Genetic and physiological architecture of early vigor in Aegilops tauschii, the D-genome donor of hexaploid wheat. A quantitative trait loci analysis

Plant growth can be studied at different organizational levels, varying from cell, leaf, and shoot to the whole plant. The early growth of seedlings is important for the plant's establishment and its eventual success. Wheat (Triticum aestivum, genome AABBDD) seedlings exhibit a low early growth rate or early vigor. The germplasm of wheat is limited. Wild relatives constitute a source of genetic variation. We explored the physiological and genetic relationships among a range of early vigor traits in Aegilops tauschii, the D-genome donor. A genetic map was constructed with amplified fragment-length polymorphism and simple sequence repeat markers, and quantitative trait loci (QTL) analysis was performed on the F(4) population of recombinant inbred lines derived from a cross between contrasting accessions. The genetic map consisted of 10 linkage groups, which were assigned to the seven chromosomes and covered 68% of the D genome. QTL analysis revealed 87 mapped QTLs (log of the odds >2.65) in clusters, 3.1 QTLs per trait, explaining 32% of the phenotypic variance. Chromosomes 1D, 4D, and 7D harbored QTLs for relative growth rate, biomass allocation, specific leaf area, leaf area ratio, and unit leaf rate. Chromosome 2D covered QTLs for rate and duration of leaf elongation, cell production rate, and cell length. Chromosome 5D harbored QTLs for the total leaf mass and area and growth rate of the number of leaves and tillers. The results show that several physiological correlations between growth traits have a genetic basis. Genetic links between traits are not absolute, opening perspectives for identification of favorable alleles in A. tauschii to improve early vigor in wheat.     

Genetic linkage map construction and QTL identification of juvenile growth traits in Torreya grandis

Mei, Prunus mume Sieb. et Zucc., is an ornamental plant popular in East Asia and, as an important member of genus Prunus, has played a pivotal role in systematic studies of the Rosaceae. However, the genetic architecture of botanical traits in this species remains elusive. This paper represents the first genome-wide mapping study of quantitative trait loci (QTLs) that affect stem growth and form, leaf morphology and leaf anatomy in an intraspecific cross derived from two different mei cultivars. Genetic mapping based on a high-density linkage map constricted from 120 SSRs and 1,484 SNPs led to the detection of multiple QTLs for each trait, some of which exert pleiotropic effects on correlative traits. Each QTL explains 3-12% of the phenotypic variance. Several leaf size traits were found to share common QTLs, whereas growth-related traits and plant form traits might be controlled by a different set of QTLs. Our findings provide unique insights into the genetic control of tree growth and architecture in mei and help to develop an efficient breeding program for selecting superior mei cultivars.     

Molecular genetics of growth and development in Populus (Salicaceae). v. mapping quantitative trait loci affecting leaf variation

We examined the genetic variation of leaf morphology and development in the 2-yr-old replicated plantation of an interspecific hybrid pedigree of Populus trichocarpa T. & G. and P. deltoides Marsh. via both molecular and quantitative genetic methods. Leaf traits chosen were those that show pronounced differences between the original parents, including leaf size, shape, orientation, color, structure, petiole size, and petiole cross section. Leaves were sampled from the current terminal, proleptic, and sylleptic branches. In the F2 generation, leaf traits were all significantly different among genotypes, but with significant effects due to genotype X crown-position interaction. Variation in leaf pigmentation, petiole length. And petiole length proportion appeared to be under the control of few quantitative trait loci (QTLs). More QTLs were associated with single leaf area, leaf shape, lamina angle, abaxial color, and petiole flatness, and in these traits the number of QTLs varied among crown positions. In general, the estimates of QTL numbers from Wright's biometric method were close to those derived from molecular markers. For those traits with few underlying QTLs, a single marker interval could explain from 30 to 60% of the observed phenotypic variance. For multigenic traits, certain markers contributed more substantially to the observed variation than others. Genetic cluster analysis showed developmentally related traits to be more strongly associated with each other than with unrelated traits. This finding was also supported by the QTL mapping. For example, the same chromosomal segment of linkage group L seemed to account for 20% of the phenotypic variation of all dimension-related traits, leaf size, petiole length. and midrib angle. In both traits. the P. deltoides alleles had positive effects and were dominant to the P. trichocarpa alleles. Similar relationships were also found for lamina angle. abaxial greenness, and petiole.     

Gene actions of QTLs affecting several agronomic traits resolved in a recombinant inbred rice population and two testcross populations

To understand the types of gene action controlling seven quantitative traits in rice, QTL mapping was performed to dissect the main effect (M-QTLs) and digenic epistatic (E-QTLs) QTLs responsible for the trait performance of 254 recombinant inbred lines (RILs) of "Lemont/Teqing", and two testcross (TC) F(1) populations derived from these RILs. The correlation analyses reveal a general pattern, i.e. trait heritability in the RILs was negatively correlated to trait heterosis in the TC hybrids. A large number of M-QTLs and E-QTLs affecting seven traits, including heading date (HD), plant height (PH), flag leaf length (FLL), flag leaf width (FLW), panicle length (PL), spikelet number per panicle (SN) and spikelet fertility (SF), were identified and could be classified into two predominant groups, additive QTLs detected primarily in the RILs, and overdominant QTLs identified exclusively in the TC populations. There is little overlap between QTLs identified in the RILs and in the TC populations. This result implied that additive gene action is largely independent from non-additive gene action in the genetic control of quantitative traits of rice. The detected E-QTLs collectively explained a much greater portion of the total phenotypic variation than the M-QTLs, supporting prior findings that epistasis has played an important role in the genetic control of quantitative traits in rice. The implications of these results to the development of inbred and hybrid cultivars were discussed.     

Genetic mapping of QTLs affecting tree growth and architecture in Populus: implication for ideotype breeding

 A segregated F₂ progeny derived from two highly divergent poplar species, Populus trichocarpa and P. deltoides, was used to evaluate the genetic basis of canopy structure and function in a clonally replicated plantation. The QTLs of large effect on growth, branch, and leaf traits were identified using the Populus linkage map constructed by 343 molecular markers. Stem height and harvest index appeared to be under the control of few QTLs with major effects, whereas variation in stem basal area, volume, and dry weight might be due to many more QTLs. Branch and leaf traits on sylleptics tended to include more QTLs with major effects than those on proleptics. In the environment where the pedigree was tested, sylleptics were very frequent in the P. trichocarpa parent but rare in the P. deltoides parent. For sylleptic traits for which two or more QTLs were identified, however, increases in the trait values were conditioned not only by the P. trichocarpa alleles, but also by the P. deltoides alleles. Similar findings were found for traits on proleptics that were differently expressed between the two parents. For both sylleptic and proleptic branch types, dominance (ranging from partial to over) was observed. The QTLs on specific linkage groups were found to be responsible for relationships between stem growth and its developmental components. Similar QTL clustering was also observed for morphological or developmental integration in poplar, i.e., traits with similar developmental origins are more strongly correlated with one another than traits with different developmental origins. The implications of these molecular genetic results for ideotype breeding of poplars are discussed. Received: 15 July 1997/Accepted: 19 August 1997     

Genome dissection of traits related to domestication in azuki bean (Vigna angularis) and comparison with other warm-season legumes

BACKGROUND: The objective of this study was to dissect into quantitative trait loci (QTLs) the large morphological and physiological differences between cultivated azuki bean (Vigna angularis) and a wild relative and to infer the commonalities of the QTLs for domestication-related traits across the Asian Vigna and with other warm-season legumes. METHODS: Two linkage maps, for the BC(1)F(1) and F(2) populations, respectively, from the same cross between azuki bean and V. nepalensis were developed. Using these linkage maps QTLs for 33 domestication-related traits were analysed and mapped. The location of mapped QTLs was compared with locations of similar QTLs in other warm-season legumes. KEY RESULTS: QTLs were detected for seed-, pod-, stem- and leaf-related traits. Most traits were controlled by between two and nine QTLs but several traits, such as pod dehiscence, were controlled by single genes. QTLs for domestication-related traits were restricted to particular regions of the azuki bean genome, especially linkage groups 1, 2, 4, 7 and 9. Linkage groups 1 and 2 had QTLs for a suite of traits including pod size, germination, seed size and lower stem length. QTLs on linkage groups 7 and 9 were associated with upper stem length, maximum leaf size and pod and seed size. Pleiotropy or close linkage of genes for domestication-related traits is suggested in these regions. While some QTLs are common to azuki bean and other warm-season legumes, many are recorded for the first time in azuki bean. CONCLUSIONS: QTLs for a large number of domestication-related traits have been mapped for the first time in azuki bean. QTLs with unexpected effect and new QTLs for traits such as seed size have been found. The results provide a foundation that will be useful for improvement of azuki bean and related legumes.     

Mapping QTLs for root morphological traits inBrassica rapa L. based on AFLP and RAPD markers

Root growth and thickening plays a key role in the final productivity and even the quality of storage roots in root crops. This study was conducted to identify and map quantitative trait loci (QTLs) affecting root morphological traits in Brassica rapa by using molecular markers. An F2 population was developed from a cross between Chinese cabbage (Brassica rapa ssp. chinensis) and turnip (B. rapa ssp. rapifera), which differed greatly in root characters. A genetic map covering 1837.1 cM, with 192 marker loci and 11 linkage groups, was constructed by using this F2 population. The F3 families derived from F2 plants were grown in the field and evaluated for taproot traits (thickness, length, and weight). QTL analysis via simple interval mapping detected 18 QTLs for the 3 root traits, including 7 QTLs for taproot thickness, 5 QTLs for taproot length, and 6 QTLs for taproot weight. Individually, the QTLs accounted for 8.4-27.4% of the phenotypic variation. The 2 major QTLs, qTRT4b for taproot thickness and qTRW4 for taproot weight, explained 27.4% and 24.8% of the total phenotypic variance, respectively. The QTLs for root traits, firstly detected in Brassica crops, may provide a basis for marker-assisted selection to improve productivity in root-crop breeding.     

Molecular mapping of quantitative trait loci for drought tolerance in maize plants

Drought tolerance is one of the most important but complex traits of crops. We looked for quantitative trait loci (QTLs) that affect drought tolerance in maize. Two maize inbreds and their advanced lines were evaluated for drought-related traits. A genetic linkage map developed using RFLP markers was used to identify QTLs associated with drought-related traits. Twenty-two QTLs were detected, with a minimum of one and a maximum of nine for drought-related traits. A single-QTL was detected for sugar concentration accounting for about 52.2% of the phenotypic variation on chromosome 6. A single-QTL was also identified for each of the traits root density, root dry weight, total biomass, relative water content, and leaf abscisic acid content, on chromosomes 1 and 7, contributing to 24, 0.2, 0.4, 7, and 19% of the phenotypic variance, respectively. Three QTLs were identified for grain yield on chromosomes 1, 5, and 9, explaining 75% of the observed phenotypic variability, whereas four QTLs were detected for osmotic potential on chromosomes 1, 3, and 9, together accounting for 50% of the phenotypic variance. Nine QTLs were detected for leaf surface area on chromosomes 3 and 9, with various degrees of phenotypic variance, ranging from 25.8 to 42.2%. Four major clusters of QTLs were identified on chromosomes 1, 3, 7, and 9. A QTL for yield on chromosome 1 was found co-locating with the QTLs for root traits, total biomass, and osmotic potential in a region of about 15 cM. A cluster of QTLs for leaf surface area were coincident with a QTL for osmotic potential on chromosome 3. The QTLs for leaf area also clustered on chromosome 9, whereas QTLs for leaf abscisic acid content and relative water content coincided on chromosome 7, 10 cM apart. Co-location of QTLs for different traits indicates potential pleiotropism or tight linkage, which may be useful for indirect selection in maize improvement for drought tolerance.     

Quantitative trait loci for resistance to herbivores in willow: field experiments with varying soils and climates

As a basis for genetic improvement of willow (Salix spp.) for use in wood biomass production, quantitative trait loci (QTLs) responsible for resistance to herbivores have been identified in a tetraploid hybrid F₂ population originating from a cross between Salix dasyclados (Wimm.) and Salix viminalis (L.) (Salicaceae). Symptoms of herbivory, caused by various insects and game, and, in addition, leaf rust, were assessed in three field locations with varying soils and climates. Eleven damage traits (lost leaf area, leaf discoloration, leaf blisters, leaf‐mite symptoms, leaf‐margin cuts, and various estimates of shoot‐tip damage by a gall midge, game, and lepidopterans) were submitted to QTL analysis. A composite interval mapping approach was used to estimate the number of QTLs, the magnitude of the QTLs, and their position on genetic linkage maps. Most of the identified QTLs were specific for each trait and location, but a few QTLs common across the locations were also detected. Each QTL explained between 8 and 24% of the phenotypic variation, depending on damage trait and field location. Clusters of QTLs for different traits were found at several linkage groups, indicating either a common genetic base or tightly linked QTL. Our results emphasize the need for verification of QTL studies over different environments.     

Phenotype/genotype associations for yield and salt tolerance in a barley mapping population segregating for two dwarfing genes

Barley traits related to salt tolerance are mapped in a population segregating for a dwarfing gene associated with salt tolerance. Twelve quantitative trait loci (QTLs) were detected for seven seedling traits in doubled haploids from the spring barley cross Derkado x B83-12/21/5 when given saline treatment in hydroponics. The location of QTLs for seedling growth stage (leaf appearance rate), stem weight prior to elongation, and tiller number are reported for the first time. In addition, four QTLs were found for the mature plant traits grain nitrogen and plot yield. In total, seven QTLs are co-located with the dwarfing genes sdw1, on chromosome 3H, and ari-e.GP, on chromosome 5H, including seedling leaf response (SGa) to gibberellic acid (GA(3)). QTLs controlling the growth of leaves (GS2) on chromosomes 2H and 3H and emergence of tillers (TN2) and grain yield were independent of the dwarfing genes. Field trials were grown in eastern Scotland and England to estimate yield and grain composition. A genetic map was used to compare the positions of QTLs for seedling traits with the location of QTLs for the mature plant traits. The results are discussed in relation to the study of barley physiology and the location of genes for dwarf habit and responses to GA.     

Quantitative resistance against Bemisia tabaci in Solanum pennellii: Genetics and metabolomics

The whitefly Bemisia tabaci is a serious threat in tomato cultivation worldwide as all varieties grown today are highly susceptible to this devastating herbivorous insect.Many accessions of the tomato wild relative Solanum pennellii show a high resistance towards B. tabaci. A mapping approach was used to elucidate the genetic background of whiteflyresistance related traits and associated biochemical traits in this species. Minor quantitative trait loci(QTLs) for whitefly adult survival(AS) and oviposition rate(OR) were identified and some were confirmed in an F2BC1 population, where they showed increased percentages of explained variance(more than 30%). Bulked segregant analyses on pools of whiteflyresistant and-susceptible F2 plants enabled the identification of metabolites that correlate either with resistance or susceptibility. Genetic mapping of these metabolites showed that a large number of them co-localize with whiteflyresistance QTLs. Some of these whitefly-resistance QTLs are hotspots for metabolite QTLs. Although a large number of metabolite QTLs correlated to whitefly resistance or susceptibility, most of them are yet unknown compounds and further studies are needed to identify the metabolic pathways and genes involved. The results indicate a direct genetic correlation between biochemical-based resistance characteristics and reduced whitefly incidence in S. pennellii.     

A genetic analysis of seed and berry weight in grapevine.

Fruit size and seedlessness are highly relevant traits in many fruit crop species, and both are primary targets of breeding programs for table grapes. In this work we performed a quantitative genetic analysis of size and seedlessness in an F1 segregating population derived from the cross between a classical seeded (Vitis vinifera L. 'Dominga') and a newly bred seedless ('Autumn Seedless') cultivar. Fruit size was scored as berry weight (BW), and for seedlessness we considered both seed fresh weight (SFW) and the number of seeds and seed traces (SN) per berry. Quantitative trait loci (QTL) analysis of BW detected 3 QTLs affecting this trait and accounting for up to 67% of the total phenotypic variance. QTL analysis for seedlessness detected 3 QTLs affecting SN (explaining up to 35% of total variance) and 6 affecting SFW (explaining up to 90% of total variance). Among them, a major effect QTL explained almost half of the phenotypic variation for SFW. Comparative analysis of QTLs for these traits reduced the number of grapevine genomic regions involved, one of them being a major effect QTL for seedlessness. Association analyses showed that microsatellite locus VMC7F2, closely linked to this QTL, is a useful marker for selection of seedlessnes.     

Quantitative trait loci affecting growth-related traits in wild barley (Hordeum spontaneum) grown under different levels of nutrient supply

The genetic basis of phenotypic plasticity of relative growth rate (RGR), its components and associated morphological traits was studied in relation to nutrient limitation. In all, 140 F(3) lines from a cross, made between two Hordeum spontaneum (wild barley) accessions sampled in Israel, were subjected to growth analysis under two nutrient levels. Quantitative trait loci (QTLs) were detected for RGR and three of its components, leaf area ratio (LAR), specific leaf area and leaf mass fraction (LMF). Indications for close linkage (potential pleiotropy) were found, for example, for LAR and LMF. An interesting case is on chromosome 6, at which QTLs for RGR and seed mass were detected in the same region. These QTLs had opposite additive effects, supporting earlier results that plants growing from lighter seeds had a higher RGR. Only two QTLs were significant under both nutrient conditions, suggesting large QTL x environment interactions for most traits. For 21 out of 26 QTLs, however, the additive genetic effect was of identical sign in both nutrient environments, but reached the significance threshold in only one of them. Nevertheless, some QTLs detected in one of the two environments had virtually no effect in the other, and QTLs for plasticity were detected for RGR, LAR and LMF, as well as for some morphological traits. QTLs with opposite effects under high and low nutrients were not found. Thus, at the genetic level, there was no evidence for a trade-off between faster growth at high versus low nutrient levels.     

A dissection model for mapping complex traits

Many quantitative traits are composites of other traits that contribute differentially to genetic variation. Quantitative trait locus (QTL) mapping of these composite traits can benefit by incorporating the mechanistic process of how their formation is mediated by the underlying components. We propose a dissection model by which to map these interconnected components traits under a joint likelihood setting. The model can test how a composite trait is determined by pleiotropic QTLs for its component traits or jointly by different sets of QTLs each responsible for a different component. The model can visualize the pattern of time‐varying genetic effects for individual components and their impacts on composite traits. The dissection model was used to map two composite traits, stemwood volume growth decomposed into its stem height, stem diameter and stem form components for Euramerican poplar adult trees, and total lateral root length constituted by its average lateral root length and lateral root number components for Euphrates poplar seedlings. We found the pattern of how QTLs for different components contribute to phenotypic variation in composite traits. The detailed understanding of the genetic machineries of composite traits will not only help in the design of molecular breeding in plants and animals, but also shed light on the evolutionary processes of quantitative traits under natural selection.     

Quantitative genetic analysis of flowering time in tomato.

Artificial selection of cultivated tomato (Solanum lycopersicum L.) has resulted in the generation of early-flowering, day-length-insensitive cultivars, despite its close relationship to other Solanum species that need more time and specific photoperiods to flower. To investigate the genetic mechanisms controlling flowering time in tomato and related species, we performed a quantitative trait locus (QTL) analysis for flowering time in an F2 mapping population derived from S. lycopersicum and its late-flowering wild relative S. chmielewskii. Flowering time was scored as the number of days from sowing to the opening of the first flower (days to flowering), and as the number of leaves under the first inflorescence (leaf number). QTL analyses detected 2 QTLs affecting days to flowering, which explained 55.3% of the total phenotypic variance, and 6 QTLs for leaf number, accounting for 66.7% of the corresponding phenotypic variance. Four of the leaf number QTLs had not previously been detected for this trait in tomato. Colocation of some QTLs with flowering-time genes included in the genetic map suggests PHYB2, FALSIFLORA, and a tomato FLC-like sequence as candidate genes that might have been targets of selection during the domestication of tomato.     

Linkage and Association Mapping for Two Major Traits Used in the Maritime Pine Breeding Program: Height Growth and Stem Straightness

BackgroundIncreasing our understanding of the genetic architecture of complex traits, through analyses of genotype-phenotype associations and of the genes/polymorphisms accounting for trait variation, is crucial, to improve the integration of molecular markers into forest tree breeding. In this study, two full-sib families and one breeding population of maritime pine were used to identify quantitative trait loci (QTLs) for height growth and stem straightness, through linkage analysis (LA) and linkage disequilibrium (LD) mapping approaches.ResultsThe populations used for LA consisted of two unrelated three-generation full-sib families (n = 197 and n = 477). These populations were assessed for height growth or stem straightness and genotyped for 248 and 217 markers, respectively. The population used for LD mapping consisted of 661 founders of the first and second generations of the breeding program. This population was phenotyped for the same traits and genotyped for 2,498 single-nucleotide polymorphism (SNP) markers corresponding to 1,652 gene loci. The gene-based reference genetic map of maritime pine was used to localize and compare the QTLs detected by the two approaches, for both traits. LA identified three QTLs for stem straightness and two QTLs for height growth. The LD study yielded seven significant associations (P ≤ 0.001): four for stem straightness and three for height growth. No colocalisation was found between QTLs identified by LA and SNPs detected by LD mapping for the same trait.ConclusionsThis study provides the first comparison of LA and LD mapping approaches in maritime pine, highlighting the complementary nature of these two approaches for deciphering the genetic architecture of two mandatory traits of the breeding program.     

QTL analysis of intraspecific differences between two Silene vulgaris ecotypes

BACKGROUND AND AIMS: Serpentine soils provide a highly selective substrate for plant colonization and growth and represent an ideal system for studying the evolution of plant-ecotypes. In the present study the aim was to identify the genetic architecture of morphological traits distinguishing serpentine and non-serpentine ecotypes of Silene vulgaris. METHODS: Using an F(2) mapping population derived from an intraspecific cross between a serpentine and a non-serpentine ecotype of S. vulgaris, the genetic architecture of 12 morphological traits was explored using a quantitative trait locus (QTL) analysis. KEY RESULTS: The QTL analysis identified a total of 49 QTLs, of which 24 were classified as major QTLs. The mean number of QTLs per trait category was found to correspond well with numbers reported in the literature for similar crosses. Clustering of QTLs for different traits was found on several linkage groups. CONCLUSIONS: Morphological traits that differentiate the two ecotypes are strongly correlated, presumably as a consequence of the joint effects of extensive linkage of QTLs for different traits and directional selection. The signature of consistent directional selection was found for leaf and shoot trait divergence. Intraspecific ecotype differences in S. vulgaris were found to be distributed across the entire genome. The study shows that QTL analyses on non-model organisms can provide novel insights into the genetic basis of plant diversification.     

QTL mapping for fiber yield-related traits by constructing the first genetic linkage map in ramie (Boehmeria nivea L. Gaud)

Ramie fiber extracted from stem bast is one of the most important natural fibers. The fiber yield of ramie is a valuable trait and is decided by several components, including stem number per plant (SN), the fiber yield per stem (FYPS), stem length (SL), stem diameter (SD), and bark thickness (BT). All of these fiber yield-related traits are inherited in a quantitative manner. The genetic basis for these traits is still uncharacterized, which has hindered the improvement of yield traits through selective ramie breeding. In this study, an F₂ population derived from two ramie varieties, Zhongzhu 1 and Qingyezhuma, with striking differences in fiber yield-related traits, was used for cutting propagation and to develop an F₂ agamous line (FAL) population. A genetic linkage map with 132 DNA loci spanning 2,265.1 cM was first constructed. The analysis of quantitative trait locus (QTL) for fiber yield-related traits was performed in ramie for the first time. Finally, a total of 6, 9, 5, 7, and 6 QTLs for FYPS, SL, SN, SD, and BT, respectively, were identified in the FAL population in two environments. Among these 33 QTLs, 9 QTLs were detected in both environments and 24 QTLs exhibited overdominance. The overdominance of these QTLs possibly contributed to the heterosis of these yield-related traits in ramie. Moreover, there were 7 QTL clusters identified. The identification of the QTLs for fiber yield-related traits will be helpful for improving the fiber yield in ramie breeding programs.     

Molecular-Marker-Facilitated Investigations of Quantitative-Trait Loci in Maize. I. Numbers, Genomic Distribution and Types of Gene Action

Individual genetic factors which underlie variation in quantitative traits of maize were investigated in each of two F2 populations by examining the mean trait expressions of genotypic classes at each of 17-20 segregating marker loci. It was demonstrated that the trait expression of marker locus classes could be interpreted in terms of genetic behavior at linked quantitative trait loci (QTLs). For each of 82 traits evaluated, QTLs were detected and located to genomic sites. The numbers of detected factors varied according to trait, with the average trait significantly influenced by almost two-thirds of the marked genomic sites. Most of the detected associations between marker loci and quantitative traits were highly significant, and could have been detected with fewer than the 1800-1900 plants evaluated in each population. The cumulative, simple effects of marker-linked regions of the genome explained between 8 and 40% of the phenotypic variation for a subset of 25 traits evaluated. Single marker loci accounted for between 0.3% and 16% of the phenotypic variation of traits. Individual plant heterozygosity, as measured by marker loci, was significantly associated with variation in many traits. The apparent types of gene action at the QTLs varied both among traits and between loci for given traits, although overdominance appeared frequently, especially for yield-related traits. The prevalence of apparent overdominance may reflect the effects of multiple QTLs within individual marker-linked regions, a situation which would tend to result in overestimation of dominance. Digenic epistasis did not appear to be important in determining the expression of the quantitative traits evaluated. Examination of the effects of marked regions on the expression of pairs of traits suggests that genomic regions vary in the direction and magnitudes of their effects on trait correlations, perhaps providing a means of selecting to dissociate some correlated traits. Marker-facilitated investigations appear to provide a powerful means of examining aspects of the genetic control of quantitative traits. Modifications of the methods employed herein will allow examination of the stability of individual gene effects in varying genetic backgrounds and environments.     

Genetic architecture of leaf ecophysiological traits in Helianthus

We investigated quantitative trait loci (QTLs) for several leaf chemistry traits in early-generation hybrids between Helianthus annuus and Helianthus petiolaris, the parental species of the ancient diploid hybrid sunflower species Helianthus anomalus, Helianthus deserticola, and Helianthus paradoxus. We grew individuals of a second-generation backcross (BC(2)) toward H. petiolaris under optimum conditions in a glass house experiment. Trait values were measured once for each individual. In addition, genotypic data previously determined for each individual were employed for composite interval mapping of QTLs. We detected QTLs for leaf carbon concentration, leaf nitrogen concentration, leaf nitrogen per unit area, and photosynthetic nitrogen use efficiency. Leaf carbon isotope discrimination (delta(13)C) and leaf nitrogen isotopic composition (delta(15)N) were analyzed, but no significant QTLs were found for these traits. Interestingly, two neighboring loci explained a relatively large percentage of the variation in leaf nitrogen per unit area. This was notable because leaf nitrogen has been shown to strongly affect the fitness of early-generation sunflower hybrids in the H. anomalus habitat, and QTLs of large effect are expected to respond relatively quickly to selection. We speculate that the genetic architecture underlying leaf nitrogen may have facilitated the colonization of active desert sand dunes by H. anomalus.