Seed yield in perennial ryegrass (<Emphasis Type="Italic">Lolium perenne</Emphasis> L.): comparative importance of component traits and detection of seed-yield-related QTL

The aims of our study were to evaluate relationships amongst morphological traits associated with seed production in a perennial ryegrass biparental population and to identify genomic regions associated with phenotypic variation in those traits using QTL analysis. This was achieved using data from two field experiments at Palmerston North and Lincoln, New Zealand, in 2003, and days to heading (DTH), reassessed in 2004. Trait association was determined for the Palmerston North experiment where measured traits included seed yield per plant (SYPlant), seed yield per spike (SYSp), reproductive tiller number (RTiller), spikelets per spike (SpktSp), florets per spikelet (FSpkt), 1000 seed weight (TSW), spike length (SpLen), florets per spike (FSp), floret site utilization (FSUtil), spread of heading (SOH) and plant growth habit (PGHabit). Traits contributing to SYPlant in order of descending value were FSpkt, FSUtil, and RTiller. High TSW was only weakly linked to SYPlant. FSUtil, SOH and RTiller were identified as valuable breeding targets for improving seed yield potential in perennial ryegrass. QTL were identified for all traits except for RTiller. QTL for SYPlant occurred on linkage groups (LG) 2 and 6. Both were co-located with QTL for SYSp and sets of SYPlant components or related traits (FSpkt, FSp; FSUtil and TSW). Major QTL for DTH were identified on LG2 and LG4 and minor QTL on LG7 in consecutive years. There was a strong genotype-by-environment interaction for SYPlant that was reflected in a lack of consistent QTL across environments, while QTL for SYSp and DTH were stable across environments. Identification of component traits and QTL important for seed yield may accelerate genetic improvement in perennial ryegrass through conventional and marker-assisted breeding, respectively.     

Quantitative trait loci for early maturity and their potential in breeding for earliness in Brassica juncea

A doubled haploid population of Brassica juncea, developed from a cross between two parental lines differing for days to maturity, was used to study the efficiency of indirect selection for a primary trait through selection of secondary trait(s) over direct selection for the primary trait when quantitative trait loci information is available for both primary and secondary traits, and applied. Days to maturity was considered as primary trait, while days to first flowering, days to end of flowering, flowering period and plant height were considered as secondary traits. An RFLP linkage map was employed for QTL analysis of maturity and maturity-determinant traits, and a stable QTL B6 simultaneously affecting these two types of traits was identified. This linked QTL explained 11.7% phenotypic variation for days to maturity, 20.7% variation for days to first flowering, 24.3% variation for days to end of flowering and 14.4% variation for plant height. Phenotypic evaluation of maturity and/or maturity-determinant traits, viz. days to first flowering, days to end of flowering and plant height revealed that limited genetic advance for early maturity can be achieved through phenotypic selection of the primary and/or the secondary trait(s). However, the estimates of genetic advance for early maturity based on combined phenotypic evaluation and linked QTL data was found to be, at least, three times higher compared to genetic advance based on phenotypic evaluation only, demonstrating the potential of marker-assisted selection in breeding for early maturity in B. juncea.     

Several stably expressed QTL for spike density of common wheat (Triticum aestivum) in multiple environments

Spike density (SD), an important spike morphological trait associated with wheat yield, is the spikelet number per spike (SNS) divided by spike length (SL). In this study, phenotypic data from eight environments were collected and a recombinant inbred line population (RIL) constructed by the wheat line 20828 and the cultivar 'Chuannong16' and a Wheat55K SNP array-based constructed genetic linkage map were used to identify SD quantitative trait locus (QTL). Correlation between SD and other agronomic traits was calculated. Genes associated with plant growth and development for major loci were predicted. The results showed that 24 QTLs associated with SD were detected in eight environments. Among them, three major QTL, namely QSd.sicau-5B.2, QSd.sicau-2D.3 and QSd.sicau-4B.1, explained up to 35.62%, 14.21% and 11.23% of phenotypic variation, respectively. The positive alleles of them were all derived from 'Chuannong16'. The significant relationships between SD and other agronomic traits were detected and discussed. Taken together, the stably expressed SD QTL under different environments identified in this study provided theoretical guidance for further fine mapping and germplasm improvement.     

Identification of QTL for oil content,seed yield,and flowering time in oilseed rape (<Emphasis Type="Italic">Brassica napus</Emphasis>)

A genetic map was constructed with 353 sequence-related amplified polymorphism and 34 simple sequence repeat markers in oilseed rape (Brassica napus L.). The map consists of 19 linkage groups and covers 1,868 cM of the rapeseed genome. A recombinant doubled haploid (DH) population consisting of 150 lines segregating for oil content and other agronomic traits was produced using standard microspore culture techniques. The DH lines were phenotyped for days to flowering, oil content in the seed, and seed yield at three locations for 3 years, generating nine environments. Data from each of the environments were analyzed separately to detect quantitative trait loci (QTL) for these three phenotypic traits. For oil content, 27 QTL were identified on 14 linkage groups; individual QTL for oil content explained 4.20–30.20% of the total phenotypic variance. For seed yield, 18 QTL on 11 linkage groups were identified, and the phenotypic variance for seed yield, as explained by a single locus, ranged from 4.61 to 24.44%. Twenty-two QTL were also detected for days to flowering, and individual loci explained 4.41–48.28% of the total phenotypic variance.     

Genome-wide association study of differences in 14 agronomic traits under low- and high-density planting models based on the 660k SNP array for common wheat

One hundred and ninety-seven wheat accessions from Yellow and Huai Winter Wheat Region (YHW) were evaluated for differences of 14 agronomic traits under low- and high-density plantings. Compared with the high-density plantings, plant height, neck length, uppermost internode length, flag leaf angle and number of sterile spikelets under the low-density plantings reduced, while heading date, flowering date, flag leaf length and width, spike length, number of fertile spikelets, grain number per spike, thousand-kernel weight and grain weight per spike increased. A total of 1,118 markers were detected based on GWAS, and seven QTLs were confirmed. One QTL on chromosomes 5BL and two other QTLs on 5Dl were all tightly associated with flowering date difference. Bioinformatics analysis revealed that two haploblocks in 5Dl were involved, and the Vrn-D1 locus was located in this interval. A region on chromosome 5B at around 531.5 Mb was significantly associated with plant height difference. Two QTLs including AX-94840438 (7BL) and AX-94563647 (7DS) were responsible for neck length or uppermost internode length difference.     

Chromosomal locations and genetic relationships of tiller and spike characters in wheat

Number of tillers per plant, plant growth habit in seedling and adult stages, and spike and spikelet characters are agronomically important features of the gross morphology of wheat. To localize to wheat chromosomes the genes for these traits, we scored them in a set of wheat recombinant-inbred mapping lines already well genotyped with molecular markers. Quantitative-trait analysis revealed a region near Gli-A2 (Xpsr10) on the short arm of chromosome 6A strongly affecting tiller number and the correlated trait of seedling growth habit. Genes with opposing effects on adult plant type were localized on the short arms of chromosomes2A and 3A, while genes affecting spike development were assigned to several A- and B-genome chromosomes. None of these genes showed synteny with counterpart QTLs reported to affect the same traits in rice. In the chromosome 2D region containing the photoperiod-insensitivity gene Ppd-D1, the major determinant of heading date in these autumn-sown lines, earliness alleles reduced tiller and spikelet numbers and increased erect seedling growth habit, but showed no influence on adult plant type or spike length. Though several of these morphological traits are generally considered to be associated with winter hardiness and their phenotypic intercorrelations were consistent with the genetic mapping evidence, no association was found between newly identified loci and known vernalization-response or frost-resistance loci. This revised version was published online in August 2006 with corrections to the Cover Date.     

Identification of a novel QTL for the number of spikelets per panicle using a cross between indica‐ and japonica‐type high‐yielding rice cultivars in Japan

Yield is a complex trait. To improve it, the accumulation of the favourable alleles of valuable genes is required for each yield‐related trait. In this study, we used two high‐yielding rice cultivars developed in Japan, indica‐type ‘Takanari’ and japonica‐type ‘Momiroman’, for a genetic analysis of the sink capacity‐related traits. An F₂ population showed transgressive segregation for the number of spikelets per panicle. Quantitative trait locus (QTL) analysis detected four QTLs for the trait. Two of the QTLs were most likely identical to previously cloned GN1a and APO1, and their Takanari alleles had positive effects. The Momiroman alleles of the other two QTLs had positive effects, and one of these QTLs was most likely identical to SPIKE/GPS. The QTL on the long arm of chromosome 3 appeared to be novel; it clustered with QTLs for grain length and days‐to‐heading. Substitution mapping revealed that the close linkage of QTLs caused the clustering. These results suggest that the combination of the favourable alleles of detected QTLs could lead to greater sink capacity than that of the parental cultivars.     

Durum wheat landraces from syria. II. patterns of variation

Summary Phenotypic variation components were estimated with respect to days to heading, flag leaf length and width, plant height, awn and spike length, awn and spike colour, spikelets per spike and seed shrivelling of 84 Syrian durum wheat landrace populations. Multivariate patterns of variation were established through principal component analysis to describe relationships between landrace groups and regions of collection. Agroecological site characteristics and plant traits were compared with respect to patterns of geographical variation. Grouping on the basis of landrace groups proved more discriminative than on the basis of regions of origin. Landraces originating from sites characterized by favourable growth conditions tended to be later in heading and to have longer spikes with longer awns but with less spikelets. The observed relation between favourable growth conditions in the regions of origin and smaller flag leaves may be caused by genotype × environment interaction.Among populations variation was high, and amounted to 96% of the total variation, whereas the remnant 4% was contributed to differences within populations and among lines. Variation among landrace groups and among regions was calculated as 79% and 75% of total variation, respectively.     

QTL mapping and analysis of epistatic interactions for grain yield and yield-related traits in <Emphasis Type="Italic">Triticum turgidum</Emphasis> L. var. <Emphasis Type="Italic">durum</Emphasis>

A quantitative trait loci (QTL) analysis of grain yield and yield-related traits was performed on 93 durum wheat recombinant inbred lines derived from the cross UC1113 × Kofa. The mapping population and parental lines were analyzed considering 19 traits assessed in different Argentine environments, namely grain yield, heading date, flowering time, plant height, biomass per plant, and spikelet number per ear, among others. A total of 224 QTL with logarithm of odds ratio (LOD) ≥ 3 and 47 additional QTL with LOD > 2.0 were detected. These QTL were clustered in 35 regions with overlapping QTL, and 12 genomic regions were associated with only one phenotypic trait. The regions with the highest number of multi-trait and stable QTL were 3BS.1, 3BS.2, 2BS.1, 1BL.1, 3AL.1, 1AS, and 4AL.3. The effects of epistatic QTL and QTL × environment interactions were also analyzed. QTL putatively located at major gene loci (Rht, Vrn, Eps, and Ppd) as well as additional major/minor QTL involved in the complex genetic basis of yield-related traits expressed in Argentine environments were identified. Interestingly, the 3AL.1 region was found to increase yield without altering grain quality or crop phenology.     

QTL analysis of seed dormancy in rice (Oryza sativa L.)

Effective cumulative temperature (ECT) after heading would be a more reasonable parameter for seed sampling of pre-harvest sprouting/seed dormancy (SD) tests in segregating populations than the days after flowering. SD is an important agronomic trait associated with grain yielding, eating quality and seed quality. To identify genomic regions affecting SD at different grain-filling temperatures, and to further examine the association between SD and ECT during grain-filling, 127 double haploid (DH) lines derived from a cross between ZYQ8 (indica)/JX17 (japonica) by anther culture were analyzed. The quantitative trait loci (QTLs) and their digenic epistasis for SD were identified using a molecular linkage map of this population. A total of four putative QTLs for SD (qSD-3, qSD-5, 6 and 11) were detected on chromosomes 3, 5, 6 and 11, together explaining 41.4% of the phenotypic variation. Nine pairs of digenic epistatic loci were associated with SD on all but chromosome 9, and their contributions to phenotypic variation varied from 2.87%–8.73%. The SD QTL on chromosome 3 was identical to the QTLs found in other mapping populations with different genetic backgrounds, which could be a desirable candidate for gene cloning and marker-assisted selection in rice breeding.     

Quantitative trait loci mapping of seed colour,hairy leaf,seedling anthocyanin,leaf chlorosis and days to flowering in F2 population of Brassica rapa L.

A diversity arrays technology (DArT) map was constructed to identify quantitative trait loci (QTL) affecting seed colour, hairy leaf, seedling anthocyanin, leaf chlorosis and days to flowering in Brassica rapa using a F₂ population from a cross between two parents with contrasting traits. Two genes with dominant epistatic interaction were responsible for seed colour. One major dominant gene controls the hairy leaf trait. Seedling anthocyanin was controlled by a major single dominant gene. The parents did not exhibit leaf chlorosis; however, 32% F₂ plants showed leaf chlorosis in the population. A distorted segregation was observed for days to flowering in the F₂ population. A linkage map was constructed with 376 DArT markers distributed over 12 linkage groups covering 579.7 cM. The DArT markers were assigned on different chromosomes of B. rapa using B. rapa genome sequences and DArT consensus map of B. napus. Two QTL (RSC1‐2 and RSC12‐56) located on chromosome A8 and chromosome A9 were identified for seed colour, which explained 19.4% and 18.2% of the phenotypic variation, respectively. The seed colour marker located in the ortholog to Arabidopsis thaliana Transparent Testa2 (AtTT2). Two QTL RLH6‐0 and RLH9‐16 were identified for hairy leaf, which explained 31.6% and 20.7% phenotypic variation, respectively. A single QTL (RSAn‐12‐157) on chromosome A7, which explained 12.8% of phenotypic variation was detected for seedling anthocyanin. The seedling anthocyanin marker is found within the A. thaliana Transparent Testa12 (AtTT12) ortholog. A QTL (RLC6‐04) for leaf chlorosis was identified, which explained 55.3% of phenotypic variation. QTL for hairy leaf and leaf chlorosis were located 0–4 cM apart on the same chromosome A1. A single QTL (RDF‐10‐0) for days to flowering was identified, which explained 21.4% phenotypic variation.     

Identification of quantitative trait loci for grain yield, seed protein concentration and maturity in field pea (Pisum sativum L.)

Breeding efforts to improve grain yield, seed protein concentration and early maturity in pea (Pisum sativum L.) have proven to be difficult. The use of molecular markers will improve our understanding of the genetic factors conditioning these traits and is expected to assist in selection of superior genotypes. This study was conducted to identify genetic loci associated with grain yield, seed protein concentration and early maturity in pea. A population of 88 recombinant inbred lines (RILs) that was developed from a cross between 'Carneval' and 'MP1401' was evaluated at 13 environments across the provinces of Alberta, Manitoba and Saskatchewan, Canada in 1998, 1999 and 2000. A linkage map consisting of 193 AFLPs (amplified fragment length polymorphism), 13 RAPDs (random amplified polymorphic DNA) and one STS (sequence tagged site) marker was used to identify putative quantitative trait loci (QTL) for grain yield, seed protein concentration and early maturity. Four QTL were identified each for grain yield and days to maturity, and three QTL were identified for seed protein concentration. A multiple QTL model for each trait showed that these genomic regions accounted for 39%, 45% and 35% of the total phenotypic variation for grain yield, seed protein concentration and days to maturity, respectively. The consistency of these QTL across environments and their potential for marker-assisted selection are discussed in this report.     

QTL identification of yield-related traits and their association with flowering and maturity in soybean

Two soybean recombinant inbred line populations, Jinpumkong 2 × SS2-2 (J × S) and Iksannamulkong × SS2-2 (I x S) showed population-specific quantitative trait loci (QTLs) for days to flowering (DF) and days to maturity (DM) and these were closely correlated within population. In the present study, we identified QTLs for six yield-related traits with simple sequence repeat markers, and biological correlations between flowering traits and yield-related traits. The yield-related traits included plant height (PH), node numbers of main stem (NNMS), pod numbers per plant (PNPP), seed numbers per pod (SNPP), 100-seed weight (SW), and seed yield per plant (SYPP). Eighteen QTLs for six yield-related traits were detected on nine chromosomes (Chrs), containing four QTLs for PH, two for NNMS, two for PNPP, three for SNPP, five for SW, and two for SYPP. Two highly significant QTLs for PH and NNMS were identified on Chr 6 (LG C2) in both populations where the major flowering gene, E1, and two DF and DM QTLs were located. One other PNPP QTL was also located on this region, explaining 12.9% of phenotypic variation. Other QTLs for yield-related traits showed population-specificity. Two significant SYPP QTLs potentially related with QTLs for SNPP and PNPP were found on the same loci of Chrs 8 (Satt390) and 10 (Sat_108). Also, highly significant positive phenotypic correlations (P < 0.01) were found between DF with PH, NNMS, PNPP, and SYPP in both populations, while flowering was negatively correlated with SNPP and SW in the J × S (P < 0.05) and I × S (P < 0.01) populations. Similar results were also shown between DM and yield-related traits, except for one SW. These QTLs identified may be useful for marker-assisted selection by soybean breeders.     

小麦籽粒产量及穗部相关性状的QTL定位

由小麦品种花培3号和豫麦57杂交获得DH群体168个株系,种植于3个环境中,利用305个SSR标记对籽粒产量和穗部相关性状(穗长、穗粒数、总小穗数、可育小穗数、小穗着生密度、千粒重和粒径)进行了QTL定位。利用基于混合线性模型的QTLNetwork 2.0软件,共检测到27个加性效应和13对上位效应位点,其中 8个加性效应位点具有环境互作效应。相关性高的性状间有一些共同的QTL位点,表现出一因多效或紧密连锁效应。5D染色体区段Xwmc215–Xgdm63,检测到控制籽粒产量、穗粒数、总小穗数、可育小穗数和小穗着生密度5个性状的QTL位点,各位点的遗传贡献率较大且遗传效应方向相同,增效等位基因均来源于豫麦57,适用于分子标记辅助育种和聚合育种。控制千粒重与穗粒数的QTL位于染色体不同区段,有利于实现穗粒数与粒重的遗传重组。     

Molecular markers for yield components in Brassica juncea – do these assist in breeding for high seed yield?

A population of 112 F₁-derived doubled haploid lines was produced from a reciprocal cross of Brassica juncea. The parents differed for seed quality, seed color and many agronomic traits. A detailed RFLP linkage map of this population, comprising 316 loci, had been constructed, and was used to map quantitative trait loci (QTL) for seed yield and yield components, viz. siliqua length, number of seeds per siliqua, number of siliques per main raceme and 1000-seed weight. Stable and significant QTLs were identified for all these yield components except seed yield. For yield components, a selection index based on combined phenotypic and molecular data (QTL effects) could double up the efficiency of selection compared to the expected genetic advance by phenotypic selection. Selection indices for high seed yield, based on the phenotypic data of yield and yield components, could only improve the efficiency of selection by 4% of the genetic advance that can be expected from direct phenotypic selection for yield alone. Inclusion of molecular data together with the phenotypic data of yield components in the selection indices did not improve the efficiency of selection for higher seed yield. This is probably due to often negative relationships among the yield components. Most of the QTLs for yield components were compensating each other, probably due to linkage, pleiotropy or developmentally induced relationships among them. The breeding strategy for B. juncea and challenges to marker assisted selection are discussed.     

燕大1817/北农6号重组自交系群体穗部性状的QTL定位

小麦穗部性状与单株产量密切相关。本研究以小麦骨干亲本燕大1817与优良品系北农6号衍生的269个重组自交系为材料,通过在北京和河北石家庄的2年田间试验数据,利用本实验室已构建的高密度SNP和SSR遗传连锁图谱进行穗长、穗粒数和穗粒重QTL定位。采用完备复合区间作图法共检测到29个穗部性状加性效应QTL,其中10个穗长QTL分布于1B、2D、3A、3B、4A、5A、5B、6A和7D染色体上,解释的表型变异率为2.96%~9.63%,QSl.cau-4A.2在所有5个环境中均能被检测到,解释的表型变异为5.89%~9.62%,另有7个QTL能在2个或2个以上环境中被检测到;8个穗粒数相关QTL分布于1A、3A、3D、4A和5B染色体上,解释的表型变异为4.06%~11.17%,为单个环境QTL。11个与穗粒重相关QTL分布于1A、1B、2A、2D、3A、4D、5A、5B和6B染色体上,解释的表型变异为2.79%~16.12%,其中QGws.cau-1B、QGws.cau-3A和QGws.cau-6B.2在2个或者2个以上环境中能被检测到。另外,鉴定出6个分布于1A、2D、3A、4A和5B染色体上的QTL富集区段。     

Co-location of seed oil content,seed hull content and seed coat color QTL in three different environments in <Emphasis Type="Italic">Brassica napus</Emphasis> L.

Increasing seed oil content is an important breeding goal for Brassica napus L. (B. napus). The identification of quantitative trait loci (QTL) for seed oil content and related traits is important for efficient selection of B. napus cultivars with high seed oil content. To get better knowledge on these traits, a molecular marker linkage map for B. napus was constructed with a recombinant inbred lines (RIL) population. The length of the map was 1,589 cM with 451 markers distributed over 25 linkage groups. QTL for seed oil content, seed hull content and seed coat color in three environments were detected by composite interval mapping (CIM) tests. Eleven QTL accounted for 5.19–13.57% of the variation for seed oil content. Twelve QTL associated with seed hull content were identified with contribution ranging from 5.80 to 22.71% and four QTL for seed coat color accounted for 5.23–15.99% of the variation. It is very interesting to found that co-localization between QTL for the three traits were found on N8. These results indicated the possibility to combine favorable alleles at different QTL to increase seed oil content, as well as to combine information about the relationship between seed oil content and other traits.     

Multivariate analysis of variation among wild oat accessions — seed traits

Summary A sample of 457 (Avena sterilis L.) accessions from the World Oat Collection maintained at the Germplasm Laboratory, Beltsville, Maryland, USA, was studied by multivariate analyses to explain the phenotypic relationships among the entries. According to these analyses, patterns of seed trait associations suggest that diversity may have geographic patterning. For example, the accessions from countries and islands of the western Mediterranean tend to have a common set of traits. Number of spikelets per panicle and groat weight and dimensions were important traits that discriminated the A. sterilis accessions into the group. It was concluded that region-specific adaptations are responsible for occurrence of certain traits and trait associations in specific geographic regions.     

QTL analysis for brown spot resistance in American rice cultivar ‘Dawn’

Rice brown spot (BS), caused by Bipolaris oryzae, causes yield loss and deterioration of grain quality. Using single-nucleotide polymorphism (SNP) markers, we conducted quantitative trait locus (QTL) analysis of BS resistance in backcross inbred lines (BILs) from a cross between an American rice cultivar, ‘Dawn’ (resistant), and ‘Koshihikari’ (susceptible). Four QTLs for BS resistance were detected in a three-year field evaluation, and ‘Dawn’ contributed the resistance alleles at all QTLs. The QTL with the greatest effect, qBSR6-kd, explained 15.1% to 20.3% of the total phenotypic variation. Although disease score and days to heading (DTH) were negatively correlated in all three years, qBSR6-kd was located near a QTL for DTH at which the ‘Dawn’ allele promoted heading. Another BS resistance QTL (qBSR3.1-kd) was unlinked to the QTLs for DTH. Therefore, these two QTLs are likely to be useful for breeding BS-resistant varieties without delaying heading. The other two BS resistance QTLs (qBSR3.2-kd and qBSR7-kd) were located near DTH QTLs at which the ‘Dawn’ alleles delayed heading. The QTLs reported here will be good candidates for developing BS-resistant cultivars.