Achieving yield gains in wheat

Wheat provides 20% of calories and protein consumed by humans. Recent genetic gains are <1% per annum (p.a.), insufficient to meet future demand. The Wheat Yield Consortium brings expertise in photosynthesis, crop adaptation and genetics to a common breeding platform. Theory suggest radiation use efficiency (RUE) of wheat could be increased ～50%; strategies include modifying specificity, catalytic rate and regulation of Rubisco, up-regulating Calvin cycle enzymes, introducing chloroplast CO(2) concentrating mechanisms, optimizing light and N distribution of canopies while minimizing photoinhibition, and increasing spike photosynthesis. Maximum yield expression will also require dynamic optimization of source: sink so that dry matter partitioning to reproductive structures is not at the cost of the roots, stems and leaves needed to maintain physiological and structural integrity. Crop development should favour spike fertility to maximize harvest index so phenology must be tailored to different photoperiods, and sensitivity to unpredictable weather must be modulated to reduce conservative responses that reduce harvest index. Strategic crossing of complementary physiological traits will be augmented with wide crossing, while genome-wide selection and high throughput phenotyping and genotyping will increase efficiency of progeny screening. To ensure investment in breeding achieves agronomic impact, sustainable crop management must also be promoted through crop improvement networks.     

Elucidating the genetic basis of biomass accumulation and radiation use efficiency in spring wheat and its role in yield potential

One of the major challenges for plant scientists is increasing wheat (Triticum aestivum) yield potential (YP). A significant bottleneck for increasing YP is achieving increased biomass through optimization of radiation use efficiency (RUE) along the crop cycle. Exotic material such as landraces and synthetic wheat has been incorporated into breeding programmes in an attempt to alleviate this; however, their contribution to YP is still unclear. To understand the genetic basis of biomass accumulation and RUE, we applied genome‐wide association study (GWAS) to a panel of 150 elite spring wheat genotypes including many landrace and synthetically derived lines. The panel was evaluated for 31 traits over 2 years under optimal growing conditions and genotyped using the 35K wheat breeders array. Marker‐trait association identified 94 SNPs significantly associated with yield, agronomic and phenology‐related traits along with RUE and final biomass (BM_PM) at various growth stages that explained 7%–17% of phenotypic variation. Common SNP markers were identified for grain yield, BM_PM and RUE on chromosomes 5A and 7A. Additionally, landrace and synthetic derivative lines showed higher thousand grain weight (TGW), BM_PM and RUE but lower grain number (GM2) and harvest index (HI). Our work demonstrates the use of exotic material as a valuable resource to increase YP. It also provides markers for use in marker‐assisted breeding to systematically increase BM_PM, RUE and TGW and avoid the TGW/GM2 and BM_PM/HI trade‐off. Thus, achieving greater genetic gains in elite germplasm while also highlighting genomic regions and candidate genes for further study.     

Sink-limitation to yield and biomass: a summary of some investigations in spring wheat

Yield potential can be expressed as a product of light interception, radiation use efficiency (RUE), and the partitioning of biomass to grain yield, or harvest index (HI). Traits related to early or late light interception have not been shown to be associated with genetic improvement of spring wheat yield in favourable environments. It is, however, well established that yield improvement is largely a result of increased HI, although the most recent studies comparing genetic progress in HI over time in spring wheat indicate that it has not made any additional progress since the mid 1980s. These observations suggest that future genetic progress in yield will most likely be achieved by focusing on constraints to RUE. Considering the possibility that RUE may be influenced indirectly by sink limitation, it is apparent that biomass may be increased by increasing grain number, for example. Experiments with high yielding spring wheat lines containing the alien translocation 7DL.7Ag showed increased grains m‐2 (15%), yield (12%), and biomass (9%) compared with controls. The translocation was also associated with a larger investment in spike mass at anthesis (15%), more grains/spike (10%), and increased flag‐leaf photosynthetic rate during grain‐filling (20%). The data suggest that increased biomass in 7DL.7Ag lines was due to significantly increased RUE post‐anthesis, as a result of a larger kernel number (sink) that increased the demand for photosynthesis during grain‐filling. The hypothesis that increased photosynthesis and RUE may respond directly to a larger number of grains/spike was tested experimentally by imposing a light treatment during boot stage. The treatment was associated with a small increase (5%) in the proportion of biomass invested in spike mass at anthesis, reflected by on average three extra grains/spike at maturity. The treatment was associated with 25% more yield and 22% more biomass than controls, while carbon assimilation rate measured on flag‐leaves during grainfilling was 10% higher than controls. The results suggest that RUE can be increased indirectly by increasing sink strength and that the current yield limiting process in spring wheat is the determination of kernel number. Experimental data are presented on how spike fertility may be increased through breeding, for example by introgression of the multi‐ovary trait to increase grain number per spikelet. In addition, results of analysis of the physiological bases of genotype × year interaction in high yield environments are presented in the context of how such information can provide a focus for genetic studies of sink limitation.     

Raising yield potential of wheat. III. Optimizing partitioning to grain while maintaining lodging resistance

A substantial increase in grain yield potential is required, along with better use of water and fertilizer, to ensure food security and environmental protection in future decades. For improvements in photosynthetic capacity to result in additional wheat yield, extra assimilates must be partitioned to developing spikes and grains and/or potential grain weight increased to accommodate the extra assimilates. At the same time, improvement in dry matter partitioning to spikes should ensure that it does not increase stem or root lodging. It is therefore crucial that improvements in structural and reproductive aspects of growth accompany increases in photosynthesis to enhance the net agronomic benefits of genetic modifications. In this article, six complementary approaches are proposed, namely: (i) optimizing developmental pattern to maximize spike fertility and grain number, (ii) optimizing spike growth to maximize grain number and dry matter harvest index, (iii) improving spike fertility through desensitizing floret abortion to environmental cues, (iv) improving potential grain size and grain filling, and (v) improving lodging resistance. Since many of the traits tackled in these approaches interact strongly, an integrative modelling approach is also proposed, to (vi) identify any trade-offs between key traits, hence to define target ideotypes in quantitative terms. The potential for genetic dissection of key traits via quantitative trait loci analysis is discussed for the efficient deployment of existing variation in breeding programmes. These proposals should maximize returns in food production from investments in increased crop biomass by increasing spike fertility, grain number per unit area and harvest index whilst optimizing the trade-offs with potential grain weight and lodging resistance.     

栽培模式对冬小麦光能利用和产量的影响

以‘泰农18’小麦为材料,于2012—2013年进行大田试验,设置当地农民习惯栽培模式(FP)、超高产栽培模式(SH)和高产高效栽培模式(HH)3个处理,研究了不同栽培模式对小麦光能截获量、光能利用效率、干物质积累量、收获指数、籽粒产量和肥料偏生产力的影响.结果表明: SH模式小麦全生育期的光能截获量、光能利用效率、干物质积累量和籽粒产量显著高于FP模式.相对于FP模式,虽然HH模式的小麦全生育期光能截获量较低,但其光能利用效率、干物质积累量及收获指数均显著提高,从而使其籽粒产量显著提高.相对于SH模式,虽然HH模式的籽粒产量在高、低肥力水平下分别降低3.8%和2.8%,而氮、磷、钾肥的偏生产力在两肥力水平下分别提高26.4%、68.5%、92.6%.在本试验条件下,综合考虑籽粒产量和养分利用效率,以“降肥、增密、延播”为技术特点的高产高效栽培模式为推荐的优化栽培模式.     

小麦地方品种繁殖更新过程中的遗传多样性变化分析

采用分别保存于长期库及中期库的3个小麦地方品种的6份材料,进行了9项农艺性状及35个与农艺性状相关的微卫星标记检测,每份材料选取30个单株进行遗传多样性与遗传结构分析。结果表明:(1)更新前,3个小麦地方品种均为遗传异质性群体,在SSR位点上的异质度分别为57.14%、48.57%和5.71%。(2)在农艺性状表现上,只有温泉小麦3在株高和穗粒数上更新后比更新前显著增加,其他材料无显著差异。(3)在SSR位点多态性表现上,3个品种在更新后均发生了遗传多样性变化,8个与粒重、产量、生育期性状相关位点存在等位位点丢失现象,其中2个与粒重、生育期相关位点频率变化显著。(4)综合农艺性状调查与SSR分子标记检测结果发现,3个品种更新前后在多样性指数上无显著差异,遗传分化系数Gst分别为0.0269、0.0324和0.0380,即更新前后遗传差异分别为2.69%、3.24%和3.80%。上述结果建议,经繁殖更新的小麦种质资源能够比较完好地保持其遗传多样性和遗传结构。对于遗传异质性小麦地方品种在繁殖更新后存在遗传多样性丢失的危险,为了保证更新前后的遗传完整性,建议在繁殖更新过程中每个品种至少应保持300个单株的群体。     

Genetic dissection of drought and heat‐responsive agronomic traits in wheat

High yield and wide adaptation are principal targets of wheat breeding but are hindered by limited knowledge on genetic basis of agronomic traits and abiotic stress tolerances. In this study, 277 wheat accessions were phenotyped across 30 environments with non‐stress, drought‐stressed, heat‐stressed, and drought‐heat‐stressed treatments and were subjected to genome‐wide association study using 395 681 single nucleotide polymorphisms. We detected 295 associated loci including consistent loci for agronomic traits across different treatments and eurytopic loci for multiple abiotic stress tolerances. A total of 22 loci overlapped with quantitative trait loci identified by biparental quantitative trait loci mapping. Six loci were simultaneously associated with agronomic traits and abiotic stress tolerance, four of which fell within selective sweep regions. Selection in Chinese wheat has increased the frequency of superior marker alleles controlling yield‐related traits in the four loci during past decades, which conversely diminished favourable genetic variation controlling abiotic stress tolerance in the same loci; two promising candidate paralogous genes colocalized with such loci, thereby providing potential targets for studying the molecular mechanism of stress tolerance–productivity trade‐off. These results uncovering promising alleles controlling agronomic traits and/or multiple abiotic stress tolerances, providing insights into heritable covariation between yield and abiotic stress tolerance, will accelerate future efforts for wheat improvement.     

Putative zeatin O‐glucosyltransferase OscZOG1 regulates root and shoot development and formation of agronomic traits in rice

As a ubiquitous reaction, glucosylation controls the bioactivity of cytokinins in plant growth and development.Here we show that genetic manipulation of zeatin-Oglucosylation regulates the formation of important agronomic traits in rice by manipulating the expression of OscZOG1 gene,encoding a putative zeatin O-glucosyltransferase. We found that OscZOG1 was preferentially expressed in shoot and root meristematic tissues and nascent organs. The growth of lateral roots was stimulated in the overexpression lines, but inhibited in RNA interference lines. In shoots, knockdown of OscZOG1 expression by RNA interference significantly improved tillering, panicle branching, grain number per panicle and seed size, which are important agronomic traits for grain yield. In contrast, constitutive expression of OscZOG1 leads to negative effects on the formation of the grain-yielding traits with a marked increase in the accumulation levels of cis-zeatin O-glucoside(c ZOG) in the transgenic rice plants. In this study,our findings demonstrate the feasibility of improving the critical yield-determinant agronomic traits, including tiller number, panicle branches, total grain number per panicle and grain weight by downregulating the expression level of OscZOG1. Our results suggest that modulating the levels of cytokinin glucosylation can function as a fine-tuning switch in regulating the formation of agronomic traits in rice.     

Heat Stress in Wheat during Reproductive and Grain-Filling Phases

Ambient temperatures have increased since the beginning of the century and are predicted to continue rising under climate change. Such increases in temperature can cause heat stress: a severe threat to wheat production in many countries, particularly when it occurs during reproductive and grain-filling phases. Heat stress reduces plant photosynthetic capacity through metabolic limitations and oxidative damage to chloroplasts, with concomitant reductions in dry matter accumulation and grain yield. Genotypes expressing heat shock proteins are better able to withstand heat stress as they protect proteins from heat-induced damage. Heat tolerance can be improved by selecting and developing wheat genotypes with heat resistance. Wheat pre-breeding and breeding may be based on secondary traits like membrane stability, photosynthetic rate and grain weight under heat stress. Nonetheless, improvement in grain yield under heat stress implies selecting genotypes for grain size and rate of grain filling. Integrating physiological and biotechnological tools with conventional breeding techniques will help to develop wheat varieties with better grain yield under heat stress during reproductive and grain-filling phases. This review discusses the impact of heat stress during reproductive and grain-filling stages of wheat on grain yield and suggests strategies to improve heat stress tolerance in wheat.     

陕西关中地区杂种小麦强优势组合优势型研究

选用在陕西关中地区表现优良且某一农艺性状表现突出的11个冬小麦品种(系),根据农艺性状分为大粒型、多穗型、穗重型和中间型4类,组成6种NCII杂交组合模式,研究不同组合模式的杂种优势,探讨根据农艺性状划分杂种小麦亲本优势型在杂种小麦强优势组合选配中的应用.结果表明:在所组配的6种组合模式中,以大粒×多穗型组合模式表现突出,其杂种优势、超亲优势和超标优势均明显高于其它组合,分别达到了59.9%、37.1%和22.5%,因此,依据杂种小麦亲本农艺性状进行亲本优势型划分可以作为杂种小麦亲本选配的依据之一.在陕西关中地区以大粒×多穗型为杂种小麦强优势组合组配模式.     

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.     

Next-generation sequencing applications for wheat crop improvement

? Bread wheat (Triticum aestivum; Poaceae) is a crop plant of great importance. It provides nearly 20% of the world's daily food supply measured by calorie intake, similar to that provided by rice. The yield of wheat has doubled over the last 40 years due to a combination of advanced agronomic practice and improved germplasm through selective breeding. More recently, yield growth has been less dramatic, and a significant improvement in wheat production will be required if demand from the growing human population is to be met. ? Next-generation sequencing (NGS) technologies are revolutionizing biology and can be applied to address critical issues in plant biology. Technologies can produce draft sequences of genomes with a significant reduction to the cost and timeframe of traditional technologies. In addition, NGS technologies can be used to assess gene structure and expression, and importantly, to identify heritable genome variation underlying important agronomic traits. ? This review provides an overview of the wheat genome and NGS technologies, details some of the problems in applying NGS technology to wheat, and describes how NGS technologies are starting to impact wheat crop improvement.     

作物光能利用效率和收获指数时空变化研究进展

1972年Monteith提出的光能利用效率模型是当前大多数作物生长和产量形成模拟研究以及遥感估产所采用的主要方法.光能利用效率(radiation use efficiency, RUE)和收获指数(harvest index, HI)是其中的两个基本参量.鉴于目前作物RUE和HI研究与应用中仍存在着一些问题,本文综述了相关研究进展,总结了不同尺度上作物RUE和HI的研究方法;介绍了当前遥感估产应用中对RUE和HI两个关键参数的处置概况;建议今后研究应在点尺度开展作物RUE和HI研究的基础上,寻求其在区域尺度上定量评估的可行性途径,切实有效地发挥作物RUE和HI研究在作物实际生产管理中和遥感产量估算方面的应用价值及潜力.     

A Study on Genotype-by-Environment Interaction Analysis for Agronomic Traits of Maize Genotypes Across Huang-Huai-Hai Region in China

Facing the trend of increasing population, how to increase maize grain yield is a very important issue to ensure food security. In this study, 28 nationally approved maize hybrids were evaluated across 24 different climatic conditions for two consecutive years (2018–2019). The purpose of this study was to select high-yield with stable genotypes and identify important agronomic traits for maize breeding program improvement. The results of this study showed that the genotype ╳ environment interaction effects of the 12 evaluated agronomic traits was highly significant (P < 0.001). We introduced a novel multi-trait genotype-ideotype distance index (MGIDI) to select genotypes based on multiple agronomic traits. The selection process exhibited by this method is unique and easy to understand, so the MGIDI index will have more and more important applications in future multi-environment trials (METs) research. The genotypes selected by the MGIDI index were G22, G10, G12 and G1 as the high yielding and stable genotypes. The parents of these selected genotypes have the ability to play a greater role as the basic germplasm in the breeding process. A new form of genotype (G) main effects and genotype (G) -by-environment (E) interaction (GGE) technician, genotype*yield*trait (GYT) biplot, based on multiple traits for genotypes selection was also applied in this study. The GYT biplot ranked genotypes by combining grain yield with other evaluated agronomic traits, and displayed the distribution of their traits, namely strengths and weaknesses.     

江苏省沿海地区小麦品种更替与演变分析

摘要：对江苏省沿海地区12个县市（滨海县、大丰市、东台市、赣榆县、灌云县、海安县、海门市、启东市、如东县、射阳县、通州市、响水县）8个时间节点（1975年、1980年、1985年、1990年、1995年、2000年、2005年、2010年）的小麦种植面积、总产量、品种种类和数量及重要农艺性状的变化等进行了全面普查,对普查数据进行整理分析,结果如下：（1）江苏沿海地区小麦植面积呈现波动减少的趋势。（2）小麦总产量和平均单产均呈现逐渐提高的趋势,在总产提高的过程中,单产的平均贡献率均远大于面积的平均贡献率,优异小麦品种的推广是平均单产提高的决定因素。随着时间的推移,单产提高的难度越来越大,其对总产的贡献率逐渐下降。（3）对江苏沿海地区小麦品种的演变分析发现：①淮北淮南麦区的小麦品种均大致经历了四次品种更新,地方品种已经全部被改良品种取代,主推品种始终以江苏省自主选育的品种为主,且随着时间的推移,占主推品种的比重越来越高;在淮北麦区,对其影响较大的外省市主要是山东省和河南省的小麦品种。②综合性状好、适应性广的品种维持时间长,推广面积大。③株高的降低,千粒重、穗粒数及穗长的增加均对小麦品种的单产提高起到了一定的促进作用,培育兼抗多种病害的小麦品种可能是今后小麦育种需要关注的重要方向。     

Advanced backcross QTL analysis for the identification of quantitative trait loci alleles from wild relatives of wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.)

Advanced backcross QTL (AB-QTL) analysis was used to identify quantitative trait loci (QTLs) for yield and yield components in a BC(2)F(2) population derived from a cross between the German winter wheat variety 'Prinz' and the synthetic wheat line W-7984 developed by CIMMYT. Two hundred and ten microsatellite markers were employed to genotype 72 pre-selected BC(2)F(2) plants and phenotypic data were collected for five agronomic traits from corresponding BC(2)F(3) families that were grown at four locations in Germany. Using single-marker regression and interval mapping, a total of 40 putative QTLs derived from W-7984 were detected, of which 11 were for yield, 16 for yield components, eight for ear emergence time and five for plant height. For 24 (60.0%) of them, alleles from the synthetic wheat W-7984 were associated with a positive effect on agronomic traits, despite the fact that synthetic wheat was overall inferior with respect to agronomic appearance and performance. The present study indicated that favorable QTL alleles could be transferred from wild relatives of wheat into an elite wheat variety for improvement of quantitative trait loci like yield by the advanced backcross QTL strategy and molecular breeding. To our knowledge, the results presented here were the first report on AB-QTL analysis in wheat.     

QTL Analysis for Grain Pentosans and Hardness Index in a Chinese 1RS.1BL × non-1RS.1BL Wheat Cross

Pentosans are quantitatively minor constituents of wheat (Triticum aestivum L.), but they influence the end-use and nutritional qualities of grain cereals. Although several studies on the physicochemical properties and genetic basis of pentosans and pentosan-associated traits have been reported, the genetic architecture and determinants of pentosans remained ambiguous. In this research, 107 QTLs for pentosans and pentosan-associated traits were detected based on a population of recombinant inbred lines (RILs) derived from a 1RS/1BL translocation line?×?non-1RS/1BL translocation line. We identified seven stable expression QTLs for pentosans and one major QTL (Q.HI.scau-7D linked to xwmc634-7D) for hardness index (HI). Q.HI.scau-7D may be the gene responsible for the bimodal HI distribution map. More than one genetic location harbored QTLs for water-soluble pentosan (WSP), and the ratio of WSP content to water-insoluble pentosans (WIP) content (RWW) varied simultaneously, meaning that these loci harbored pleiotropic genes for WSP and RWW. WSP was the main factor in pentosans affecting wheat HI. This study gives a better understanding of the molecular genetics of pentosans and pentosan-related traits, and provides a certain theoretical basis for molecular genetics and breeding for pentosans.     

Quantitative trait loci associated with drought tolerance at reproductive stage in rice

Drought is a major constraint to rice (Oryza sativa) yield and its stability in rainfed and poorly irrigated environments. Identifying genomic regions influencing the response of yield and its components to water deficits will aid in our understanding of the genetics of drought tolerance and development of more drought tolerant cultivars. Quantitative trait loci (QTL) for grain yield and its components and other agronomic traits were identified using a subset of 154 doubled haploid lines derived from a cross between two rice cultivars, CT9993-510 to 1-M and IR62266-42 to 6-2. Drought stress treatments were managed by use of a line source sprinkler irrigation system, which provided a linearly decreasing level of irrigation coinciding with the sensitive reproductive growth stages. The research was conducted at the Ubon Rice Research Center, Ubon, Thailand. A total of 77 QTL were identified for grain yield and its components under varying levels of water stress. Out of the total of 77 QTL, the number of QTL per trait were: 7-grain yield (GY); 8-biological yield (BY); 6-harvest index (HI); 5-d to flowering after initiation of irrigation gradient (DFAIG); 10-total spikelet number (TSN); 7-percent spikelet sterility (PSS); 23-panicle number (PN); and 11-plant height (PH). The phenotypic variation explained by individual QTL ranged from 7.5% to 55.7%. Under well-watered conditions, we observed a high genetic association for BY, HI, DFAIG, PSS, TSN, PH, and GY. However, only BY and HI were found to be significantly associated with GY under drought treatments. QTL flanked by markers RG104 to RM231, EMP2_2 to RM127, and G2132 to RZ598 on chromosomes 3, 4, and 8 were associated with GY, HI, DFAIG, BY, PSS, and PN under drought treatments. The aggregate effects of these QTL on chromosomes 3, 4, and 8 resulted in higher grain yield. These QTL will be useful for rainfed rice improvement, and will also contribute to our understanding of the genetic control of GY under drought conditions at the sensitive reproductive stage. Close linkage or pleiotropy may be responsible for the coincidence of QTL detected in this experiment. Digenic interactions between QTL main effects for GY, BY, HI, and PSS were observed under irrigation treatments. Most (but not all) DH lines have the same response in measure of productivity when the intensity of water deficit was increased, but no QTL by irrigation treatment interaction was detected. The identification of genomic regions associated with GY and its components under drought stress will be useful for marker-based approaches to improve GY and its stability for farmers in drought-prone rice environments.