Dissection of two QTL clusters underlying yield-related heterosis in the cabbage founder parent 01–20

Cabbage has significant heterosis and most commercial cultivars are hybrids.To explore genetic basis of cabbage heterosis and promote cabbage heterosis utilization,we constructed two populations by crossing 100 DH lines derived from a cabbage hybrid 01–20×96–100 with two female parents.Hybrids exhibited different extents of heterosis,the mean value of economic yield was 2.6 times bigger than parents.We identified 66 and 73 QTLs associated with mid-parent heterosis and transgressive heterosis of twelve yield-related traits,respectively.Some QTLs could be detected under the two-year experiment existed in two populations with different testers,showing relatively high phenotypic contribution rate(15.8%–20.0%).Heterosis QTLs exhibited clustered distribution in several cabbage chromosome regions.Two dominant genetic regions,mk300–316 and mk258–268,originated from the elite parent 01–20,exhibited significant genetic effects for yield-related heterosis,which were first identified.Three elite DH lines(D22,D46,D83)harboring these two dominant regions were selected as having strong heterosis in cabbage production.Candidate gene analysis revealed that some genes participating in biosynthetic processes of carbohydrates and some responses to auxin might affect cabbage yield heterosis.QTL identification and genetic dissection of yield-related traits provide new insights into the genetic effects of cabbage heterosis.

Introgression of QTL from Aegilops tauschii enhances yield-related traits in common wheat

To break the narrow diversity bottleneck of the wheat D genome, a set of Aegilops tauschii-wheat introgression(A-WI) lines was developed by crossing Ae. tauschii accession T015 with common wheat elite cultivar Zhoumai 18(Zhou18). A high-density genetic map was constructed based on Single Nucleotide Polymorphism(SNP) markers and 15 yield-related traits were evaluated in 11 environments for detecting quantitative trait loci(QTL). A total of 27 environmentally stable QTL were identified in at least five environments, 20 of which were derived from Ae. tauschii T015, explaining up to 24.27% of the phenotypic variations. The major QTL for kernel length(KL), QKl-2D.5, was delimited to a physical interval of approximately 2.6 Mb harboring 52 candidate genes. Three Kompetitive Allele Specific PCR(KASP)markers were successfully developed based on nonsynonymous nucleotide mutations of candidate gene AetT093_2Dv1G100900.1 and showed that A-WI lines with the T015 haplotype had significantly longer KL than the Zhou18 haplotype across all 11 environments. Four primary valuable A-WIs with good trait performance and carrying yield-related QTL were selected for breeding improvement. The results will facilitate the efficient transfer of beneficial genes from Ae. tauschii into wheat cultivars to improve wheat yield and other traits.

Identification of tolerance to high density and lodging in short petiolate germplasm M657 and the effect of density on yield-related phenotypes of soybean

Soybean yield has traditionally been increased through high planting density,but investigating plant height and petiole traits to select for compact architecture,lodging resistance,and high yield varieties is an underexplored option for further improving yield.We compared the relationships between yield-related traits,lodging resistance,and petioleassociated phenotypes in the short petiole germplasm M657 with three control accessions during 2017–2018 in four locations in the Huang–Huai region,China.The results showed that M657 exhibited stable and high tolerance to high planting density and resistance to lodging,especially at the highest density(8×105 plants ha–1).The regression analysis indicated that a shorter petiole length was significantly associated with increased lodging resistance.The yield analysis showed that M657 achieved higher yields under higher densities,especially in the northern part of the Huang–Huai region.Among the varieties,there were markedly different responses to intra-and inter-row spacing designs with respect to both lodging and yield that were related to location and density.Lodging was positively correlated with planting density,plant height,petiole length,and number of effective branches,but negatively correlated with stem diameter,seed number per plant,and seed weight per plant.The yield of soybean was increased by appropriately increasing the planting density on the basis of the current soybean varieties in the Huang–Huai region.This study provides a valuable new germplasm resource for the introgression of compact architecture traits that are amenable to providing a high yield in high density planting systems,and it establishes a high-yield model of soybean in the Huang–Huai region.

Genome-wide association study exposed the pleiotropic genes for yield-related and oil quality traits in Brassica napus L.

Oilseed rape(Brassica napus L.) is an allotetraploid(AACC,2n=38) crop,valued for its edible oil and protein content.seed yield and nutritional composition of rapeseed are influenced by its yield and oil quality traits.However,the genetic basis of yield-related and oil-quality traits remain ambiguous.A panel of 266 diversified oilseed rape accessions was genotyped using 223 simple sequence repeat(SSR) markers covering all 19 chromosomes to identify significant markers associated with yield and quality traits.Twelve yield-related and six quality traits were investigated in two consecutive years(2014 and 2015),with three replications in two environments(Changshun,CS;and Qinghe,QH).Using the model GLM with population structure and kinship(Q+K),a total of 25 significant SSR markers(P <0.001) were detected to be associated with these twelve yield-related and six quality traits,explaining 4.56%-19.17% of the phenotypic variation for each trait.Based on these markers,BnaA03g23490D, BnaC09g46370D,BnaA07g37150D,BnaA01g32590D, and BnaC09g37280D were identified as pleiotropic genes controlling multiple traits.These candidate genes illustrated the potential for the genetic understanding of yield and oil quality traits.Most importantly,these significant markers can be used for marker-assisted breeding of oilseed rape in different environments.

Identification of QTLs for Yield-Related Traits in the Recombinant Inbred Line Population Derived from the Cross Between a Synthetic Hexaploid Wheat-Derived Variety Chuanmai 42 and a Chinese Elite Variety Chuannong 16

Synthetic hexaploid wheat （SHW） represents a valuable source of new resistances to a range of biotic and abiotic stresses. A recombinant inbred line （RIL） population with 127 recombinant inbred lines derived from a SHW-derived variety Chuanmai 42 crossing with a Chinese spring wheat variety Chuannong 16 was used to map QTLs for agronomic traits including grain yield, grains per square meter, thousand-kernel weight, spikes per square meter, grain number per spike, grains weight per spike, and biomass yield. The population was genotyped using 184 simple-sequence repeat （SSR） markers and 34 sequence-related amplified polymorphism （SRAP） markers. Of 76 QTLs （LOD〉2.5） identified, 42 were found to have a positive effect from Chuanmai 42. The QTL QGy.saas-4D.2 associated with grain yield on chromosome 4D was detected in four of the six environments and the combined analysis, and the mean yield, across six environments, of individuals carrying the Chuanmai 42 allele at this locus was 8.9% higher than that of those lines carrying the Chuannong 16 allele. Seven clusters of the yield-coincident QTLs were detected on 1A, 4A, 3B, 5B, 4D, and 7D.

Optimal yield-related attributes of irrigated rice for high yield potential based on path analysis and stability analysis

Improvement of yield in rice(Oryza sativa L.) is vital for ensuring food security in China. Both rice breeders and growers need an improved understanding of the relationship between yield and yield-related traits. New indica cultivars(53 in 2007 and 48 in 2008) were grown in Taoyuan,Yunnan province, to identify important components contributing to yield. Additionally, two standard indica rice cultivars with similar yield potentials, II You 107(a large-panicle type) and Xieyou 107(a heavy-panicle type), were planted in Taoyuan, Yunnan province and Nanjing,Jiangsu province, from 2006 to 2008 to evaluate the stability of yield and yield-related attributes.Growth duration(GD), leaf area index(LAI), panicles per m2(PN), and spikelets per m2(SM) were significantly and positively correlated with grain yield(GY) over all years. Sequential path analysis identified PN and panicle weight(PW) as important first-order traits that influenced grain yield. All direct effects were significant, as indicated by bootstrap analysis. Yield potential varied greatly across locations but not across years. Plant height(PH), days from heading to maturity(HM), and grain weight(GW) were stable traits that showed little variation across sites or years, whereas GD(mainly the pre-heading period, PHP) and PN varied significantly across locations. To achieve a yield of 15 t ha-1, a cultivar should have a PH of 110–125 cm, a long GD with HM of approximately 40 days, a PN of 300–400 m-2, and a GW of 29–31 mg.

Response of phenology- and yield-related traits of maize to elevated temperature in a temperate region

Extreme high temperatures detrimental to maize production are projected to occur more frequently with future climate change.Phenology and yield-related traits were investigated under several levels of elevated temperature in two early-maturing hybrid cultivars:Junda 6(grown in northeastern China)and Chalok 1(grown in South Korea).They were cultivated in plastic houses in Suwon,Korea(37.27°N,126.99°E)held at target temperatures of ambient(AT),AT+1.5°C,AT+3°C,and AT+5°C at one sowing date in 2013 and three different sowing dates in 2014.Vegetative and reproductive growth durations showed variation depending on sowing date,experimental year,and cultivar.Growth duration tended to decrease,but not necessarily,with temperature elevation,but somewhat increased again above a certain temperature.High temperature-dependent variation was greater during grain filling than in the vegetative period before anthesis.Elevated temperature showed no significant effects on duration or peak dates of silking and anthesis,and thus on anthesis–silking interval.Grain yield tended to decrease with temperature elevation above ambient,showing a sharper linear decrease with mean growing season temperature increase in Junda 6 than in Chalok 1.The decrease in kernel number accounted for a much greater contribution to the yield reductions due to temperature elevation than did the decrease in individual kernel weight in both cultivars.Individual harvestable kernel weight was not significantly affected by temperature elevation treatments.Kernel number showed a linear decrease with mean growth temperature from early ear formation to early grain-filling stage,with Junda 6 showing a much severer decrease than Chalok 1.Kernel number reduction due to temperature elevation was attributable more to the decrease in differentiated ovule number than to the decrease in kernel set in Chalok 1,but largely to the decrease of kernel set in Junda 6.

Identifying Genomic Regions Conferring Morphological and Yield-related Traits in Soybean Based On A Four-way Recombinant Inbred Line Population

Improvement of seed yield of soybean(Glycine max(L.)Merr.)is generally achieved by combining morphological and yield-related traits,such as plant height(PH),node number on main stem(NN),pod number per plant(NP),seed number per plant(NS),100-seed weight(HSW)and seed weight per plant(SWPP).Identifying quantitative trait loci(QTLs)for morphological and yield-related traits is therefore important for breeding.In this study,a four-way recombinant inbred line population comprising 160 lines derived from the cross(Kenfeng14×Kenfeng15)×(Heinong48×Kenfeng19)was planted in five different environments and morphological and yield-related trait data were used to identify QTLs by the inclusive composite interval mapping method.Totally 38 QTLs for PH,40 QTLs for NN,26 QTLs for NP,10 QTLs for NS,26 QTLs for HSW and 49 QTLs for SWPP were detected in 125 genomic regions.Single QTLs explained 2.17%-14.60%,2.00%-10.04%,2.37%-9.77%,2.62%-8.61%,0.47%-6.51%and 0.14%-12.39%of the phenotypic variation for PH,NN,NP,NS,HSW and SWPP,respectively.Among these 125 genomic regions,120 were newly associated with morphological and yield-related traits.The results would facilitate the molecular breeding of morphological and yield-related traits in soybean.

Association mapping and domestication analysis to dissect genetic improvement process of upland cotton yield-related traits in China

Background: Cotton fiber yield is a complex trait,which can be influenced by multiple agronomic traits.Unravelling the genetic basis of cotton fiber yield-related traits contributes to genetic improvement of cotton.Results: In this study,503 upland cotton varieties covering the four breeding stages(BS1–BS4,1911–2011)in China were used for association mapping and domestication analysis.One hundred and forty SSR markers significantly associated with ten fiber yield-related traits were identified,among which,29 markers showed an increasing trend contribution to cotton yield-related traits from BS1 to BS4,and 26 markers showed decreased trend effect.Four favorable alleles of 9 major loci(R^(2)≥3)were strongly selected during the breeding stages,and the candidate genes of the four strongly selected alleles were predicated according to the gene function annotation and tissue expression data.Conclusions :The study not only uncovers the genetic basis of 10 cotton yield-related traits but also provides genetic evidence for cotton improvement during the cotton breeding process in China.

Development of Oryza rufipogon and O. sativa Introgression Lines and Assessment for Yield-related Quantitative Trait Loci

Introgression lines population was effectively used in mapping quantitative trait loci （QTLs）, identifying favorable genes, discovering hidden genetic variation, evaluating the action or interaction of QTLs in multiple conditions and providing the favorable experimental materials for plant breeding and genetic research. In this study, an advanced backcross and consecutive selfing strategy was used to develop introgression lines （ILs）, which derived from an accession of Oryza rufipogon Griff. collected from Yuanjiang County, Yunnan Province of China, as the donor, and an elite indica cultivar Teqing （O. sativa L.）, as the recipient. Introgression segments from O. rufipogon were screened using 179 polymorphic simple sequence repeats （SSR） markers in the genome of each IL. Introgressed segments carried by the introgression lines population contained 120 ILs covering the whole O. rufipogon genome. The mean number of homozygous O. rufipogon segments per introgression line was about 3.88. The average length of introgressed segments was approximate 25.5 cM, and about 20.8% of these segments had sizes less than 10 cM. The genome of each IL harbored the chromosomal fragments of O. rufipogon ranging from 0.54% to 23.7%, with an overall average of 5.79%. At each locus, the ratio of substitution of O. rufipogon alleles had a range of 1.67-9.33, with an average of 5.50. A wide range of alterations in morphological and yield-related traits were also found in the introgression lines population. Using single-point analysis, a total of 37 putative QTLs for yield and yield components were detected at two sites with 7%-20% explaining the phenotypic variance. Nineteen QTLs （51.4%） were detected at both sites, and the alleles from O. rufipogon at fifteen loci （40.5%） improved the yield and yield components in the Teqing background. These O. rufipogon-O, sativa introgression lines will serve as genetic materials for identifying and using favorable genes from common wild rice.

Analysis of QTLs for yield-related traits in Yuanjiang common wild rice (Oryza rufipogon Griff.)

Using an accession of common wild rice （Oryza rufipogon Griff.） collected from Yuanjiang County, Yunnan Province, China, as the donor and an elite cultivar 93-11, widely used in two-line indica hybrid rice production in China, as the recurrent parent, an advanced backcross populations were developed. Through genotyping of 187 SSR markers and investigation of six yield-related traits of two gen- erations （BC4F2 and BC4F4）, a total of 26 QTLs were detected by employing single point analysis and interval mapping in both genera-tions. Of the 26 QTLs, the alleles of 10 （38.5%） QTLs originating from O. rufipogon had shown a beneficial effect for yield-related traits in the 93-11 genetic background. In addition, five QTLs controlling yield and its components were newly identified, indicating that there are potentially novel alleles in Yuanjiang common wild rice. Three regions underling significant QTLs for several yield-related traits were detected on chromosome 1, 7 and 12. The QTL clusters were founded and corresponding agronomic traits of those QTLs showed highly significant correlation, suggesting the pleiotropism or tight linkage. Fine-mapping and cloning of these yield-related QTLs from wild rice would be helpful to elucidating molecular mechanism of rice domestication and rice breeding in the future.

Fine mapping of a major QTL qHYF_B06 for peanut yield

High yield is a major objective for peanut(Arachis hypogaea L.) breeding worldwide. However, fewer yield-related quantitative trait loci(QTL) have been reported in peanut than in other staple food crops such as rice(Oryza sativa), wheat(Triticum aestivum), and maize(Zea mays). This study aimed to identify stable major-effect QTL associated with pod yield per plant, hundred-pod weight for double-seeded pods,hundred-seed weight, shelling percentage, and pod number per plant, allowing us to predict candidate genes by means of transcriptome and genome sequencing. To this end, we used a population of recombinant inbred lines comprising 192 F9:11families derived from a JH6 × KX01-6 cross to construct a highresolution genetic map(1705.7 c M) consisting of 2273 polymorphic SNPs, with 0.75 c M(on average)between adjacent SNPs. We identified two high-confidence, yield-related QTL, qHYF_A08 and qHYF_B06, explaining 5.78%–31.40% of phenotypic variation and with LOD values of 5.10–24.48, in six environments. qHYF_A08 mainly explained the variation in shelling percentage, whereas qHYF_B06explained variation in hundred-pod weight and hundred-seed weight and accounted for 8.77%–31.40%of the variation in effective pod number per plant, pod number per plant, and shelling percentage. We narrowed down qHYF_B06 to an 890-kb interval using an advanced mapping population.Transcriptome and genome analyses revealed that only Arahy.129FS0 and Arahy.3R9A5K in the candidate mapping interval were differentially expressed between JH6 and KX01-6, with substantial structural variations in their promoter and coding regions. Genotypes of 208 peanut accessions determined using a diagnostic CAPS marker suggested that the two haplotypes of Arahy.3R9A5K were highly associated with hundred-seed weight and hundred-pod weight;this diagnostic CAPs marker could therefore be useful for selecting high-yielding lines during peanut breeding. Overall, our results provide valuable information for cloning alleles with favorable effects on peanut yield.

Major Gene Identification and Quantitative Trait Locus Mapping for YieldRelated Traits in Upland Cotton(Gossypium hirsutum L.)

Segregation analysis of the mixed genetic model of major gene plus polygene was used to identify the major genes for cotton yield-related traits using six generations P1, P2, F1, B1, B2, and F2 generated from the cross of Baimian 1 x TM-1. In addition to boll size and seed index, the major genes for the other five traits were detected： one each for seed yield, lint percentage, boll number, lint index; and two for lint yield. Quantitative trait locus/loci （QTL） mapping was performed in the F2 and F2：3 populations of above cross through molecular marker technology, and a total of 50 QTL （26 suggestive and 24 significant） for yield-related traits were detected. Four common QTL were discovered： qLP-3b（F2）/qLP-3（F2：3） and qLP-19b （F2）/qLP-19（F2：3） for lint percentage, qBN-17（F2）/qBN-17（F2：3） for boll number, and qBS-26b（F2）/qBS-26（F2：3） for boll size. Especially, qLP- 3b（Fz）/qLP-3（F2：3）, not only had LOD scores 〉3 but also exceeded the permutation threshold （5.13 and 5.29, respectively）, correspondingly explaining 23.47 and 29.55% of phenotypic variation. This QTL should be considered preferentially in marker assisted selection （MAS）. Segregation analysis and QTL mapping could mutually complement and verify, which provides a theoretical basis for genetic improvement of cotton yield-related traits by using major genes （QTL）.

qRgls1.06, a major QTL conferring resistance to gray leaf spot disease in maize

Gray leaf spot(GLS)caused by Cercospora zeae-maydis and C.zeina is an extremely devastating leaf disease that limits maize production annually.The use of GLS-resistant maize hybrids is the most cost-effective approach for reducing losses.Resistance to GLS is quantitatively inherited in maize(Zea mays L.)and further sources of resistance remain to be analyzed.Here,we detected qRgls1.06,a major quantitative trait locus for GLS resistance in bin 1.06 that explained approximately 55%of the phenotype variance.Fine mapping over 2 consecutive years localized qRgls1.06 to a 2.38-Mb region.Homozygous qRgls1.06^(WGR/WGR) plants in DZ01 background displayed higher GLS resistance and 100-grain weight than DZ01 plants.The GLS responses of several susceptible elite inbred lines were improved by the introduction of qRgls1.06 by marker-assisted backcrossing.Our findings extend the understanding of the genetic basis of resistance to GLS and provide a set of resistant germplasm for genetic improvement of resistance to GLS in maize.

Cloning and Expression Level Analysis of Two BnaANT Candidate Genes in Brassica napus

AINTEGUMENTA （ANT） gene has been proved to have a novel function on controlling the organ size and the seed weight in Arabidopsis thaliana, the research on its orthologous gene in Brassica napus will be of potential interest to elucidate the molecular mechanism of rapeseed development and increase the rapeseed output. A SMART cDNA library of the flower bud from B. napus cultivar Zhongshuang 9 was constructed, and the full-length cDNAs of two homologous BnaANT candidate genes were cloned by PCR in B. napus, namely BnaX.ANT.a （GenBank accession no. DQ211969） and BnaX.ANT.b （accession no. DQ211970）. They shared 87.1 and 83.4% amino acid identity with ANT of A. thaliana, respectively. There is 87.4% amino acid identity between BnaX.ANT.a and BnaX.ANT.b. By quantitative real-time PCR, the obviously differential expression levels of the two BnaANT candidate genes were detected in the 28 DAF seeds of the big and small grains B. napus lines, the expression abundance of BnaANT in the 28 DAF seed of 9311 was over three times compared to that of 9260, which indicates that these two BnaANT genes are also likely related to the floral organ size and the seed weight in B. napus.

Influence of Rind Hardness on Sugarcane Quality

Sugar recovery in a factory depends upon regular crushing and quality of varieties besides various other working parameters. Strategies for increased productivity and sugar recovery, varietal improvement can bring out through selection of parents to get maximum gains. These are governed by sugar content, fiber, rind hardness and cane yield. In order to assess the breeding value of different traits including rind hardness, study was undertaken at Sugarcane Research Institute, Shahjahanpur, during 2011-2012. Phenotypic difference and correlations among agronomic characters, basic quality characters and rind hardness in 120 segregating genotypes derived from contrast bi-parental cross (UP9530 × Co86011) of sugarcane cultivar were studied in this experiment. All the traits observed were found statistically significant among the mapping populations. The mean value of rind hardness was 5.91 with a range from 2.08 to 12.10 among 120 segregating population including their parents. Rind hardness (RHD) showed significant positive genotypic correlation with sugar yield-related traits viz;cane weight (CW, r = 0.325), cane yield (CYLD, r = 0.380) and sugar yield (SUYLD, r = 0.248). Significant negative correlation was observed for rind hardness with commercial cane sugar percent (CCS%, r = -0.388), number of millable cane (NMC, r = -0.216), hand refractometer brix %-October (HBR, r = -0.154), Brix% in juice (BR, r = -0.119) and Sucrose% in juice (SUC, r = -0.080). The simple correlation analysis showed that RHD, CW, CYLD and SUYLD showed significant positive correlation for agronomic characters, but for basic quality characters had negative correlations. Result indicated that rind hardness was found to have economically important trait due to showing significant positive and negative correlation with sugar yield and sugar content, respectively. The study also revealed that with the reduction in rind hardness, sugar recovery may increase. Present findings indicated that the rind hardness could play as a key role in the selection of elite genotypes in breeding program to develop high sugar, high yielding, erect, non-lodging, disease and insect resistant varieties. The details of the study were discussed in the manuscript.

Estimating wheat spike-leaf composite indicator(SLI)dynamics by coupling spectral indices and machine learning

The contribution of spike photosynthesis to grain yield(GY)has been overlooked in the accurate spectral prediction of yield.Thus,it’s essential to construct and estimate a yield-related phenotypic trait considering spike photosynthesis.Based on field and spectral reflectance data from 19 wheat cultivars under two nitrogen fertilization conditions in two years,our objectives were to(i)construct a yield-related phenotypic trait(spike–leaf composite indicator,SLI)accounting for the contribution of the spike to photosynthesis,(ii)develop a novel spectral index(enhanced triangle vegetation index,ETVI3)sensitive to SLI,and(iii)establish and evaluate SLI estimation models by integrating spectral indices and machine learning algorithms.The results showed that SLI was sensitive to nitrogen fertilizer and wheat cultivar variation as well as a better predictor of yield than the leaf area index.ETVI3 maintained a strong correlation with SLI throughout the growth stage,whereas the correlations of other spectral indices with SLI were poor after spike emergence.Integrating spectral indices and machine learning algorithms improved the estimation accuracy of SLI,with the most accurate estimates of SLI showing coefficient of determination,root mean square error(RMSE),and relative RMSE values of 0.71,0.047,and 26.93%,respectively.These results provide new insights into the role of fruiting organs for the accurate spectral prediction of GY.This high-throughput SLI estimation approach can be applied for wheat yield prediction at whole growth stages and may be assisted with agronomical practices and variety selection.