Identification and functional analysis of a locus for improvement of lodging resistance in rice

We identified a new locus responsible for increased pushing resistance of the lower part of rice (Oryza sativa) and analyzed its physiological function to understand how to improve lodging resistance in rice. Quantitative trait loci (QTLs) controlling pushing resistance of the lower part were analyzed in a population of backcross inbred lines of japonica Nipponbare x indica Kasalath plants cut out at 40 cm to exclude the effect of the weight of the upper parts. Five QTLs for pushing resistance were detected; only one QTL from Kasalath on chromosome 5 (prl5) had a positive effect. The likelihood odds ratio curve of prl5 echoed that for lodging resistance by typhoon. We selected three near-isogenic lines (NILs) in which the chromosomal region of prl5 was substituted with that of Kasalath in the Nipponbare background. The dry weights and densities and the contents of accumulated carbohydrate in stems below 40 cm (lower stems) in each NIL were significantly higher than those of Nipponbare. There was no difference between Nipponbare and the NILs in yield, root characteristics, or the weights of the upper parts. Pushing resistance of the lower part and lodging resistance in the NILs were up to twice as high as in Nipponbare. These results suggest that prl5 might affect the characteristics of the lower stems of the NILs, thus increasing lodging resistance.     

Improvement of lodging resistance with QTLs for stem diameter in rice (Oryza sativa L.)

Varietal differences among ten rice cultivars showed that stem diameter is a key factor in lodging resistance (measured in terms of pushing resistance). Two near-isogenic lines (NILs) were selected from a series of chromosome segment substitution lines developed between cultivars Nipponbar and Kasalath, one containing a single stem diameter QTL (sdm8; NIL114), and another with four stem diameter QTLs (sdm1, sdm7, sdm8, sdm12; NIL28). Compared with the Nipponbare control, stem diameters were larger in NIL114 and NIL28 by about 7 and 39%, respectively. Pushing resistance in NIL28 was significantly greater than in Nipponbare, but NIL114 was similar to Nipponbare. The two NILs had greater weight of lower stem and culm wall thickness than Nipponbare. NIL28 had higher plant height, which is a negative effect on lodging resistance, than Nipponbare. The non-structural carbohydrate contents of NIL stems were higher than that of Nipponbare, whereas the silicon contents were lower in the NILs, and cellulose contents were lower only in NIL28. The basal internodes of the two NILs were significantly stiffer than those of Nipponbare. These results suggest that increasing stem diameter in rice breeding programs would improve lodging resistance, although the combination of multiple QTLs would be necessary to produce thicker stems with higher pushing resistance, whereas the higher plant height could also result from the combination of multiple QTLs.     

水稻茎秆形态结构特征和化学成分与抗倒伏关系综述

水稻茎秆形态结构特征和化学成分与其抗倒伏性能相关。本文对水稻茎秆高度、茎秆基部第1、2节间长度、茎秆粗细、茎壁厚度、厚壁组织(机械组织)数量和强度、维管束数量、细胞壁纤维素和木质素含量、细胞中碳水化合物积累的数量、硅与钾的含量以及茎秆抗倒伏相关的QTL等与水稻的抗倒伏性的相互关系进行了综述,为水稻抗倒伏优良品种性状的选育提供参考。     

超高产杂交稻不同生育期的光合色素含量、净光合速率和水分利用效率

 较为系统地研究了两个超高产杂交稻‘两优培九’、‘华安3号’和多年来大面积推广的常规杂交稻‘汕优63’不同生育期的光合色素含量、净光合速率和水分利用效率。结果表明,在苗期,3个杂交稻的单位叶面积的叶绿素（Chl）含量差别不大,类胡萝卜素（Car）的含量以‘汕优63’为最高。然而,随着发育阶段的推进,到分蘖期,尤其是抽穗期剑叶的单位叶面积Chl和Car含量,两个超高产杂交稻高于‘汕优63’。从苗期到抽穗期,超高产杂交稻‘两优培九’和‘华安3号’的净光合速率（Pn）都比‘汕优63’高,而在苗期的午间强光条件下和分蘖期的早晨以及抽穗期的早晚相对弱光条件下其Pn的差别尤为显著。在苗期,‘汕优63’的水分利用效率（WUE）略高于‘两优培九’和‘华安3号’;到分蘖期和抽穗期,在上午10时之前和下午14时以后的时间段,则是两个超高产杂交稻明显高于‘汕优63’。这些结果说明,超高产杂交稻‘两优培九’和‘华安3号’不仅有较高的Pn和较强的抗光抑制能力,而且还能充分利用早晨和傍晚较弱的光强进行光合作用,这些是它们之所以高产的重要生理基础。此外,超高产杂交稻在生长发育的中后期（分蘖期和抽穗期）具有较高的WUE,有利于节约农业用水。     

Identification and physiological analyses of a locus for rice yield potential across the genetic background

A new locus responsible for increased yield potential across the genetic background in rice (Oryza sativa L.) was identified and evaluated. Quantitative trait loci (QTLs) were analysed for the ratio of filled grains, a yield component, in backcrossed inbred lines of a japonica 'Nipponbare'xindica 'Kasalath' cross for 3 years. Only one QTL (rg5), with a positive Kasalath allele, was detected across environments (years). The physiological functions of rg5 were clarified in a near-isogenic line (NILrg5) with a Kasalath chromosome segment containing rg5 in a Nipponbare genetic background. In NILrg5, carbohydrate storage capacity before heading or sink activity in the first or last stages of the reproductive phase was significantly higher than in Nipponbare (control). The ratio of filled grains and yield per plant were significantly higher in NILrg5 than in Nipponbare, by 5% (P <0.01) and 15% (P <0.05), respectively. These results suggest that rg5 improves carbohydrate storage capacity and keeps sink activity higher in the reproductive stage, and consequently increases yield potential. Greater capacity to accumulate carbohydrate is the main target for increasing rice yield potential; therefore, rg5 might function under other genetic backgrounds. Substitution of the rice cv. Kasalath chromosome segment containing rg5 gave higher yield potential in the top premium rice cv. Koshihikari. These results suggest that rg5 might be able to affect yield under different genetic backgrounds, and physiological analyses of the targeted locus might reveal these effects.     

Quantitative trait loci for sucrose, starch, and hexose accumulation before heading in rice.

We studied the storage of sucrose, starch, and hexose before heading in rice (Oryza sativa L.) plants by quantitative trait locus (QTL) analysis with a population of backcross inbred lines (BILs) of japonica cv. Nipponbare x indica cv. Kasalath. Carbohydrates are accumulated in the rice plant before heading and are translated to the panicle after heading. A higher capacity for accumulation is thus a main target for improvement in yield. The form of carbohydrate (sucrose, starch, or hexose) differs depending on the organ in which it is stored. There was no correlation between starch and sucrose or hexose contents in BILs, and the positions of QTLs controlling starch differed from those for sucrose and hexose accumulation. These results suggest that the genetic control of accumulation differs between starch and sugars. QTLs that control the ratio of sucrose to starch content were detected, suggesting the existence of a mechanism(s) that determines this ratio. On chromosome 1, sucrose-phosphate synthase 1, the key enzyme in sucrose synthesis was close to the peaks of the likelihood odds ratios in QTLs for sucrose or hexose content. These results suggest that SPS1 is related to conversion of carbohydrate to sucrose as accumulated form in a plant before heading.     

不同CO2浓度和N肥水平对粳稻茎鞘非结构性碳水化合物含量与积累的影响

为了解CO2浓度升高和N肥水平对水稻茎鞘内非结构性碳水化合物(NSC)含量和积累量的影响,利用开顶式气室(OTC),以常规粳稻"南粳9108"为试验材料,设置3个CO2浓度水平:对照T0(背景大气)、T0+120μmol·mol-1(T1)和T0+200μmol·mol-1(T2)。在OTC内采用盆栽方式,设置3个氮(N)肥水平:10 g N·m^-2(N1)、20 g N·m^-2(N2)和30g N·m^-2(N3)。分别于水稻抽穗期、灌浆期(抽穗后20 d)和成熟期对地上部分各器官生物量、茎鞘NSC含量以及顶部四张叶片的N含量进行分析。结果表明:CO2浓度升高对抽穗期叶N含量总体无显著影响,但显著降低灌浆期N2和N3水平的叶N含量;CO2浓度升高对抽穗期茎鞘NSC含量和积累量无显著影响,抽穗期置换到高CO2浓度环境使灌浆期茎鞘NSC积累显著增加,置换到低CO2浓度环境使NSC积累显著减少。同一CO2浓度条件下,NSC含量和积累量均为N1>N2>N3,且N1处理均显著高于N3处理,CO2浓度升高和N水平的交互作用对灌浆期茎鞘NSC含量影响显著。水稻产量在不同CO2浓度水平间无显著差异,但随施氮水平的提高而增加。抽穗期与灌浆期水稻茎鞘NSC含量和积累量与茎鞘干重呈极显著正相关,与叶N含量呈极显著负相关;叶N衰减越慢,灌浆期水稻茎鞘NSC残留比(RNSC)越低;结实率和产量与RNSC呈显著负相关,RNSC越大,茎鞘NSC转移的越少,结实率和产量越低。     

qUVR-10, a major quantitative trait locus for ultraviolet-B resistance in rice, encodes cyclobutane pyrimidine dimer photolyase

Rice qUVR-10, a quantitative trait locus (QTL) for ultraviolet-B (UVB) resistance on chromosome 10, was cloned by map-based strategy. It was detected in backcross inbred lines (BILs) derived from a cross between the japonica variety Nipponbare (UV resistant) and the indica variety Kasalath (UV sensitive). Plants homozygous for the Nipponbare allele at the qUVR-10 locus were more resistant to UVB compared with the Kasalath allele. High-resolution mapping using 1850 F(2) plants enabled us to delimit qUVR-10 to a <27-kb genomic region. We identified a gene encoding the cyclobutane pyrimidine dimer (CPD) photolyase in this region. Activity of CPD photorepair in Nipponbare was higher than that of Kasalath and nearly isogenic with qUVR-10 [NIL(qUVR-10)], suggesting that the CPD photolyase of Kasalath was defective. We introduced a genomic fragment containing the CPD photolyase gene of Nipponbare to NIL(qUVR-10). Transgenic plants showed the same level of resistance as Nipponbare did, indicating that the qUVR-10 encoded the CPD photolyase. Comparison of the qUVR-10 sequence in the Nipponbare and Kasalath alleles revealed one probable candidate for the functional nucleotide polymorphism. It was indicated that single-base substitution in the CPD photolyase gene caused the alteration of activity of CPD photorepair and UVB resistance. Furthermore, we were able to develop a UV-hyperresistant plant by overexpression of the photolyase gene.     

New target for rice lodging resistance and its effect in a typhoon

We demonstrated the new target for lodging resistance in rice (Oryza sativa L.) by the analysis of physiological function of a locus for lodging resistance in a typhoon (lrt5) with the near isogenic line under rice “Koshihikari” genetic background (tentatively named S1). The higher lodging resistance of S1 was observed during a typhoon in September 2004 (28 days after heading), when most other plants in “Koshihikari” became lodged. Visual observations showed that bending of the upper stems triggered lodging during the typhoon; the upper stem of “Koshihikari” buckled completely, whereas that of S1 remained straight. In addition to the strong rain and winds during the typhoon, the weight of the buckled upper plant parts increased the pressure on adjacent plants and caused a domino effect in “Koshihikari”. Young’s modulus, an indicator of the rigidity of the culm, was significantly higher in S1 than in “Koshihikari”. In the upper culm, the starch content in S1 was 4.8 times the value in “Koshihikari”, and senescence was delayed in the upper leaves of S1. These results suggest that the rigidity of the upper culm by the higher starch content (as a result of delayed senescence in the upper leaves) may be responsible for the higher lodging resistance during a typhoon in rice. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

不同密度下臭氧胁迫对Ⅱ优084水稻光合作用和物质生产的影响—FACE研究

依托中国稻田臭氧FACE(free air ozone concentration enrichment)技术平台,以超级稻Ⅱ优084为供试材料,臭氧设置当前大气臭氧浓度和高臭氧浓度(比前者高50%),移栽密度设置低密度(16穴·m^-2)、中密度(24穴·m^-2)和高密度(32穴·m^-2),研究不同移栽密度条件下近地层臭氧浓度升高对水稻光合作用、物质生产以及茎鞘非结构性碳水化合物浓度和含量的影响.结果表明: 臭氧浓度升高使水稻移栽后63 d、77 d和86 d剑叶SPAD值分别下降6%、11%和13%,均达显著或极显著水平.臭氧胁迫下结实期叶片净光合速率、气孔导度和蒸腾速率的降幅亦随时间推移而明显增加.高臭氧浓度使水稻抽穗至成熟期的物质生产量平均下降46%,从而使最终生物产量下降25%,均达显著水平.臭氧浓度升高使水稻拔节后茎鞘可溶性糖和淀粉的浓度和含量均显著降低,但使抽穗前茎鞘贮藏同化物的转运率大幅增加.方差分析表明,臭氧与密度间的互作对水稻所有测定参数均无显著影响.综上,近地层臭氧浓度升高使超级稻Ⅱ优084生育中后期的光合和生长均明显受抑,但这种抑制作用不受移栽密度的影响.
     

Changes in the Chlorophyll Contents of Leaves and in Levels of Cytokinins in Root Exudates during Ripening of Rice Cultivars Nipponbare and Akenohoshi

Changes in fluxes of cytokinins in exudates transported viathe xylem from roots of rice plants cvs. Nipponbare (a standardJapanese cultivar) and Akenohoshi (a slowly senescing cultivar)were measured by mass spectrometry with deuterium-labeled standards.The fluxes of zeatin (Z), trans-ribosylzeatin (trans-RZ), N⁶-isopentenyladenine(iP), and "conjugated Z" (Z in the hydrolysates of highly polarfractions) decreased from heading to the late ripening stagein both cultivars. At the late ripening stage, iP and Z couldno longer be detected, while the flux of N⁶-isopentenyladenosine(iPA) increased slightly. In Akenohoshi, conjugated Z was thepredominant cytokinin from heading to the middle of the ripeningstage. The flux of each of the cytokinins in Akenohoshi washigher than that in Nipponbare at every time point, with theexception of the flux of iPA just after heading. The total concentrationof cytokinins in the xylem exudate of Akenohoshi was higherthan that of Nipponbare after the middle of ripening stage.The chlorophyll content of the third leaves, which were senescingrapidly, was significantly correlated with the flux of totalcytokinins per plant or per unit leaf area. These results suggestthat the larger amounts of cytokinins, in particular conjugatedZ, transported from the roots to the shoots caused the slowsenescence of leaves in Akenohoshi during the ripening stage. (Received May 9, 1994; Accepted July 1, 1995)     

Comparison of lodging safety factor of untreated and succinic Acid 2,2-dimethylhydrazide-treated shoots of mulberry tree

This study examined the lodging resistance of mulberry tree (Morus bombycis Koidz. cv Kenmochi) shoots treated or not treated with succinic acid 2,2-dimethylhydrazide (SADH). The lodging safety factor, an indicator of lodging resistance, was defined as the ratio of critical lodging load to the leaf fresh weight observed, provided that the distribution of the critical lodging load along the stem was similar to that of the leaf fresh weight observed. The critical lodging load was experimentally estimated by loading weights onto the stems. In the untreated trees, the lodging safety factor was maintained at about 3.2. In the SADH-treated trees, the stem elongation was inhibited to about 80% of that in the untreated trees, and the percentage of shoot dry matter partitioned into the leaves was always larger than that of the untreated trees. This dwarfing of the stem caused by SADH increased the critical lodging load supported by the unit stem dry weight, while this large investment of materials in leaves increased the leaf fresh weight supported by the unit stem dry weight. Since the increments canceled each other, the lodging safety factor of the SADH-treated shoots was similar to that of the untreated ones. These results suggest that the shoot formation of the mulberry tree is controlled to maintain the lodging safety factor at a constant level.     

Changes at panicle emergence in the water relations of a wetland and a dryland Japonica rice cultivar under wetland conditions

The diurnal and seasonal changes in plant water relations of two Japonica rice ( Oryza sativa L.) cultivars, Nipponbare and Tachiminori, were studied under flooded conditions at Kyoto University. The dryland cv. Tachiminori maintained higher predawn and midday leaf osmotic potentials relative to the wetland cv. Nipponbare during the vegetative stage, but the ranking was reversed after flowering. The relationship between leaf water potential and leaf osmotic potential showed that prior to panicle emergence Nipponbare was able to adjust osmotically to maintain turgor, whereas after heading there was little turgor maintenance. Tachiminori showed little difference in osmotic adjustment before and after panicle emergence. Fertilizer treatment during panicle development also helped to maintain the degree of osmotic adjustment in both cultivars.     

Assessment of leaf/stem ratio in wheat straw feedstock and impact on enzymatic conversion

The composition of wheat straw leaf and stem fractions were characterized using traditional strong acid hydrolysis, and monoclonal antibodies using comprehensive microarray polymer profiling (CoMPP). These results are then related to high throughput lignocellulose pretreatment and saccharification screening data. Pure leaf fraction of wheat straw was the least recalcitrant compared to pure stem and easily digested by commercial cellulases after moderate hydrothermal pretreatment; 63% and 31% (w/w) of glucan, 88% and 61% of xylan were released from the leaf and stem fractions, respectively. By preparing samples of various leaf‐to‐stem (L/S) ratios, we found shifting conversion behavior as processing parameters were modified. Increasing the enzyme dosage, pretreatment temperature and pretreatment time all significantly improved conversion rates in samples with more than 50% leaf content, whereas less impact was observed on samples with less than 50% leaf content. Enzyme affinity, desorption and readsorption with leaf and stem fractions may affect the sugar yield in wheat straw saccharification. The data suggest that the L/S ratio is an important parameter when adjusting or optimizing conversion processes and additionally in feedstock breeding. Furthermore, this highlights the need for rapid techniques for determining L/S ratio in wheat straw harvests. The CoMPP data on specific carbohydrates and leaf pectin highlight carbohydrate epitopes that may be useful as markers in the development of novel screening techniques; especially pectin or arabinogalactan proteins related epitopes are promising.     

Contrasting Characteristics of Lodging Resistance in Two Super-Rice Hybrids Differing in Harvest Index

The “super rice” breeding program in China has been successful in developing high-yielding hybrids, including few with high harvest index values. However, there is limited information on the relationship between lodging resistance and harvest index, and the mechanisms underlying the relationship in super-rice hybrids. In this study, a two-year field experiment was conducted to compare lodging resistance and its related traits between two super-rice hybrids differing in harvest index, i.e., Guiliangyou 2 (G2) with a high harvest index and Y-liangyou 1 (Y1) with a typical harvest index of modern high-yielding rice varieties. Results showed that compared to Y1, G2 was lower in plant height due to its lower aboveground N uptake, and its higher stem breaking resistance (i.e., lower stem breaking index) resulted from a lower stem height at its center of gravity. Consequently, G2 had a higher lodging resistance (i.e., lower plant lodging index) than Y1. This study suggests that developing super-rice hybrids with high harvest index values is a possible way to achieve both high grain yield and strong lodging resistance in rice.     

甘蓝型油菜抗茎倒材料的筛选与候选基因挖掘

该研究采用茎秆抗折力指标评价了200份甘蓝型油菜种质资源抗茎倒伏能力,据此筛选出极端抗茎倒材料和不抗茎倒材料各1份,随后测定了2份材料成熟期茎秆的理化组分含量,并对2份极端抗茎倒差异材料蕾苔期、盛花期茎秆进行转录组测序分析,为甘蓝型油菜抗茎倒伏的遗传改良奠定基础。结果表明:(1)200份甘蓝型油菜种质资源的茎秆抗折力和茎粗均为正态分布,均属于数量遗传性状,并依据茎秆抗折力和农艺性状筛选出生育期相近,株型相似,茎秆粗度差异不显著,但茎秆抗折力差异显著的极端不抗茎倒材料GY172和抗茎倒材料GY199。(2)GY199的韧皮部比GY172更加致密,而GY172成熟期茎秆中的半纤维素、木质素、中性洗涤纤维及总可溶性糖含量均显著高于GY199,而其纤维素含量极显著低于GY199,即成熟期茎秆纤维素含量与这2个材料的茎秆抗折力呈正相关。(3)蕾苔期、盛花期茎秆转录组测序发现,碳代谢、碳固定、磷酸戊糖途径、氨基酸的生物合成、糖酵解/糖异生等途径的14个基因(BnaA10G0056100ZS、BnaC08G0455100ZS、BnaA08G0262400ZS、BnaC08G0239700ZS、BnaA07G0362300ZS、BnaC02G0081300ZS、BnaC04G0273000ZS等)以及纤维素合成相关的9个基因(BnaA05G0152200ZS、BnaA01G0411100ZS、BnaA03G0018900ZS、BnaA03G0037800ZS等)在抗茎倒材料GY199中显著上调表达,这些基因可能参与调控了茎秆强度性状,可作为油菜抗茎倒候选基因。     

我国不同年代玉米自交系茎秆性状演替规律

茎秆是玉米植株重要的组成部分,与植株倒伏、籽粒产量密切相关.研究我国玉米茎秆性状变化趋势,探讨不同年代茎秆性状演变规律,为我国玉米抗倒伏及高产品种选育提供参考.本研究以20世纪60-90年代65个代表性玉米自交系为材料,调查植株茎高、穗位高、茎节数、穗位节、茎粗、茎秆含水量、茎秆含糖量7个茎秆性状,及其衍生的穗位系数(...     

Characterising the morphological characters and carbohydrate metabolism of oat culms and their association with lodging resistance

• Lodging resistance can be improved by enhancing the mechanical strength of culms, and culm carbohydrates could improve this mechanical strength. Culm carbohydrates can regulate development of the culm and affect its toughness.
• The present study determines the relationship between lodging and carbohydrate content in oat culms. Field experiments were conducted in alpine regions in 2017 and 2018 using three oat varieties with different lodging resistance. Lodging‐related morphological characteristics were directly determined and culm carbohydrate content and enzyme activity related to cellulose synthesis and sucrose metabolism were evaluated with ultraviolet spectrophotometry.
• Results showed that the lower the gravity height or the lower ratio of gravity height to plant height, the stronger the lodging resistance of the varieties. Higher culm nonstructural (NSC) and structural (SC) carbohydrate content contributed to the ability of culms to resist lodging, especially the content of cellulose and sucrose. PCA showed that sucrose metabolism and SC content were closely related to lodging resistance. Correlation analysis showed that the lodging index (LI) was significantly negatively correlated with NSC. Sucrose content was highly and significantly positively correlated with NSC. Additionally, the activities of sucrose phosphate synthase (SPS) and sucrose synthase (SS) were highly and significantly positively correlated with sucrose and cellulose content.
• The relationship between field characters and oat lodging, as well as the regulatory mechanism of carbohydrate content on lodging resistance of the culm are discussed.