ADP‐glucose pyrophosphorylase genes,associated with kernel weight,underwent selection during wheat domestication and breeding

ADP‐glucose pyrophosphorylase, comprising two small subunits and two large subunits, is considered a key enzyme in the endosperm starch synthesis pathway in wheat (Triticum aestivum L.). Two genes, TaAGP‐S1‐7A and TaAGP‐L‐1B, were investigated in this study. Haplotypes of these genes were associated with thousand kernel weight (TKW) in different populations. Mean TKWs of favoured haplotypes were significantly higher than those of nonfavoured ones. Two molecular markers developed to distinguish these haplotypes could be used in molecular breeding. Frequencies of favoured haplotypes were dramatically increased in cultivars released in China after the 1940s. These favoured haplotypes were also positively selected in six major wheat production regions globally. Selection of AGP‐S1 and AGP‐L‐1B in wheat mainly occurred during and after hexaploidization. Strong additive effects of the favoured haplotypes of with other genes for starch synthesis were also detected in different populations.     

Identifying loci influencing 1,000-kernel weight in wheat by microsatellite screening for evidence of selection during breeding

Chinese wheat mini core collection (262 accessions) was genotyped at 531 microsatellite loci representing a mean marker density of 5.1 cM. One-thousand-kernel weights (TKW) of lines were measured in five trials (three environments in four growing seasons). Structure analysis based on 42 unlinked SSR loci indicated that the materials formed two sub-populations, viz., landraces and modern varieties. A large difference in TKW (7.08 g, P<0.001) was found between the two sub-groups. Therefore, TKW is a major yield component that was improved in the past 6 decades; it increased from a mean 31.5 g in the 1940s to 44.64 g in the 2000s, representing a 2.19 g increase in each decade. Analyses based on a mixed linear model (MLM), population structure (Q) and relative kinship (K) revealed 22 SSR loci that were significantly associated with mean TKW (MTKW) of the five trials estimated by the best linear unbiased predictor (BLUP) method. They were mainly distributed on chromosomes of homoeologous groups 1, 2, 3, 5 and 7. Six loci, cfa2234-3A, gwm156-3B, barc56-5A, gwm234-5B, wmc17-7A and cfa2257-7A individually explained more than 11.84% of the total phenotypic variation. Favored alleles for breeding at the 22 loci were inferred according to their estimated effects on MTKW based on mean difference of varieties grouped by genotypes. Statistical simulation showed that these favored alleles have additive genetic effects. Frequency changes of alleles at loci associated with TKW are much more dramatic than those at neutral loci between the sub-groups. The numbers of favored alleles in modern varieties indicate there is still considerable genetic potential for their use as markers for genome selection of TKW in wheat breeding. Alleles that can be used globally to increase TKW were inferred according to their distribution by latitude and frequency of changes between landraces and the modern varieties.     

Assessment of differences in morphological and physiological leaf lodging characteristics between two cultivars of Hippeastrum rutilum

Kernel weight and morphology are important traits affecting cereal yields and quality. Dissecting the genetic basis of thousand kernel weight (TKW) and its related traits is an effective method to improve wheat yield. In this study, we performed quantitative trait loci (QTL) analysis using recombinant inbred lines derived from the cross ‘PuBing3228 × Gao8901’ (PG-RIL) to dissect the genetic basis of kernel traits. A total of 17 stable QTLs related to kernel traits were identified, notably, two stable QTLs QTkw.cas-1A.2 and QTkw.cas-4A explained the largest portion of the phenotypic variance for TKW and kernel length (KL), and the other two stable QTLs QTkw.cas-6A.1 and QTkw.cas-7D.2 contributed more effects on kernel width (KW). Conditional QTL analysis revealed that the stable QTLs for TKW were mainly affected by KW. The QTLs QTkw.cas-7D.2 and QKw.cas-7D.1 associated with TKW and KW were delimited to the physical interval of approximately 3.82 Mb harboring 47 candidate genes. Among them, the candidate gene TaFT-D1 had a 1 bp insertions/deletion (InDel) within the third exon, which might be the reason for diversity in TKW and KW between the two parents. A Kompetitive Allele-Specific PCR (KASP) marker of TaFT-D1 allele was developed and verified by PG-RIL and a natural population consisted of 141 cultivar/lines. It was found that the favorable TaFT-D1 (G)-allele has been positively selected during Chinese wheat breeding. Thus, these results can be used for further positional cloning and marker-assisted selection in wheat breeding programs. Seventeen stable QTLs related to kernel traits were identified. The stable QTLs for thousand kernel weight were mainly affected by kernel width. TaFT-D1 could be the candidate gene for QTLs QTkw.cas-7D.2 and QKw.cas-7D.1.     

Single nucleotide polymorphism tightly linked to a major QTL on chromosome 7A for both kernel length and kernel weight in wheat

Thousand-kernel weight (TKW) is one of the major components of grain yield in wheat (Triticum aestivum). Identifying major quantitative trait loci (QTLs) for TKW and developing effective markers are prerequisite for success in marker-assisted selection (MAS) to improve wheat yield through breeding. This study mapped a major QTL, designated as TaTKW-7AL, for increasing TKW on the long arm of chromosome 7A of ‘Clark’ to a 1.3-cM interval between single nucleotide polymorphism (SNP) markers IWB13913 and IWA5913. This QTL explained 19.7 % of the phenotypic variation for TKW. A QTL for increasing kernel length (KL), one of the major components of TKW, was mapped in the same interval as TaTKW-7AL, suggesting that increased TKW by the QTL in ‘Clark’ is most likely due to the increased KL. Association analysis on a diversity panel of 200 US winter wheat accessions also identified a haplotype of three SNP markers (IWB13913, IWB6693 and IWA5913) that were tightly associated with the both KL and TKW. The analysis of allele frequencies of the haplotype in the diversity panel suggested that the favorable allele of TaTKW-7AL has not been strongly selected for in practice and has potential to be used to improve grain yield in US hard winter wheat breeding. Two user-friendly flanking KASPar markers, IWB13913 and IWA5913, were developed for MAS of TaTKW-7AL.     

TaTPP-7A positively feedback regulates grain filling and wheat grain yield through T6P-SnRK1 signalling pathway and sugar–ABA interaction

Grain size and filling are two key determinants of grain thousand-kernel weight (TKW) and crop yield, therefore they have undergone strong selection since cereal was domesticated. Genetic dissection of the two traits will improve yield potential in crops. A quantitative trait locus significantly associated with wheat grain TKW was detected on chromosome 7AS flanked by a simple sequence repeat marker of Wmc17 in Chinese wheat 262 mini-core collection by genome-wide association study. Combined with the bulked segregant RNA-sequencing (BSR-seq) analysis of an F₂ genetic segregation population with extremely different TKW traits, a candidate trehalose-6-phosphate phosphatase gene located at 135.0 Mb (CS V1.0), designated as TaTPP-7A, was identified. This gene was specifically expressed in developing grains and strongly influenced grain filling and size. Overexpression (OE) of TaTPP-7A in wheat enhanced grain TKW and wheat yield greatly. Detailed analysis revealed that OE of TaTPP-7A significantly increased the expression levels of starch synthesis- and senescence-related genes involved in abscisic acid (ABA) and ethylene pathways. Moreover, most of the sucrose metabolism and starch regulation-related genes were potentially regulated by SnRK1. In addition, TaTPP-7A is a crucial domestication- and breeding-targeted gene and it feedback regulates sucrose lysis, flux, and utilization in the grain endosperm mainly through the T6P-SnRK1 pathway and sugar–ABA interaction. Thus, we confirmed the T6P signalling pathway as the central regulatory system for sucrose allocation and source–sink interactions in wheat grains and propose that the trehalose pathway components have great potential to increase yields in cereal crops.     

A genome-wide association study of wheat yield and quality-related traits in southwest China

Wheat (Triticum aestivum L.) is one of the most productive and important crops and its yield potential and quality characteristics are tightly linked with the global food security. In this study, genome-wide association study (GWAS) was conducted for yield and quality-related traits. Based on the high-density wheat 90K Illumina iSelect SNP Array, 192 bread wheat lines from southwest China, including 25 synthetic hexaploid wheat lines, 80 landraces, and 87 cultivars were analyzed. Association analysis results indicated that there were 57, 27, 30, and 34 SNPs associated with plant height (PH), grain protein content (GPC), thousand kernel weight (TKW), and SDS sedimentation volume (SSV) have been detected, respectively. Then, integrating RNA-Seq with bioinformatics analysis, 246 candidate genes (102 for GPC, 52 for TKW, and 92 for SSV) were found. Further analysis indicated that one up-regulated and two down-regulated expression genes affect GPC. Additionally, two haplotypes significantly affecting PH were detected in a 2.2-Mb genome region encompassing a gene which encoded an ubiquitin-specific protease, TaUBP24. The functional markers of TaUBP24 have been developed, which could be used for marker-assisted selection to improve wheat quality and yield.     

Terminal Heat Tolerance in Bread Wheat Determined by Agronomical Traits and SSR Markers

Since global warming affects wheat cropping systems, more has yet to be indicated on the parameters, which control terminal heat tolerance, and severely influence wheat (Triticum aestivum L.) productivity. Identification of tolerant wheat genotypes by heat tolerance-linked molecular markers is a rapid and cost-effective screening tool in plant breeding. Accordingly, in a four-year field experiment (2015–2019), 44 wheat genotypes were selected out of 100 genotypes, and were examined in timely and late planting (mid-January resulting in heat stress). Stress decreased yield components, including 1000-kernel weight (TKW), grains per spike, and plants per square meter, and the physiological traits, including days to heading and days to maturity, grain filling duration, and greenness, and eventually decreased grain yield up to?~?28%. The early maturity genotypes resulted in higher yields under stress conditions by a stress-avoidance mechanism. Among 14 SSR markers, GWM577 was positively correlated with yield, and WMS3062, GWM261, and WMS1025 had positive correlations with longevity under stress. Accordingly, WMS3062 and GWM261 can be used to determine high yield and early maturity genotypes. Furthermore, GWM114 showed a positive correlation with TKW, indicating their usefulness for grouping wheat genotypes and for identifying heat-related markers. Since the crossing of the genetically distant genotypes can create more diverse populations, the results could be applied to plan breeding projects to establish more diverse populations for different chromosomal locations and traits under heat stress conditions. Moreover, our findings demonstrated that the morphological and molecular analyses could be useful for describing wheat genetic variation of heat tolerance.

     

A haplotype block associated with thousand-kernel weight on chromosome 5DS in common wheat (Triticum aestivum L.)

Spike number per unit area, number of grains per spike, and thousand-kernel weight (TKW) are important yield components for wheat (Triticum aestivum L.). TKW has the highest heritability among the thre...     

Wheat kernel dimensions: how do they contribute to kernel weight at an individual QTL level?

Kernel dimensions (KD) contribute greatly to thousand-kernel weight (TKW) in wheat. In the present study, quantitative trait loci (QTL) for TKW, kernel length (KL), kernel width (KW) and kernel diameter ratio (KDR) were detected by both conditional and unconditional QTL mapping methods. Two related F(8:9) recombinant inbred line (RIL) populations, comprising 485 and 229 lines, respectively, were used in this study, and the trait phenotypes were evaluated in four environments. Unconditional QTL mapping analysis detected 77 additive QTL for four traits in two populations. Of these, 24 QTL were verified in at least three trials, and five of them were major QTL, thus being of great value for marker assisted selection in breeding programmes. Conditional QTL mapping analysis, compared with unconditional QTL mapping analysis, resulted in reduction in the number of QTL for TKW due to the elimination of TKW variations caused by its conditional traits; based on which we first dissected genetic control system involved in the synthetic process between TKW and KD at an individual QTL level. Results indicated that, at the QTL level, KW had the strongest influence on TKW, followed by KL, and KDR had the lowest level contribution to TKW. In addition, the present study proved that it is not all-inclusive to determine genetic relationships of a pairwise QTL for two related/causal traits based on whether they were co-located. Thus, conditional QTL mapping method should be used to evaluate possible genetic relationships of two related/causal traits.     

TaCKX6-D1, the ortholog of rice OsCKX2, is associated with grain weight in hexaploid wheat

The cytokinin oxidase/dehydrogenase (CKX) gene plays a principal role in controlling cytokinin levels and has been shown to be a major quantitative trait locus (QTL) affecting grain number in rice. However, the function and evaluation of the haplotypes of the wheat CKX gene have yet to be illustrated. In this study, TaCKX6-D1, a wheat ortholog of rice OsCKX2, was cloned and its haplotype variants were determined to be significantly associated with the 1000-grain weight on the basis of linkage mapping, association analysis and gene expression analysis. Five TaCKX6-D1 haplotypes, designated a-e, were identified. An indel marker was developed to identify haplotype a, which was associated with higher grain weight. Haplotype a showed decreased expression relative to haplotype b in seeds at 8 d after pollination. Sequence variations among modern cultivars, landraces and wild species suggest a significant domestication signature at the TaCKX6-D1 locus in Chinese wheat germplasm. TaCKX6-D1 may serve as a useful gene for the breeding of high-yielding wheat. A strategy for allele mining and utilization of TaCKX6-D1 was proposed. Our study also sheds light on the mechanisms of grain development and domestication of wheat, as well as the functional divergence of orthologs in comparative genomics.     

Rye Pm8 and wheat Pm3 are orthologous genes and show evolutionary conservation of resistance function against powdery mildew

The improvement of wheat through breeding has relied strongly on the use of genetic material from related wild and domesticated grass species. The 1RS chromosome arm from rye was introgressed into wheat and crossed into many wheat lines, as it improves yield and fungal disease resistance. Pm8 is a powdery mildew resistance gene on 1RS which, after widespread agricultural cultivation, is now widely overcome by adapted mildew races. Here we show by homology‐based cloning and subsequent physical and genetic mapping that Pm8 is the rye orthologue of the Pm3 allelic series of mildew resistance genes in wheat. The cloned gene was functionally validated as Pm8 by transient, single‐cell expression analysis and stable transformation. Sequence analysis revealed a complex mosaic of ancient haplotypes among Pm3‐ and Pm8‐like genes from different members of the Triticeae. These results show that the two genes have evolved independently after the divergence of the species 7.5 million years ago and kept their function in mildew resistance. During this long time span the co‐evolving pathogens have not overcome these genes, which is in strong contrast to the breakdown of Pm8 resistance since its introduction into commercial wheat 70 years ago. Sequence comparison revealed that evolutionary pressure acted on the same subdomains and sequence features of the two orthologous genes. This suggests that they recognize directly or indirectly the same pathogen effectors that have been conserved in the powdery mildews of wheat and rye.     

Kernel weight per spike: what contributes to it at the individual QTL level?

Spike length (SL), spikelet number (SPN) per spike, kernel number per spike (KNPS), and thousand-kernel weight (TKW) have strong genetic associations with kernel weight per spike (KWPS) in wheat. To investigate their genetic relationships at the individual quantitative trait locus (QTL) level, both unconditional and conditional QTL mapping for KWPS with respect to SL, SPN, KNPS, and TKW were conducted. Two related F_8:9 recombinant inbred line populations, comprising 485 and 229 lines, respectively, were used. The trait phenotypic performances of each population were evaluated in four different environments. Unconditional QTL mapping analysis identified 22 putative additive QTL for KWPS, five of which were stable QTL, and only QKwps-WJ-1B.2 showed significant additive-by-environment interaction effects. In comparison with unconditional QTL mapping analysis, conditional QTL mapping analysis indicated that, at the QTL level, KNPS and TKW contributed more to KWPS than did SL and SPN. Any unconditional QTL for KWPS detected in this study was associated with at least one of its four related traits. The present study will provide assistance in the understanding of the genetic relationships between KWPS and its related traits.     

Quantitative trait loci mapping reveals pleiotropic effect for grain iron and zinc concentrations in wheat

Malnutrition because of the deficiency of minerals such as iron (Fe) and zinc (Zn) afflicts over 2 billion people worldwide. Wheat is a major staple crop, providing 20% of dietary energy and protein consumption worldwide. Breeding wheat with elevated levels of grain Zn and Fe concentrations (GZn and GFe) represents a significant opportunity to increase the intake of these micronutrients for the resource poor people who depend on it as a major source of dietary energy. Synthetic hexaploid wheats (SHWs) have large genetic variation for GZn and GFe, which can be exploited for developing wheat varieties with higher concentrations of these minerals. The objective of this study was to localise genomic regions associated with GZn and GFe, thousand kernel weight (TKW) and test weight (TW) in a mapping population derived from the cross of Seri M82 and the SHW CWI76364. Major quantitative trait loci (QTL) on chromosome 4BS were detected for GZn and GFe; the QTL explained up to 19.6% of the total phenotypic variation for GZn and showed pleiotropic effects on GFe. This indicates that simultaneous improvement of GZn and GFe is feasible. Three and five QTL for TW and TKW were detected, respectively. One of the QTL for TKW was also located on chromosome 4BS. Positive correlations between plant height and GZn/GFe were observed. The 4BS QTL is of great interest for breeding biofortified wheat by means of marker‐assisted selection.     

Association and Validation of Yield-Favored Alleles in Chinese Cultivars of Common Wheat (Triticumaestivum L.)

Common wheat is one of the most important crops in China, which is the largest producer in the world. A set of 230 cultivars was used to identify yield-related loci by association mapping. This set was tested for seven yield-related traits, viz. plant height (PH), spike length (SL), spikelet number per spike (SNPS), kernel number per spike (KNPS), thousand-kernel weight (TKW), kernel weight per spike (KWPS), and sterile spikelet number (SSN) per plant in four environments. A total of 106 simple sequence repeat (SSR) markers distributed on all 21 chromosomes were used to screen the set. Twenty-one and 19 of them were associated with KNPS and TKW, respectively. Association mapping detected 73 significant associations across 50 SSRs, and the phenotypic variation explained (R²) by the associations ranged from 1.54 to 23.93%. The associated loci were distributed on all chromosomes except 4A, 7A, and 7D. Significant and potentially new alleles were present on 8 chromosomes, namely1A, 1D, 2A, 2D, 3D, 4B, 5B, and 6B. Further analysis showed that genetic effects of associated loci were greatly influenced by association panels, and the R² of crucial loci were lower in modern cultivars than in the mini core collection, probably caused by strong selection in wheat breeding. In order to confirm the results of association analysis, yield-related favorable alleles Xgwm135-1A₁₃₈, Xgwm337-1D₁₈₆, Xgwm102-2D₁₄₄, and Xgwm132-6B₁₂₈ were evaluated in a double haploid (DH) population derived from Hanxuan10 xLumai14.These favorable alleles that were validated in various populations might be valuable in breeding for high-yield.     

不同生态环境与播种期下小麦籽粒品质变异规律的研究

选用 6个不同品质类型小麦品种在 4个生态点进行分期播种试验, 系统分析了不同生态环境下小麦籽粒产量与品质的变异特征及其与主要气候生态因子间的关系。结果表明 :生态点、品种以及地点×品种互作对籽粒产量、千粒重、蛋白质、湿面筋和淀粉含量、沉降值与降落值的影响均达到显著水平 ;不同播种期处理对产量与淀粉含量的影响达到极显著水平, 而播种期×品种互作对千粒重、降落值、淀粉含量及沉降值的效应达到显著水平 ;地点×播种期×品种互作仅对产量、湿面筋、淀粉含量与沉降值有显著的影响。在 4个不同生态点中, 南京点的籽粒蛋白质与湿面筋含量最低, 但淀粉含量最高 ;徐州点的产量和千粒重最大 ;泰安点的蛋白质含量与湿面筋含量最高, 沉降值最小 ;保定点的产量、千粒重最小, 但沉降值最大。不同播种期处理下, 适播与晚播的籽粒蛋白质含量、湿面筋含量、淀粉含量、沉降值与降落值都显著高于早播, 而早播期的产量和千粒重最大。各小麦品种在不同地点与播种期下产量与品质性状的变异中以降落值的变异系数为最大, 淀粉的变异为最小。开花至成熟期的日均温与淀粉含量呈线性负相关, 与产量、蛋白质和湿面筋含量及降落值呈二次曲线相关关系 ;日温差与产量和千粒重呈二次曲线相关关系, 籽粒蛋白质和湿面筋含量、沉降值及降落值则随昼温差的增加线性递增 ;开花至成熟期降雨量与产量、千粒重都呈现先升后降的二次曲线相关关系, 而与籽粒蛋白质含量和降落值呈现线性负相关关系 ;籽粒蛋白质和湿面筋含量与降落值随开花至成熟期的累计日照时数呈现线性正相关关系。     

Fitness analyses of Arabidopsis thaliana mutants depleted of FtsH metalloproteases and characterization of three FtsH6 deletion mutants exposed to high light stress, senescence and chilling

Glutamine synthetase (GS) plays a key role in the growth, nitrogen (N) use and yield potential of cereal crops. Investigating the haplotype variation of GS genes and its association with agronomic traits may provide useful information for improving wheat N-use efficiency and yield. We isolated the promoter and coding region sequences of the plastic glutamine synthetase isoform (GS2) genes located on chromosomes 2A, 2B and 2D in bread wheat. By analyzing nucleotide sequence variations of the coding region, two, six and two haplotypes were distinguished for TaGS2-A1 (a and b), TaGS2-B1 (a-f) and TaGS2-D1 (a and b), respectively. By analyzing the frequency data of different haplotypes and their association with N use and agronomic traits, four major and favorable TaGS2 haplotypes (A1b, B1a, B1b, D1a) were revealed. These favorable haplotypes may confer better seedling growth, better agronomic performance, and improved N uptake during vegetative growth or grain N concentration. Our data suggest that certain TaGS2 haplotypes may be valuable in breeding wheat varieties with improved agronomic performance and N-use efficiency.     

Haplotype variation of Green Revolution gene Rht-D1 during wheat domestication and improvement

Green Revolution made a substantial contribution to wheat yields worldwide in the 1960 s and 1970 s. It is of great importance to analyze the haplotype variation of Rht-D1, the Green Revolution gene, during wheat(Triticum aestivum L.)domestication and breeding to understand its evolution and function in wheat breeding history. In this study, the Rht-D1 and its flanking regions were sequenced and single nucleotide polymorphisms were detected based on a panel of 45 accessions of Aegilops tauschii, 51 accessions of landraces and 80 accessions of commercial varieties. Genetic diversity in the wild accessions was much higher than that in the varieties and higher than that reported previously. Seven haplotypes(Hapl I to Hapl VII) of Rht-D1 were identified and their evolutionary relationships were proposed. In addition to the well-known Green Revolution allele Rht-D1b, Hapl VII(an allele Rht-D1k) was identified in early breeding varieties, which reduced plant height by 16%. The results suggested that Rht-D1k had been used in breeding before the Green Revolution and made a great contribution to wheat production worldwide.Based on the breeding history and molecular evidence, we proposed that the wheat Green Revolution in China and International Maize and Wheat Improvement Center(CIMMYT)occurred independently.     

春小麦千粒重相关性状的QTL定位及其耐热性分析

春小麦灌浆中后期正逢高温天气,适于发掘与耐高温相关的QTL.本研究利用春小麦Avocet/Sujata重组自交系群体,自2016-2018年在石家庄藁城区和天津两地4个环境下种植,进行千粒重(TKW)、粒长(KL)和粒宽(KW)等性状QTL定位,探讨这些QTL与灌浆期高温和品种适应性的关系.结果显示:在4个环境下共检测...     

QTL for yield and associated traits in the Seri/Babax population grown across several environments in Mexico, in the West Asia, North Africa, and South Asia regions

Heat and drought adaptive quantitative trait loci (QTL) in a spring bread wheat population resulting from the Seri/Babax cross designed to minimize confounding agronomic traits have been identified previously in trials conducted in Mexico. The same population was grown across a wide range of environments where heat and drought stress are naturally experienced including environments in Mexico, West Asia, North Africa (WANA), and South Asia regions. A molecular genetic linkage map including 475 marker loci associated to 29 linkage groups was used for QTL analysis of yield, days to heading (DH) and to maturity (DM), grain number (GM2), thousand kernel weight (TKW), plant height (PH), canopy temperature at the vegetative and grain filling stages (CTvg and CTgf), and early ground cover. A QTL for yield on chromosome 4A was confirmed across several environments, in subsets of lines with uniform allelic expression of a major phenology QTL, but not independently from PH. With terminal stress, TKW QTL was linked or pleiotropic to DH and DM. The link between phenology and TKW suggested that early maturity would favor the post—anthesis grain growth periods resulting in increased grain size and yields under terminal stress. GM2 and TKW were partially associated with markers at different positions suggesting different genetic regulation and room for improvement of both traits. Prediction accuracy of yield was improved by 5 % when using marker scores of component traits (GM2 and DH) together with yield in multiple regression. This procedure may provide accumulation of more favorable alleles during selection.