高浓度CO2对稻穗不同位置籽粒结实和米质性状的影响

不断升高的大气CO₂浓度影响水稻颖花发育、灌浆结实和品质形成,但这种影响是否与籽粒在稻穗上的着生部位有关尚不清楚.利用稻田FACE (Free-Air CO₂ Enrichment)平台,以优质丰产粳稻‘武运粳23’为材料,CO₂处理设背景CO₂浓度(Ambient)和高CO₂浓度(增200 μmol·mol^-1, FACE)两个水平,研究开放大田条件下高浓度CO₂对水稻颖花密度、籽粒结实能力、稻米外观和食味品质的影响及其与稻穗不同着生位置的关系.结果表明:FACE处理使武运粳23籽粒产量平均增加18.3%,从产量构成因素看,穗数和饱粒重分别增加21.4%、9.4%,每穗颖花数、饱粒率平均减少9.0%、2.2%.FACE水稻饱粒率下降主要与稻穗不同部位空粒率大幅增加有关.FACE水稻每穗颖花数减少主要与稻穗上部、中部二次枝梗现存颖花大幅减少有关,而其他位置颖花数均无显著变化;稻穗不同位置饱粒重和饱粒率对FACE的响应无显著差异.FACE处理使绿粒率下降,但糙米长度和宽度均增加,稻穗不同部位趋势一致.FACE使垩白粒率(增幅59%)、垩白度(增幅55%)均极显著增加,增幅表现为稻穗一次枝梗>二次枝梗、上部>中部>下部.FACE使稻穗不同位置稻米直链淀粉含量略增,使最高粘度、热浆粘度、崩解值、最终粘度和消减值略降,但多未达显著水平.FACE使稻米糊化温度显著下降,弱势粒的降幅大于强势粒.综上,高浓度CO₂环境下武运粳23产量增加主要与穗数增多和籽粒增重有关,而稻穗明显变小;高浓度CO₂使稻米绿粒率减少,垩白增多,而对蒸煮食味品质影响较少;颖花着生位置对高浓度CO₂环境下水稻颖花发育、结实和品质的影响因不同测定指标而异.     

Hormonal changes in the grains of rice subjected to water stress during grain filling

Lodging-resistant rice (Oryza sativa) cultivars usually show slow grain filling when nitrogen is applied in large amounts. This study investigated the possibility that a hormonal change may mediate the effect of water deficit that enhances whole plant senescence and speeds up grain filling. Two rice cultivars showing high lodging resistance and slow grain filling were field grown and applied with either normal or high amount nitrogen (HN) at heading. Well-watered and water-stressed (WS) treatments were imposed 9 days post anthesis to maturity. Results showed that WS increased partitioning of fixed (14)CO(2) into grains, accelerated the grain filling rate but shortened the grain filling period, whereas the HN did the opposite way. Cytokinin (zeatin + zeatin riboside) and indole-3-acetic acid contents in the grains transiently increased at early filling stage and WS treatments hastened their declines at the late grain filling stage. Gibberellins (GAs; GA(1) + GA(4)) in the grains were also high at early grain filling but HN enhanced, whereas WS substantially reduced, its accumulation. Opposite to GAs, abscisic acid (ABA) in the grains was low at early grain filling but WS remarkably enhanced its accumulation. The peak values of ABA were significantly correlated with the maximum grain filling rates (r = 0.92**, P < 0.01) and the partitioning of fixed (14)C into grains (r = 0.95**, P < 0.01). Exogenously applied ABA on pot-grown HN rice showed similar results as those by WS. Results suggest that an altered hormonal balance in rice grains by water stress during grain filling, especially a decrease in GAs and an increase in ABA, enhances the remobilization of prestored carbon to the grains and accelerates the grain filling rate.     

Hormones in the grains in relation to sink strength and postanthesis development of spikelets in rice

Inferior spikelets usually exhibit a slower grain filling rate and lower grain weight than superior spikelets in a rice (Oryza sativa L.) panicle. This study investigated whether the variations in grain filling between the two kinds of spikelets were attributed to their sink strength and whether the sink strength was regulated by the hormonal levels in the grains. Using two field-grown rice genotypes, the division rate of endosperm cells, hormonal levels in the grains, and grain weight of both superior and inferior spikelets were determined during the grain filling period. The results showed that superior spikelets had dominance over inferior spikelets in endosperm cell division rate and cell number, grain filling and grain weight. Changes in zeatin (Z) and zeatin riboside (ZR) contents paralleled and were very significantly correlated with the cell division rate and cell number. Cell division rate and the content of indole-3-acetic acid (IAA) in the grains were also significantly correlated. Gibberellin (GAs; GA₁+ GA₄) content of the grains was high but ABA levels were low at the early grain filling stage. ABA increased substantially during the linear phase of grain growth and was very significantly correlated with grain dry weight during this period. Application of kinetin at 2 through 6 days post anthesis (DPA) significantly increased cell number, while spraying ABA at 11 through 15 DPA significantly increased the grain filling rate. The results suggest that differences in sink strength are responsible for variations in grain filling between superior and inferior spikelets. Both cytokinins and IAA in the grains may mediate cell division in rice endosperm at early grain filling stages, and therefore regulate the sink size of the grain, whereas ABA content correlates with sink activity during the linear period of grain growth.     

Apoplasmic assimilates and grain growth of contrasting rice cultivars differing in grain dry mass and size

Apical dominance in assimilate filling impacts grain growth in basal spikelets of rice panicle. In this study, organic materials of the pericarp, apoplasmic space and endosperm of the apical and basal caryopses, and photosynthesis of the flag leaf were measured during early part of grain development in three types of rice cultivars with similar phenology, but difference in grain weight and size in the dry and wet seasons of 2006 and 2007, respectively. Photosynthetic activity of the flag leaf was consistently low in small-seeded cultivars. Rates of grain filling and cell division of endosperm and concentration of assimilates, starch, proteins and chlorophylls of the caryopsis were lower, but spikelet ethylene production and peroxidase activity were higher in a small-seeded cultivar compared to a big-seeded cultivar. Similar disparities in grain filling and other attributes were noticed for the inferior basal spikelets of the panicle compared to the superior apical spikelets, except the assimilate concentration of the pericarp and endosperm. Temporal fluctuation in assimilate concentration of the organs were similar between the cultivars. Concentration of apoplasmic assimilates mostly exhibited negative correlation with that of pericarp and endosperm. Compared to the apical spikelets, correlation was more negative for the basal spikelets. Conversely, correlation was positive between the concentration of apoplasmic assimilates and endosperm cell number and grain weight of the cultivars. Ethylene released from the spikelets at anthesis affected growth and cell division rates of endosperm and enhanced protein and chlorophyll degradation and peroxidase activity of the caryopsis. It was concluded that variation in spikelet ethylene production may be responsible for differences in size or weight of grains among rice cultivars and spikelets at different locations of the panicle. The concentration of apoplasmic assimilates could be an indicator for grain filling capacity, and ethylene regulated the concentration by affecting pericarp activity for assimilate unloading.     

Ethylene control of seed coat development in low and high sterile semidwarf indica rice cultivars

Although basally positioned inferior spikelets of rice panicles emerge late from the flag leaf enclosure (boot), they mature early which precludes adequate grain filling. It is assumed that extended exposure to ethylene inside the boot restricts assimilate partitioning to the endosperm in basal spikelets by affecting the functions of seed coat. In the present study, ethylene concentration inside the boot was measured in two high yielding rice cultivars differing in percentage of spikelet sterility. To manipulate the concentration/action of ethylene, silver nitrate, aminoethoxyvinyl glycine and 2-chloroethylphosphonic acid were injected into the boot. The effect of these chemicals on the concentration of photosynthetic pigments, lipid peroxidation and peroxidase activity of the seed coat and lemma and palea were measured to monitor development. Ethylene reduced development during the juvenile phase but accelerated degradation of the photosynthetic tissues of the spikelets in the senescent phase. Boot ethylene correlated positively with number of barren spikelets in the panicle and negatively with concentration of photosynthetic pigments of the seed coat of inferior spikelets. The concentration of ethylene was higher in the high sterile cultivar Mahalaxmi than that of the low sterile Mahanadi. Inhibition of ethylene action or synthesis improved grain filling. The chemicals were most effective on the inferior spikelets. It was concluded that ethylene retarded seed coat development during the prestorage phase and reduced grain filling of basal spikelets.     

籼型常规早稻穗分化期低温对颖花形成和籽粒充实的影响

以籼型常规早稻中嘉早17为材料,于盆栽条件下采用人工气候箱控温,在水稻穗分化一次枝梗原基分化期(Ⅱ)与花粉母细胞减数分裂期(Ⅵ)进行17和20 ℃的低温胁迫处理,研究不同低温对水稻枝梗、颖花分化与退化及籽粒充实的影响.结果表明： 与对照相比,不同低温处理均显著降低每穗枝梗及颖花分化数和现存数,颖花现存数降幅为7.2%～12.4%,同时增加了枝梗和颖花的退化数,影响了花粉活性、花药开裂等花器官发育,导致籽粒充实不良,以17 ℃低温胁迫效应更明显.穗分化Ⅵ期低温处理总枝梗和颖花分化数与现存数低于穗分化Ⅱ期,但二次枝梗和颖花退化数较多,颖花退化数较穗分化Ⅱ期高11.6%;穗分化Ⅱ期低温处理穗部籽粒结实率显著低于穗分化Ⅵ期,降幅达3.7%,主要与花粉粒活性、柱头花粉散落数、花药开裂系数和籽粒充实度受低温影响较大有关.另外,穗分化Ⅱ、Ⅵ两时期受17 ℃低温胁迫效应大于20 ℃.综合穗分化两时期低温胁迫效应的差异,生产中需加强相应栽培措施的调控.     

川东南高温伏旱区杂交中稻超稀栽培对稻米整精米率的影响与组合间库源结构的关系

以18个杂交中稻组合材料,研究了超稀栽培与传统栽培间稻米整精米率的变异与组合间库源结构的关系。结果表明:中小穗型组合在超稀栽培条件下的整精米率会明显改善,而大穗型组合的整精米率反而降低。其原因在于中小穗型组合在传统栽培条件下的叶粒比和灌浆速率较高,改为超稀栽培后,叶粒比和灌浆速率明显降低,相对提高了籽粒容重,近而提高了整精米率。大穗型组合在传统栽培条件下的叶粒比和灌浆速率较低,改为超稀栽培后,叶粒比和灌浆速率进一步明显降低,此时因库源比例失调,源明显不足以致籽粒充实度差,从而相对降低了籽粒容重和整精米率。超稀栽培应选用传统栽培条件下着粒数不超过177粒的组合为宜。     

Demand for Assimilates Determines the Productivity of Intensive and Extensive Rice Crops in Primorskii Krai

Intensive and extensive rice crops (Oryza sativa L.), regionally cultivated in Primorskii krai (maritime territory), were grown under full and 50% attenuated solar radiation. Plants of different varieties were used to examine the supply of newly synthesized and reutilized ¹⁴C-assimilates to caryopses and to estimate the dry weight dynamics of whole plants, vegetative organs, and grains. Cultivar-specific differences were revealed with respect to the sink capacity of caryopses, the export of photosynthates from the upper leaf and their delivery to the panicle, and the contributions of newly produced and reutilized assimilates to grain filling. In rice plants of all varieties grown under full insolation, the amount of photosynthates produced during grain filling was insufficient to satisfy the demand of caryopses; one-fourth or one-fifth of this demand was satisfied at the expense of mobilization of stored metabolites. The mobilization was accelerated by the elevated demand for assimilates and by attenuated insolation. In artificially shaded plants of intensive varieties, the pool of newly produced assimilates was lower and reutilization of previously gained assimilates started earlier than in shaded plants of extensive varieties. It is concluded that the higher grain yield of intensive rice varieties, cultivated in Primorskii krai, is determined by a higher demand for assimilates and by a higher production and accelerated supply of newly formed photosynthates to caryopses during the first half of the grain-filling stage. The potential productivity of these varieties is constrained by the deficit of assimilates during the second half of grain-filling stage. The low grain productivity of extensive varieties is caused by the insufficient number of grains in panicles and by low demand for assimilates throughout the period of grain filling.     

Involvement of polyamines in the post-anthesis development of inferior and superior spikelets in rice

Early-flowered superior spikelets usually exhibit a faster grain filling rate and heavier grain weight than late-flowered inferior spikelets in rice (Oryza sativa L.). But the intrinsic factors responsible for the variations between the two types of spikelets are unclear. This study investigated whether and how polyamines (PAs) are involved in regulating post-anthesis development of rice spikelets. Six rice genotypes differing in grain filling rate were field grown, and PA levels and activities of the enzymes involved in PA biosynthesis were measured in both superior and inferior spikelets. The results showed that superior spikelets exhibited higher levels of free spermidine (Spd) and free spermine (Spm) and higher activities of arginine decarboxylase (ADC, EC 4.1.1.19), S-adenosylmethionine decarboxylase (SAMDC, EC 4.1.1.50) and Spd synthase (EC 2.5.1.16) than inferior spikelets at the early endosperm cell division and grain filling stage. The maximum concentrations of free Spd and free Spm and the maximum activities of ADC, SAMDC and Spd synthase were significantly correlated with the maximum cell division and grain filling rates, maximum cell number and grain weight. Application of Spd and Spm to panicles resulted in significantly higher rates of endosperm cell division and grain filling in inferior spikelets along with the activities of sucrose synthase (EC 2.4.1.13), ADP glucose pyrophosphorylase (EC 2.7.7.27) and soluble starch synthase (EC 2.4.1.21), suggesting that these PAs are involved in the sucrose-starch metabolic pathway. The results indicate that the poor development of inferior spikelets is attributed, at least partly, to the low PA level and its low biosynthetic activity.     

转拟南芥NPR1基因桂99 T3代植株的抗病性及农艺性状表现

以转拟南芥AtNPR1基因的恢复系品种桂99T3代纯合株系为材料,考查其农艺性状及其抗病性,并比较转基因植株与桂99侵染水稻白叶枯病菌后的农艺性状。结果表明:转基因植株表现出对水稻白叶枯病的抗性显著增强77%以上;穗长、剑叶长、有效穗数、一次枝梗数、每穗实粒数、单株产量和谷粒宽等农艺性状与未转基因桂99无显著差别。在受到水稻白叶枯病菌侵染后,转基因植株的一次枝梗数、每穗粒数、每穗实粒数和单株产量等方面均比对照桂99高出13%～78%。说明AtNPR1基因增强了水稻的抗病能力,从而降低了病害引起的产量损失。转基因植株的恢复力不受影响,稻米品质比桂99更加优良。本工作为转基因水稻抗病育种的研究奠定了基础。     

水稻籼粳交DH群体籽粒充实度的遗传分析

籽粒充实度差是限制亚种间杂交稻产量潜力发挥的重要因素。作者对籼粳交（圭630／02428）DH群体的籽粒充实度进行了研究,结果表明,籽粒充实度是受多基因控制的数量性状,在该DH群体中发生分离的基因估计数为5－6对。通过估测三级（偏度）和四级（峰度）统计量,检测到控制籽粒充实度的多基因间存在重叠作用。DH群体中籽粒充实度表现出明显的超亲分离,表明基因重组可实现控制籽粒充实度的增效基因的聚合。籽粒充实度与单穗产量、穗均实粒数和干物质／总库容之比呈显著或极显著正相关,与千粒重、穗均颖花数和穗均总库容相关不显著。     

Comparative mapping of QTLs for agronomic traits of rice across environments using a doubled haploid population

We report here the RFLP mapping of quantitative triat loci (QTLs) that affect some important agronomic traits in cultivated rice. An anther culturederived doubled haploid (DH) population was established from a cross between an indica and a japonica rice variety. On the basis of this population a molecular linkage map comprising 137 markers was constructed that covered the rice genome at intervals of 14.8cM on average. Interval mapping of the linkage map was used to locate QTLs for such important agronomic traits as heading date, plant height, number of spikelets per panicle, number of grains Per panicle, 1000-grain weight and percentage of seed set. Evidence of genotype-byenvironment interaction was found by comparing QTL maps of the same population grown in three diverse environments. A total of 22 QTLs for six agronomic traits were detected that were significant in at least one environment, but only 7 were significant in all three environments, 7 were significant in two environments and 8 could only be detected in a single environment. However, QTL-by-environment interaction was traitdependent. QTLs for spikelets and grains per panicle were common across environments, while traits like heading date and plant height were more sensitive to environment.     

Comparative mapping of QTLs for agronomic traits of rice across environments by using a doubled-haploid population

We report here the RFLP mapping of quantitative trait loci (QTLs) which affect some important agronomic traits in cultivated rice. An anther culture-derived doubled-haploid (DH) population was established from a cross between indica and japonica rice varieties. A molecular linkage map comprising 137 markers was constructed based on this population which covered the rice genome at intervals of 14.8 cM on average. The linkage map was used to locate QTLs for such important agronomic traits as heading date, plant height, number of spikelets per panicle, number of grains per panicle, 1 000-grain weight and the percentage of seed set, by interval mapping. Evidence of genotype-by-environment interaction was found by comparing QTL maps of the same population grown in three diverse environments. A total of 22 QTLs for six agronomic traits was detected which were significant in at least one environment, but only seven were significant in all three environments; seven were significant in two environments and eight could only be detected in a single environment. However, QTLs-by-environment interaction was trait dependent. QTLs for spikelets and grains per panicle were common across environments while traits like heading date and plant height were more sensitive to environment. Received: 22 February 1996 / Accepted: 10 May 1996     

Dynamic Changes of Endogenous Plant Hormones in Rice Grains in Different Parts of Panicle at Grain filling Stage

The grain fresh weight and grain-filling rate at early grain developing stage in rice ( Oryza sativa L. ) were higher in the upper than in the lower part of panicle during grain filling. The grains in the upper and/or the lower part of panicle were analyzed for the endogenous plant hormones. The results indicated that the peak content of ABA and/or GA1 + 3 in the developing grains appeared at about the same stage when the highest increasing rate of grain appeared in the upper and in the lower part of the panicle respectively. After heading, the IAA content in developing grains steadily decreased in both the upper and the lower part of the panicle, GAn + 7 content decreased at first and then increased, and again decreased. At the early grain-filling stage, the ABA and GA1 + 3 contents in the developing grains were higher in the upper than in the lower part of the panicle. The role that the hormones play in the developing grains and the reason why the grains seed set better in the upper than in the lower part of the panicle were discussed.     

水稻结实过程中穗不同部位谷粒中内源激素的动态变化

水稻（ＯｒｙｚａｓａｔｉｖａＬ．）结实期间,穗上部谷粒的粒重比穗下部谷粒的高,灌浆前期的增重速率比穗下部谷粒的高,在整个灌浆过程中增重速率的变化比穗下部谷粒的快。利用ＥＬＩＳＡ方法对穗上部谷粒和穗下部谷粒中各种内源激素含量的分析表明,在灌浆过程中,谷粒中未检测出ｉＰＡｓ;在穗上部谷粒和穗下部谷粒中,ＡＢＡ和ＧＡ(１＋３)的含量的最大值出现的时间分别与穗上部谷粒和穗下部谷粒增重最快的时间基本一致,无论穗上部谷粒还是穗下部谷粒中,ＩＡＡ的含量基本上均呈下降趋势;ＧＡ(４＋７)的含量是先上升、后下降、再上升;在结实的前中期,谷粒中ＡＢＡ、ＧＡ(１＋３)的含量基本上均是穗上部谷粒高于穗下部谷粒。探讨了各种激素在谷粒灌浆过程中的作用及从激素的角度探讨了穗上部谷粒比穗下部谷粒结实性好的原因。     

Transport of Recently Assimilated15N Nitrogen to Individual Spikelets in Spring Wheat Grown in Culture Solution

Two cultivars of spring wheat ( Triticum aestivum L.), Sport(high protein) and WL4 (low protein), were grown to maturityin culture solution. Nitrogen in the form of nitrate was addedin daily doses at stepwise-decreasing relative rates to ensurenormal development, and both cultivars received the same totalamount of N during development. At weekly intervals from anthesisto maturity the daily nitrate dose was, for selected groupsof plants, labelled with¹⁵N. After the labelling period theselected plants were harvested and analysed. The cultivar WL4produced more biomass than Sport, as well as more spikeletsand more grains per ear, with a higher mean grain weight, suchthat grain yield of WL4 was 57% greater than Sport. The earsof both cultivars were heterogenous: mean grain weight was highestin middle spikelets, which also contained more grains; the Ncontent followed the pattern of dry weight with more N in themiddle spikelets; but the N concentration was practically thesame in all spikelets (2.15% of d. wt in WL4 and 3.33% in Sport).The distribution of¹⁵N showed that the main stem ear maturedmuch earlier than tiller ears. The results of this nitrogen-labellingexperiment show that, late in development, substantial amountsof recently-absorbed N were immediately assimilated and transportedto the ears. Transport of¹⁵N decreased earlier to the top spikeletsthan to the bottom spikelets. As both cultivars were grown underidentical conditions and both received the same amount of Nit was concluded that the difference in grain N concentrationwas not caused by differences in the capacity of N assimilationand translocation but rather by different rates of accumulationof non-nitrogenous dry matter in the grains. Ear; grain; nitrate; nitrogen transport; Triticum aestivum L.; yield     

Most susceptible stage of rice panicle development to Oebalus pugnax (Hemiptera: Pentatomidae)

Oebalus pugnax (F.) (Hemiptera: Pentatomidae) damage to rice, Orya savita L., reduces rough and head rice yields, and grain quality. O. pugnax feeds on developing kernels, introducing pathogenic fungi and causing a discoloration of the grain known as "peck." The objective of this study was to determine the stage of rice panicle development most susceptible to O. pugnax attack. During 2005 and 2006, in greenhouse and field experiments, rice plants were caged at the boot stage and then infested with adult or nymphal O. pugnax. Plants were infested during one of three stages of panicle development: heading, milk, or soft dough. Insects were allowed to feed on the plants for the duration of each stage and then killed. After maturation, panicles were harvested, and grain was hulled and milled. Grain weight, percentage of pecky grain, and percentage of whole grain after milling were recorded. No differences were found in the weight of rough, brown, or milled rice infested with O. pugnax during different stages of panicle development. Number of filled grains per cage was not affected by O. pugnax, and number of empty grains per cage was affected in two of four experiments. Higher percentage of peck was found in grain from panicles infested during dough and milk than in grain from panicles infested during heading. Adult O. pugnax caused higher percentage of peck than nymphs in all stages of panicle development. An inverse relationship was found between percentage of peck and percentage of whole grain weight only in one of the experiments.     

重穗型杂交水稻籽粒灌浆过程中强势和弱势颖花内源IAA、ABA和GA水平的动态状况

研究了重穗型杂交水稻培矮 6 4s/E3 2的灌浆过程和强、弱势颖花中内源IAA、ABA和GA1 GA3水平的动态状况。籽粒发育过程中不同内源激素水平高低依次为 :IAA >GA1 GA3>ABA。IAA和ABA水平在强势颖花中较高而GA1 GA3水平在弱势颖花中较高。 3种激素水平的变化与谷粒增重速率之间均存在正相关 ,两个最高的相关系数值分别存在于单位鲜重样本的IAA含量(ng/gFW ) 与籽粒鲜重的增重速率之间 (r =0 .82 1 8 )和单个籽粒IAA含量 (ng/grain)与籽粒干重的增重速率之间 (r =0 .8485 )。推测启动和维持籽粒灌浆过程可能需要较高的IAA水平 ;ABA可能具有促进籽粒中同化物的累积和种子成熟的作用 ;GA1 GA3可能具有保持弱势颖花活性的特殊作用