Favorable Alleles for Stem Water-Soluble Carbohydrates Identified by Association Analysis Contribute to Grain Weight under Drought Stress Conditions in Wheat

Drought is a major environmental constraint to crop distribution and productivity. Stem water-soluble carbohydrates (WSC) buffer wheat grain yield against conditions unfavorable for photosynthesis during the grain filling stage. In this study, 262 winter wheat accessions and 209 genome-wide SSR markers were collected and used to undertake association analysis based on a mixed linear model (MLM). The WSC in different internodes at three growth stages and 1000-grain weight (TGW) were investigated under four environmental regimes (well-watered, drought stress during the whole growth period, and two levels of terminal drought stress imposed by chemical desiccation under the well-watered and drought stress during the whole growth period conditions). Under diverse drought stress conditions, WSC in lower internodes showed significant positive correlations with TGW, especially at the flowering stage under well-watered conditions and at grain filling under drought stress. Sixteen novel WSC-favorable alleles were identified, and five of them contributed to significantly higher TGW. In addition, pyramiding WSC favorable alleles was not only effective for obtaining accessions with higher WSC, but also for enhancing TGW under different water regimes. During the past fifty years of wheat breeding, WSC was selected incidentally. The average number of favorable WSC alleles increased from 1.13 in the pre-1960 period to 4.41 in the post-2000 period. The results indicate a high potential for using marker-assisted selection to pyramid WSC favorable alleles in improving WSC and TGW in wheat.     

不同供水条件下冬小麦叶与非叶绿色 器官光合日变化特征

为揭示小麦叶片与非叶绿色器官的光合活性在一日中的变化特性及其在器官间的差异性,探讨群体及不同器官光合日变化对不同供水条件的响应特征,在田间设置生育期不灌水(I0)、灌2水(I2,拔节水＋开花水)和灌4水(I4,起身水+孕穗水+开花水+灌浆水)3个处理,于灌浆期测定了群体光合与呼吸速率的日变化, 旗叶片、叶鞘、穗、穗下节间各器官光合速率、蒸腾速率、气孔导度及叶绿素荧光参数的日变化。结果表明,灌浆期小麦穗和穗下节间光合速率日变化呈单峰曲线,而旗叶叶片与旗叶鞘光合速率均呈双峰型,表现出不同程度的午休。随着灌水次数减少,各器官光合速率降低,叶片对严重水分亏缺的反应大于各非叶器官。器官光合速率的日变化与F_v/F_m变化相一致,而与气孔导度日变化有较大差异。各器官上午的累积光合量均高于下午,上午光合量占日总光合量的比例为51％-62％,随着灌水次数减少而增大。不同灌水处理群体光合速率、呼吸速率日变化均未出现午休现象。春季浇2水处理与春浇4水处理相比,灌浆期群体光合速率及日光合积累量没有显著差异。综合研究认为,小麦叶与非叶器官光合性能及其日变化特征有较大不同,非叶光合对水分亏缺的敏感性低于叶片,生育期浇2水可以获得与浇4水相似的群体日光合积累量。     

Association Analysis of Genomic Loci Important for Grain Weight Control in Elite Common Wheat Varieties Cultivated with Variable Water and Fertiliser Supply

Grain weight, an essential yield component, is under strong genetic control and markedly influenced by the environment. Here, by genome-wide association analysis with a panel of 94 elite common wheat varieties, 37 loci were found significantly associated with thousand-grain weight (TGW) in one or more environments differing in water and fertiliser levels. Five loci were stably associated with TGW under all 12 environments examined. Their elite alleles had positive effects on TGW. Four, two, three, and two loci were consistently associated with TGW in the irrigated and fertilised (IF), rainfed (RF), reduced nitrogen (RN), and reduced phosphorus (RP) environments. The elite alleles of the IF-specific loci enhanced TGW under well-resourced conditions, whereas those of the RF-, RN-, or RP-specific loci conferred tolerance to the TGW decrease when irrigation, nitrogen, or phosphorus were reduced. Moreover, the elite alleles of the environment-independent and -specific loci often acted additively to enhance TGW. Four additional loci were found associated with TGW in specific locations, one of which was shown to contribute to the TGW difference between two experimental sites. Further analysis of 14 associated loci revealed that nine affected both grain length and width, whereas the remaining loci influenced either grain length or width, indicating that these loci control grain weight by regulating kernel size. Finally, the elite allele of Xpsp3152 frequently co-segregated with the larger grain haplotype of TaGW2-6A, suggesting probable genetic and functional linkages between Xpsp3152 and GW2 that are important for grain weight control in cereal plants. Our study provides new knowledge on TGW control in elite common wheat lines, which may aid the improvement of wheat grain weight trait in further research.     

Genome-Wide Association of Stem Water Soluble Carbohydrates in Bread Wheat

Water soluble carbohydrates (WSC) in stems play an important role in buffering grain yield in wheat against biotic and abiotic stresses; however, knowledge of genes controlling WSC is very limited. We conducted a genome-wide association study (GWAS) using a high-density 90K SNP array to better understand the genetic basis underlying WSC, and to explore marker-based breeding approaches. WSC was evaluated in an association panel comprising 166 Chinese bread wheat cultivars planted in four environments. Fifty two marker-trait associations (MTAs) distributed across 23 loci were identified for phenotypic best linear unbiased estimates (BLUEs), and 11 MTAs were identified in two or more environments. Liner regression showed a clear dependence of WSC BLUE scores on numbers of favorable (increasing WSC content) and unfavorable alleles (decreasing WSC), indicating that genotypes with higher numbers of favorable or lower numbers of unfavorable alleles had higher WSC content. In silico analysis of flanking sequences of trait-associated SNPs revealed eight candidate genes related to WSC content grouped into two categories based on the type of encoding proteins, namely, defense response proteins and proteins triggered by environmental stresses. The identified SNPs and candidate genes related to WSC provide opportunities for breeding higher WSC wheat cultivars.     

Fructan contents and dry matter deposition in different tissues of the wheat grain during development

The role of fructan metabolism in the assimilate relations of the grain of wheat (Triticum aestivum L.) was investigated by determination of the dry matter and fructan content of grain components at short intervals during grain filling. During the initial phase of rapid expansion, most of the assimilates entering the grain were partitioned to the outer pericarp. A large fraction of these assimilates were used for the synthesis of fructan. Dry matter deposition and fructan synthesis in the outer pericarp ceased at about 5d after anthesis. At the same time, the endosperm and the inner pericarp and testa started to accumulate dry matter at a fast rate. This was also associated with significant fructan synthesis in the latter tissues. The outer pericarp lost about 45% of its former maximum dry weight between 9 and 19 d after anthesis. This loss was due almost entirely to the near complete disappearance of water-soluble carbohydrates, most of which was fructan. The inner pericarp and testa accumulated dry matter until about mid-grain filling. The fructan contents of the inner pericarp and testa and the endosperm decreased slowly towards the end of grain filling. Most of the fructans in the inner pericarp and testa and the endosperm had a low molecular weight, whereas higher molecular weight fructans predominated in the outer pericarp. The embryo did not contain fructan. The presence of low molecular weight fructans in the endosperm cavity at mid-grain filling was confirmed. It is suggested that fructan synthesis is closely linked to growth-related water deposition in the different tissues of the wheat grain and serves to sequester the surplus of imported sucrose.     

Wheat Grain Filling Is Limited by Grain Filling Capacity rather than the Duration of Flag Leaf Photosynthesis: A Case Study Using NAM RNAi Plants

It has been proposed that delayed leaf senescence can extend grain filling duration and thus increase yields in cereal crops. We found that wheat (Triticum aestivum) NAM RNAi plants with delayed senescence carried out 40% more flag leaf photosynthesis after anthesis than control plants, but had the same rate and duration of starch accumulation during grain filling and the same final grain weight. The additional photosynthate available in NAM RNAi plants was in part stored as fructans in the stems, whereas stem fructans were remobilised during grain filling in control plants. In both genotypes, activity of starch synthase was limiting for starch synthesis in the later stages of grain filling. We suggest that in order to realise the potential yield gains offered by delayed leaf senescence, this trait should be combined with increased grain filling capacity.     

Contribution of the wheat (Triticum aestivum L.) flag leaf to grain yield in response to plant growth regulators

In West-Europe, intensive cereal management uses plant growth regulators (PGRs) especially for wheat. A green-house experiment compared the effects of two PGRs on flag leaf characteristics and yield of winter wheat. Chlormequat chloride + choline chloride (CCC) and chlormequat chloride + choline chloride + imazaquin (CCC+I) were applied to winter wheat at growth stage 5 (Feekes Large scale). CCC and CCC+I significantly increased flag leaf surface area at anthesis. Both treatments also enhanced chlorophyll content of the main stem flag leaf. The grain filling period was extended with PGR application by 2 days. CCC and CCC+I significantly increased net CO₂ assimilation rates during the flag leaf life. No effects of PGR spraying were observed on the pattern of ¹⁴C labelled assimilate distribution. Increased grain yield was due to the increase in average grain weight. The results indicate that PGR treatments increased flag leaf contribution to grain filling. The addition of imazaquin (I) to chlormequat (CCC) improved the effects of CCC.     

Ven、K型小麦雄性不育系春性F1杂种的优势表现及与叶绿素含量和同工酶谱的关系

利用Ven、K型小麦雄性不育系与北方春性普通小麦杂交,研究了Ven、K型春性F1杂种的优势表现及其与叶绿素含量、同工酶谱的关系。结果表明：Ven型和K型杂种F1存在显著的个体杂种优质,在主要农艺性状中尤其以单株穗数、千粒重、单株生产力的杂种优势明显,其中单株穗数对单株生产力的贡献最大;相关分析表明：单株生产力与单株穗数呈正相关,相关系数为0.7122;与每穗粒数和千粒重的相关系数分别为－0.1183和0.4941。灌浆期旗叶、倒二叶叶绿素含量与千粒重的相关性最大,呈正相关。杂种F1优势强的组合其种子和苗期的过氧化物同工酶和酯酶的酶谱一般呈双亲互补型或“杂种型”酶带,该特征可作为杂种优势预测的指标。     

潘庆民1,于振文2,王月福2

采用盆栽和水泥池栽研究了追氮时期对小麦光合作用、14C同化物运转分配和硝酸还原酶(NR)活性的影响.结果表明,拔节(雌雄蕊原基形成)期较起身(二棱)期追施氮肥,显著提高了小麦开花后的旗叶叶绿素含量和单叶光合速率;灌浆期旗叶14C同化物向籽粒转移比例显著提高,而在营养器官的滞留比例显著降低;旗叶和根系中硝酸还原酶(NR)活性亦显著提高.小麦穗粒数、粒重和产量增加,蛋白质含量提高.     

干旱胁迫下冬小麦光合产物分配格局及其与产量的关系

研究于2005-2006年在中国科学院封丘农田生态试验站防雨棚下进行,在充足供水(W1)、轻度干旱(W2)和严重干旱(W3)条件下,研究冬小麦周麦18(Zhoumai18)和济麦20(Jimai20)干物质积累和分配、可溶性碳水化合物(WSC)含量及其与产量形成的关系。结果显示,随着干旱胁迫程度加重,两个品种冬小麦干物质积累下降,光合产物分配格局改变。干旱胁迫导致花前叶片分配指数下降、茎和叶鞘分配指数上升,花后穗分配指数上升。干旱胁迫下冬小麦拔节期茎WSC减少,但孕穗期、开花期各器官WSC均增加。相关分析表明,冬小麦千粒重与孕穗期、开花期茎和根WSC,与开花期叶片干重显著正相关(P0.05);穗粒数与拔节期茎WSC极显著正相关(P0.01),与开花期茎和叶片干重显著正相关(P0.05)。研究认为拔节期是冬小麦需水关键期,干旱胁迫能促进光合产物向当时生长中心分配,不同生育期茎WSC是冬小麦抗旱育种的重要生理生态指标之一。     

The Relationship between Polyamines and Hormones in the Regulation of Wheat Grain Filling

The grain weight of wheat is strongly influenced by filling. Polyamines (PA) are involved in regulating plant growth. However, the effects of PA on wheat grain filling and its mechanism of action are unclear. The objective of the present study was to investigate the relationship between PAs and hormones in the regulation of wheat grain filling. Three PAs, spermidine (Spd), spermine (Spm), and putrescine (Put), were exogenously applied, and the grain filling characteristics and changes in endogenous PA and hormones, i.e., indole-3-acetic acid (IAA), zeatin (Z) + zeatin riboside (ZR), abscisic acid (ABA), ethylene (ETH) and gibberellin 1+4 (GAs), were quantified during wheat grain filling. Exogenous applications of Spd and Spm significantly increased the grain filling rate and weight, but exogenous Put had no significant effects on these measures. Exogenous Spd and Spm significantly increased the endogenous Spd, Spm, Z+ZR, ABA, and IAA contents and significantly decreased ETH evolution in grains. The endogenous Spd, Spm and Z+ZR contents were positively and significantly correlated with the grain filling rate and weight of wheat, and the endogenous ETH evolution was negatively and significantly correlated with the wheat grain filling rate and weight. Based upon these results, we concluded that PAs were involved in the balance of hormones that regulated the grain filling of wheat.     

Contributions of <Emphasis Type="Italic">TaSUTs</Emphasis> to grain weight in wheat under drought

Key message

The homologous genes to OsSUT1-5 in wheat were identified and detailed analysed. TaSUT1 was the predominant sucrose transporter group and it illustrated the genotypic variations towards drought during grain filling.

Abstract

Sucrose transporters (SUT) play crucial roles in wheat stem water soluble carbohydrate (WSC) remobilization to grain. To determine the major functional SUT gene groups in shoot parts of wheat during grain development, drought tolerant varieties, Westonia and Kauz, were investigated in field drought experiments. Fourteen homologous genes to OsSUT1-5 were identified on five homeologous groups, namely TaSUT1_4A, TaSUT1_4B, TaSUT1_4D; TaSUT2_5A, TaSUT2_5B, TaSUT2_5D; TaSUT3_1A, TaSUT3_1D; TaSUT4_6A, TaSUT4_6B, TaSUT4_6D; TaSUT5_2A, TaSUT5_2B, and TaSUT5_2D, and their gene structures were analysed. Wheat plants above the ground were harvested from pre-anthesis to grain maturity and the stem, leaf sheath, rachis, lemma and developing grain were used for analysing TaSUT gene expression. Grain weight, thousand grain weight, kernel number per spike, biomass and stem WSC were characterized. The study showed that among the five TaSUT groups, TaSUT1 was the predominant sucrose transporting group in all organs sampled, and the expression was particularly high in the developing grain. In contrast to TaSUT1, the gene expression levels of TaSUT2, TaSUT3 and TaSUT4 were lower, except for TaSUT3 which showed preferential expression in the lemma before anthesis. The TaSUT5 gene group was very weakly expressed in all tissues. The upregulated gene expression of TaSUT1 Westonia type in stem and grain reveal a crucial role in stem WSC remobilization to grain under drought. The high TaSUT1 gene expression and the significant correlations with thousand grain weight (TGW) and kernel number per spike demonstrated the contribution in Kauz’s high grain yield in an irrigated environment and high TGW in Westonia under drought stress. Further molecular level identification is required for gene marker development.

     

Effect of Heat Stress During Grain Filling on Phosphoenolpyruvate Carboxylase and Ribulose-1,5-bisphosphate Carboxylase/Oxygenase Activities of Various Green Organs in Winter Wheat

In two winter wheat (Triticum aestivum L.) cultivars differing in their response to high temperature, JD8 (tolerant) and J411 (sensitive) we studied the effect of heat stress on the activities of phosphoenolpyruvate carboxylase (PEPC) and ribulose-1,5-bisphosphate carboxylase (RuBPC) in green organs during grain-filling. There were significantly higher PEPC activities and lower RuBPC activities in each of the non-leaf organs (awn, glume, lemma, peduncle, and sheath) than in the flag leaf blade. Under heat stress for 12 d, the activity of RuBPC quickly declined and the activity of PEPC first increased and later declined in all organs, resulting in a great increase of the PEPC/RuBPC ratios in the organs, particularly in non-leaf organs which had a higher PEPC/RuBPC than the flag leaf blade in all times. The PEPC activity and PEPC/RuBPC ratio in every organ of JD8 were higher than those in the same organ of J411. Thus the differences in PEPC activities and PEPC/RuBPC may be associated with the differences in photosynthetic heat tolerance among the organs of the same plant or between the two cultivars.     

4种作物部分非叶器官气孔频度及其在光合作用中的意义（英文

 通过对小麦（Triticum aestivum L.）、大豆 (Glycine max (L.) Merrill)、 油菜（Brassica napus L.）和玉米（Zea mays L.）等4种作物部分非叶器官(油菜和大豆的豆荚;小麦的外稃和玉米的苞叶)的气孔频度、气孔大小和气孔指数进行了比较研究。结果发现,上述作物非叶器官的气孔频度均较对应叶低;而气孔大小和气孔指数则变化较大。其中油菜和大豆非叶器官气孔的直径通常比对应叶大,小麦和玉米非叶器官的气孔直径则较小;大豆和油菜的非叶器官气孔指数比对应叶小,     

Anthesis date mainly explained correlations between post-anthesis leaf senescence, grain yield, and grain protein concentration in a winter wheat population segregating for flowering time QTLs

The genetic variability of the duration of leaf senescence during grain filling has been shown to affect both carbon and nitrogen acquisition. In particular, maintaining green leaves during grain filling possibly leads to increased grain yield, but its associated effect on grain protein concentration has not been studied. The aim of this study was to dissect the genetic factors contributing to correlations observed at the phenotypic level between leaf senescence during grain filling, grain protein concentration, and grain yield in winter wheat. With this aim in view, an analysis of quantitative trait locus (QTL) co-locations for these traits was carried out on a doubled haploid mapping population grown in a large multienvironment trial network. Pleiotropic QTLs affecting leaf senescence and grain yield and/or grain protein concentration were identified on chromosomes 2D, 2A, and 7D. These were associated with QTLs for anthesis date, showing that the phenotypic correlations with leaf senescence were mainly explained by flowering time in this wheat population. Study of the allelic effects of these pleiotropic QTLs showed that delaying leaf senescence was associated with increased grain yield or grain protein concentration depending on the environments considered. It is proposed that this differential effect of delaying leaf senescence on grain yield and grain protein concentration might be related to the nitrogen availability during the post-anthesis period. It is concluded that the benefit of using leaf senescence as a selection criterion to improve grain protein concentration in wheat cultivars may be limited and would largely depend on the targeted environments, particularly on their nitrogen availability during the post-anthesis period.     

Rapid Identification of Major QTLs Associated with Rice Grain Weight and Their Utilization

To uncover the genetics of rice grain weight, we constructed an RIL population derived from a cross between a large grain accession M201 and a small size variety JY293. Specific Locus Amplified Fragment Sequencing (SLAF-Seq) technology was used to genotype two bulked DNA pools made from individual DNA of the heaviest 30 lines and the lightest 30 lines according to the 1000 grain weight (TGW). Bulked segregant analysis (BSA) was used to identify SLAFs strongly associated with TGW. Two marker-intensive regions at 24,600,000–24,850,000 bp and 25,000,000–25,350,000 bp on chromosome 3 were identified tightly related to the TGW. Then a linkage map of chromosome 3 was constructed with SSR markers and some SLAF derived single nucleotide polymorphisms (SNPs). Quantitative trait locus (QTL) mapping for TGW, grain length, grain width, and grain thickness revealed one major QTL in the second hot-region and two other minor QTLs for grain weight. These three QTLs displayed hierarchical effects on grain length and grain weight in order of qTGW3.2 (qGL3) qTGW3.1 (GS3) qTGW3.3. Multiple comparisons of means among the eight combinations of 3 QTLs revealed that the lines with two of three QTLs deriving from M201 displayed a large grain weight phenotype (TGW 40.2g, average data of three years) and lines with both qTGW3.1 and qTGW3.3 alleles from M201 (42.5g) had similar grain weight to the qTGW3.2 (40.8g) alone. Two strategies with similar effectiveness were proposed to improve grain weight by marker-assisted selection (MAS). One is to introduce the novel qTGW3.2 allele alone, and the other is to pyramid qTGW3.1 and qTGW3.3 alleles together. One new allele of GS3 (39 bp deletion in intron 1) and two SNPs in coding sequence of qGL3 identified in this study from M201 are useful in pyramiding elite alleles for molecular breeding for improvement of rice yield.     

QTL mapping for grain filling rate and yield-related traits in RILs of the Chinese winter wheat population Heshangmai × Yu8679

A set of 142 winter wheat recombinant inbred lines (RILs) deriving from the cross Heshangmai x Yu8679 were tried in four ecological environments during the seasons 2006 and 2007. Nine agronomic traits comprising mean grain filling rate (GFR(mean)), maximum grain filling rate (GFR(max)), grain filling duration (GFD), grain number per ear (GNE), grain weight per ear (GWE), flowering time (FT), maturation time (MT), plant height (PHT) and thousand grain weight (TGW) were evaluated in Beijing (2006 and 2007), Chengdu (2007) and Hefei (2007). A genetic map comprising 173 SSR markers and two EST markers was generated. Based on the genetic map and phenotypic data, quantitative trait loci (QTL) were mapped for these agronomic traits. A total of 99 putative QTLs were identified for the nine traits over four environments except GFD, PHT and MT, measured in two environments (BJ07 and CD07), respectively. Of the QTL detected, 17 for GFR(mean), 16 for GFR(max), 21 for TGW and 10 for GWE involving the chromosomes 1A, 1B, 2A, 2D, 3A, 3B, 3D, 4A, 4D, 5A, 5B, 6D and 7D were identified. Moreover, 13 genomic regions showing pleiotropic effects were detected in chromosomes 1A, 1B, 1D, 2A, 2B, 2D, 3A, 3B, 4B, 4D, 5B, 6D and 7D; these QTL revealing pleiotropic effects may be informative for a better understanding of the genetic basis of grain filling rate and other yield-related traits, and represent potential targets for multi-trait marker aided selection in wheat.     

晚播小麦叶片衰老代谢和粒重变化的比较研究

对７个小麦品种在晚播条件下的叶片衰老生理特性和粒重变化进行了比较研究。根据小麦叶片的衰老特征相差差异,将７个小麦品种区分为３个类型;后健型、早衰型和中间型。在小麦旗叶的衰老过程中,后健型小麦品种旗叶叶绿素和类胡萝卜素含量显著高于早衰型,脂质过氧化产物ＭＤＡ含量显著低于早衰型,小麦粒重降幅依次为早衰型〉中间型〉后健型,并讨论了活性氧代谢在小麦叶片衰老过程中可能作用。     

Nighttime Warming Will Increase Winter Wheat Yield Through Improving Plant Development and Grain Growth in North China

A better understanding of the actual impacts of nighttime warming on winter wheat growth will assist in breeding new varieties and agronomic innovation for food security under future climates. A 3-year experiment was conducted over an entire growth period of winter wheat using a passive warming facility in North China. An increase of 1.1 °C in mean nighttime temperature promoted wheat development, causing a 6-day reduction of the preanthesis period but a 5-day extension of the postanthesis period. This warming significantly stimulated the rate of leaf respiration at nighttime, resulting in higher carbohydrate depletion compared to that of the unwarmed control. However, stimulation of nighttime respiration and carbohydrate depletion could be compensated for by warming-led promotion of daytime photosynthesis and carbohydrate assimilation. Meanwhile, the flag leaf area per plant and the total green leaves area were significantly higher in the warmed plots than in the unwarmed plots. Besides extending the duration of grain filling, nighttime warming significantly promoted the filling rates of the superior and inferior grains, resulting in a significant increase in the 1,000-grain weight by 6.3 %. Consequently, this moderate increase in nighttime air temperature significantly increased wheat aboveground biomass and grain yield by 12.3 and 12.0 % (p < 0.05), respectively. A moderate warming at nighttime can improve the sink-source balance of winter wheat for higher yield. Our results suggest that climatic warming may benefit winter wheat production through improvement of plant development and grain growth in North China.     

影响小麦粒重的农艺性状,生理指标的主成分分析

本研究对长江中游麦区11个小麦品种（系）的13个与粒重密切相关的农艺性状和生理指标进行了主成分分析。13个原始指标综合成为4个独立的新指标;主成分1,光合同化因子;主成分2,籽粒灌浆速率与构成因子;主成分3,籽粒灌浆持续期与产量构成因子;主成分4,籽粒灌浆参数综合因子。主成分值与粒重相关分析表明,主成分4及主成分2与粒重有极显著的正相关。通过逐步回归分析,建立了小麦粒重与主成分值之间的回归方程。以主成分值为指标的系数聚类,可将十一个品种分为三大类型。不同类型品种主成分值的大小和相互协调方式不同,直接导致品种（系）粒重不同。