超级稻花后强、弱势粒淀粉合成相关酶活性和激素含量变化及其与籽粒灌浆的关系

 种植4个超级稻品种\[两优培九和Ⅱ优084（杂交籼稻）、淮稻9号和武粳15（粳稻）\]和2个高产对照品种\[汕优63（杂交籼稻）和扬辐粳8号（粳稻）\],观察其结实期强、弱势粒中蔗糖合酶（SuSase）、腺苷二磷酸葡萄糖焦磷酸化酶（AGPase）、淀粉合酶（StSase）和淀粉分支酶（SBE）活性及玉米素+玉米素核苷（Z+ZR）、3 吲哚乙酸（IAA）和脱落酸（ABA）含量的变化,并测定了强、弱势粒灌浆速率。结果表明,超级稻品种强势粒的最大灌浆速率、到达最大灌浆速率的时间、平均灌浆速率和糙米重与对照品种差异较小,超级稻品种弱势粒的灌浆速率和糙米重显著低于对照。灌浆期强、弱势粒的SuSase、AGPase 、StSase 和SBE活性变化及Z+ZR、IAA 和ABA 含量变化均呈单峰曲线。弱势粒的SuSase、AGPase、StSase 和SBE的峰值活性和平均活性及其Z+ZR 和IAA的峰值含量和平均含量均低于强势粒。弱势粒的ABA 峰值含量和平均含量显著高于强势粒,超级稻品种高于对照品种。籽粒灌浆速率与SuSase、AGPase和StSase 活性及Z+ZR 和IAA含量均呈显著或极显著正相关,与SBE活性及ABA含量的相关不显著。说明超级稻品种弱势粒中较低的SuSase、AGPase和StSase活性及较低的Z+ZR和IAA含量是其灌浆速率小、粒重轻的一个重要原因。     

Changes in Enzyme Activities Involved in Starch Synthesis and Hormone Concentrations in Superior and Inferior Spikelets and Their Association with Grain Filling of Super Rice

The changes in activities of key enzymes involved in sucrose-to-starch conversion and concentrations of hormones in superior and inferior spikelets of super rice were investigated and their association with grain filling was analyzed.Four super rice cultivars,Liangyoupeijiu,IIyou 084,Huaidao 9 and Wujing 15,and two high-yielding and elite check cultivars,Shanyou 63 and Yangfujing 8,were used.The activities of sucrose synthase (SuSase),adenosine diphosphoglucose pyrophosphorylase (AGPase),starch synthase (StSase) and starch branching enzyme (SBE),and the concentrations of zeatin + zeatin riboside (Z + ZR),indole-3-acetic acid (IAA) and abscisic acid (ABA) in superior and inferior spikelets were determined during the grain filling period and their relationships with grain filling rate were analyzed.Maximum grain filling rate,the time reaching the maximum grain-filling rate,mean grain filling rate and brown rice weight for superior spikelets showed a slight difference between the super and check rice cultivars,but were significantly lower in the super rice than in the check rice for inferior spikelets.Changes of enzyme activities and hormone concentrations in grains exhibited single peak curves during the grain filling period.The peak values and the mean activities of SuSase,AGPase,StSase and SBE were lower in inferior spikelets than in superior ones,as well as the peak values and the mean concentrations of Z + ZR and IAA.However,the peak value and the mean concentration of ABA were significantly higher in inferior spikelets than in superior ones and greater in the super rice than in the check rice.The grain filling rate was positively and significantly correlated with the activities of SuSase,AGPase and StSase and the concentrations of Z + ZR and IAA.The results suggested that the low activities of SuSase,AGPase and StSase and the low concentrations of Z + ZR and IAA might be important physiological reasons for the slow grain filling rate and light grain weight of inferior spikelets in super rice.     

Research Advances in High-Yielding Cultivation and Physiology of Super Rice

In 1996,China launched a program to breed super rice or super hybrid rice by combining intersubspecific heterosis with ideal plant types.Today,approximately 80 super rice varieties have been released and some of them show high grain yields of 12-21 t/hm2 in field experiments.The main reasons for the high yields of super rice varieties,compared with those of conventional varieties,can be summarized as follows:more spikelets per panicle and larger sink size (number of spikelets per square meter);larger leaf area index,longer duration of green leaf,greater photosynthetic rate,higher lodging resistance,greater dry matter accumulation before the heading stage,greater remobilization of pre-stored carbohydrates from stems and leaves to grains during the grain-filling period;and larger root system and greater root activity.However,there are two main problems in super rice production:poor grain-filling of the later-flowering inferior spikelets (in contrast to earlier-flowering superior spikelets),and low and unstable seed-setting rate.Here,we review recent research advances in the crop physiology of super rice,focusing on biological features,formation of yield components,and population quality.Finally,we suggest further research on crop physiology of super rice.     

超级稻高产栽培生理研究进展

 我国于1996年启动了理想株型与亚种间杂种优势相结合为技术路线的超级稻育种计划,目前已育成约80个超级稻品种。这些品种产量高,兼顾品质与抗性,在试验示范区或特定气候条件下产量可达到12～21 t/hm2,展示了超级稻的巨大增产潜力。对于超级稻品种高产的原因,可概括为以下几个方面：与常规高产品种相比,超级稻品种每穗颖花数多、库容量大(即单位面积颖花数多);叶面积指数大、绿叶面积持续期长、光合速率高、茎秆抗倒性强;  抽穗前干物质累积量高, 结实期茎叶中碳水化合物转运到籽粒中的量大;  根量大、根系活性强。但是,超级稻品种在生产上也存在一些问题,突出地表现在两个方面：一是强势粒(着生在穗子中上部、开花较早的籽粒)、弱势粒(着生在穗子基部、开花较迟的籽粒)充实整齐性差;二是结实率低且不稳定。综述了超级稻的生物特性、产量结构、群体质量特征以及相关生理机理方面的研究进展,提出深入开展超级稻栽培生理研究的建议。     

吉林省47年来粳稻品种遗传改良过程中某些农艺性状的变化

对吉林省1958-2005年间育成的25个粳稻品种的产量及农艺性状的研究结果表明,随着粳稻品种的改良,产量呈线性增加。根据回归方程计算,产量由1958年的6 198.68 kg/hm2增加至2005年的12 682.8 kg/hm2,平均每年增加137.96 kg/hm2,增长率为2.23%。每穗粒数、结实率、单穗重和总穗数随育成年代推移显著增加,并与产量呈正相关,其中每穗粒数与产量的相关系数最大（r=0.85**）。株高随育成年代推移显著降低,茎叶夹角显著减小,灌浆期叶面积指数增加,抽穗期推迟。结果表明,产量的增加主要与品种改良过程中总库容的扩大有关,其中每穗粒数的增加是关键。     

Association of Phytate Formation with Grain Filling in Rice

The grain-filling characteristics of six rice varieties (combinations) and the relationships between their relative biochemistry composition during phytate synthesizing and grain plumpness were studied. Regarding results for ISHR1,ISHR2, R198 and JW21, with good grain plumpness, the two-step-filling in superior spikelets and inferior spikelets was not clear, while for ISHR3 and 559, with poor grain plumpness, it was very clear. From booting stage to flowering stage, the contents of phytate and inositol in varieties with good grain plumpness was obviously higher than those in varieties with poor grain plumpness. While at grain filling stage, the content of inorganic phosphorus in varieties with poor grain plumpness was obviously higher than that in varieties with good grain plumpness. The contents of phytate and inositol from booting stage to flowering stage was highly significantly correlated with the initial filling power (RO), the mean filling rate (RM) and grain filling percentage (PGF), and the content of inorganic phosphorus at grain filling stage was negatively significantly correlated with R0, FM and PGF. Furthermore, effective approach to improving grain filling was put forward.     

不同氮利用率粳稻品种的碳氮积累与转运特征及其生理机制

【目的】探明不同氮利用率水稻品种的氮素积累与转运特征及其机制。【方法】2个氮高效品种(武运粳30号和连粳7号)和2个氮低效品种(扬粳4038和宁粳1号)种植于大田,设置2个施氮量：全生育期不施氮(0N)和全生育期施氮180kg/hm²(180N),比较分析了不同氮利用率粳稻品种干物质生产、氮素积累与转运差异及其机制。【结果】与氮低效品种相比,氮高效品种具有较高的产量、氮肥利用率、总颖花量和结实率,较高的花前干物质转运量和花后干物质积累量,分蘖至穗分化始期和抽穗至成熟期较高的净同化率和作物生长率,抽穗期较高的糖花比,灌浆期较高的籽粒库活性、籽粒中脱落酸与1-氨基环丙烷-1-羧酸含量的比值和茎鞘中较高的非结构性碳水化合物的转运和蔗糖合成相关酶活性以及蔗糖转运蛋白基因的表达量,抽穗后较高的氮转运、氮素吸收量,灌浆期较高的比叶氮含量、叶片中细胞分裂素含量、氮代谢酶活性以及氮素转运相关基因的表达量。【结论】氮高效品种穗分化前和抽穗后较高的物质生产效率以及灌浆期较高的碳氮转运与积累是产量和氮肥利用率协同提高的重要机制。     

Association of Phytate Formation with Grain Filling in Rice

The grain-filling characteristics of six rice varieties (combinations) and the relationships between their relative biochemistry composition during phytate synthesizing and grain plumpness were studied. Regarding results for ISHR1,ISHR2, R198 and JW21, with good grain plumpness, the two-step-filling in superior spikelets and inferior spikelets was not clear, while for ISHR3 and 559, with poor grain plumpness, it was very clear. From booting stage to flowering stage, the contents of phytate and inositol in varieties with good grain plumpness was obviously higher than those in varieties with poor grain plumpness. While at grain filling stage, the content of inorganic phosphorus in varieties with poor grain plumpness was obviously higher than that in varieties with good grain plumpness. The contents of phytate and inositol from booting stage to flowering stage was highly significantly correlated with the initial filling power (R0), the mean filling rate (RM) and grainfilling percentage (PGF), and the content of inorganic phosphorus at grain filling stage was negatively significantly correlated with R0, FM and PGF. Furthermore, effective approach to improving grain filling was put forward.     

重穗型杂交稻D优527的栽培生理特点及调控对策研究

对重穗型杂交稻D优527与中等穗型的杂交稻组合汕优63和R527的栽培生理特性进行了比较。重穗型杂交稻D优527秆硬抗倒、穗大粒多、库容量大,单穗重大,产量高,且具很高的增产潜力。其茎鞘物质在齐穗后向穗部的转运量大、转运率和转换率高,但后期光合生产量相对较低,从而导致库虽大但由于源的相对不足而使库的充实度偏低。因此,增强后期光合生产力以增加源的供应是提高其籽粒充实度,进一步提高其产量的关键。D优527已具备高质量群体结构特征,对进一步发挥其高质量群体优势的高产栽培技术途径进行了探讨。     

内源IAA对杂交稻强、弱势粒灌浆增重的影响

以超级杂交稻协优9308和杂交稻协优9312、Ⅱ优162及其相应恢复系为材料,对灌浆初期籽粒中内源IAA、糖分的变化,以及IAA抑制剂对IAA以及3H 标记葡萄糖吸收分配和籽粒灌浆的影响进行了研究。结果表明在籽粒灌浆初期强势粒的IAA含量以及增重的速度始终高于弱势粒。在开花受精7 d以前,强势粒中果糖、葡萄糖及蔗糖的含量高于弱势粒,之后相反。强、弱势粒间存在的这种灌浆势差异,杂交组合大于其相应的恢复系。IAA合成及运输抑制剂明显降低3H 葡萄糖在籽粒中的分配。外源IAA对幼穗的处理,缩小了强、弱势粒间结实率的差异,使整穗的结实率也有所提高。     

Activities of Enzymes for Sucrose-Starch Conversion in Developing Endosperm of Rice and Their Association with Grain Filling in Extra-Heavy Panicle Types

Abstract

Rice cultivars with numerous spikelets per panicle (extra-heavy panicle types) frequently fail to exhibit their high yield potential due to low grain filling. Existing genetic variation in grain filling, however, opens possibilities for genetic improvement for this trait. We studied the correlation between grain filling and the activities of enzymes for sucrose-starch conversion in developing endosperm. The activity of sucrose synthase (EC 2.4.1.13, SuSy) and ADPglucose pyrophosphorylase (EC 2.7.7.27, AGPase), were measured in three extra-heavy panicle types and a standard cultivar grown at two locations under different environmental conditions. The proportions of grains with definite specific gravities and the rate of grain filling were adopted as the parameters related to grain filling. AGPase activity, but not that of SuSy, was consistently correlated to high proportions of high-density grains (specific gravity > 1.20) and high rates of grain growth in spikelets, particularly in those on secondary branches in which low grain filling is the rule. Such correlation was also detected in spikelets on primary branches which generally show better grain filling, but only early stages. Therefore, a high activity of AGPase might contribute to the reduction of the sucrose concentration by accelerating sucrose metabolism at the developing seed, a sink terminus of the phloem. Thus the sink-directed phloem transport of sucrose would be promoted, resulting in improved grain filling of extra-heavy panicle types. SuSy would play some roles in such a cultivar difference in grain filling, but depending on environments.     

烯效唑对杂交水稻籽粒灌浆及产量的影响研究

以杂交稻组合汕优63为材料,探讨了不同施药时期(浸种、孕穗期喷施)和不同质量浓度(0,10,20,40,80 mg/L)烯效唑处理对籽粒生长过程及产量的影响.结果表明,烯效唑对强势粒生长有一定抑制作用,但同时却促进了弱势粒的生长,在一定程度上具有协调强、弱势粒灌浆的作用,降低成熟期强势粒单粒重而提高弱势粒单粒重;使强势粒的终极生长量降低,达最大生长速率的时间延后,从而延长弱势粒的灌浆时间,提高弱势粒的灌浆速率及最大干重;由于烯效唑对弱势粒生长的促进作用,显著提高了其穗实粒数,进而提高了单穗籽粒干重及水稻产量.处理时期不同,其最佳浓度不同;浸种处理以质量浓度为40mg/L时产量最高,喷施处理以20mg/L时的产量最高.     

不同氮肥水平下结实期灌溉方式对水稻弱势粒灌浆及产量的影响

【目的】旨在阐明氮肥和灌溉方式对水稻产量、籽粒灌浆及生理特性的影响。【方法】以大穗型品种甬优2640和中穗型品种淮稻5号为供试材料进行盆钵试验,大田育秧移栽后设置3种氮肥水平,即0 N(不施氮)、MN(2 g N/盆)、HN(4 g N/盆);抽穗至成熟期设置3种灌溉方式,即CI(保持水层灌溉)、WMD(轻干湿交替灌溉,土壤水势-15 k Pa时复水)、WSD(重干湿交替灌溉,土壤水势-30 k Pa时复水)。【结果】在CI下,两个品种产量均以MN水平最高;WMD处理下,两个品种产量均以HN水平最高,但与MN下差异不显著,WSD处理下两个品种产量均以HN最高;而在籽粒灌浆上,两个品种强势粒的灌浆速率和最终粒重在各个水氮处理间无显著差异,弱势粒的灌浆速率和最终粒重在良好水势条件CI和轻度水分胁迫WMD下,分别在0 N和MN水平下表现较优;但在重度水分胁迫WSD下,0N水平表现最低,HN最高,但与MN差异不显著。以上都表明产量与弱势粒的灌浆在水氮间存在着明显的交互作用。在品种间,大穗型籼粳杂交稻甬优2640弱势粒灌浆速率及粒重都低于中穗型常规粳稻淮稻5号,其产量优势主要源自较高的每穗粒数。最后,WMD+MN处理下有较高的氮肥利用率,较少的施氮量获得较高的产量,达到节水节氮增产的效果,其次也增加了根系生理活性和叶片光合性能,非结构性碳水化合物(NSC)转运率,促进了地上部的生长发育,同时也加强了物质运转,促进了灌浆中后期弱势粒籽粒的充实,最终达到产量增加的目的,成为本研究最佳水氮运筹方式。     

南方晚粳杂交稻与常规稻籽粒灌浆及后期叶片光合特性的差异

以南方杂交晚粳秀优5号和常规晚粳秀水110为材料,应用Richards方程对两个品种的籽粒灌浆过程进行拟合,研究了两个水稻品种的灌浆特性,并测定分析了两类品种的后期叶片光合特性。 秀优5号的起始灌浆势、最大灌浆速率和平均灌浆速率均略低于秀水110,但活跃灌浆时间明显长于秀水110,说明杂交晚粳灌浆更为平缓、灌浆期长,有利于获得较高粒重,也有利于缓解其穗型较大引起的籽粒间物质的激烈竞争。齐穗后10 d,秀优5号的光合和叶绿素荧光参数与秀水110无显著差异,但齐穗后25 d,秀优5号除初始荧光值的增加幅度未达显著水平外,其他参数均显著高于秀水110。研究结果表明,南方杂交粳稻在灌浆后期仍保持较高的物质生产能力,是它具有较长籽粒灌浆持续期,大穗与较高粒重相协调,最终实现高产的重要生理基础。     

水分和氮肥对大穗型水稻籽粒灌浆结实的影响与生理分析

大穗型水稻具有较大的库容,增产潜力大。灌浆结实期的水分和氮肥运筹对水稻产量和品质形成起关键作用,研究此阶段水分和氮肥对大穗型水稻生理机制的影响,对于指导大穗型水稻的高产优质栽培具有现实意义。本文综述了有关大穗型水稻籽粒灌浆结实特点、水分和氮肥对大穗型水稻灌浆结实的影响及其生理机制的研究结果,并提出了一些目前大穗型水稻研究中存在的问题以及今后的研究方向。     

水氮管理对不同氮效率水稻根系性状、氮素吸收利用及产量的影响

【目的】探究水氮管理措施对不同氮效率水稻根系构型、氮素吸收利用和产量形成的影响,以及根系性状特征与氮素吸收利用和产量关系。【方法】试验采用三因素裂裂区设计,主区为2个不同氮效率水稻品种德香4103(氮高效型)和宜香3724(氮低效型),裂区设置"常规灌溉"和"控制性交替灌溉"2种水分管理方式,裂裂区为SPAD指导施肥、优化施肥以及农民习惯施肥3种施氮模式,运用岭回归分析方法探究根系构型与氮素吸收利用和产量的关系。【结果】水稻抽穗期根系性状、产量、每穗粒数、千粒重及总颖花量均存在显著的基因型差异。氮高效品种德香4103每穗粒数多,群体库容量大,产量较氮低效品种宜香3724高0.24%~11.31%;控制性交替灌溉有利于水稻千粒重的增长,常规灌溉则对水稻有效穗数、每穗粒数及群体颖花量提高更为有利;SPAD指导施肥和优化施肥处理能够提高水稻有效穗数和每穗粒数,扩大群体颖花量以保证其对农民习惯施肥的产量优势;由于水氮互作效应的存在,控制性交替灌溉下施氮处理与空白处理水稻千粒重的差距比常规灌溉的大幅降低,使得控制性交替灌溉下施用氮肥的增产效果更佳。德香4103的氮肥生理利用率较宜香3724平均高8.69%,常规灌溉下水稻氮积累量较高,控制性交替灌溉下氮肥回收率、农学利用率、生理利用率均较优;与农民习惯施肥处理相比,SPAD指导施肥与优化施肥模式更有利于水稻氮素吸收利用效率的提高。拔节期、抽穗期和成熟期水稻根系构型与产量岭回归方程的决定系数范围为0.4198~0.9028,其中,抽穗期根系性状与产量关系最为密切,氮高效和氮低效品种的决定系数均超过0.9。在拔节期,水稻细分枝根长对产量影响最大;在抽穗期,氮高效和氮低效品种存在差异,前者是粗分枝根长,后者是细分枝根表面积对产量影响最大;在成熟期,不定根长与产量关系最为密切。水稻抽穗期根系构型对氮积累量变化的解释程度较高,岭回归方程决定系数均接近0.7。就水氮管理措施而言,氮高效和氮低效水稻均应采用常规灌溉配套SPAD指导施肥或控制性交替灌溉结合优化施肥来实现产量的提高。【结论】水稻抽穗期根系构型与产量、氮积累量关系密切,采用合理的水氮管理措施能够实现水稻产量和氮素吸收利用效率的同步提高。     

Effects of high temperature and shading on grain abscisic acid content and grain filling pattern in rice (Oryza sativa L.)

ABSTRACT

Abscisic acid (ABA) is a key factor regulating starch biosynthesis genes and is involved in assimilate partitioning to individual spikelets. The objective of this study was to clarify the effects of high temperature and shading during grain filling on grain ABA content and the grain filling pattern of spikelets located at different positions in a panicle. We grew the rice cultivar ‘Koshihikari’ in pots in 2009 under two temperature regimes and two light conditions during grain filling. We periodically measured grain dry weight and grain ABA content (pmol per grain) and concentration (pmol per grain dry weight). Shading increased a grain weight difference between superior and inferior spikelets while high temperature decreased the difference regardless of light condition. High temperature decreased ABA content and concentration in grains. There was a close correlation between mean grain ABA content and mean grain-filling rate averaged over the first half of grain filling.     

产量构成因素及穗部性状对籼稻品种库容的影响

 在群体水培条件下,以国内外不同年代育成的88份（2001年）和122份（2002年）常规籼稻代表品种为材料,测定它们的茎蘖动态、产量及其构成因素、穗部性状等,采用组内最小平方和的动态聚类方法将供试品种按库容量从低到高依次分为A、B、C、D、E、F共6类,研究各类品种产量及其构成因素、穗部性状的基本特点。结果表明：1）供试品种间库容量的差异很大,库容量最大的品种比最小的品种高426%（2001年）、817%（2002年）,F类品种比A、B、C、D、E类品种分别高18471%、8893%、5745%、3432%、1405%（2001年）和30505%、15354%、9028%、5124%、2129%（2002年）;2）随着库容量水平的提高,产量呈显著增加趋势;3）大库容类型籼稻品种单位面积穗数、每穗粒数、千粒重均显著大于小库容类型品种;4）大库容籼稻品种平均穗长、着粒密度、每穗一次枝梗数、每穗二次枝梗数较大;5）提高每穗粒数、单位面积穗数均可促进单位面积库容量的提高,两者对库容量作用的大小相仿,但均显著大于千粒重对库容量的影响。着粒密度、穗长对每穗粒数的影响显著大于每穗二次枝粳数,以及一、二次枝粳数比值对每穗粒数的影响。     

机直播杂交籼稻结实期物质转运与不同粒位米质的关系

选用8个杂交籼稻品种为试验材料,比较机直播不同品种间结实期物质转运、产量及穗部不同粒位（强、弱势粒,一次、二次枝梗,上、中、下部）米质的差异,并探讨结实期物质转运与不同粒位米质的关系,为机直播杂交籼稻品种筛选提供理论依据。结果表明,具有较高的颖花数和千粒重,且结实期具有较高的茎鞘和叶片物质输出与转换率,是机直播高产品种的主要特征。强势粒、一次枝梗籽粒垩白度和垩白粒率均低于弱势粒及二次枝梗籽粒,食味值均高于弱势粒及二次枝梗籽粒;穗上部籽粒米质特征均不同程度高于穗中部和下部籽粒,但穗中部和下部籽粒米质特征品种间存在差异;宜香优2115不同粒位食味值均不同程度高于其他品种。相关分析表明,结实期茎鞘物质输出率及转换率与不同粒位垩白均存在显著或极显著负相关（r=-0.66^*~-0.85^**）,与不同粒位食味值呈显著或极显著正相关（r=0.67^*~0.86^**）。提高机直播杂交籼稻结实期茎鞘物质转运率及转换率可改善不同粒位籽粒食味品质,宜香优2115作机直播在产量及米质上综合表现好,尤其在食味品质方面优势明显,适宜作为机直播水稻品种应用。     

水稻颖花与籽粒发育模拟的初步研究

在文献资料与田间试验研究的基础上,建立了基于过程的水稻颖花原基分化与退化数、结实粒数以及灌浆动态预测的机理模型。模型量化了穗部器官发育与生理发育时间（PDT）及环境因子的动态关系,确定了穗部籽粒发育的不均衡性与开花时间和同化物供应状况的关系,进一步模拟了不同类型品种强势粒与弱势粒不同粒位的灌浆动态。模型验证结果表明,此模型对颖花动态和籽粒灌浆过程有较好的预测性。