Quantitative Trait Loci Alleles Research Articles

Transcriptome Profiling of Embryonic Development Rate in Rainbow Trout Advanced Backcross Introgression Lines

In rainbow trout (Oncorhynchus mykiss) and other fishes, embryonic development rate is an ecologically and evolutionarily important trait that is closely associated with survival and physiological performance later in life. To identify genes differentially regulated in fast and slow-developing embryos of rainbow trout, we examined gene expression across developmental time points in rainbow trout embryos possessing alleles linked to a major quantitative trait loci (QTL) for fast versus slow embryonic development rate. Whole genome expression microarray analyses were conducted using embryos from a fourth generation backcross family, whereby each backcross generation involved the introgression of the fast-developing alleles for a major development rate QTL into a slow-developing clonal line of rainbow trout. Embryos were collected at 15, 19, and 28days post-fertilization; sex and QTL genotype were determined using molecular markers, and cDNA from 48 embryos were used for microarray analysis. A total of 183 features were identified with significant differences between embryonic development rate genotypes. Genes associated with cell cycle growth, muscle contraction and protein synthesis were expressed significantly higher in embryos with the fast-developing allele (Clearwater) than those with the slow-developing allele (Oregon State University), which may associate with fast growth and early body mass construction in embryo development. Across time points, individuals with the fast-developing QTL allele appeared to have earlier onset of these developmental processes when compared to individuals with the slow development alleles, even as early as 15days post-fertilization. Differentially expressed candidate genes chosen for linkage mapping were localized primarily to regions outside of the major embryonic development rate QTL, with the exception of a single gene (very low-density lipoprotein receptor precursor).

Identification of a major QTL allele from wild soybean (Glycine soja Sieb. & Zucc.) for increasing alkaline salt tolerance in soybean

Salt-affected soils are generally classified into two main categories, sodic (alkaline) and saline. Our previous studies showed that the wild soybean accession JWS156-1 (Glycine soja) from the Kinki area of Japan was tolerant to NaCl salt, and the quantitative trait locus (QTL) for NaCl salt tolerance was located on soybean linkage group N (chromosome 3). Further investigation revealed that the wild soybean accession JWS156-1 also had a higher tolerance to alkaline salt stress. In the present study, an F(6) recombinant inbred line mapping population (n = 112) and an F(2) population (n = 149) derived from crosses between a cultivated soybean cultivar Jackson and JWS156-1 were used to identify QTL for alkaline salt tolerance in soybean. Evaluation of soybean alkaline salt tolerance was carried out based on salt tolerance rating (STR) and leaf chlorophyll content (SPAD value) after treatment with 180 mM NaHCO(3) for about 3 weeks under greenhouse conditions. In both populations, a significant QTL for alkaline salt tolerance was detected on the molecular linkage group D2 (chromosome 17), which accounted for 50.2 and 13.0% of the total variation for STR in the F(6) and the F(2) populations, respectively. The wild soybean contributed to the tolerance allele in the progenies. Our results suggest that QTL for alkaline salt tolerance is different from the QTL for NaCl salt tolerance found previously in this wild soybean genotype. The DNA markers closely associated with the QTLs might be useful for marker-assisted selection to pyramid tolerance genes in soybean for both alkaline and saline stresses.

Quantitative Trait Loci and Trait Correlations for Maize Stover Cell Wall Composition and Glucose Release for Cellulosic Ethanol

In cellulosic ethanol production, the efficiency of converting maize (Zea mays L.) stover into fermentable sugars partly depends on the stover cell wall structure. Breeding for improved stover quality for cellulosic ethanol may benefit from the use of molecular markers. However, limited quantitative trait loci (QTL) studies have been published for maize stover cell wall components, and no QTL study has been published for glucose release (GLCRel) from stover by a cellulosic ethanol conversion process. Our objectives were to characterize the relationships among stover cell wall components and GLCRel, and to identify QTL with major effects, if any, influencing stover cell wall composition and GLCRel. Testcrosses of 223 intermated B73 × Mo17 recombinant inbreds and the parent lines were analyzed for cell wall composition and GLCRel after acid pretreatment and enzymatic hydrolysis. As expected, glucose (GLC), xylose (XYL), and Klason lignin (KL) composed the bulk (∼72%) of the stover dry matter. Significant genetic variance and moderate heritability were observed for all traits. Genetic and phenotypic correlations among traits were generally in the favorable direction but also reflected the complexity of maize stover cell wall composition. We found 152 QTL, mostly with small effects, for GLCRel and cell wall components on both a dry matter and cell wall basis. Because no major QTL were found, methods that increase the frequency of favorable QTL alleles or that predict performance based on markers would be appropriate in marker‐assisted breeding for maize stover quality for cellulosic ethanol.

Detection of Quantitative Trait Loci for Growth and Carcass Traits on BTA6 in a Hanwoo Population

The purpose of this study was to detect quantitative trait loci (QTL) for growth and carcass quality traits on BTA6 in a population of Hanwoo cattle. Three hundred and sixty one steers were produced from 39 sires that were sired by 17 grandsires in the two Hanwoo farming branches of the National Livestock Research Institute of Korea, between Spring 2000 and Fall 2002. DNA samples were collected for all of the steers, sires and grandsires, and the phenotypes for six growth and carcass quality traits were measured at 24 months of age. Twelve microsatellite markers were chosen on BTA6 and a linkage map was constructed by using seven of the twelve markers. Then, a chromosome-wide QTL scan was performed by applying an Animal Model, in which effects of QTL alleles within the grand sires were fitted as a random term. Three QTL were detected at the 5% chromosome-wise level for backfat thickness, average daily gain, and final weight. The most likely positions for the QTL were in the proximal region, i.e. 0 cM, 35 cM, and 63 cM, respectively. Also, another QTL for longissimus dorsi muscle area was detected at the 10% chromosome-wise level at 67 cM. These results were, in general, consistent with our previous report, in which candidate gene analyses showed that a SNP near ILSTS035 flanked by BM4621 (62.5 cM) and BMS2460 (81.3 cM) was associated with final weight, carcass weight, average daily gain, and longissimus dorsi muscle area in the same Hanwoo population.

QTL alleles from a winter feed type can improve malting quality in barley

AbstractMapping quantitative trait loci (QTLs) responsible for malting quality traits in barley populations has been the main genetic approach to malting quality breeding. A ‘winter × spring’ doubled haploid barley population ‘Nure’ × ‘Tremois’, where such traits were segregating, has been recently developed. Our objective was to map QTLs for malting quality from 2 years of trials in two contrasting locations. QTLs were found on six chromosomes, with a main cluster on chromosome 1H. For wort viscosity and malt extract, favourable alleles at two loci on chromosome 5H were carried by the winter feeding parent ‘Nure’. Doubled‐haploids with higher quality than the spring malting cultivar ‘Tremois’ showed either a facultative or a winter growth habit and a level of frost tolerance comparable with that of the winter tolerant parent ‘Nure’. Markers and QTLs of quality traits were further validated on a separate set of DH lines, coming from the same cross, by means of marker‐assisted selection. This showed that, at least in the present cross combination, positive contributions to malting quality can be found in winter feed barley.

Identification of genomic regions carrying QTL for agronomic and quality traits in rye (Secale cereale) introgression libraries

AbstractIntrogression libraries can be used to localize genomic regions carrying quantitative trait loci (QTL). We used this approach to detect QTL regions affecting the per se performance of agronomic and quality traits with two rye (Secale cereale L.) introgression libraries. Our objectives were to detect candidate introgression lines (pre‐ILs) that have a different per se performance than the recurrent parent and to identify the underlying QTL regions. The introgression libraries containing 40 BC2S3 lines each were established with marker‐assisted backcrossing from crosses of the heterozygous Iranian primitive rye accession Altevogt 14160 and the elite inbred line L2053‐N. To assess the phenotypic effect of the donor chromosome segments (DCS) the pre‐ILs were evaluated for grain yield, plant height, thousand‐kernel weight, test weight, falling number and protein content in replicated field trials at five locations in Germany over 2 years. In total, 58 significant (P < 0.05) differences between pre‐ILs and L2053‐N were observed in each introgression library. The DCS in pre‐ILs differing from the recurrent parent possess most likely the responsible QTL. Genomic regions carrying favourable QTL alleles were detected for test weight, thousand‐kernel weight and protein content. We conclude that Altevogt 14160 can not only be used to enrich the genetic variation of the restricted hybrid rye gene pools but will also allow the breeder to efficiently detect favourable QTL for marker‐assisted selection.

Identification and characterization of large-effect quantitative trait loci for grain yield under lowland drought stress in rice using bulk-segregant analysis

An F(4:5) population of 490 recombinant inbred lines (RILs) from the cross Apo/(2*)Swarna was used to detect quantitative trait loci (QTL) with large effects on grain yield under drought stress using bulk-segregant analysis (BSA). Swarna is an important rainfed lowland rice variety grown on millions of hectares in Asia, but is highly susceptible to drought and aerobic soil conditions. Apo is an aerobic-adapted variety with moderate tolerance to drought. Two rice microsatellite (RM) markers, RM324, and RM416, located on chromosomes 2 and 3, respectively, were shown via BSA to be strongly associated with yield under lowland drought stress. The effects of these QTL were tested in a total of eight hydrological environments over a period of 3 years. The QTL linked to RM416 (DTY(3.1)) had a large effect on grain yield under severe lowland drought stress, explaining about 31% of genetic variance for the trait (P < 0.0001). It also explained considerable variance for yield under mild stress in lowland conditions and aerobic environments. To our knowledge this is the first reported QTL that has a large effect on yield in both lowland drought and aerobic environments. The QTL linked to RM324 (DTY(2.1)) had a highly significant effect on grain yield in lowland drought stress (R(2) = 13-16%) and in two aerobic trials. The effect of these QTL on grain yield was verified to be not mainly due to phenology differences. Effects of DTY(3.1) on yield under stress have been observed in several other rice mapping populations studied at IRRI. Results of this study indicate that BSA is an effective method of identifying QTL alleles with large effects on rice yield under severe drought stress. The Apo alleles for these large-effect QTL for grain yield under drought and aerobic conditions may be immediately exploited in marker-assisted-breeding to improve the drought tolerance of Swarna.

Genetic dissection of Lactuca saligna nonhost resistance to downy mildew at various lettuce developmental stages

This study used the pathosystem of lettuce (Lactuca spp.) and downy mildew (Bremia lactucae) as a model to investigate the inheritance of nonhost resistance, and focused on the contribution of quantitative trait loci (QTLs) to nonhost resistance at various developmental stages in the lettuce life cycle. A set of 28 backcross inbred lines (BILs) of L. saligna CGN05271 (nonhost) introgressions in a L. sativa cv. Olof (host) background identified 16 introgressions that contributed to resistance at various plant developmental stages: seedlings, young plants, adult plants in the greenhouse and adult plants in the field. This paper provisionally considered these introgressions to be 16 QTLs. Of these 16 QTLs, seven were identified previously and nine were new. For 15 QTLs (Rbq1, Rbq2, rbq3–7 and Rbq8–15), the resistance alleles were derived from the nonhost L. saligna; the resistance allele of the other QTL (Rbq16) was from the susceptible L. sativa cv. Olof. Of the 15 QTLs in L. saligna, only two, rbq5 and rbq7, were found to be effective at every plant developmental stage; the other 13 QTLs were only effective at certain developmental stages. Experiments with seven B. lactucae races did not provide evidence that any QTL was race‐specific. The data suggest that nonhost resistance in L. saligna is the result of cumulative effects of many resistance QTLs operating at various developmental stages.

Genome‐wide identification of quantitative trait loci for carcass composition and meat quality in a large‐scale White Duroc × Chinese Erhualian resource population

Carcass and meat quality traits are economically important in pigs. In this study, 17 carcass composition traits and 23 meat quality traits were recorded in 1028 F(2) animals from a White Duroc x Erhualian resource population. All pigs in this experimental population were genotyped for 194 informative markers covering the entire porcine genome. Seventy-seven genome-wide significant quantitative trait loci (QTL) for carcass traits and 68 for meat quality were mapped to 34 genomic regions. These results not only confirmed many previously reported QTL but also revealed novel regions associated with the measured traits. For carcass traits, the most prominent QTL was identified for carcass length and head weight at 57 cM on SSC7, which explained up to 50% of the phenotypic variance and had a 95% confidence interval of only 3 cM. Moreover, QTL for kidney and spleen weight and lengths of cervical vertebrae were reported for the first time in pigs. For meat quality traits, two significant QTL on SSC5 and X were identified for both intramuscular fat content and marbling score in the longissimus muscle, while three significant QTL on SSC1 and SSC9 were found exclusively for IMF. Both LM and the semimembranous muscle showed common QTL for colour score on SSC4, 5, 7, 8, 13 and X and discordant QTL on other chromosomes. White Duroc alleles at a majority of QTL detected were favourable for carcass composition, while favourable QTL alleles for meat quality originated from both White Duroc and Erhualian.

Characterization of a QTL region affecting clinical mastitis and protein yield on BTA6

Quantitative trait loci affecting clinical mastitis were detected and fine mapped to a narrow region on bovine chromosome 6 in the Norwegian Red cattle population. The region includes the casein gene cluster and several candidate genes thought to influence clinical mastitis. The most significant results were found for SNPs within the Mucin 7 gene. This gene encodes an antimicrobial peptide and constitutes part of the first line of defence for the mucosal immune system. Detection of long haplotypes extending several Mb may indicate that artificial selection has influenced the haplotype structures in the region. A search for selection sweeps supports this observation and coincides with association results found both by single SNP and haplotype analyses. Our analyses identified haplotypes carrying quantitative trait loci alleles associated with high protein yield and simultaneously fewer incidences of clinical mastitis. The fact that such haplotypes are found in relative high frequencies in Norwegian Red may reflect the combined breeding goal that is characterized by selection for both milk production and disease resistance. The identification of these haplotypes raises the possibility of overcoming the unfavourable genetic correlation between these traits through haplotype-assisted selection.

Evaluation of QTL alleles from exotic sources for hybrid seed yield in the original and different genetic backgrounds of spring-type Brassica napus L.

Previously identified alleles at quantitative trait loci (QTL) for hybrid seed yield were re-evaluated in the same genetic background (in hybrid combination with the same tester) as the original QTL mapping study and also evaluated in a different genetic background (in hybrid combination with two different testers). The QTL were identified from wide crosses of exotic germplasm sources with spring-type Brassica napus L., in which alleles from the exotic germplasm sources increased hybrid seed yield. Results from the re-evaluation of six QTL, in the same genetic background and hybrid combination, indicate that several of the exotic donor QTL alleles did increase hybrid seed yield and could be successfully used for improving the original single-cross hybrid. However, results from the evaluation of seven QTL (including the same six previous QTL) in a new genetic background, in combination with two new testers, indicate that the exotic QTL alleles were often no different or produced significantly lower hybrid seed yield than the spring QTL alleles. In all studies, the QTL were also very sensitive to environmental interactions. Thus, our results indicate that although these exotic sources contain favorable QTL alleles when introgressed into one spring hybrid background, the effects are not predictive of other genetic backgrounds or hybrid combinations. Although QTL affecting hybrid seed yield have been identified, comparisons of multiple QTL alleles are needed to determine the most favorable allele at each locus. Characterization of QTL complementation across testers will be required to predict their effects in multiple hybrid combinations.

Identification of genomic regions determining the phenological development leading to floral transition in wheat (Triticum aestivum L.)

Autumn-seeded winter cereals acquire tolerance to freezing temperatures and become vernalized by exposure to low temperature (LT). The level of accumulated LT tolerance depends on the cold acclimation rate and factors controlling timing of floral transition at the shoot apical meristem. In this study, genomic loci controlling the floral transition time were mapped in a winter wheat (T. aestivum L.) doubled haploid (DH) mapping population segregating for LT tolerance and rate of phenological development. The final leaf number (FLN), days to FLN, and days to anthesis were determined for 142 DH lines grown with and without vernalization in controlled environments. Analysis of trait data by composite interval mapping (CIM) identified 11 genomic regions that carried quantitative trait loci (QTLs) for the developmental traits studied. CIM analysis showed that the time for floral transition in both vernalized and non-vernalized plants was controlled by common QTL regions on chromosomes 1B, 2A, 2B, 6A and 7A. A QTL identified on chromosome 4A influenced floral transition time only in vernalized plants. Alleles of the LT-tolerant parent, Norstar, delayed floral transition at all QTLs except at the 2A locus. Some of the QTL alleles delaying floral transition also increased the length of vegetative growth and delayed flowering time. The genes underlying the QTLs identified in this study encode factors involved in regional adaptation of cold hardy winter wheat.

Marker-assisted selection can reduce true as well as pedigree-estimated inbreeding

This study investigated whether selection using genotype information reduced the rate and level of true inbreeding, that is, identity by descent, at a selectively neutral locus as well as a locus under selection compared with traditional BLUP selection. In addition, the founder representation at these loci and the within-family selection at the nonneutral locus were studied. The study was carried out using stochastic simulation of a population resembling the breeding nucleus of a dairy cattle population for 25 yr. Each year, 10 proven bulls were selected across herds along with 100 dams from within each of 40 herds. Selection was performed using BLUP, marker-assisted, or gene-assisted selection for a trait with low heritability (h2=0.04) only expressed in females, mimicking a health trait. The simulated genome consisted of 2 chromosomes. One biallelic quantitative trait loci (QTL) with an initial frequency of the favorable allele of 0.1, and initially explaining 25% of the genetic variance as well as 4 markers were simulated in linkage disequilibrium, all positioned at chromosome 1. Chromosome 2 was selectively neutral, and consisted of a single neutral locus. The results showed that in addition to reducing pedigree-estimated inbreeding, the incorporation of genotype information in the selection criteria also reduced the level and rate of true inbreeding. In general, true inbreeding in the QTL was greater than pedigree-estimated inbreeding with respect to both the level and rate of inbreeding, as expected. Also as expected, true and pedigree-estimated inbreeding in the neutral locus were the same. Furthermore, after 25 yr, or approximately 5 generations, the pedigree-estimated level of inbreeding was reduced by 11 and 24% compared with BLUP in gene- and marker-assisted selection, respectively, and the level of true inbreeding in the QTL was reduced by 22 and 13%, respectively. The difference between selection scenarios was found to be caused by a larger number of founders being represented at the QTL when using genotype information in the selection criteria. This in turn was caused by an increased selection of individuals sharing the favorable QTL allele rather than individuals sharing genes on average, which was shown by a higher Mendelian selection differential in the QTL. Hence, even though the selection pressure was increased at the QTL, more variation was retained. The results suggest that marker-assisted selection is a useful selection strategy.

Simultaneous selection of major and minor genes: use of QTL to increase selection efficiency of coleoptile length of wheat (Triticum aestivum L.)

Plant breeders simultaneously select for qualitative traits controlled by one or a small number of major genes, as well as for polygenic traits controlled by multiple genes that may be detected as quantitative trait loci (QTL). In this study, we applied computer simulation to investigate simultaneous selection for alleles at both major and minor gene (as QTL) loci in breeding populations of two wheat parental lines, HM14BS and Sunstate. Loci targeted for selection included six major genes affecting plant height, disease resistance, and grain quality, plus 6 known and 11 "unidentified" QTL affecting coleoptile length (CL). Parental line HM14BS contributed the target alleles at two of the major gene loci, while parental line Sunstate contributed target alleles at four loci. The parents have similar plant height, but HM14BS has a longer coleoptile, a desirable attribute for deep sowing in rainfed environments. Including the wild-type allele at the major reduced-height locus Rht-D1, HM14BS was assumed to have 13 QTL for increased CL, and Sunstate four; these assumptions being derived from mapping studies and empirical data from an actual HM14BS/Sunstate population. Simulation indicated that compared to backcross populations, a single biparental F(1) cross produced the highest frequency of target genotypes (six desired alleles at major genes plus desired QTL alleles for long CL). From 1,000 simulation runs, an average of 2.4 individuals with the target genotype were present in unselected F(1)-derived doubled haploid (DH) or recombinant inbred line (RIL) populations of size 200. A selection scheme for the six major genes increased the number of target individuals to 19.1, and additional marker-assisted selection (MAS) for CL increased the number to 23.0. Phenotypic selection (PS) of CL outperformed MAS in this study due to the high heritability of CL, incompletely linked markers for known QTL, and the existence of unidentified QTL. However, a selection scheme combining MAS and PS was equally as efficient as PS and would result in net savings in production and time to delivery of long coleoptile wheats containing the six favorable alleles.

A major QTL for resistance of rice to the parasitic plant Striga hermonthica is not dependent on genetic background.

The use of Striga-resistant germplasm is likely to be a cost-effective control strategy for preventing loss of yield owing to Striga. Previously, the authors identified quantitative trait loci (QTL) for resistance in rice to Striga hermonthica (Del.) Benth. in backcross inbred lines (BILs) derived from a cross between two cultivars Nipponbare and Kasalath. It is essential to validate QTL in different environments and/or genetic backgrounds to develop molecular markers linked to resistance QTL for use in marker-assisted selection (MAS) programmes. This study aimed to establish whether a large-effect Kasalath-derived resistance QTL allele on chromosome 4 of rice also conferred resistance in a different mapping population derived from a cross between Koshihikari and Kasalath, and to identify any further Striga resistance QTL. Three Striga resistance QTL were detected in Koshihikari-Kasalath BILs, two of which were derived from the Kasalath allele and one from the Koshihkari allele. The largest QTL (Kasalath allele) explained 16% of the variation in the mapping population and was located on chromosome 4. Comparison between these data and those of the authors' previous analysis revealed that the confidence intervals of the chromosome-4 QTL in the Nipponbare-Kasalath cross and the Kasalath-Koshihikari cross overlapped between 6.5 Mbp and 8 Mbp on the physical rice genome assembly. This study has both verified and narrowed down the position of a Striga resistance QTL of major effect, and demonstrated that it may be a tractable target for MAS.

Two distinct major QTL for resistance to fire blight co-localize on linkage group 12 in apple genotypes ‘Evereste’ andMalus floribundaclone 821

Fire blight, caused by the bacterium Erwinia amylovora, is one of the most destructive diseases of apple (Malus xdomestica) worldwide. No major, qualitative gene for resistance to this disease has been identified so far in apple. A quantitative trait locus (QTL) analysis was performed in two F1 progenies derived from two controled crosses: one between the susceptible rootstock cultivar 'MM106' and the resistant ornamental cultivar 'Evereste' and the other one between the moderately susceptible cultivar 'Golden Delicious' and the wild apple Malus floribunda clone 821, with unknown level of fire blight resistance. Both progenies were inoculated in the greenhouse with the same reference strain of E. amylovora. The length of stem necrosis was scored 7 and 14 days after inoculation. A strong QTL effect was identified in both 'Evereste' and M. floribunda 821 at a similar position on the distal region of linkage group 12 of the apple genome. From 50% to 70% of the phenotypic variation was explained by the QTL in 'Evereste' progeny according to the scored trait. More than 40% of the phenotypic variation was explained by the M. floribunda QTL in the second progeny. It was shown that 'Evereste' and M. floribunda 821 carried distinct QTL alleles at that genomic position. A small additional QTL was identified in 'Evereste' on linkage group 15, which explained about 6% of the phenotypic variation. Although it was not possible to confirm whether or not 'Evereste' and M. floribunda QTL belonged to the same locus or two distinct closely related loci, these QTL can be valuable targets in marker-assisted selection to obtain fire blight resistant apple cultivars and form a starting point for discovering the function of the genes controlling apple fire blight resistance.

GENETIC ARCHITECTURE FOR THE ADAPTIVE ORIGIN OF ANNUAL WILD RICE,ORYZA NIVARA

The wild progenitors of cultivated rice, Oryza nivara and Oryza rufipogon, provide an experimental system for characterizing the genetic basis of adaptation. The evolution of annual O. nivara from a perennial ancestor resembling its sister species, O. rufipogon, was associated with an ecological shift from persistently wet to seasonally dry habitats. Here we report a quantitative trait locus (QTL) analysis of phenotypic differentiation in life history, mating system, and flowering time between O. nivara and O. rufipogon. The exponential distribution of effect sizes of QTL fits the prediction of a recently proposed population genetic model of adaptation. More than 80% of QTL alleles of O. nivara acted in the same direction of phenotypic evolution, suggesting that they were fixed under directional selection. The loss of photoperiod sensitivity, which might be essential to the survival of the ancestral populations of O. nivara in the new environment, was controlled by QTL of relatively large effect. Mating system evolution from cross- to self-fertilization through the modification of panicle and floral morphology was controlled by QTL of small-to-moderate effect. The lack of segregation of the recessive annual habit in the F(2) mapping populations suggested that the evolution of annual from perennial life form had a complex genetic basis. The study captured the genetic architecture for the adaptive origin of O. nivara and provides a foundation for rigorous experimental tests of population genetic theories of adaptation.

Confirmation and fine-mapping of a major QTL for resistance to infectious pancreatic necrosis in Atlantic salmon (Salmo salar): population-level associations between markers and trait.

BackgroundInfectious pancreatic necrosis (IPN) is one of the most prevalent and economically devastating diseases in Atlantic salmon (Salmo salar) farming worldwide. The disease causes large mortalities at both the fry- and post-smolt stages. Family selection for increased IPN resistance is performed through the use of controlled challenge tests, where survival rates of sib-groups are recorded. However, since challenge-tested animals cannot be used as breeding candidates, within-family selection is not performed and only half of the genetic variation for IPN resistance is being exploited. DNA markers linked to quantitative trait loci (QTL) affecting IPN resistance would therefore be a powerful selection tool. The aim of this study was to identify and fine-map QTL for IPN-resistance in Atlantic salmon, for use in marker-assisted selection to increase the rate of genetic improvement for this trait.ResultsA genome scan was carried out using 10 large full-sib families of challenge-tested Atlantic salmon post-smolts and microsatellite markers distributed across the genome. One major QTL for IPN-resistance was detected, explaining 29% and 83% of the phenotypic and genetic variances, respectively. This QTL mapped to the same location as a QTL recently detected in a Scottish Atlantic salmon population. The QTL was found to be segregating in 10 out of 20 mapping parents, and subsequent fine-mapping with additional markers narrowed the QTL peak to a 4 cM region on linkage group 21. Challenge-tested fry were used to show that the QTL had the same effect on fry as on post-smolt, with the confidence interval for QTL position in fry overlapping the confidence interval found in post-smolts. A total of 178 parents were tested for segregation of the QTL, identifying 72 QTL-heterozygous parents. Genotypes at QTL-heterozygous parents were used to determine linkage phases between alleles at the underlying DNA polymorphism and alleles at single markers or multi-marker haplotypes. One four-marker haplotype was found to be the best predictor of QTL alleles, and was successfully used to deduce genotypes of the underlying polymorphism in 72% of the parents of the next generation within a breeding nucleus. A highly significant population-level correlation was found between deduced alleles at the underlying polymorphism and survival of offspring groups in the fry challenge test, parents with the three deduced genotypes (QQ, Qq, qq) having mean offspring mortality rates of 0.13, 0.32, and 0.49, respectively. The frequency of the high-resistance allele (Q) in the population was estimated to be 0.30. Apart from this major QTL, one other experiment-wise significant QTL for IPN-resistance was detected, located on linkage group 4.ConclusionThe QTL confirmed in this study represents a case of a major gene explaining the bulk of genetic variation for a presumed complex trait. QTL genotypes were deduced within most parents of the 2005 generation of a major breeding company, providing a solid framework for linkage-based MAS within the whole population in subsequent generations. Since haplotype-trait associations valid at the population level were found, there is also a potential for MAS based on linkage disequilibrium (LD). However, in order to use MAS across many generations without reassessment of linkage phases between markers and the underlying polymorphism, the QTL needs to be positioned with even greater accuracy. This will require higher marker densities than are currently available.

CONNECTING QTLS TO THE G-MATRIX OF EVOLUTIONARY QUANTITATIVE GENETICS

Evolutionary quantitative genetics has recently advanced in two distinct streams. Many biologists address evolutionary questions by estimating phenotypic selection and genetic (co)variances (G matrices). Simultaneously, an increasing number of studies have applied quantitative trait locus (QTL) mapping methods to dissect variation. Both conceptual and practical difficulties have isolated these two foci of quantitative genetics. A conceptual integration follows from the recognition that QTL allele frequencies are the essential variables relating the G-matrix to marker-based mapping experiments. Breeding designs initiated from randomly selected parental genotypes can be used to estimate QTL-specific genetic (co)variances. These statistics appropriately distill allelic variation and provide an explicit population context for QTL mapping estimates. Within this framework, one can parse the G-matrix into a set of mutually exclusive genomic components and ask whether these parts are similar or dissimilar in their respective features, for example the magnitude of phenotypic effects and the extent and nature of pleiotropy. As these features are critical determinants of sustained response to selection, the integration of QTL mapping methods into G-matrix estimation can provide a concrete, genetically based experimental program to investigate the evolutionary potential of natural populations.

Mapping of resistance to spot blotch disease caused by Bipolaris sorokiniana in spring wheat

Spot blotch caused by Bipolaris sorokiniana is a destructive disease of wheat in warm and humid wheat growing regions of the world. The development of disease resistant cultivars is considered as the most effective control strategy for spot blotch. An intervarietal mapping population in the form of recombinant inbred lines (RILs) was developed from a cross 'Yangmai 6' (a Chinese source of resistance) x 'Sonalika' (a spot blotch susceptible cultivar). The 139 single seed descent (SSD) derived F(6), F(7), F(8) lines of 'Yangmai 6' x 'Sonalika' were evaluated for resistance to spot blotch in three blocks in each of the 3 years. Joint and/or single year analysis by composite interval mapping (CIM) and likelihood of odd ratio (LOD) >2.2, identified four quantitative trait loci (QTL) on the chromosomes 2AL, 2BS, 5BL and 6DL. These QTLs were designated as QSb.bhu-2A, QSb.bhu-2B, QSb.bhu-5B and QSb.bhu-6D, respectively. A total of 63.10% of phenotypic variation was explained by these QTLs based on the mean over years. Two QTLs on chromosomes 2B and 5B with major effects were consistent over 3 years. All QTL alleles for resistance were derived from the resistant parent 'Yangmai 6'.