scholarly journals Genetic Variation Among and Within Populations in Swedish Species of Sorbus aucuparia L. and Prunus padus L. Assessed in a Nursery Trial

2005 ◽  
Vol 54 (1-6) ◽  
pp. 1-8 ◽  
Author(s):  
V. Baliuckas ◽  
T. Lagerström ◽  
L. Norell ◽  
G. Eriksson
Keyword(s):  
Genetic Variation ◽  
Growth Traits ◽  
Genetic Correlations ◽  
Population Variation ◽  
Sorbus Aucuparia ◽  
Population Effect ◽  
Prunus Padus ◽  
Bud Flushing ◽  
Over Time ◽  
Leaf Colouring

Abstract Seedlings originating from open-pollinated offspring of six and four populations of Prunus padus and Sorbus aucuparia, respectively, were studied with respect to phenology and growth traits for 3-4 years in a nursery. There were no replications at the population levels since the experiments should be converted to seedling seed orchards. Therefore, a special statistical model for analysis of the population effect was developed making use of neighbour performances. This model was also used for derivation of heritabilities. The heritabilities for phenology traits were in many cases high in P. padus, > 0.40, while they varied in the range 0.07-0.62 in S. aucuparia. The population effect was significant for all growth rhythm traits in P. padus and for a majority of traits in S. aucuparia. In both species the heritability for height decreased over time. Only bud flushing in P. padus indicated a relationship with population latitudinal origin in some cases. The genetic correlations between bud flushing different years were relatively strong in both species while the corresponding correlations for leaf colouring were moderate in P. padus and weak in S. aucuparia. In conclusion, the observed structure suggests that the pattern of seed dispersal may have an influence on the among- and within-population variation.

scholarly journals 211 Changes in genetic parameters of fitness and growth traits under genomic selection in pigs

2019 ◽  
Vol 97 (Supplement_3) ◽  
pp. 41-41
Author(s):  
Jorge Hidalgo ◽  
Shogo Tsuruta ◽  
Daniela Lourenco ◽  
Yijian Huang ◽  
Kent Gray ◽  
...  
Keyword(s):  
Genetic Variation ◽  
Genomic Selection ◽  
Variance Components ◽  
Genetic Parameters ◽  
Growth Traits ◽  
Average Daily Gain ◽  
Genetic Correlations ◽  
Data Set ◽  
Daily Gain ◽  
Over Time

Abstract Genomic selection increases intensity of selection and decreases generation interval. However, intensive selection reduces genetic variation and can strengthen undesirable genetic correlations. The purpose of this study was to investigate changes in genetic parameters of litter size (LS), number born alive (NBA), number born dead (NBD) and average daily gain (ADG) and weight at off-test (WT) in pigs over time. The data set contained 20,086 (LS), 21,230 (NBA), 21,230 (NBD), 144,717 (ADG) and 144,718 (WT) phenotypic records. Pedigree file included 369,776 animals born between 2001 and 2018, of which 39,038 were genotyped. Two trait models were evaluated (LS-ADG, LS-WT, NBA-ADG, NBA-WT, NBD-ADG and NBD-WT) using 3-year sliding subsets starting from 2010. Variance components and genetic parameters were estimated using GIBBS2F90 program. Computations were performed with (BLUP) or without (ssGBLUP) genotypes. For BLUP (ssGBLUP), the changes in heritability from 2010–2012 to 2015–2018 were 0.08 to 0.09 (0.08 to 0.06) for LS, 0.33 to 0.24 (0.37 to 0.16) for ADG, 0.11 to 0.07 (0.10 to 0.07) for NBD, and 0.32 to 0.24 (0.38 to 0.17) for WT. Differences were also observed for genetic correlations as they were -0.23 to -0.73 (-0.31 to -0.58) for LS-ADG, -0.24 to -0.74 (-0.31 to -0.50) for LS-WT, -0.33 to -0.65 (-0.41 to -0.53) for NBA-ADG, -0.35 to -0.66 (-0.42 to -0.45) for NBA-WT, 0.12 to 0.04 (0.12 to 0.16) for NBD-ADG, and 0.11 to 0.05 (0.11 to 0.22) for NBD-WT. Genomic selection in pigs reduced heritabilities and emphasized the antagonistic relationship that are under strong selection. Heritabilities estimated from ssGBLUP declined more than those by BLUP while changes in the genetic correlations were smaller and more gradual by ssGBLUP. Differences between ssGBLUP and BLUP could be caused by genomic pre-selection unaccounted for by BLUP.

scholarly journals Characterizing the oligogenic architecture of plant growth phenotypes informs genomic selection approaches in a common wheat population

BMC Genomics ◽  
2021 ◽  
Vol 22 (1) ◽  
Author(s):  
Noah DeWitt ◽  
Mohammed Guedira ◽  
Edwin Lauer ◽  
J. Paul Murphy ◽  
David Marshall ◽  
...  
Keyword(s):  
Genetic Variation ◽  
Plant Growth ◽  
Genomic Selection ◽  
Plant Height ◽  
Prediction Models ◽  
Growth Traits ◽  
Heading Date ◽  
Additive Genetic Variation ◽  
Moderate Effect ◽  
Over Time

Abstract Background Genetic variation in growth over the course of the season is a major source of grain yield variation in wheat, and for this reason variants controlling heading date and plant height are among the best-characterized in wheat genetics. While the major variants for these traits have been cloned, the importance of these variants in contributing to genetic variation for plant growth over time is not fully understood. Here we develop a biparental population segregating for major variants for both plant height and flowering time to characterize the genetic architecture of the traits and identify additional novel QTL. Results We find that additive genetic variation for both traits is almost entirely associated with major and moderate-effect QTL, including four novel heading date QTL and four novel plant height QTL. FT2 and Vrn-A3 are proposed as candidate genes underlying QTL on chromosomes 3A and 7A, while Rht8 is mapped to chromosome 2D. These mapped QTL also underlie genetic variation in a longitudinal analysis of plant growth over time. The oligogenic architecture of these traits is further demonstrated by the superior trait prediction accuracy of QTL-based prediction models compared to polygenic genomic selection models. Conclusions In a population constructed from two modern wheat cultivars adapted to the southeast U.S., almost all additive genetic variation in plant growth traits is associated with known major variants or novel moderate-effect QTL. Major transgressive segregation was observed in this population despite the similar plant height and heading date characters of the parental lines. This segregation is being driven primarily by a small number of mapped QTL, instead of by many small-effect, undetected QTL. As most breeding populations in the southeast U.S. segregate for known QTL for these traits, genetic variation in plant height and heading date in these populations likely emerges from similar combinations of major and moderate effect QTL. We can make more accurate and cost-effective prediction models by targeted genotyping of key SNPs.

Genetic evaluation of crossbred lamb production. 4. Genetic parameters for first-cross animal performance

2007 ◽  
Vol 58 (8) ◽  
pp. 839 ◽  
Author(s):  
V. M. Ingham ◽  
N. M. Fogarty ◽  
A. R. Gilmour ◽  
R. A. Afolayan ◽  
L. J. Cummins ◽  
...  
Keyword(s):  
Genetic Variation ◽  
Meat Quality ◽  
Mixed Models ◽  
Fixed Effects ◽  
Growth Traits ◽  
Genetic Correlations ◽  
Fibre Diameter ◽  
Quality Traits ◽  
Meat Quality Traits ◽  
Lamb Growth

The study estimated heritability for lamb growth and carcass performance, hogget ewe wool production, and worm egg count among crossbred progeny of maternal breed sires, as well as the genetic and phenotypic correlations among the traits. The data were from crossbred progeny of 91 sires from maternal breeds including Border Leicester, East Friesian, Finnsheep, Coopworth, White Suffolk, Corriedale, and Booroola Leicester. The sires were mated to Merino ewes at 3 sites over 3 years (and also Corriedale ewes at one site), with 3 common sires used at each site and year to provide genetic links. These sheep comprised part of the national maternal sire central progeny test program (MCPT) to evaluate the genetic variation for economically important production traits in progeny of maternal and dual-purpose (meat and wool) sires and the scope for genetic improvement. The matings resulted in 7846 first-cross lambs born, with 2964 wether lambs slaughtered at an average age of 214 days, and wool data from 2795 hogget ewes. Data were analysed using univariate mixed models containing fixed effects for site, year, sex and type of birth and rearing, dam source and sire breed, and random terms for sire and dam effects. Heritabilities and genetic correlations were estimated based on variances from progeny of 70 sires by fitting the same mixed models using a REML procedure in univariate and multivariate analyses. Estimates of heritability were low for lamb growth traits (0.07–0.29), meat colour and meat pH (0.10–0.23), and faecal worm egg count (0.10), moderate for carcass fat and muscle traits (0.32–0.47), and moderate to high for wool traits (0.36–0.55). Estimates of direct genetic correlations among liveweights at various ages were high and positive (0.41–0.77) and those between liveweights and most carcass and meat quality traits were small and varied in sign. Liveweights were moderately to highly positively correlated with most wool traits, except fibre diameter (–0.28–0.08). The study indicates that there is genetic variation for wool, growth, carcass, and meat quality traits, as well as for faecal worm egg count, with scope for selection within Australian maternal sire breeds of sheep.

scholarly journals The role of genetic diversity in the evolution and maintenance of environmentally-cued, male alternative reproductive tactics

2019 ◽  
Author(s):  
KA Stewart ◽  
R Draaijer ◽  
MR Kolasa ◽  
IM Smallegange
Keyword(s):  
Genetic Variation ◽  
Genetic Differentiation ◽  
Genetic Correlations ◽  
Alternative Reproductive Tactics ◽  
Population Variation ◽  
Reproductive Tactics ◽  
Genetic Profiling ◽  
Significant Difference ◽  
The Face ◽  
Bulb Mite

AbstractBackgroundAlternative reproductive tactics (ARTs) are taxonomically pervasive strategies adopted by individuals to maximize reproductive success within populations. Even for conditionally-dependent traits, consensus postulates most ARTs involve both genetic and environmental interactions (GEIs), but to date, quantifying genetic variation underlying the threshold disposing an individual to switch phenotypes in response to an environmental cue has been a difficult undertaking. Our study aims to investigate the origins and maintenance of ARTs within environmentally disparate populations of the microscopic bulb mite, Rhizoglyphus robini, that express ‘fighter’ and ‘scrambler’ male morphs mediated by a complex combination of environmental and genetic factors.ResultsUsing never-before-published individual genetic profiling, we found all individuals across populations are highly inbred with the exception of scrambler males in stressed environments. In fact within the poor environment, scrambler males and females showed no significant difference in genetic differentiation (Fst) compared to all other comparisons, and although fighters were highly divergent from the rest of the population in both poor or rich environments (e.g., Fst, STRUCTURE), fighters demonstrated approximately three times less genetic divergence from the population in poor environments. AMOVA analyses further corroborated significant genetic differentiation across subpopulations, between morphs and sexes, and among subpopulations within each environment.ConclusionOur study provides new insights into the origin of ARTs in the bulb mite, highlighting the importance of GEIs: genetic correlations, epistatic interactions, and sex-specific inbreeding depression across environmental stressors. Asymmetric reproductive output, coupled with the purging of highly inbred individuals during environmental oscillations, also facilitates genetic variation within populations, despite evidence for strong directional selection. This cryptic genetic variation also conceivably facilitates stable population persistence even in the face of spatially or temporally unstable environmental challenges. Ultimately, understanding the genetic context that maintains thresholds, even for conditionally-dependent ARTs, will enhance our understanding of within population variation and our ability to predict responses to selection.

scholarly journals Multi-Level Genetic Variation and Selection Strategy of Neolamarckia cadamba in Successive Years

Forests ◽  
2021 ◽  
Vol 12 (11) ◽  
pp. 1455
Author(s):  
Qingmin Que ◽  
Chunmei Li ◽  
Buye Li ◽  
Huiyun Song ◽  
Pei Li ◽  
...  
Keyword(s):  
Genetic Variation ◽  
Tree Species ◽  
Growth Traits ◽  
Genetic Correlations ◽  
Wood Products ◽  
Early Selection ◽  
Selection Efficiency ◽  
Age Related ◽  
Multi Level ◽  
Variation And Selection

Neolamarckia cadamba (Roxb.) Bosser is a tropical evergreen broadleaf tree species that could play an important role in meeting the increasing demand for wood products. However, multi-level genetic variation and selection efficiency for growth traits in N. cadamba is poorly characterized. We therefore investigated the efficiency of early selection in N. cadamba by monitoring the height (HT), diameter at breast height (DBH), and tree volume (V) in 39 half-sib families from 11 provenances at ages 2, 3, 4, 5, and 6 years in a progeny test. Age-related trends in growth rate, genetic parameters in multi-level, efficiency of early selection, and realized gain in multi-level for growth traits were analyzed. The result showed that genetic variation among families within provenances was higher than that among provenances. The estimated individual heritability values for the growth traits ranged from 0.05 to 0.26, indicating that the variation of growth traits in N. cadamba was subject to weak or intermediate genetic control. The age–age genetic correlations for growth traits were always positive and high (0.51–0.99), and the relationships between the genetic/phenotypic correlations and the logarithm of the age ratio (LAR) were described well by linear models (R2 > 0.85, except the fitting coefficient of genetic correlation and LAR for HT was 0.35). On the basis of an early selection efficiency analysis, we found that it is the best time to perform early selection for N. cadamba at age 5 before half-rotation, and the selection efficiencies were 157.28%, 151.56%, and 127.08% for V, DBH, and HT, respectively. Higher realized gain can be obtained by selecting superior trees from superior families. These results can be expected to provide theoretical guidance and materials for breeding programs in N. cadamba and can even be a reference for breeding strategies of other fast-growing tree species.

scholarly journals Novel insights into the genetic relationship between growth and disease resistance in Pacific salmon

2018 ◽  
Author(s):  
Agustin Barría ◽  
Andrea. B. Doeschl-Wilson ◽  
Jean P. Lhorente ◽  
Ross. D. Houston ◽  
José. M. Yáñez
Keyword(s):  
Genetic Variation ◽  
Disease Resistance ◽  
Survival Time ◽  
Genetic Relationship ◽  
Coho Salmon ◽  
Growth Traits ◽  
Bacterial Load ◽  
Average Daily Gain ◽  
Genetic Correlations ◽  
Piscirickettsia Salmonis

AbstractBackgroundBreeding for disease resistance has become a highly desirable strategy for mitigating infectious disease problems in aquaculture. However, knowledge of the genetic relationship between resistance and other economically important traits, such as growth, is important to assess prior to including disease resistance into the breeding goal. Our study assessed the genetic correlations between growth and survival traits in a large bacterial infection challenge experiment. A population of 2,606 coho salmon individuals from 107 full-sibling families were challenged with the bacteria Piscirickettsia salmonis. Growth was measured as average daily gain prior (ADG0) and during (ADGi) the experimental infection and as harvest weight (HW). Resistance was measured as Survival time (ST) and binary survival (BS). Furthermore, individual measures of bacterial load (BL) were assessed as new resistance phenotypes and to provide an indication of genetic variation in tolerance in salmonid species.ResultsSignificant moderate heritabilities were estimated for ADG0 (0.30 ± 0.05), HW (0.38 ± 0.03), and for the survival traits ST (0.16 ± 0.03) and BS (0.18 ± 0.03). In contrast, heritabilities for ADGi and log-transformed BL were low (0.07 ± 0.02 (significant) and 0.04 ± 0.03, respectively), although these increased to moderate significant levels (0.20 ± 0.09 and 0.12 ± 0.05, respectively) when traits were assessed in survivors only. Significant and favorable genetic correlations were found between ADG0 and the growth traits ADGi (0.40 ± 0.16) and HW (0.64 ± 0.09), as well as with resistance as ST (0.43 ± 0.18), indicating that fish with higher genetic growth rate early on and prior to infection not only tend to maintain their genetic growth advantage until harvest, but also tend to grow faster and survive longer during infection. Furthermore, no robust unfavorable genetic correlations between ADG0 and any of the other traits considered in this study, in particular BL, was identified. Adding log BL as covariates into the models for growth under infection and survival provided an indication for genetic variation in tolerance.ConclusionsThese results suggest that selective breeding for early growth would be expected to simultaneously increase survival time and growth performance during an infection with Piscirickettsia salmonis after accounting for variation in bacterial load, and harvest weight in this coho salmon population, without negatively impacting on pathogen burden.

scholarly journals Genetic Parameters of Growth Traits and Wood Density in Eucalyptus grandis Progenies Planted in Argentina

2009 ◽  
Vol 58 (1-6) ◽  
pp. 11-19 ◽  
Author(s):  
J. J. Vargas Hernández ◽  
J. López Upton ◽  
G. Ramirez Valverde ◽  
L. Harrand
Keyword(s):  
Genetic Variation ◽  
Wood Density ◽  
Genetic Parameters ◽  
Growth Traits ◽  
Genetic Correlations ◽  
Eucalyptus Grandis ◽  
Breeding Population ◽  
The Other ◽  
Stem Form ◽  
Pilodyn Penetration

Abstract To determine the genetic parameters of growth traits and wood density (estimated by Pilodyn penetration) in a population of Eucalyptus grandis formed by families from 11 seed origins in NE Queensland, Australia and a local land race, 8.5-year-old progenies established in three sites in NE Argentina were evaluated. Local trees exhibited higher growth rate at two sites and better stem form, but also deeper Pilodyn penetration, at all sites; thus, there is potential to infuse new genetic variation into the current breeding population in the region. A significant amount of genetic variation was found for all traits assessed, with growth traits [diameter (DBH), height (HT) and volume (VOL)] showing different genetic structure and degree of genetic control as compared to stem straightness (FORM), bark thickness (BARK) and Pilodyn penetration (PILO). The effect of origin was negligible for growth traits while it was important for the other traits, so it should be considered in the selection strategy; in addition, individual heritability for growth traits (0.11-0.16 in the combined analyses) was generally lower than for the other traits (0.20-0.35). Genetic correlations between sites for all traits were high enough to indicate that genotype-environment interaction would not affect the selection program by using a single breeding population for this region. Genetic correlations between growth traits were high (rg > 0.80), and negligible between these and FORM; the only unfavorable genetic correlation was between PILO and BARK (rg = -0.44). Age-age correlations for growth traits and FORM were very high (rg > 0.90). Results suggest that all traits evaluated could be handled in a breeding program, either independently or combined, without negative correlated effects. Early selection for growth traits and stem form is also possible, with major savings in time.

scholarly journals Non-Destructive Assessment of Wood Stiffness in Scots Pine (Pinus sylvestris L.) and its Use in Forest Tree Improvement

Forests ◽  
2019 ◽  
Vol 10 (6) ◽  
pp. 491 ◽  
Author(s):  
Irena Fundova ◽  
Tomas Funda ◽  
Harry X. Wu
Keyword(s):  
Scots Pine ◽  
Growth Traits ◽  
Wood Quality ◽  
Genetic Correlations ◽  
Forest Tree ◽  
Wood Stiffness ◽  
Standing Trees ◽  
Genetic And Phenotypic Correlations ◽  
Sib Families ◽  
Using Data

Wood stiffness is an important wood mechanical property that predetermines the suitability of sawn timber for construction purposes. Negative genetic correlations between wood stiffness and growth traits have, however, been reported for many conifer species including Scots pine. It is, therefore, important that breeding programs consider wood stiffness and growth traits simultaneously. The study aims to (1) evaluate different approaches of calculating the dynamic modulus of elasticity (MOE, non-destructively assessed stiffness) using data from X-ray analysis (SilviScan) as a benchmark, (2) estimate genetic parameters, and (3) apply index selection. In total, we non-destructively measured 622 standing trees from 175 full-sib families for acoustic velocity (VEL) using Hitman and for wood density (DEN) using Resistograph and Pilodyn. We combined VEL with different wood densities, raw (DENRES) and adjusted (DENRES.TB) Resistograph density, Pilodyn density measured with (DENPIL) and without bark (DENPIL.B), constant of 1000 kg·m−3 (DENCONST), and SilviScan density (DENSILV), to calculate MOEs and compare them with the benchmark SilviScan MOE (MOESILV). We also derived Smith–Hazel indices for simultaneous improvement of stem diameter (DBH) and wood stiffness. The highest additive genetic and phenotypic correlations of the benchmark MOESILV with the alternative MOE measures (tested) were attained by MOEDENSILV (0.95 and 0.75, respectively) and were closely followed by MOEDENRES.TB (0.91 and 0.70, respectively) and MOEDENCONST and VEL (0.91 and 0.65, respectively for both). Correlations with MOEDENPIL, MOEDENPIL.B, and MOEDENRES were lower. Narrow-sense heritabilities were moderate, ranging from 0.39 (MOESILV) to 0.46 (MOEDENSILV). All indices revealed an opportunity for joint improvement of DBH and MOE. Conclusions: MOEDENRES.TB appears to be the most efficient approach for indirect selection for wood stiffness in Scots pine, although VEL alone and MOEDENCONST have provided very good results too. An index combining DBH and MOEDENRES.TB seems to offer the best compromise for simultaneous improvement of growth, fiber, and wood quality traits.

Genetic Parameters and Genotype-by-Environment Interactions in Regional Progeny Tests of Pinus taeda L. in the Southern USA

2020 ◽  
Author(s):  
Edwin Lauer ◽  
Andrew Sims ◽  
Steven McKeand ◽  
Fikret Isik
Keyword(s):  
Genetic Variation ◽  
Pinus Taeda ◽  
Genetic Parameters ◽  
Genetic Correlations ◽  
Breeding Program ◽  
Model Fit ◽  
Genotype By Environment Interactions ◽  
Genotype By Environment ◽  
Progeny Tests ◽  
Pinus Taeda L

Abstract Genetic parameters were estimated using a five-series multienvironment trial of Pinus taeda L. in the southern USA. There were 324 half-sib families planted in five test series across 37 locations. A set of six variance/covariance matrices for the genotype-by-environment (G × E) effect for tree height and diameter were compared on the basis of model fit. In single-series analysis, extended factor analytical models provided generally superior model fit to simpler models for both traits; however, in the combined-series analysis, diameter was optimally modeled using simpler variance/covariance structures. A three-way compound term for modeling G × E interactions among and within series yielded substantial improvements in terms of model fit and standard errors of predictions. Heritability of family means ranged between 0.63 and 0.90 for both height and diameter. Average additive genetic correlations among sites were 0.70 and 0.61 for height and diameter, respectively, suggesting the presence of some G × E interaction. Pairs of sites with the lowest additive genetic correlations were located at opposite ends of the latitude range. Latent factor regression revealed a small number of parents with large factor scores that changed ranks significantly between southern and northern environments. Study Implications Multienvironmental progeny tests of loblolly pine (Pinus taeda L.) were established over 10 years in the southern United States to understand the genetic variation for the traits of economic importance. There was substantial genetic variation between open-pollinated families, suggesting that family selection would be efficient in the breeding program. Genotype-by-environment interactions were negligible among sites in the deployment region but became larger between sites at the extremes of the distribution. The data from these trials are invaluable in informing the breeding program about the genetic merit of selection candidates and their potential interaction with the environment. These results can be used to guide deployment decisions in the southern USA, helping landowners match germplasm with geography to achieve optimal financial returns and conservation outcomes.

scholarly journals The Molecular Basis of Quantitative Genetic Variation in Central and Secondary Metabolism in Arabidopsis

Genetics ◽  
1998 ◽  
Vol 149 (2) ◽  
pp. 739-747 ◽  
Author(s):  
Thomas Mitchell-Olds ◽  
Deana Pedersen
Keyword(s):  
Enzyme Activity ◽  
Genetic Variation ◽  
Secondary Metabolism ◽  
Genetic Correlations ◽  
Theoretical Models ◽  
Model Systems ◽  
Activity Levels ◽  
Significant Qtls ◽  
Cis Acting ◽  
Quantitative Genetic

Abstract To find the genes controlling quantitative variation, we need model systems where functional information on physiology, development, and gene regulation can guide evolutionary inferences. We mapped quantitative trait loci (QTLs) influencing quantitative levels of enzyme activity in primary and secondary metabolism in Arabidopsis. All 10 enzymes showed highly significant quantitative genetic variation. Strong positive genetic correlations were found among activity levels of 5 glycolytic enzymes, PGI, PGM, GPD, FBP, and G6P, suggesting that enzymes with closely related metabolic functions are coregulated. Significant QTLs were found influencing activity of most enzymes. Some enzyme activity QTLs mapped very close to known enzyme-encoding loci (e.g., hexokinase, PGI, and PGM). A hexokinase QTL is attributable to cis-acting regulatory variation at the AtHXK1 locus or a closely linked regulatory locus, rather than polypeptide sequence differences. We also found a QTL on chromosome IV that may be a joint regulator of GPD, PGI, and G6P activity. In addition, a QTL affecting PGM activity maps within 700 kb of the PGM-encoding locus. This QTL is predicted to alter starch biosynthesis by 3.4%, corresponding with theoretical models, suggesting that QTLs reflect pleiotropic effects of mutant alleles.

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