Mapping quantitative trait loci controlling seed dormancy and heading date in rice, Oryza sativa L., using backcross inbred lines

 To detect quantitative trait loci (QTLs) controlling seed dormancy, 98 BC₁F₅ lines (backcross inbred lines) derived from a backcross of Nipponbare (japonica)/Kasalath (indica)//Nipponbare were analyzed genetically. We used 245 RFLP markers to construct a framework linkage map. Five putative QTLs affecting seed dormancy were detected on chromosomes 3, 5, 7 (two regions) and 8, respectively. Phenotypic variations explained by each QTL ranged from 6.7% to 22.5% and the five putative QTLs explained about 48% of the total phenotypic variation in the BC₁F₅ lines. Except for those of the QTLs on chromosome 8, the Nipponbare alleles increased the germination rate. Five putative QTLs controlling heading date were detected on chromosomes 2, 3, 4, 6 and 7, respectively. The phenotypic variation explained by each QTL for heading date ranged from 5.7% to 23.4% and the five putative QTLs explained about 52% of the total phenotypic variation. The Nipponbare alleles increased the number of days to heading, except for those of two QTLs on chromosomes 2 and 3. The map location of a putative QTL for heading date coincided with that of a major QTL for seed dormancy on chromosome 3, although two major heading-date QTLs did not coincide with any seed dormancy QTLs detected in this study. Received: 10 October 1997 / Accepted: 12 January 1998     

Genetic characterisation of seed yield and fertility traits in perennial ryegrass (<Emphasis Type="Italic">Lolium perenne</Emphasis> L.)

Seed yield is a trait of major interest for the key grassland species Lolium perenne L. An F2 mapping population of perennial ryegrass (VrnA), recently characterised for vernalisation response, was assessed in a glasshouse for traits related to seed yield based on a lattice design with four replications over 2 years. The traits heading date, plant height, length of panicles, number of panicles per plant, seed yield per panicle, flag leaf length, flag leaf width and seed yield per plant revealed repeatabilities ranging from 41 to 76% and a considerable amount of genetic variation in the VrnA population. Path analysis partitioned the direct and indirect effects of seed yield components on seed yield per plant. Seed yield per panicle showed the highest effect on total seed yield. The adjusted mean values of each trait and a genetic linkage map consisting of 97 anonymous and 85 gene associated DNA markers were used for quantitative trait loci (QTL) analysis. Of particular interest were two QTL on linkage group (LG) 1 and LG 2, explaining 41 and 18%, respectively, of the observed phenotypic variation for the trait seed yield per panicle. Both QTL co-located with two major QTL for total seed yield per plant possibly representing the S and Z loci of the gametophytic self incompatibility (SI) system of perennial ryegrass. The diversity of SI alleles in mapping parents and the degree of heterozygosity at SI loci in the full sib progeny determines the interference of self incompatibility with seed production.     

Genes for Dwarfness in Wheat, TRITICUM AESTIVUM L

The genetic control of plant height was studied in crosses of four spring wheats involving the standard height variety Ramona 50 and short-statured selections Olesen, D6301, and D6899. Data from parent, F₁, F₂, and F₃ populations indicated that four independently segregating loci account for most of the differences among the four varieties. Two major genes of a highly recessive nature condition reduced height in Olesen and the Norin 10 derivative D6301. Olesen also carries a third dwarfing gene which is partially dominant in its effects over genes for tallness. This gene, or a gene that acts in a similar manner, is also present in the standard height variety Ramona 50. Dwarfing in D6899, a derivative of Tom Thumb, is controlled primarily by a single gene with mainly additive effects which is not present in any of the other three varieties.

Genetic components estimated from generation means (parental, F₁, F₂, F₃, and backcross) indicated that additive gene effects were the major component of variation in four of the six crosses, and of similar magnitude to dominance effects in another cross. The primary source of genetic variation in the cross Olesen x D6899 was due to epistasis with both additive x additive and dominance x dominance effects of major importance. The results of the generation mean analyses were consistent with the models for major-gene control of plant height based on segregation patterns.

     

Pleiotropism of the Photoperiod-Insensitive Allele of Hd1 on Heading Date,Plant Height and Yield Traits in Rice

Five populations segregated in isogenic backgrounds and three sets of near isogenic lines (NILs) overlapping in a 362.3-kb region covering heading date gene Hd1 were developed from the indica rice cross Zhenshan97 (ZS97)/Milyang 46 (MY46). They were used to analyze the effects of Hd1 on heading date, plant height and yield traits. In a background of the parental mixtures, the photoperiod-sensitive allele derived from ZS97 functioned in promoting and delaying flowering in the natural short-day and long-day conditions, respectively. In the background of ZS97, no response to the photoperiod was observed, whereas the photoperiod-insensitive allele derived from MY46 functioned in delaying flowering, increasing plant height, and enhancing grain productivity. The additive effects estimated in two NIL sets were 6.14 and 6.14 d for heading date, 4.46 and 5.55 cm for plant height, 10.82 and 11.54 for the number of spikelets per panicle, 6.82 and 8.00 for the number of grains per panicle, and 2.16 and 2.23 g for grain yield per plant, which explained 94.1% and 96.3%, 70.5% and 84.8%, 52.4% and 55.2%, 28.9% and 39.2%, and 36.5% and 26.9% of the phenotypic variances, respectively. Since the photoperiod-insensitive allele of Hd1 confers a long vegetative phase, it is a good candidate for breeding rice varieties with high yielding potential for low latitudes.     

Detection of epistatic interactions between exotic alleles introgressed from wild barley (H. vulgare ssp. spontaneum)

The expression of a quantitative phenotype can be controlled through genotype, environment and genotype by environment interaction effects. Further, genotype effects can be attributed to major genes, quantitative trait loci (QTL) and gene by gene interactions, which are also termed epistatic interactions. The present study demonstrates that two-way epistatic interactions can play an important role for the expression of domestication-related traits like heading date, plant height and yield. In the BC₂DH population S42, carrying wild barley introgressions in the genetic background of the spring barley cultivar Scarlett, 13, 8 and 12 marker by marker interaction effects could be detected for the traits heading date, plant height and yield, respectively. Significant allelic combinations at interacting loci coincided for heading date, plant height and yield suggesting the presence of pleiotropic effects rather than several linked QTL. The mode of epistasis observed was primarily characterised by either (1) compensatory effects, where allelic combinations from the same genotype buffered the phenotype, or (2) augmented effects, where only the combination of the exotic allele at both interacting loci caused an altered phenotype. The present study shows that estimates of main effects of QTL can be confounded by interactions with background loci, suggesting that the identification of epistatic effects is important for gene cloning and marker-assisted selection. Furthermore, interaction effects between loci and putative candidate genes detected in the present study reveal potential functional relationships, which can be used to further elucidate gene networks in barley.     

Identification of quantitative trait loci and epistatic interactions for plant height and heading date in rice

Appropriate heading date and plant height are prerequisites for attaining the desired yield level in rice breeding programs. In this study, we analyzed the genetic bases of heading date and plant height at both single- locus and two-locus levels, using a population of 240 F_2:3 families derived from a cross between two elite rice lines. Measurements for the traits were obtained over 2 years in replicated field trials. A linkage map was constructed with 151 polymorphic marker loci, based on which interval mapping was performed using Mapmaker/QTL. The analyses detected six QTLs for plant height and six QTLs for heading date; collectively the QTLs for heading date accounted for a much greater amount of phenotypic variation than did the QTLs for plant height. Two-way analyses of variance, with all possible two-locus combinations, detected large numbers (from 101 to 257) of significant digenic interactions in the 2 years for both traits involving markers distributed in the entire genome; 22 and 39 were simultaneously detected in both years for plant height and heading date, respectively. Each of the interactions individually accounted for only a very small portion of the phenotypic variation. The majority of the significant interactions involved marker loci that did not detect significant effects by single-locus analyses, and many of the QTLs detected by single-locus analyses were involved in epistatic interactions. The results clearly demonstrated the importance of epistatic interactions in the genetic bases of heading date and plant height. Received: 5 May 2001 / Accepted: 3 August 2001     

Mapping and Isolation of Quantitative Trait Loci Controlling Plant Height and Heading Date in Rice

Quantitative trait loci (QTL) were identified for heading date and plant height in rice ( Oryza sativa L.) using a recombinant inbred line population consisting of 241 lines. Totally 4 QTLs for heading date and 4 QTLs for plant height were detected in three years. The QTL with large effects located in the interval C1023-R1440 on chromosome 7 was simultaneously detected in three years for both traits. In order to distinguished the interval whether contained one QTL with pleiotropy effect or two close linked QTLs, a recombinant line RIL50, whose genetic background was high similar to Zhenshan 97 except the regions covered the major QTL from Minghui 63, was selected to cross with Zhenshan 97. A BC1F2 population from the cross, which could be regarded as near isogenic lines (NIL) with the targeted QTL (QTL-NIL), was used to collect heading date and plant height data. The frequency distribution of the two traits in the BC1F2 population was bimodal, and their segregation ratios were in accordance with the expected Mendelian inheritance ratios. Normally, the short plants flowered early in the population, the high plants with late heading date, but the relationships between the plant height and the heading date of 6 plants conflicted with the case. The above results clearly demonstrated that QTL could be treated as single Mendelian factor. Moreover, there are two close linked genes controlling the heading date and the plant height on chromosome 7, respectively.     

Homoeologous loci control the accumulation of seed glucosinolates in oilseed rape (Brassica napus).

The genetic control of seed glucosinolate content in oilseed rape was investigated using two intervarietal backcross populations. Four QTLs segregating in the population derived from a Brassica napus L. 'Victor' x Brassica napus L. 'Tapidor' cross, together accounting for 76% of the phenotypic variation, were mapped. Three of these loci also appeared to control the accumulation of seed glucosinolates in a Brassica napus L. 'Bienvenu' x 'Tapidor' cross, and accounted for 86% of the phenotypic variation. The three QTLs common to both populations mapped to homoeologous regions of the B. napus genome, suggesting that seed glucosinolate accumulation is controlled by duplicate genes. It was possible to extend the comparative analysis of QTLs controlling seed glucosinolate accumulation by aligning the published genetic maps generated by several research groups. This comparative mapping demonstrated that high-glucosinolate varieties often carry low-glucosinolate alleles at one or more of the loci controlling seed glucosinolate accumulation.     

Improving rice yield and quality by QTL pyramiding

To facilitate marker-assisted transfer of desirable genes for improvement of yield traits, we used a set of backcross recombinant inbred lines (BRIL) derived from two elite parental lines, ‘Zhenshan97’ and ‘93-11’, to resolve a quantitative trait loci (QTL) cluster for heading date and yield-related traits in rice. Four main-effect QTL (qHD6.1, qHD6.2, qHD7, and qHD8) and four epistatic QTL affecting heading date in the BRIL were detected in two experimental trials. The major QTL (qHD8) was confirmed in three heterogeneous inbred families (HIF) that segregated for this target region, and narrowed down to a 20-kb segment in a large HIF-derived population. qHD8 was found to interact with qHD7 and had a pleiotropic effect responsible for heading date and yield components. To test usability of the identified QTL in rice improvement, we further developed near-isogenic lines (NIL) containing one or more target genes by marker-assisted transfer of ‘93-11’ alleles at qHD8, qHD7, and qHD6.1, and the GS3 gene for grain size into ‘Zhenshan97’. The pyramid line NIL(qHD8 + GS3) had higher yield potential, longer grains, and a more suitable heading date than ‘Zhenshan97’. Comparison of the NIL showed existence of epistasis between alleles at different loci and background effect on qHD8, which are very important for pyramiding of desirable alleles at the target QTL. These results will be particularly useful not only to understand the genetic basis of yield-related traits but also to improve the efficiency of marker-assisted selection for favorable loci in rice breeding programs.     

Inheritance of seed protein content and other agronomic characters in long bean (Vigna sesquipedalis Fruw.)

Summary Seven varieties of long bean, which included three local and four exotic, were crossed in a complete diallel. This was an attempt to study the inheritance of crude protein content, protein yield, flowering date, pod yield and yield components.Both additive and non-additive gene effects were responsible for the genetic variation in the diallel population. However, dominance variance was more important than additive variance in crude protein content, number of pods per plant and number of seeds per pod. For seed weight and pod length, additive variance was more important.The crude protein content, protein yield and number of pods per plant appeared to be controlled by overdominance effects. Partial dominance seemed to be the case for flowering date, pod length and seed weight; complete to overdominance for pod yield. High protein appeared to be associated with recessive genes whereas there was a general trend of high yielding parents carrying more dominant genes.     

A multiparental cross population for mapping QTL for agronomic traits in durum wheat (Triticum turgidum ssp. durum)

Multiparental cross designs for mapping quantitative trait loci (QTL) provide an efficient alternative to biparental populations because of their broader genetic basis and potentially higher mapping resolution. We describe the development and deployment of a recombinant inbred line (RIL) population in durum wheat (Triticum turgidum ssp. durum) obtained by crossing four elite cultivars. A linkage map spanning 2664 cM and including 7594 single nucleotide polymorphisms (SNPs) was produced by genotyping 338 RILs. QTL analysis was carried out by both interval mapping on founder haplotype probabilities and SNP bi‐allelic tests for heading date and maturity date, plant height and grain yield from four field experiments. Sixteen QTL were identified across environments and detection methods, including two yield QTL on chromosomes 2BL and 7AS, with the former mapped independently from the photoperiod response gene Ppd‐B1, while the latter overlapped with the vernalization locus VRN‐A3. Additionally, 21 QTL with environment‐specific effects were found. Our results indicated a prevalence of environment‐specific QTL with relatively small effect on the control of grain yield. For all traits, functionally different QTL alleles in terms of direction and size of genetic effect were distributed among parents. We showed that QTL results based on founder haplotypes closely matched functional alleles at known heading date loci. Despite the four founders, only 2.1 different functional haplotypes were estimated per QTL, on average. This durum wheat population provides a mapping resource for detailed genetic dissection of agronomic traits in an elite background typical of breeding programmes.     

Variability and covariability for plant height,heading date,and seed weight in wheat crosses

Summary Variability, covariability, heritability, and expected genetic gains from selection for heading date, plant height, and kernel weight were estimated in progenies derived from six wheat crosses. The crosses differed in the magnitude of the genetic variabilities of their progenies, but all crosses had significant variabilities for all traits. Heritability estimates were calculated by the variance components method. The estimates of heritability were relatively high for all three traits and averaged 86% for heading date, 77% for plant height, and 70% for kernel weight. The presence of significant genetic variabilities and high heritability estimates indicated that selection would be effective for the three traits.The segregates derived from crosses between medium tall parents showed transgressive segregation that would permit isolation of short-statured types. Transgressive segregation also occurred for heading date and kernel weight.Heading date and plant height were positively and highly significantly correlated in four crosses out of six. But both plant height and heading date had, in general, low negative correlations with kernel weight which would not preclude the development of short wheats with high kernel weight from these crosses. The association between characters was mostly genetic in cause.     

Identifying genetic components controlling fertility in the outcrossing grass species perennial ryegrass (Lolium perenne) by quantitative trait loci analysis and comparative genetics

Mutational load and resource allocation factors and their effects on limiting seed set were investigated in ryegrass by comparative mapping genomics and quantitative trait loci (QTL) analysis in two perennial ryegrass (Lolium perenne) mapping families sharing common genetic markers. Quantitative trait loci for seed-set were identified on chromosome (LG) 7 in both families and on LG4 of the F2/WSC family. On LG7, seed-set and heading date QTLs colocalized in both families and cannot be unequivocally resolved. Comparative genomics suggests that the LG7 region is syntenous to a region of rice LG6 which contains both fertility (S5(n)) and heading date (Hd1, Hd3a) candidate genes. The LG4 region is syntenous to a region of rice LG3 which contains a fertility (S33) candidate gene. QTL maxima for seed-set and heading date on LG4 in the F2/WSC family are separated by c. 8 cm, indicating distinct genetic control. Low seed set is under the control of recessive genes at both LG4 and LG7 locations. The identification of QTLs associated with seed set, a major component of seed yield in perennial ryegrass, indicates that mutational load associated with these genomic regions can be mitigated through marker-assisted selection.     

A comparative study of seed yield parameters in Arabidopsis thaliana mutants and transgenics

Because seed yield is the major factor determining the commercial success of grain crop cultivars, there is a large interest to obtain more understanding of the genetic factors underlying this trait. Despite many studies, mainly in the model plant Arabidopsis thaliana, have reported transgenes and mutants with effects on seed number and/or seed size, knowledge about seed yield parameters remains fragmented. This study investigated the effect of 46 genes, either in gain- and/or loss-of-function situations, with a total of 64 Arabidopsis lines being examined for seed phenotypes such as seed size, seed number per silique, number of inflorescences, number of branches on the main inflorescence and number of siliques. Sixteen of the 46 genes, examined in 14 Arabidopsis lines, were reported earlier to directly affect in seed size and/or seed number or to indirectly affect seed yield by their involvement in biomass production. Other genes involved in vegetative growth, flower or inflorescence development or cell division were hypothesized to potentially affect the final seed size and seed number. Analysis of this comprehensive data set shows that of the 14 lines previously described to be affected in seed size or seed number, only nine showed a comparable effect. Overall, this study provides the community with a useful resource for identifying genes with effects on seed yield and candidate genes underlying seed QTL. In addition, this study highlights the need for more thorough analysis of genes affecting seed yield.     

水稻抗褐飞虱种质Swarnalata抽穗期和休眠性相关QTL检测

为有效利用抗褐飞虱水稻Swarnalata,对2013年南京种植的Swarnalata/02428 F2分离群体进行抽穗期和种子休眠性考察,利用172个分子标记构建了Swarnalata/02428 F2的分子遗传连锁图谱,图谱全长为3311.4c M,标记间平均图距为19.22c M。利用Windows QTL Cartographer V2.5软件对该分离群体进行抽穗期和种子休眠性相关QTL检测,共检测到7个抽穗期相关QTL,分别位于第2、3、6、11染色体,其中位于第11染色体的q HD-11-1贡献率最高,为28.85%;检测到3个种子休眠性相关QTL,分别位于第3、6、9染色体,其中位于第9染色体的q Sd-9贡献率最高,为22.11%。分析表明,本研究检测到的抽穗期QTL与种子休眠QTL所在位置不同,说明该群体中种子休眠与抽穗期没有直接关系,它们分别由不同基因控制。本研究不仅为水稻休眠基因的精细定位及克隆奠定基础,也为更有效利用Swarnalata中的抗褐飞虱基因提供基础和一些优良的中间材料。     

普通野生稻中增产相关QTL的发掘

普通野生稻(Oryza Rufipogon)是重要的遗传资源,发掘其优良等位基因将对水稻遗传改良产生重要影响。文章从以珍汕97为轮回亲本,普通野生稻为供体的BC2F1群体中选择一个与珍汕97表型明显不同的单株BC2F1-15,经过连续自交获得回交重组自交系BC2F5群体。均匀分布于12条染色体的126个多态性SSR(Simplesequence repeats)标记基因型分析,发现BC2F1-15单株在30%的标记位点为杂合基因型;利用该群体共检测到4个抽穗期、3个株高、4个每穗颖花数、2个千粒重和1个单株产量QTL。在第7染色体RM481-RM2区间,检测到抽穗期、每穗颖花数和产量QTL,野生稻等位基因表现增效作用;其他3个每穗颖花数QTL位点,野生稻等位基因也均具有增效作用。结果表明野生稻携带有增产相关的等位基因,这些有利等位基因无疑是水稻遗传改良可资利用的新资源。     

几个水稻品种抽穗期主效基因与微效基因的定位研究

在构建２张ＲＦＬＰ图谱的基础上,定位分析了控制水稻抽穗期的主效基因和微效基因。在特三矮２号／Ｃ．Ｂ．群体中定位到２个主效基因和２个微效基因。该２个主基因分别位于第３、８染色体上,累加贡献率约达５０％,加性效应值分别为７天和６天,而分别位于第１、１２染色体的２个微效基因的贡献率仅分别为８．３％和９．６％,加性效应值仅为３天和４天。在外引２号／Ｃ．Ｂ．群体中定位了２个连锁于第６染色体的主效基因和１个位于第８染色体的微效基因,该２个主效基因的贡献率分别为３５．５％和２７．４％,来自外引２号的该２个基因其效应均为明显推迟抽穗,因而可推测它们为感光性基因,微效基因的贡献率仅为８．９％,基因效应值较小。     

水稻上部节间长度等数量性状基因的定位及其遗传效应分析

对水稻籼粳杂交（窄叶青８号×京系１７）Ｆ＿１的花药进行离体培养,建立一个含１３３个ＤＨ系的作图群体,通过构建分子连锁图谱,对水稻上部节间长度、株高和抽穗期的ＱＴＬ进行区间作图,定位了影响上部节间长度的１２个ＱＴＬ、株高的４个ＱＴＬ和抽穗期的１个ＱＴＬ。对这些ＱＴＬ的遗传效应分析的结果表明,控制抽穗期的１个ＱＴＬ即Ｈｄ８ａ为主效基因,其余的１６个ＱＴＬ为微效基因。控制上部节间长度单个的ＱＴＬ对表型的贡献率介于８－１８％,其加性效应可使所控制的节间长度增加大约１．６－３．６ｃｍ。值得注意的是,一些控制相关性状的、作用方向相同的ＱＴＬ定位于同一染色体的相同或邻近区段上。这一结果揭示了这些性状相关的遗传基础,在水稻育种中运用这些ＱＴＬ将有助于对株高进行精细的遗传调控。     

Effect of a rye dwarfing gene on plant height,heading stage,and Fusarium head blight in triticale (×Triticosecale Wittmack)

Key message

The rye-derived dwarfing gene Ddw1 on chromosome 5R acts in triticale in considerably reducing plant height, increasing FHB severity and delaying heading stage.

Abstract

Triticale, an amphiploid hybrid between durum wheat and rye, is an European cereal mainly grown in Germany, France, Poland, and Belarus for feeding purposes. Dwarfing genes might further improve the genetic potential of triticale concerning lodging resistance and yield. However, they might have pleiotropic effects on other, agronomically important traits including Fusarium head blight. Therefore, we analyzed a population of 199 doubled haploid (DH) lines of the cross HeTi117-06 × Pigmej for plant height, heading stage, and FHB severity across 2 locations and 2 years. The most prominent QTL was detected on chromosome 5R explaining 48, 77, and 71 % of genotypic variation for FHB severity, plant height, and heading stage, respectively. The frequency of recovery in cross validation was ≥90 % for all three traits. Because the markers that detect dwarfing gene Ddw1 in rye are also in our population the most closely linked markers, we assume that this major QTL resembles Ddw1. For FHB severity two, for plant height three, and for heading stage five additional QTL were detected. Caused by the considerable genetic variation for heading stage and FHB severity within the progeny with the dwarfing allele, short-strawed, early heading and FHB-resistant lines can be developed when population size is large enough.     

Genetic architecture of quantitative trait loci associated with morphological and agronomic trait differences in a wild by cultivated barley cross.

Hordeum vulgare subsp. spontaneum is the progenitor of cultivated barley (Hordeum vulgare L.). Domestication combined with plant breeding has led to the morphological and agronomic characteristics of modern barley cultivars. The objective of this study was to map the genetic factors that morphologically and agronomically differentiate wild barley from modern barley cultivars. To address this objective, we identified quantitative trait loci (QTLs) associated with plant height, flag leaf width, spike length, spike width, glume length in relation to seed length, awn length, fragility of ear rachis, endosperm width and groove depth, heading date, flag leaf length, number of tillers per plant, and kernel color in a Harrington/OUH602 advanced backcross (BC2F8) population. This population was genotyped with 113 simple sequence repeat markers. Thirty QTLs were identified, of which 16 were newly identified in this study. One to 4 QTLs were identified for each of the traits except glume length, for which no QTL was detected. The portion of phenotypic variation accounted for by individual QTLs ranged from about 9% to 54%. For traits with more than one QTL, the phenotypic variation explained ranged from 25% to 71%. Taken together, our results reveal the genetic architecture of morphological and agronomic traits that differentiate wild from cultivated barley.