selective neutrality is largely irrelevant, that mutation-selection balance seems best at explaining genetic variance in intelligence, and that balancing selection by environmental heterogeneity seems best at explaining genetic variance in personality traits. We propose a general model of heritable personality differences that conceptualises intelligence as fitness components and personality traits as individual reaction norms of genotypes across environments, with different fitness consequences in different environmental niches. We also discuss the place of mental health in the model. This evolutionary genetic framework highlights the role of gene-environment interactions in the study of personality, yields new insight into the person-situation-debate and the structure of personality, and has practical implications for both quantitative and molecular genetic studies of personality. Copyright © 2007 John Wiley & Sons, Ltd. [source] EMPIRICAL COMPARISON OF G MATRIX TEST STATISTICS: FINDING BIOLOGICALLY RELEVANT CHANGEEVOLUTION, Issue 10 2009Brittny Calsbeek A central assumption of quantitative genetic theory is that the breeder's equation (R=GP,1S) accurately predicts the evolutionary response to selection. Recent studies highlight the fact that the additive genetic variance,covariance matrix (G) may change over time, rendering the breeder's equation incapable of predicting evolutionary change over more than a few generations. Although some consensus on whether G changes over time has been reached, multiple, often-incompatible methods for comparing G matrices are currently used. A major challenge of G matrix comparison is determining the biological relevance of observed change. Here, we develop a "selection skewers"G matrix comparison statistic that uses the breeder's equation to compare the response to selection given two G matrices while holding selection intensity constant. We present a bootstrap algorithm that determines the significance of G matrix differences using the selection skewers method, random skewers, Mantel's and Bartlett's tests, and eigenanalysis. We then compare these methods by applying the bootstrap to a dataset of laboratory populations of Tribolium castaneum. We find that the results of matrix comparison statistics are inconsistent based on differing a priori goals of each test, and that the selection skewers method is useful for identifying biologically relevant G matrix differences. [source] IS INBREEDING DEPRESSION LOWER IN MALADAPTED POPULATIONS?EVOLUTION, Issue 7 2009A QUANTITATIVE GENETICS MODEL Despite abundant empirical evidence that inbreeding depression varies with both the environment and the genotypic context, theoretical predictions about such effects are still rare. Using a quantitative genetics model, we predict amounts of inbreeding depression for fitness emerging from Gaussian stabilizing selection on some phenotypic trait, on which, for simplicity, genetic effects are strictly additive. Given the strength of stabilizing selection, inbreeding depression then varies simply with the genetic variance for the trait under selection and the distance between the mean breeding value and the optimal phenotype. This allows us to relate the expected inbreeding depression to the degree of maladaptation of the population to its environment. We confront analytical predictions with simulations, in well-adapted populations at equilibrium, as well as in maladapted populations undergoing either a transient environmental shift, or gene swamping in heterogeneous habitats. We predict minimal inbreeding depression in situations of extreme maladaptation. Our model provides a new basis for interpreting experiments that measure inbreeding depression for the same set of genotypes in different environments, by demonstrating that the history of adaptation, in addition to environmental harshness per se, may account for differences in inbreeding depression. [source] WATER STRESS ALTERS THE GENETIC ARCHITECTURE OF FUNCTIONAL TRAITS ASSOCIATED WITH DROUGHT ADAPTATION IN AVENA BARBATAEVOLUTION, Issue 3 2009Mark E. Sherrard Environmental stress can alter genetic variation and covariation underlying functional traits, and thus affect adaptive evolution in response to natural selection. However, the genetic basis of functional traits is rarely examined in contrasting resource environments, and consequently, there is no consensus regarding whether environmental stress constrains or facilitates adaptive evolution. We tested whether resource availability affects genetic variation for and covariation among seven physiological traits and seven morphological/performance traits by growing the annual grass Avena barbata in dry and well-watered treatments. We found that differences in the overall genetic variance,covariance (G) matrix between environments were driven by physiological traits rather than morphology and performance traits. More physiological traits were heritable in the dry treatment than the well-watered treatment and many of the genetic correlations among physiological traits were environment dependent. In contrast, genetic variation and covariation among the morphological and performance traits did not differ across treatments. Furthermore, genetic correlations between physiology and performance were stronger in the dry treatment, which contributed to differences in the overall G -matrix. Our results therefore suggest that physiological adaptation would be constrained by low heritable variation in resource-rich environments, but facilitated by higher heritable variation and stronger genetic correlations with performance traits in resource-poor environments. [source] AGE-SPECIFIC GENETIC AND MATERNAL EFFECTS IN FECUNDITY OF PREINDUSTRIAL FINNISH WOMENEVOLUTION, Issue 9 2008Jenni E. Pettay A population's potential for evolutionary change depends on the amount of genetic variability expressed in traits under selection. Studies attempting to measure this variability typically do so over the life span of individuals, but theory suggests that the amount of additive genetic variance can change during the course of individuals' lives. Here we use pedigree data from historical Finns and a quantitative genetic framework to investigate how female fecundity, throughout an individual's reproductive life, is influenced by "maternal" versus additive genetic effects. We show that although maternal effects explain variation in female fecundity early in life, these effects wane with female age. Moreover, this decline in maternal effects is associated with a concomitant increase in additive genetic variance with age. Our results thus highlight that single over-lifetime estimates of trait heritability may give a misleading view of a trait's potential to respond to changing selection pressures. [source] A REAPPRAISAL OF BATEMAN'S CLASSIC STUDY OF INTRASEXUAL SELECTIONEVOLUTION, Issue 11 2007Brian F. Snyder Bateman's (1948) study showing greater variances in number of mates and reproductive success in male than female Drosophila melanogaster is a foundational paper in sexual selection. Here we show for the first time that his methods had flaws, including the elimination of genetic variance, sampling biases, miscalculations of fitness variances, statistical pseudo-replication, and selective presentation of data. We conclude that Bateman's results are unreliable, his conclusions are questionable, and his observed variances are similar to those expected under random mating. Despite our analysis, we do not intend this article as a criticism of Bateman; he accomplished his work without modern computational tools, and his approach was groundbreaking emphasizing the significance of fitness variance for sexual selection. However, this reanalysis has implications for what counts as evidence for sexual selection and we believe that our concerns should be of interest to contemporary students of sexual selection. We call for repetitions of Bateman's study using modern statistical and molecular methods. [source] EFFECTS OF MIGRATION ON THE GENETIC COVARIANCE MATRIXEVOLUTION, Issue 10 2007Frédéric Guillaume In 1996, Schluter showed that the direction of morphological divergence of closely related species is biased toward the line of least genetic resistance, represented by gmax, the leading eigenvector of the matrix of genetic variance,covariance (the G -matrix). G is used to predict the direction of evolutionary change in natural populations. However, this usage requires that G is sufficiently constant over time to have enough predictive significance. Here, we explore the alternative explanation that G can evolve due to gene flow to conform to the direction of divergence between incipient species. We use computer simulations in a mainland,island migration model with stabilizing selection on two quantitative traits. We show that a high level of gene flow from a mainland population is required to significantly affect the orientation of the G -matrix in an island population. The changes caused by the introgression of the mainland alleles into the island population affect all aspects of the shape of G (size, eccentricity, and orientation) and lead to the alignment of gmax with the line of divergence between the two populations' phenotypic optima. Those changes decrease with increased correlation in mutational effects and with a correlated selection. Our results suggest that high migration rates, such as those often seen at the intraspecific level, will substantially affect the shape and orientation of G, whereas low migration (e.g., at the interspecific level) is unlikely to substantially affect the evolution of G. [source] THE CHANGE IN QUANTITATIVE GENETIC VARIATION WITH INBREEDINGEVOLUTION, Issue 12 2006Josh Van Buskirk Abstract Inbreeding is known to reduce heterozygosity of neutral genetic markers, but its impact on quantitative genetic variation is debated. Theory predicts a linear decline in additive genetic variance (VA) with increasing inbreeding coefficient (F) when loci underlying the trait act additively, but a nonlinear hump-shaped relationship when dominance and epistasis are important. Predictions for heritability (h2) are similar, although the exact shape depends on the value of h2 in the absence of inbreeding. We located 22 published studies in which the level of genetic variation in [source] LOCAL HETEROZYGOSITY-FITNESS CORRELATIONS WITH GLOBAL POSITIVE EFFECTS ON FITNESS IN THREESPINE STICKLEBACKEVOLUTION, Issue 8 2006Mélissa Lieutenant-Gosselin Abstract The complex interactions between genetic diversity and evolution have important implications in many biological areas including conservation, speciation, and mate choice. A common way to study these interactions is to look at heterozygosity-fitness correlations (HFCs). Until recently, HFCs based on noncoding markers were believed to result primarily from global inbreeding effects. However, accumulating theoretical and empirical evidence shows that HFCs may often result from genes being linked to the markers used (local effect). Moreover, local effect HFCs could differ from global inbreeding effects in their direction and occurrence. Consequently, the investigation of the structure and consequences of local HFCs is emerging as a new important goal in evolutionary biology. In this study of a wild threespine stickleback (Gasterosteus aculeatus) population, we first tested the presence of significant positive or negative local effects of heterozygosity at 30 microsatellites loci on five fitness components: survival, mating success, territoriality, length, and body condition. Then, we evaluated the direction and shape of total impact of local HFCs, and estimated the magnitude of the impacts on fitness using regression coefficients and selection differentials. We found that multilocus heterozygosity was not a reliable estimator of individual inbreeding coefficient, which supported the relevance of single-locus based analyses. Highly significant and temporally stable local HFCs were observed. These were mainly positive, but negative effects of heterozygosity were also found. Strong and opposite effects of heterozygosity are probably present in many populations, but may be blurred in HFC analyses looking for global effects only. In this population, both negative and positive HFCs are apparently driving mate preference by females, which is likely to contribute to the maintenance of both additive and nonadditive genetic variance. [source] MULTILOCUS GENETICS AND THE COEVOLUTION OF QUANTITATIVE TRAITSEVOLUTION, Issue 7 2006Michael Kopp Abstract We develop and analyze an explicit multilocus genetic model of coevolution. We assume that interactions between two species (mutualists, competitors, or victim and exploiter) are mediated by a pair of additive quantitative traits that are also subject to direct stabilizing selection toward intermediate optima. Using a weak-selection approximation, we derive analytical results for a symmetric case with equal locus effects and no mutation, and we complement these results by numerical simulations of more general cases. We show that mutualistic and competitive interactions always result in coevolution toward a stable equilibrium with no more than one polymorphic locus per species. Victimexploiter interactions can lead to different dynamic regimes including evolution toward stable equilibria, cycles, and chaos. At equilibrium, the victim is often characterized by a very large genetic variance, whereas the exploiter is polymorphic in no more than one locus. Compared to related one-locus or quantitative genetic models, the multilocus model exhibits two major new properties. First, the equilibrium structure is considerably more complex. We derive detailed conditions for the existence and stability of various classes of equilibria and demonstrate the possibility of multiple simultaneously stable states. Second, the genetic variances change dynamically, which in turn significantly affects the dynamics of the mean trait values. In particular, the dynamics tend to be destabilized by an increase in the number of loci. [source] THE PHENOTYPIC VARIANCE WITHIN PLASTIC TRAITS UNDER MIGRATION-MUTATION-SELECTION BALANCEEVOLUTION, Issue 6 2006Xu-Sheng Zhang Abstract How phenotypic variances of quantitative traits are influenced by the heterogeneity in environment is an important problem in evolutionary biology. In this study, both genetic and environmental variances in a plastic trait under migration-mutation-stabilizing selection are investigated. For this, a linear reaction norm is used to approximate the mapping from genotype to phenotype, and a population of clonal inheritance is assumed to live in a habitat consisting of many patches in which environmental conditions vary among patches and generations. The life cycle is assumed to be selection-reproduction-mutation-migration. Analysis shows that phenotypic plasticity is adaptive if correlations between the optimal phenotype and environment have become established in both space and/or time, and it is thus possible to maintain environmental variance (VE) in the plastic trait. Under the special situation of no mutation but maximum migration such that separate patches form an effective single-site habitat, the genotype that maximizes the geometric mean fitness will come to fixation and thus genetic variance (VG) cannot be maintained. With mutation and/or restricted migration, VG can be maintained and it increases with mutation rate but decreases with migration rate; whereas VE is little affected by them. Temporal variation in environmental quality increases VG while its spatial variance decreases VG. Variation in environmental conditions may decrease the environmental variance in the plastic trait. [source] AN EXACT FORM OF THE BREEDER'S EQUATION FOR THE EVOLUTION OF A QUANTITATIVE TRAIT UNDER NATURAL SELECTIONEVOLUTION, Issue 11 2005John S. Heywood Abstract 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 s,2A/s,2P should be viewed as an approximation to the offspring-parent linear regression rather than the other way around. [source] THE COEVOLUTIONARY DYNAMICS OF ANTAGONISTIC INTERACTIONS MEDIATED BY QUANTITATIVE TRAITS WITH EVOLVING VARIANCESEVOLUTION, Issue 10 2005Scott L. Nuismer Abstract Quantitative traits frequently mediate coevolutionary interactions between predator and prey or parasite and host. Previous efforts to understand and predict the coevolutionary dynamics of these interactions have generally assumed that standing genetic variation is fixed or absent altogether. We develop a genetically explicit model of coevolution that bridges the gap between these approaches by allowing genetic variation itself to evolve. Analysis of this model shows that the evolution of genetic variance has improtant consequences for the dyanmics and outcome of coevolution. Of particular importance is our demonstration that coevolutionary cycles can emerge in the absence of stabilizing selection, and outcome not possible in previous models of coevolution mediated by quantitative traits. whether coevolutionary cycles evolve depends upon the strength of selection, the number of loci, and the rate of mutation in each of the interacting species. Our results also generate novel predictions for the expected sign and magnitude of linkage disequilibria in each species. [source] PERSPECTIVE: SIGN EPISTASIS AND GENETIC COSTRAINT ON EVOLUTIONARY TRAJECTORIESEVOLUTION, Issue 6 2005Daniel M. Weinreich Abstract Epistasis for fitness means that the selective effect of a mutation is conditional on the genetic background in which it appears. Although epistasis is widely observed in nature, our understanding of its consequences for evolution by natural selection remains incomplete. In particular, much attention focuses only on its influence on the instantaneous rate of changes in frequency of selected alleles via epistatic contribution to the additive genetic variance for fitness. Thus, in this framework epistasis only has evolutionary importance if the interacting loci are simultaneously segregating in the population. However, the selective accessibility of mutational trajectories to high fitness genotypes may depend on the genetic background in which novel mutations appear, and this effect is independent of population polymorphism at other loci. Here we explore this second influence of epistasis on evolution by natural selection. We show that it is the consequence of a particular form of epistasis, which we designate sign epistasis. Sign epistasis means that the sign of the fitness effect of a mutation is under epistatic control; thus, such a mutation is beneficial on some genetic backgrounds and deleterious on others. Recent experimental innovations in microbial systems now permit assessment of the fitness effects of individual mutations on multiple genetic backgrounds. We review this literature and identify many examples of sign epistasis, and we suggest that the implications of these results may generalize to other organisms. These theoretical and empirical considerations imply that strong genetic constraint on the selective accessibility of trajectories to high fitness genotypes may exist and suggest specific areas of investigation for future research. [source] MULTIVARIATE QUANTITATIVE GENETICS AND THE LEK PARADOX: GENETIC VARIANCE IN MALE SEXUALLY SELECTED TRAITS OF DROSOPHILA SERRATA UNDER FIELD CONDITIONSEVOLUTION, Issue 12 2004Emma Hine Abstract Single male sexually selected traits have been found to exhibit substantial genetic variance, even though natural and sexual selection are predicted to deplete genetic variance in these traits. We tested whether genetic variance in multiple male display traits of Drosophila serrata was maintained under field conditions. A breeding design involving 300 field-reared males and their laboratory-reared offspring allowed the estimation of the genetic variance-covariance matrix for six male cuticular hydrocarbons (CHCs) under field conditions. Despite individual CHCs displaying substantial genetic variance under field conditions, the vast majority of genetic variance in CHCs was not closely associated with the direction of sexual selection measured on field phenotypes. Relative concentrations of three CHCs correlated positively with body size in the field, but not under laboratory conditions, suggesting condition-dependent expression of CHCs under field conditions. Therefore condition dependence may not maintain genetic variance in preferred combinations of male CHCs under field conditions, suggesting that the large mutational target supplied by the evolution of condition dependence may not provide a solution to the lek paradox in this species. Sustained sexual selection may be adequate to deplete genetic variance in the direction of selection, perhaps as a consequence of the low rate of favorable mutations expected in multiple trait systems. [source] EFFECTS OF GENETIC DRIFT ON VARIANCE COMPONENTS UNDER A GENERAL MODEL OF EPISTASISEVOLUTION, Issue 10 2004N.H. Barton Abstract We analyze the changes in the mean and variance components of a quantitative trait caused by changes in allele frequencies, concentrating on the effects of genetic drift. We use a general representation of epistasis and dominance that allows an arbitrary relation between genotype and phenotype for any number of diallelic loci. We assume initial and final Hardy-Weinberg and linkage equilibrium in our analyses of drift-induced changes. Random drift generates transient linkage disequilibria that cause correlations between allele frequency fluctuations at different loci. However, we show that these have negligible effects, at least for interactions among small numbers of loci. Our analyses are based on diffusion approximations that summarize the effects of drift in terms of F, the inbreeding coefficient, interpreted as the expected proportional decrease in heterozygosity at each locus. For haploids, the variance of the trait mean after a population bottleneck is var(,z,) =where n is the number of loci contributing to the trait variance, VA(1)=VA is the additive genetic variance, and VA(k) is the kth-order additive epistatic variance. The expected additive genetic variance after the bottleneck, denoted (V*A), is closely related to var(,z,); (V*A) (1 ,F)Thus, epistasis inflates the expected additive variance above VA(1 ,F), the expectation under additivity. For haploids (and diploids without dominance), the expected value of every variance component is inflated by the existence of higher order interactions (e.g., third-order epistasis inflates (V*AA)). This is not true in general with diploidy, because dominance alone can reduce (V*A) below VA(1 ,F) (e.g., when dominant alleles are rare). Without dominance, diploidy produces simple expressions: var(,z,)==1 (2F) kVA(k) and (V*A) = (1 ,F)k(2F)k-1VA(k) With dominance (and even without epistasis), var(,z,)and (V*A) no longer depend solely on the variance components in the base population. For small F, the expected additive variance simplifies to (V*A)(1 ,F) VA+ 4FVAA+2FVD+2FCAD, where CAD is a sum of two terms describing covariances between additive effects and dominance and additive × dominance interactions. Whether population bottlenecks lead to expected increases in additive variance depends primarily on the ratio of nonadditive to additive genetic variance in the base population, but dominance precludes simple predictions based solely on variance components. We illustrate these results using a model in which genotypic values are drawn at random, allowing extreme and erratic epistatic interactions. Although our analyses clarify the conditions under which drift is expected to increase VA, we question the evolutionary importance of such increases. [source] COMPARING STRENGTHS OF DIRECTIONAL SELECTION: HOW STRONG IS STRONG?EVOLUTION, Issue 10 2004Joe Hereford Abstract The fundamental equation in evolutionary quantitative genetics, the Lande equation, describes the response to directional selection as a product of the additive genetic variance and the selection gradient of trait value on relative fitness. Comparisons of both genetic variances and selection gradients across traits or populations require standardization, as both are scale dependent. The Lande equation can be standardized in two ways. Standardizing by the variance of the selected trait yields the response in units of standard deviation as the product of the heritability and the variance-standardized selection gradient. This standardization conflates selection and variation because the phenotypic variance is a function of the genetic variance. Alternatively, one can standardize the Lande equation using the trait mean, yielding the proportional response to selection as the product of the squared coefficient of additive genetic variance and the mean-standardized selection gradient. Mean-standardized selection gradients are particularly useful for summarizing the strength of selection because the mean-standardized gradient for fitness itself is one, a convenient benchmark for strong selection. We review published estimates of directional selection in natural populations using mean-standardized selection gradients. Only 38 published studies provided all the necessary information for calculation of mean-standardized gradients. The median absolute value of multivariate mean-standardized gradients shows that selection is on average 54% as strong as selection on fitness. Correcting for the upward bias introduced by taking absolute values lowers the median to 31%, still very strong selection. Such large estimates clearly cannot be representative of selection on all traits. Some possible sources of overestimation of the strength of selection include confounding environmental and genotypic effects on fitness, the use of fitness components as proxies for fitness, and biases in publication or choice of traits to study. [source] CLIMATIC AND TEMPORAL EFFECTS ON THE EXPRESSION OF SECONDARY SEXUAL CHARACTERS: GENETIC AND ENVIRONMENTAL COMPONENTSEVOLUTION, Issue 3 2004Dany Garant Abstract Despite great interest in sexual selection, relatively little is known in detail about the genetic and environmental determinants of secondary sexual characters in natural populations. Such information is important for determining the way in which populations may respond to sexual selection. We report analyses of genetic and large-scale environmental components of phenotypic variation of two secondary sexual plumage characters (forehead and wing patch size) in the collared flycatcher Ficedula albicollis over a 22-year period. We found significant heritability for both characters but little genetic covariance between the two. We found a positive association between forehead patch size and a large-scale climatic index, the North Atlantic Oscillation (NAO) index, but not for wing patch. This pattern was observed in both cross-sectional and longitudinal data suggesting that the population response to NAO index can be explained as the result of phenotypic plasticity. Heritability of forehead patch size for old males, calculated under favorable conditions (NAO index median), was greater than that under unfavorable conditions (NAO index < median). These changes occurred because there were opposing changes in additive genetic variance (VA) and residual variance (VR) under favorable and unfavorable conditions, with VA increasing and VR decreasing in good environments. However, no such effect was detected for young birds, or for wing patch size in either age class. In addition to these environmental effects on both phenotypic and genetic variances, we found evidence for a significant decrease of forehead patch size over time in older birds. This change appears to be caused by a change in the sign of viability selection on forehead patch size, which is associated with a decline in the breeding value of multiple breeders. Our data thus reveal complex patterns of environmental influence on the expression of secondary sexual characters, which may have important implications for understanding selection and evolution of these characters. [source] |