QTL mapping for six ear leaf architecture traits under water‐stressed and well‐watered conditions in maize (Zea mays L.)

Morphological traits for ear leaf are determinant traits influencing plant architecture and drought tolerance in maize. However, the genetic controls of ear leaf architecture traits remain poorly understood under drought stress. Here, we identified 100 quantitative trait loci (QTLs) for leaf angle, leaf orientation value, leaf length, leaf width, leaf size and leaf shape value of ear leaf across four populations under drought‐stressed and unstressed conditions, which explained 0.71%–20.62% of phenotypic variation in single watering condition. Forty‐five of the 100 QTLs were identified under water‐stressed conditions, and 29 stable QTLs (sQTLs) were identified under water‐stressed conditions, which could be useful for the genetic improvement of maize drought tolerance via QTL pyramiding. We further integrated 27 independent QTL studies in a meta‐analysis to identify 21 meta‐QTLs (mQTLs). Then, 24 candidate genes controlling leaf architecture traits coincided with 20 corresponding mQTLs. Thus, new/valuable information on quantitative traits has shed some light on the molecular mechanisms responsible for leaf architecture traits affected by watering conditions. Furthermore, alleles for leaf architecture traits provide useful targets for marker‐assisted selection to generate high‐yielding maize varieties.     

Multiple‐line cross QTL mapping for grain yield and thousand kernel weight in triticale

Grain yield and its component trait thousand kernel weight are important traits in triticale breeding programmes. Here, we used a large mapping population of 647 doubled haploid lines derived from four families to dissect the genetic architecture underlying grain yield and thousand kernel weight by multiple‐line cross QTL mapping. We identified 3 QTL for grain yield and 13 for thousand kernel weight which cross‐validated explained 5.2% and 48.2% of the genotypic variance, respectively. Both traits showed a positive phenotypic correlation, and we found two QTL overlapping between them. Full two‐dimensional epistasis scans revealed epistatic QTL for both traits, suggesting that epistatic interactions contribute to their genetic architecture. Based on QTL identified in our results, we conclude that the potential for marker‐assisted selection is limited for grain yield but more promising for thousand kernel weight.     

Spraying exogenous synthetic cytokinin 6‐benzyladenine following the waterlogging improves grain growth of waterlogged maize in the field

6‐Benzyladenine (6‐BA, a synthetic cytokinin) could improve the ability to resist adversity. However, very little attention has been given to its role in alleviating waterlogging damages on grain growth. A field experiment was performed to investigate effects of exogenous 6‐BA after waterlogging for 6 days at the third leaf stage (V3‐WL + 6‐BA) or the sixth leaf stage (V6‐WL + 6‐BA) on grain filling, and endogenous hormone of summer maize hybrids DengHai605 (DH605) and ZhengDan958 (ZD958), which were planted popularly in China. Exogenous 6‐BA alleviated waterlogging damages on grain filling by improving grain weight and volume, which was beneficial to yield increase. Grain yield of DH605 under V3‐WL + 6‐BA and V6‐WL + 6‐BA increased by 16% and 12%, respectively, compared with that of V3‐WL (waterlogging at the third leaf stage) and V6‐WL (waterlogging at the sixth leaf stage), while the corresponding values for ZD958 increased by 20% and 15%, respectively. Moreover, exogenous 6‐BA alleviated waterlogging damages on grain endogenous hormone content by increasing levels of endogenous indole‐3‐acetic acid, zeatin riboside, and gibberellic acid, and decreasing level of abscisic acid in grain of waterlogged summer maize during grain‐filling periods. Clearly, exogenous 6‐BA improved grain‐filling characteristics, and endogenous hormone, resulting in a significant yield increase of waterlogged summer maize.     

Molecular mapping of quantitative trait loci for grain moisture at harvest in maize

In maize, high grain moisture (GM) at harvest causes problems in harvesting, threshing, artificial drying, storage, transportation and processing. Understanding the genetic basis of GM will be useful for breeding low‐GM varieties. A quantitative genetics approach was used to identify quantitative trait loci (QTL) related to GM at harvest in field‐grown maize. The GM of a double haploid population consisting of 240 lines derived from Xianyu335 was evaluated in three planting seasons and a high‐density genetic linkage map covering 1546.4 cM was constructed. The broad‐sense heritability of GM at harvest was 71.0%. Using composite interval mapping, six QTL for GM at harvest were identified on five chromosomes (Chr). Two QTL located on Chr1, qgm1‐1 and qgm1‐2, explained 5.0% and 10.8% of the phenotypic variation in GM at harvest, respectively. The QTL qgm2, qgm3, qgm4 and qgm5 accounted for 3.3%, 8.3%, 5.4% and 11.0% of the mean phenotypic variation, respectively. Because of their consistent detection over multiple planting seasons, the detected QTL appear to be robust and reliable for the breeding of low‐GM varieties.     

QTL mapping of nutrient components in maize kernels under low nitrogen conditions

A set of 213 F_2:3 families were used to investigate the effects of nitrogen (N) on grain yield and the concentrations of three nutrient components in maize (Zea mays L.) kernels. A genetic linkage map was constructed using 189 SSR (simple sequence repeat) markers, spanning a total of 2003 cM, including 11 linkages, and the families were evaluated under high N and low N conditions at two farm locations. The results indicate that low N conditions may induce an increase in starch concentration, but a decrease in protein levels. Twenty‐six quantitative trait loci (QTL) were detected for four measured traits in the two N treatments at both locations, including eight QTL for grain yield, seven QTL for oil content, six QTL for protein content and five QTL for starch content. The total number of QTL detected for the four measured traits under high N levels was greater than the QTL detected under low N conditions, and several QTL were identified that specifically expressed under different N conditions. These particular QTL could help provide greater understanding of the genetic basis of N‐usage efficiency.     

Soybean plant height QTL mapping and meta‐analysis for mining candidate genes

Plant height is closely related to seed yield of soybean. The goal of this study was to identify important loci affecting soybean plant height using meta‐analysis based on a reference physical map. Plant height related to QTLs was mapped across eight years with a RIL population by WinQTLCart v2.5. 182 QTLs related to plant height of soybean from database and our research were collected, and each QTL was projected onto the soybean physical map by software BioMercator v2.1. The confidence interval of meta‐QTL ranged from 0.09 to 5.07 Mb, and the mean phenotypic variance ranged from 4.9% to 73.0%. Furthermore, 4,259 candidate genes were located in these consensus QTLs, and 40 of them were involved in the plant growth and stem elongation and annotated as plant hormone signal transduction (pathway ID ko04075) in KEGG pathway. These results would lay a foundation for fine mapping of QTLs/genes related to plant height and marker‐assisted selection for breeding in soybean.     

用3种方法定位玉米萌发期和苗期的耐盐和耐碱相关性状QTL

以耐盐碱郑58和盐碱敏感昌7-2为亲本,构建包含151份F2:5重组自交系(RILs)群体。基于3K芯片对郑58、昌7-2及其F2:5家系进行基因型分析,构建了包含1407个SNP分子标记的高密度遗传连锁图谱。该图谱的各染色体标记数在84～191之间,标记间的平均距离为0.81 cM。胁迫液为200 mmol L–1 NaCl和100 mmol L–1 Na2CO3,对照液为蒸馏水或霍格兰营养液,对盐、碱胁迫和自然条件下玉米的发芽率(GP)、株高(PH)、植株干、鲜重(FW、DW)、幼苗组织含水量(TWS)、植株地上部分钠含量(SNC)、钾含量(SKC)、钠/钾含量比(NKR)、苗期耐盐率(STR)、耐碱率(ATR)10项指标,采用3种不同的作图方法同时定位研究,对加性QTL定位采用复合区间作图法(CIM)和完备区间作图法(ICIM),对加性QTL与环境互作联合分析采用混合线性模型的复合区间作图法(MCIM)。结果表明,(1)与对照条件下各性状表型值相比,耐碱相关性状的降低较耐盐相关性状明显,说明玉米对碱胁迫更加敏感和碱胁迫对玉米的伤害更严重。碱与盐胁迫下SKC相当而SNC差异较大,表明Na+、K+的吸收和运输是相互独立的两个过程,玉米盐、碱胁迫可能是两种性质不同的胁迫。(2)在自然、盐和碱胁迫条件下,运用CIM分别检测到27、28、40个加性QTL;运用ICIM分别检测到28、23、17个加性QTL;运用MCIM共检测到11个耐盐加性QTL、4个环境互作QTL以及11个耐碱加性QTL、3个环境互作QTL。(3)盐胁迫条件下的qPH-9、qSTR-8、qNKR-6、qNKR-7和碱胁迫条件下的qPH-9、qATR-3能被3种作图方法重复检测到。与前人结果比较, qPH-9、qSTR-8、qNKR-6、q-ATR-3定位在相同或邻近区域,qNKR-7尚未见报道。本研究结果为精细定位玉米耐盐碱主效基因、挖掘候选基因和开发用于标记辅助选择的实用功能标记奠定基础。     

QTL mapping for seedling traits associated with low‐nitrogen tolerance using a set of advanced backcross introgression lines of rice

Nitrogen (N) deficiency is a major yield‐limiting factor in rice production. The objective of this study was to identify putative QTLs for low‐N stress tolerance of rice, using an advanced backcross population derived from crosses between an indica cultivar ‘93‐11’ and a japonica cultivar ‘Nipponbare’ and genotyped at 250 marker loci. Plant height, maximum root length, root dry weight, shoot dry weight and plant dry weight under two N conditions and their relative traits were used to evaluate low‐N tolerance at the seedling stage. A total of 44 QTLs were identified on chromosomes 1, 2, 3, 4, 5, 6, 8 and 9. Eight intervals on five chromosomes were identified to harbour multiple QTLs, suggesting pleiotropism or multigenic effects according to the contributor of alleles. Some QTL clusters were found in the nearby regions of genes associated with N recycling in rice, indicating that the key N metabolism genes might have effects on the expression of QTLs. Several unique QTLs for relative traits were detected, which suggested the specific genetic basis of relative performance.     

QTL analysis for cooking traits of super rice with a high‐density SNP genetic map and fine mapping of a novel boiled grain length locus

Hybrid rice has contributed substantially to the improvement of grain production worldwide, yet its poor cooking and tasting characteristics have long been recognized. In this study, 132 recombinant inbred lines derived from LYPJ were used to identify quantitative trait loci (QTLs) for 12 cooking traits with the high‐density SNP linkage map recently developed by our team. We identified 17 QTLs on chromosomes 1, 2, 4, 5, 6, 7, 8, 9 and 11, which accounted for 7.50% to 23.50% of the phenotypic variations. A novel major QTL qBGL7 for boiled grain length was further fine‐mapped to an interval of 440 Kb between the two markers RM21906 and gl3 using a BC₃F₂ population. Two near‐isogenic lines with extreme boiled grain length, GX5‐176 and GX5‐101, could be directly used in improving cooking quality. We also identified a QTL for soaked grain width expansion rate, qSGWE6, in the Wx gene region on chromosome 6. The Wx differential regulation coincided with sequential variation between the two parents. Our work offered a theoretical basis for molecular breeding of high‐quality hybrid rice.     

Quantitative trait loci mapping and epistatic analysis for grain yield and yield components using molecular markers with an elite maize hybrid

Summary The aim of this investigation was to map quantitative trait loci (QTL) associated with grain yield and yield components in maize and to analyze the role of epistasis in controlling these traits. An F_2:3 population from an elite hybrid (Zong3 × 87-1) was used to evaluate grain yield and yield components in two locations (Wuhan and Xiangfan, China) using a randomized complete-block design. The mapping population included 266 F_2:3 family lines. A genetic linkage map containing 150 simple sequence repeats and 24 restriction fragment length polymorphism markers was constructed, spanning a total of 2531.6 cM with an average interval of 14.5 cM. A logarithm-of-odds threshold of 2.8 was used as the criterion to confirm the presence of one QTL after 1000 permutations. Twenty-nine QTL were detected for four yield traits, with 11 of them detected simultaneously in both locations. Single QTL contribution to phenotypic variations ranged from 3.7% to 16.8%. Additive, partial dominance, dominance, and overdominance effects were all identified for investigated traits. A greater proportion of overdominance effects was always observed for traits that exhibited higher levels of heterosis. At the P ≤ 0.005 level with 1000 random permutations, 175 and 315 significant digenic interactions were detected in two locations for four yield traits using all possible locus pairs of molecular markers. Twenty-four significant digenic interactions were simultaneously detected for four yield traits at both locations. All three possible digenic interaction types were observed for investigated traits. Each of the interactions accounted for only a small proportion of the phenotypic variation, with an average of 4.0% for single interaction. Most interactions (74.9%) occurred among marker loci, in which significant effects were not detected by single-locus analysis. Some QTL (52.2%) detected by single-locus analysis were involved in epistatic interactions. These results demonstrate that digenic interactions at the two-locus level might play an important role in the genetic basis of maize heterosis.     

Fine mapping of a QTL for the number of spikelets per panicle by using near‐isogenic lines derived from an interspecific cross between Oryza sativa and Oryza minuta

We constructed a high‐resolution physical map for the qSPP7 QTL for spikelets per panicle (SPP) on rice chromosome 7 across a 28.6‐kb region containing four predicted genes. Using a series of BC₇F₄ near‐isogenic lines (NILs) derived from a cross between the Korean japonica cultivar ‘Hwaseongbyeo’ and Oryza minuta (IRGC Acc. No. 101144), three QTLs for the number of SPP, grains per panicle and primary branches were identified in the cluster (P ≤ 0.01). All three QTLs were additive, and alleles from the O. minuta parent were beneficial in the ‘Hwaseongbyeo’ background. qSPP7 was mapped to a 28.6‐kb region between the two simple sequence repeat (SSR) markers RM4952 and RM21605. The additive effect of the O. minuta allele at qSPP7 was 23 SPP, and 43.6% of the phenotypic variance was explained by the segregation of the SSR marker RM4952. Colocalization of the three QTLs suggested that this locus was associated with panicle structure and had pleiotropic effects. The NIL populations and molecular markers are useful for cloning qspp7.     

The genetic architecture of zinc and iron content in maize grains as revealed by QTL mapping and meta-analysis

Micronutrient malnutrition, especially zinc (Zn) and iron (Fe) deficiency in diets, has aroused worldwide attention. Biofortification of food crops has been considered as a promising approach for alleviating this deficiency. Quantitative trait locus (QTL) analysis was performed to dissect the genetic mechanism of Zn and Fe content in maize grains using a total of 218 F_2:3 families derived from a cross between inbred lines 178 and P53. Meta-analysis was used to integrate genetic maps and detect Meta-QTL (MQTL) across several independent QTL researches for traits related to Zn or Fe content. Five significant QTLs and 10 MQTLs were detected. Two informative genomic regions, bins 2.07 and 2.08, showed a great importance for Zn and Fe content QTLs. The correlation between Zn and Fe level in maize grains was proposed by MQTLs as 8 of the 10 involved both traits. The results of this study suggest that QTL mapping and meta-analysis is an effective approach to understand the genetic basis of Zn and Fe accumulation in maize grains.     

Mapping and validation of the quantitative trait loci for leaf stay‐green‐associated parameters in maize

Functional stay‐green is generally regarded as a desirable trait of varieties in major crops including maize. In this study, we used an F_3:4 recombinant inbred line population with 165 lines from a cross between a stay‐green inbred line (Zheng58) and a model inbred line (B73) using 211 polymorphic simple sequence repeat markers to map quantitative trait loci for three stay‐green‐associated parameters, chlorophyll content, photosystem II photochemical efficiency and stay‐green area, at maturity stage, detected a total of 23 quantitative trait loci (QTL) on nine chromosomes. Single QTL explained 3.7–13.5% of the phenotypic variance. Additionally, we validated some important stay‐green QTL using a heterogeneous inbred family approach and found that the stay‐green‐associated parameters were significantly correlated with the plant yield. This study may contribute to a better insight into the regulatory mechanism behind leaf stay‐green in maize and a novel development of elite maize varieties with delayed leaf senescence through molecular marker‐assisted selection.     

Prediction of deoxynivalenol and zearalenone concentrations in Fusarium graminearum inoculated backcross populations of maize by symptom rating and near‐infrared spectroscopy

Gibberella ear rot (GER) caused by Fusarium graminearum is a destructive disease in maize of temperate regions resulting in yield reduction and contamination by the mycotoxins deoxynivalenol (DON) and zearalenone (ZON). We wanted to analyse whether prediction of DON and ZON concentrations is feasible either by GER severity ratings or by near‐infrared spectroscopy (NIRS). We analysed 80 and 102 lines developed by backcrossing doubled‐haploid lines from segregating populations to the resistant and susceptible parent, respectively, by artificial infection at three locations in Germany and France. Both backcross (BC) populations differed substantially in their means for all traits with significant (P < 0.01) genotypic variances. DON and ZON concentrations measured by immunotests were significantly (P < 0.01) correlated with each other and with GER severity within each BC population (0.6 ≤ r ≤ 0.9, P < 0.01). DON concentration measured by immunotest and NIRS significantly correlated (r ≈ 0.9, P < 0.01). In conclusion, DON and ZON concentrations could be reliably predicted by GER severity. Additional NIRS analysis of DON concentration might be useful for the positively selected fraction.     

Maize participatory breeding in Portugal: Comparison of farmer's and breeder's on‐farm selection

“VASO” is a Portuguese participatory maize breeding project (1984), where several maize landraces such as “Pigarro” have been selected both by a farmer's (phenotypic recurrent selection) and a breeder's approach (S2 lines recurrent selection). The objectives of this study were to determine the phenotypic and genotypic responses to participatory selection using these two different approaches, to clarify to which extent both selection methods preserve genetic diversity, and conclude what is the preferred method to apply in sustainable farming systems. The results, obtained via ANOVA, regression analyses and molecular markers, indicate that for both selection methods, genetic diversity was not significantly reduced, even with the most intensive breeder's selection. Although there were some common outputs, such as the determinated versus indeterminated ears, cob and ear weight ratio per ear and rachis 2, specific phenotypic traits evolved in opposite directions between the two selection approaches. Yield increase was only detected during farmer selection, indicating its interest on PPB. Candidate genes were identified for a few of the traits under selection as potential functional markers in participatory plant breeding.     

Genotype by environment effects and selection for drought tolerance in tropical maize. I. Two mode pattern analysis of yield

Ten trials evaluated the performance of several late tropical maize populations (La Posta Sequía, Pool 26 Sequía and Tuxpeño Sequía) selected for tolerance to drought during flowering and grain filling and also for yield potential. Families (S₁ or full-sib) had been selected recurrently for six to eight years on an index of traits. Pattern (clustering and ordination) analysis was used to analyse the relative performance of entries that included cycles of selection for drought tolerance in the populations and non-drought tolerant checks. Mean environment (E) yields ranged from 1.0 to 10.4 t ha^-1. Analysis of variance showed that 97.9% of the total sums of squares was accounted for by E, and that, of the remaining sums of squares the G × E (genotype by environment interaction) was almost 3 times that of the contribution of G alone. Cluster analysis separated the checks, the earlier maturing drought tolerant entries and the later maturing drought tolerant entries. This was verified by principal component (PC) analysis of the G × E matrix. Grouping of the environments (i.e. based on entry performance), resulted in the separation of different types of droughts, and of medium and high yielding well-watered environments. The patterns of discrimination observed indicated that the yield gains under drought would have been unlikely to occur if selection had been done only in well-watered environments. Within each population, selection improved broad adaptation (higher mean yield) to both drought and well-watered environments and cycles of selection ‘jumped’ from non-drought-tolerant to drought-tolerant groups as their specific adaptation to drought environments increased.