QTL analysis of flower and fruit traits in sour cherry

The map locations and effects of quantitative trait loci (QTLs) were estimated for eight flower and fruit traits in sour cherry (Prunus cerasus L.) using a restriction fragment length polymorphism (RFLP) genetic linkage map constructed from a double pseudo-testcross. The mapping population consisted of 86 progeny from the cross between two sour cherry cultivars, Rheinische Schattenmorelle (RS)×Erdi Botermo (EB). The genetic linkage maps for RS and EB were 398.2 cM and 222.2 cM, respectively, with an average interval length of 9.8 cM. The RS/EB linkage map that was generated with shared segregating markers consisted of 17 linkage groups covering 272.9 cM with an average interval length of 4.8 cM. Eleven putatively significant QTLs (LOD >2.4) were detected for six characters (bloom time, ripening time, % pistil death, % pollen germination, fruit weight, and soluble solids concentration). The percentage of phenotypic variation explained by a single QTL ranged from 12.9% to 25.9%. Of the QTLs identified for the traits in which the two parents differed significantly, 50% had allelic effects opposite to those predicted from the parental phenotype. Three QTLs affecting flower traits (bloom time, % pistil death, and % pollen germination) mapped to a single linkage group, EB 1. The RFLP closest to the bloom time QTL on EB 1 was detected by a sweet cherry cDNA clone pS141 whose partial amino acid sequence was 81% identical to that of a Japanese pear stylar RNase. Received: 4 March 1999 / Accepted: 27 August 1999     

QTL analysis of fruit quality in fresh market tomato: a few chromosome regions control the variation of sensory and instrumental traits

The organoleptic quality of tomato fruit involves a set of attributes (flavour, aroma, texture) that can be evaluated either by sensory analyses or by instrumental measures. In order to study the genetic control of this characteristic, a recombinant inbred line (RIL) population was developed from an intraspecific cross between a cherry tomato line with a good overall aroma intensity and an inbred line with medium flavour but bigger fruits. A total of 38 traits involved in organoleptic quality were evaluated. Physical traits included fruit weight, diameter, colour, firmness, and elasticity. Chemical traits were dry matter weight, titratable acidity, pH, and the contents of soluble solids, sugars, lycopene, carotene, and 12 aroma volatiles. A panel of trained assessors quantified sensory attributes: flavour (sweetness and sourness), aroma (overall aroma intensity, together with candy, lemon, citrus fruit, and pharmaceutical aromas) and texture (firmness, meltiness, mealiness, juiciness, and skin difficult to swallow). RILs showed a large range of variation. Molecular markers were used to map a total of 130 quantitative trait loci (QTL) for the 38 traits. They were mainly distributed in a few chromosome regions. Major QTLs (R(2) >30%) were detected for fruit weight, diameter, colour, firmness, meltiness, and for six aroma volatiles. The relationships between instrumental measures and sensory traits were analysed with regard to the QTL map. A special insight was provided about the few regions where QTLs are related to multiple traits. A few examples are shown to illustrate how the simultaneous analysis of QTL segregation for related traits may aid in understanding the genetic control of quality traits and pave the way towards QTL characterization.     

QTL analysis of fruit components in the progeny of a Rennell Island Tall coconut (Cocos nucifera L.) individual

We investigated the genetic factors controlling fruit components in coconut by performing QTL analyses for fruit component weights and ratios in a segregating progeny of a Rennell Island Tall genotype. The underlying linkage map of this population was already established in a previous study, as well as QTL analyses for fruit production, which were used to complement our results. The addition of 53 new markers (mainly SSRs) led to minor amendments in the map. A total of 52 putative QTLs were identified for the 11 traits under study. Thirty-four of them were grouped in six small clusters, which probably correspond to single pleiotropic genes. Some additional QTLs located apart from these clusters also had relatively large effects on the individual traits. The QTLs for fruit component weight, endosperm humidity and fruit production were found at different locations in the genome, suggesting that efficient marker-assisted selection for yield can be achieved by selecting QTLs for the individual components. The detected QTLs descend from a genotype belonging to the “Pacific” coconut group. Based on the known molecular and phenotypic differences between “Pacific” and “Indo-Atlantic” coconuts, we suggest that a large fraction of coconut genetic diversity is still to be investigated by studying populations derived from crosses between these groups. Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Genetic analysis of the slow-melting flesh character in peach

The slow-melting flesh (SMF) trait in peach [Prunus persica (L.) Batsch] defines a slower process of postharvest fruit-softening than the prevalent melting flesh (MF) types. This gives a longer shelf life and a delayed harvest-time resulting in better fruit quality. Unlike other known fruit texture traits, SMF is difficult to measure and has a complex inheritance. We examined this character over 2 years in the offspring of two crosses, both with “Big Top,” an SMF nectarine, as the female parent, and with a melting flesh (MF) nectarine as the male parent (“Armking” and “Nectaross”). Following harvest, a texturometer was used to provide a textural profile analysis, and fruit firmness evolution was measured with a penetrometer over a period of 5 days’ storage at 20 °C. Linkage maps were constructed with a high-density SNP chip, and a phenotype-genotype analysis allowed the detection of three independent genomic regions where most QTLs (quantitative trait loci) were located. Two of these, on linkage groups 4 and 5, explained the variability for two characters—maturity date and firmness loss—that is, the QTL on linkage group 4 found in the MF parents and that on linkage group 5 in Big Top. A third region on linkage group 6, which identified a QTL for maturity date only in Armking, has no apparent association to the softening process. The relationship between maturity date and fruit-firmness loss and a hypothesis on the inheritance of the SMF character are discussed.     

Identification and stability of QTLs for fruit quality traits in apple

Breeding for fruit quality traits is complex due to the polygenic (quantitative) nature of the genetic control of these traits. Therefore, to improve the speed and efficiency of genotype selection, attention in recent years has focused on the identification of quantitative trait loci (QTLs) and molecular markers associated with these QTLs. However, despite the huge potential of molecular markers in breeding programmes, their implementation in practice has been limited by the lack of information on the stability of QTLs across different environments and within different genetic backgrounds. Here, we present the results from a comprehensive analysis of the inheritance of fruit quality traits within a population derived from a cross between the apple cultivars ‘Telamon’ and ‘Braeburn’ over two successive seasons. A total of 74 different QTLs were identified for all the major fruit physiological traits including fruit height, diameter, weight and stiffness, flesh firmness, rate of flesh browning, acidity, the oBrix content and harvest date. Seventeen of these QTLs were ‘major’ QTLs, accounting for over 20% of the observed population variance of the trait. However, only one third (26) of the identified QTLs were stable over both harvest years, and of these year-stable QTLs only one was a major QTL. A direct comparison with published QTL results obtained using other populations (King et al., Theor Appl Genet 102:1227–1235, 2001; Liebhard et al., Plant Mol Biol 52:511–526, 2003) is difficult because the linkage maps do not share a sufficient number of common markers and due to differences in the trait evaluation protocols. Nonetheless, our results suggest that for the six fruit quality traits which were measured in all populations, nine out of a total of 45 QTLs were common or stable across all population × environments combinations. These results are discussed in the framework of the development and application of molecular markers for fruit quality trait improvement.     

Genetic mapping of QTLs affecting productivity and plant architecture in a full-sib cross from non-inbred parents in Cassava (<Emphasis Type="Italic">Manihot esculenta</Emphasis> Crantz)

An attempt was made to identify quantitative trait loci (QTLs) for several productivity and plant architecture traits in a full-sib progeny of 144 individuals from two non-inbred parents in cassava. A molecular linkage map of this cross constructed previously with over 250 markers was the source of molecular markers. The progeny were grown under field conditions at two locations (Palmira and Quilichao) in Colombia and evaluated in 2 years (1998 and 1999) for architecture and productivity traits. Architecture traits evaluated were plant height (PH), branching height (BH), branching levels (BL), branching index (BI), stem portion with leaves (SPL) and leaf area index (LAI). Productivity traits were those related to total dry matter production and distribution, namely fresh root yield (FRY), fresh shoot yield (FSY), harvest index (HI) and the number of storage roots (NR). Phenotypic evaluation of the traits in this population revealed continuous variation for all traits. Broad-sense heritability estimates, ranged from 36% (for NR) to 94% (for BH). Several significant phenotypic correlations were observed between architecture and productivity traits. Primary QTLs, using the single-QTL model, and secondary QTLs, by a primary QTL interaction model, were detected by interval mapping. A total of 30 primary QTLs and 84 secondary QTLs were detected. We identified 35% of detected QTLs in two or more trials, the other QTLs were environment-specific. These results underscore the significant genotype × environment interactions found for most of the traits. Several genomic segments affecting multiple traits were identified and were in agreement with correlation among traits. All QTLs identified for FRY were found associated with either component traits of productivity or architecture traits. This study suggests that QTLs for plant architecture can be used to improve productivity. However an exhaustive search and analysis of QTLs controlling architecture is required before marker-assisted selection (MAS) for increasing productivity can be initiated.Communicated by H. C. Becker     

Mapping of quantitative trait loci (QTLs) for oil yield using SSRs and gene-based markers in African oil palm (<Emphasis Type="Italic">Elaeis guineensis</Emphasis> Jacq.)

The identification of quantitative trait loci (QTLs) based on anchor markers, especially candidate genes that control a trait of interest, has been noted to increase the power of QTL detection. Since these markers can be scored as co-dominant data, they are also valuable for comparing and integrating the QTL linkage maps from diverse mapping populations. To estimate the position and effects of QTLs linked to oil yield traits in African oil palm, co-dominant microsatellites (SSR) and candidate gene-based sequence polymorphisms were applied to construct a linkage map for a progeny showing large differences in oil yield components. The progeny was genotyped for 97 SSR markers, 93 gene-linked markers, and 12 non-gene-linked SNP markers. From these, 190 segregating loci could be arranged into 31 linkage groups while 12 markers remained unmapped. Using the single marker linkage, interval mapping and multiple QTL methods, 16 putative QTLs on seven linkage groups affecting important oil yield related traits such as fresh fruit bunch yield (FFB), ratio of oil per fruit (OF), oil per bunch (OB), fruit per bunch (FB) and wet mesocarp per fruit (WMF) could be identified in the segregating population with estimated values for explained variance ranging from 12.4 % to 54.5 %. Markers designed from some candidate genes involved in lipid biosynthesis were found to be mapped near significant QTLs for various economic yield traits. Associations between QTLs and potential candidate genes are discussed.     

Quantitative trait loci for yield components in oil palm (Elaeis guineensis Jacq.)

The development of an oil palm RFLP marker map has enabled marker-based QTL mapping studies to be undertaken. Information from 153 RFLP markers was used in combination with phenotypic data from an F₂ population to estimate the position and effects of quantitative trait loci (QTLs) for traits including yield of fruit and its components and measures of vegetative growth. The mapping population consisted of 84 palms segregating for the major gene influencing shell thickness. Marker data were analysed to produce a linkage map consisting of 22 linkage groups. The QTL mapping analysis was carried out by interval mapping and single-marker analysis for the unlinked markers; significance thresholds were generated by permutation. Using both single-marker and interval-mapping analysis significant marker associated QTL effects were found for 11 of the 13 traits analysed. The results of interval-mapping analysis of fruit weight, petiole cross section and rachis length, and ratios of shell:fruit, mesocarp:fruit and kernel:fruit indicated significant (P<0.05) QTLs at the genome-wide threshold. The putative QTLs were associated with between 8.2% and 44.0% of the phenotypic variation, with an average of 27% for the single-marker analysis and 19% for the interval-mapping analysis. The higher percentage of phenotypic variation explained in the single-marker analysis, when compared to the interval-mapping analysis, is likely to be due to the lower stringency associated with the single-marker analysis. Large dominance deviations were associated with a sizeable proportion of the putative QTLs. The ultimate objective of mapping QTLs in commercial populations is to utilise novel breeding strategies such as marker-assisted selection (MAS). The potential impact of MAS in oil palm breeding programmes is discussed. Received: 26 June 2000 / Accepted: 24 October 2000     

Identification of quantitative trait loci involved in fruit quality traits in melon (<Emphasis Type="Italic">Cucumis melo</Emphasis> L.)

Two populations [an F₂ and a set of 77 double haploid lines (DHLs)] developed from a cross between a Piel de Sapo cultivar (PS) and the exotic Korean accession PI 161375 were used to detect QTLs involved in melon fruit quality traits: earliness (EA), fruit shape (FS), fruit weight (FW) and sugar content (SSC); and loci involved in the colour traits: external colour (ECOL) and flesh colour (FC). High variation was found, showing transgressive segregations for all traits. The highest correlation among experiments was observed for FS and the lowest for FW and SSC. Correlations among traits within experiments were, in general, not significant. QTL analysis, performed by Composite Interval Mapping, allowed the detection of nine QTLs for EA, eight for FS, six for FW and five for SSC. Major QTLs (R ²>25%) were detected for all traits. QTLs for different traits were no clearly co-localised, suggesting low pleiotropic effects at QTLs. Sixty-one per cent of them were detected in two or more experiments. QTLs for FS were detected in more trials than QTLs for FW and SSC, confirming that FS is under highly hereditable polygenic control. ECOL segregated as yellow:green in both experimental populations. The genetic control of ECOL was found to be complex, probably involving more than two loci with epistatic interactions. One of these loci was mapped on linkage group 9, but the other loci could not be clearly resolved. FC segregated as white:green:orange. The locus responsible for the green FC was mapped on linkage group 1, and it was proposed to correspond to the previously described locus gf. The genetic control of orange FC was complex: two loci in linkage groups 2 and 12 were associated with orange flesh, but larger population sizes would be necessary to elucidate completely the genetic control of orange flesh in this cross. Exotic alleles from PI161375 showed beneficial effects on EA, FW and SSC, indicating the usefulness of PI 161375 as a new source of genetic variability to improve European and American cultivars.Communicated by H.F. Linskens     

Sex-specific genetic architecture of human fatness in Chinese: the SAPPHIRe Study

To dissect the genetic architecture of sexual dimorphism in obesity-related traits, we evaluated the sex–genotype interaction, sex-specific heritability and genome-wide linkages for seven measurements related to obesity. A total of 1,365 non-diabetic Chinese subjects from the family study of the Stanford Asia–Pacific Program of Hypertension and Insulin Resistance were used to search for quantitative trait loci (QTLs) responsible for the obesity-related traits. Pleiotropy and co-incidence effects from the QTLs were also examined using the bivariate linkage approach. We found that sex-specific differences in heritability and the genotype–sex interaction effects were substantially significant for most of these traits. Several QTLs with strong linkage evidence were identified after incorporating genotype by sex (G × S) interactions into the linkage mapping, including one QTL for hip circumference [maximum LOD score (MLS) = 4.22, empirical p = 0.000033] and two QTLs: for BMI on chromosome 12q with MLS 3.37 (empirical p = 0.0043) and 3.10 (empirical p = 0.0054). Sex-specific analyses demonstrated that these linkage signals all resulted from females rather than males. Most of these QTLs for obesity-related traits replicated the findings in other ethnic groups. Bivariate linkage analyses showed several obesity traits were influenced by a common set of QTLs. All regions with linkage signals were observed in one gender, but not in the whole sample, suggesting the genetic architecture of obesity-related traits does differ by gender. These findings are useful for further identification of the liability genes for these phenotypes through candidate genes or genome-wide association analysis.     

茄子分子遗传图谱的构建及果实性状的QTL定位

以茄子(Solanum melongena)材料09-101-M和10TL-102-F4-1的重组自交系(RIL)为作图群体,构建总长度为991.7c M、共包含16个连锁群157个位点、平均图距为6.32 c M的遗传图谱。应用复合区间作图法(CIM),共检测到18个与茄子果实性状相关的QTLs,其中10个为主效QTLs,8个QTLs在两年两点的实验中能够被重复检测到。在所有QTLs中,控制果重的QTL fw1.1的效应值最大,为23.8%–31.6%,被定位在LG01(E09)上E25M34–E33M57b区域内;果长、果径与果重显著相关,且控制果长、果径与果重的QTL位于同一连锁群的相同区域。     

利用双单倍体群体剖析水稻产量及其相关性状的遗传基础

主效QTL、上位性效应和它们与环境的互作(QE)都是数量性状的重要遗传因素。利用籼粳交珍汕97／武育粳2号F1植株上的花药进行组织培养得到的190个双单倍体群体和179个微卫星标记,通过两年两重复田间试验,采用混合线性模型方法分析了9个控制水稻产量及其相关性状的遗传效应,得到57个主效QTL,41对上位性互作,8对QTL与环境的互作和7对上位性效应与环境的互作。单个主效QTL解释这些性状1．3％～25．8％的表型方差。各性状QTL的累积表型贡献率达11．5％～66．8％。大多数性状之间具有显著的表型相关性,相关性较高的性状之间常具有较多共同或紧密连锁的QTL。结果表明,基因的多效性或紧密连锁可能是性状相关的重要遗传基础。     

Genetic analysis of organoleptic quality in fresh market tomato. 1. Mapping QTLs for physical and chemical traits

Improving organoleptic quality is an important but complex goal for fresh market tomato breeders. A total of 26 traits involved in organoleptic quality variation were evaluated, in order to understand the genetic control of this characteristic. A recombinant inbred line (RIL) population was developed from an intraspecific cross between a cherry tomato line with a good overall aroma intensity and an inbred line with a common taste but with bigger fruits. Physical traits included fruit weight, diameter, color (L,a,b), firmness and elasticity. Chemical traits were dry matter weight, titratable acidity, pH, and the contents of soluble solids, sugars, lycopene, carotene and 12 aroma volatiles. RILs showed a large range of variation for most of the traits and many of them were transgressive. Some correlations between aroma volatiles were in accordance with the metabolic pathway they originated from. A total of 81 significant QTLs were detected for the 26 traits by simple and composite interval mapping. They were mainly distributed in a few regions on chromosomes 2, 3, 4, 8, 9, 11 and 12. Major QTLs (R²>30%) were detected for fruit weight, diameter, and color, and for six aroma volatiles. Co-localization of QTLs controlling correlated traits was mainly found on chromosome 2. QTLs for fruit weight and sugar content or dry matter weight were often co-localized. However, a QTL for soluble-solids content and dry matter weight have been detected on chromosome 9 in a region without fruit weight QTLs. QTLs for seven aroma volatiles, lycopene content and fruit color were also co-localized. The QTL localizations were compared with those detected in crosses between Lycopersicon esculentum and wild tomato species. Received: 19 January 2000 / Accepted: 26 May 2000     

Quantitative trait loci and underlying candidate genes controlling agronomical and fruit quality traits in octoploid strawberry <Emphasis Type="Italic">(Fragaria</Emphasis> × <Emphasis Type="Italic">ananassa</Emphasis>)

Breeding for fruit quality traits in strawberry (Fragaria × ananassa, 2n = 8x = 56) is complex due to the polygenic nature of these traits and the octoploid constitution of this species. In order to improve the efficiency of genotype selection, the identification of quantitative trait loci (QTL) and associated molecular markers will constitute a valuable tool for breeding programs. However, the implementation of these markers in breeding programs depends upon the complexity and stability of QTLs across different environments. In this work, the genetic control of 17 agronomical and fruit quality traits was investigated in strawberry using a F₁ population derived from an intraspecific cross between two contrasting selection lines, ‘232’ and ‘1392’. QTL analyses were performed over three successive years based on the separate parental linkage maps and a pseudo-testcross strategy. The integrated strawberry genetic map consists of 338 molecular markers covering 37 linkage groups, thus exceeding the 28 chromosomes. 33 QTLs were identified for 14 of the 17 studied traits and approximately 37% of them were stable over time. For each trait, 1–5 QTLs were identified with individual effects ranging between 9.2 and 30.5% of the phenotypic variation, indicating that all analysed traits are complex and quantitatively inherited. Many QTLs controlling correlated traits were co-located in homoeology group V, indicating linkage or pleiotropic effects of loci. Candidate genes for several QTLs controlling yield, anthocyanins, firmness and l-ascorbic acid are proposed based on both their co-localization and predicted function. We also report conserved QTLs among strawberry and other Rosaceae based on their syntenic location.     

Fine mapping of QTLs of chromosome 2 affecting the fruit architecture and composition of tomato

Negative correlations between quality traits and fruit size may hamper the breeding of fresh market tomato varieties for better organoleptic qualities. In a recent QTL analysis, QTLs with large effects on fruit weight, locule number and several quality traits were detected in the distal 50 cM of chromosome 2, but favorable alleles for fruit weight and locule number were unfavorable to quality traits. Substitution mapping was undertaken to determine whether the effects were due to a single QTL or to several tightly linked QTLs. Several chromosomal segments were characterized using near-isogenic lines. Five of them appeared to be involved in one or several traits. Considering the five segments from the top to the bottom of the region, the QTLs detected in each segment controlled the variation of: (1) fruit weight, (2) soluble solids content and dry matter weight, (3) fruit weight, (4) locule number and (5) fruit weight, dry matter weight, total sugars, titratable acidity and soluble solids content. This last cluster illustrates an antagonism between fruit weight and four quality traits, as favorable alleles are not conferred by the same parent in both cases. Nevertheless, several antagonistic QTLs were separated from each other in the first four segments, holding the promise for marker-assisted improvement of fruit quality traits without compromising the fruit size.     

Mapping Architectural, Phenological, and Fruit Quality QTLs in Apricot

The F1 population “Harostar”?×?“Rouge de Mauves” was used to perform a quantitative trait loci (QTL) mapping for tree architecture traits (i.e., tree diameter, total branch number, tree shape); time to initial reproductive bud break; and fruit quality traits (i.e., ground color, fruit form, soluble solid content) using data collected from 2010 to 2012. For architectural traits, interval mapping detected QTLs only in “Rouge de Mauves” on linkage group 1 for trunk diameter in 2010, on LG6 for total branch number in 2010, and on LG1 and LG5 for tree shape for both years 2010 and 2011 combined. QTLs were detected only in “Harostar” for time to initial reproductive bud break on LG1 and LG4 in 2011. For fruit quality traits, data were collected in 2011 and 2012. QTLs were identified on LG1 in 2011 and on LG4 in 2012 for soluble solid content, on LG3 for ground color in both years, on LG7 only in 2011, and on LG3 for fruit form in both years. The QTLs that we identified were compared to those previously identified in other Prunus spp.