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     

重穗型杂交水稻籽粒灌浆过程中强势和强势颖花内源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     

不同水分管理下优质稻花后植株碳氮流转与籽粒生长及品质的相关性

以桂华占、八桂香为材料,在干湿交替灌溉、亏缺灌溉、淹水灌溉3种水分条件下,研究优质稻花后植株碳氮流转与籽粒生长及品质的相关性。结果表明:不同水分管理下,桂华占和八桂香花后碳氮流转与籽粒的生长间存在密切相关。主要表现在:(1)茎鞘和叶片干物质转运对籽粒干物质积累的贡献率为16.86%～25.68%,花后茎叶干物质运转速度和运转率与籽粒起始灌浆势呈显著甚至极显著正相关;籽粒最大灌浆速率、活跃灌浆期、持续灌浆时间与叶片干物质运转速度和运转率呈极显著正相关,与茎鞘干物质运转速度和运转率呈极显著负相关;(2)茎鞘碳同化物转运对籽粒的产量和淀粉产量的贡献率则为干湿交替灌溉>亏缺灌溉>淹水灌溉;但叶片碳同化物转运对籽粒的产量和淀粉产量的贡献率则为淹水灌溉>亏缺灌溉>干湿交替灌溉;茎叶可溶性糖积累量的减少和籽粒直链淀粉含量和积累量增加是同步的,且茎叶可溶性糖积累量快速递减期(花后3～12d)与直链淀粉含量和积累量快速递增期(花后6～12d)同步;(3)茎鞘和叶片氮素转运对籽粒氮素积累的贡献率为44.05%～117.66%,叶片总氮转运对籽粒氮素积累的贡献率大于茎鞘,茎鞘和叶片氮同化物对籽粒氮素的贡献率以淹水灌溉处理的最大,亏缺灌溉处理的次之,干湿交替灌溉处理的最小。     

Grain sink strength may be related to the poor grain filling of indica-japonica rice (Oryza sativa) hybrids

The physiological and biochemical factors contributing to poor grain filling of indica-japonica rice ( Oryza sativa L.) hybrids were studied by analyzing the role of grain sink strength in dry matter accumulation of grains of two types of rice cultivars, Yayou 2 (an indica-japonica hybrid) and Yanjing 2 (a japonica cultivar). Carbon dioxide enrichment and plant hormone application were imposed at anthesis and the number of endosperm cells, dry matter accumulation and the activities of some sugar-metabolizing enzymes of grains were measured during grain filling. In Yayou 2, strong-potential grains (SPGs) accumulated dry weight much earlier than weak-potential grains (WPGs), but this difference was not obvious for Yanjing 2. Carbon dioxide enrichment imposed after heading significantly stimulated dry matter accumulation of WPGs of Yayou 2, but had little influence on WPGs of Yanjing 2 and SPGs of both cultivars. Leaf sheath dry matter decreased steadily in both cultivars during early stages of grain filling and accumulated during the later stages. Carbon dioxide enrichment increased leaf sheath dry matter. Dry matter accumulated by grains was linearly related to the increases in endosperm cell numbers and the activities of sucrose synthase (SS) and ADP-glucose pyrophosphorylase (AGPase) in SPGs and WPGs of both cultivars. Application of either 6-benzyladenine or abscisic acid had no significant influences on both endosperm cell number and grain dry matter accumulation. These results suggest that grain sink strength, determined by both cell numbers, SS and AGPase activities in the endosperm control the dry matter accumulation of grains.     

Postanthesis allocation of photosynthates and grain growth in wheat cultivars as affected by source/sink change

Two wheat cultivars, Hesheng 2 with large grain yield potential, and Shannong 505 with small grain yield potential, were used for investigating the responses of postanthesis photosynthesis, dry matter accumulation and allocation, and grain growth to source/sink changes. At the initial grain filling stage, Hesheng 2 was sensitive to source reduction leading to an increase of net photosynthetic rate (P_N) by 10 %; however, little effect of sink reduction was observed. In Shannong 505, P_N was obviously decreased by sink reduction, and changed a little after source reduction. At the rapid grain filling stage, Hesheng 2 was sensitive to both source and sink reduction resulting in the increase or decrease of P_N, respectively. However, the response of P_N in Shannong 505 to source/sink changes was similar to that in previous stage. The dry matter (DM) accumulation after anthesis was affected by source/sink changes. In Hesheng 2, the decrease in DM was higher than that in Shannong 505 after the same source or sink reduction. Source reduction caused a decrease in the allocation of DM to the sheath and stem, and promoted the reserve photosynthates to be reallocated to grain. The effect of sink reduction was contrary. The grain mass of Hesheng 2 was more easily regulated by source/sink changes than that of Shannong 505. The effect source/sink changes on grain mass was in order upper > basal > middle spikelets on spike. As for a spikelet, the effect was found mainly in the grain mass at the positions 3 and 4 from base of the spikelet. This revised version was published online in July 2006 with corrections to the Cover Date.     

Endosperm enrichment and physicochemical properties of superior and inferior grain starch in super hybrid rice

• A significant asynchronous phenomenon exists in super hybrid rice because of the differences in spike and spikelet positions, which affect the accumulation and properties of starch. However, little is known about the endosperm enrichment and physicochemical properties of starch in superior and inferior grains in super hybrid rice.
• Rice YY2640 was selected as study material to investigate the enrichment and physicochemical properties of starch in superior and inferior grains in super rice using semi‐thin sections, X‐ray diffraction and related technologies.
• Superior grain filling was a continuous process, whereas inferior grain only started 8–10 days after anthesis. The order of starch accumulation starts in the central endosperm, then in the endosperm of the proximal vascular bundle and finally in the aleurone layer. Compared with the inferior grains, the superior grains have a higher 1000‐grain weight, apparent amylose content, total starch content, average starch granule size, relative crystallinity, solubility and a resonance peak ratio at 1022/995 cm^?1, whereas the swelling power and ratio of the resonance peak at 1045/1022 cm^?1 were lower. The final degree of hydrolysis of HCl, AAG and PPA of the superior grains were significantly lower than those of the inferior grains.
• The findings indicate that the different physicochemical properties of starch were mainly related to the development order of superior and inferior grains and the spatial enrichment of starch.

     

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.     

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.     

Properties of glutelin from mature and developing rice grain

Aminograms and SDS-polyacrylamide electrophoresis of milled rice glutelin of 12 Oryza sativa samples showed similar composition and ratio of 1 : 1 : 1 for subunits with MW 38 000:25 000: 16 000, indicating little possibility of finding variants of rice glutelins. Fractionation of S-cyanoethyl glutelin of 3 rices on polyacrylamide-agarose gels gave MW subunits differing in amino acid analysis of which the subunits with MW > 38 000 had the highest lysine content. Of the solubility fractions of endosperm glutelin, the fraction extracted by 0.5 M NaCl-0.6 % β-mercapto-ethanol-0.5% SDS was closest to glutelin in properties. In the developing grain of two varieties, appearance of protein bodies and rapid synthesis of glutelin from 7 days after flowering onward coincided with a drop in lysine content and appearance of MW 38 000 and 25 000 of crude glutelin. The MW 38 000 subunit is thus unique to endo-sperm glutelin.     

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.     

不同生态环境下水稻基因型产量形成与源库特性的比较研究

以日本和IRRI的9个水稻品种为材料,分别以武香粳9号和两优培九为对照,在江苏南京和云南丽江研究了不同水稻基因型干物质积累与源库形成特征及其在不同生态环境下的差异.结果表明,生态环境对水稻产量和干物质积累量影响显著.高产水稻品种积累了高额干物质量,且干物质生产优势在中后期.高产品种的总颖花量、LAI及群体生长速率(CGR)都较高.稻谷产量随干物质积累总量的增加而提高,与齐穗后干物质积累量、总颖花量和LAI呈极显著正相关,与粒叶比呈显著正相关.与云南丽江点相比,群体LAI、单位面积颖花量和抽穗后干物质积累量少及生长速率(CGR)低是南京点稻谷产量低的关键因素.     

籼稻桂华占、八桂香花后营养器官干物质流转与籽粒灌浆关系

以桂华占和八桂香2个籼稻品种为材料,研究籼稻花后不同部位器官物质积累、运转与籽粒生长的动态特征及相互关系。结果表明:(1)叶、叶鞘、节间干物质流转存在一定差异,倒2叶鞘对籽粒的贡献超过倒3叶鞘和倒1叶鞘,倒3节间对籽粒的贡献超过倒2节间和倒1节间;(2)不同部位籽粒的灌浆速率和拐点粒重呈现UPG(上部籽粒)MPG(中部籽粒)BPG(下部籽粒)变化趋势,拐点时间和活跃灌浆时间及持续灌浆时间均呈现BPGMPGUPG变化规律,UPG启动早,灌浆速率大,BPG的灌浆速率小,灌浆时间滞后,籽粒粒重呈现UPGMPGBPG;(3)叶片、叶鞘及节间干物质运转速度和运转率都与籽粒起始灌浆势呈正相关,其中器官间与起始灌浆势的相关系数大小表现为节间叶鞘叶片,不同叶位间与起始灌浆势的相关系数大小表现为倒2叶倒3叶倒1叶,其中节间干物质运转对籽粒生长的作用大于叶鞘,叶片干物质运转与籽粒生长的相关性最小。倒1节鞘物质输出与BPG生长时间上同步,倒2节鞘与MPG生长同步,倒3节鞘与UPG生长同步。     

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.     

分期施氮与CO2浓度升高对小麦光合及其物质积累和产量的互作效应

采用开顶式气室和盆栽方法,以冬小麦品种‘小偃22’为材料,探讨了分期施氮与CO2浓度升高对小麦抽穗期和灌浆中期旗叶光合、地上部物质积累和产量的互作效应.结果显示:(1)不施氮条件下CO2浓度升高对小麦旗叶叶绿素含量(SPAD)和可溶性蛋白含量、光合能力、地上部花后干物质和氮素累积量、籽粒产量的影响不明显(P>0.05)或产生显著负效应;在施氮(300mg/kg土)条件下各指标均不同程度增加,且大多数达到显著水平.(2)与氮肥全部基施相比,分期施氮时CO2浓度升高使灌浆期旗叶光合能力、地上部花后干物质和氮素累积以及产量增加的幅度较大,其中以播前、返青期和孕穗期施氮比例为5∶3∶2时最明显.研究表明,适当分期施氮可能更有利于发挥CO2浓度升高对冬小麦的增产作用.     

Activities of key enzymes for starch synthesis in relation to growth of superior and inferior grains on winter wheat (Triticum aestivum L.) spike

Weight of individual grains is a major yield component in wheat. The non-uniform distribution of single grain weight on a wheat spike is assumed to be closely associated with starch synthesis in grains. The present study was undertaken to determine if the enzymes involved in starch synthesis cause the differences in single grain weight between superior and inferior grains on a wheat spike. Using two high-yield winter wheat (Triticum aestivum L.) varieties differing in grain weight and three nitrogen rates for one variety, the contents of amylose and amylopectin, and activities of enzymes involved in starch synthesis in both superior and inferior grains were investigated during the entire period of grain filling. Superior grains showed generally higher starch accumulation rates and activities of enzymes including SS (sucrose synthase), UDPGPPase (UDP-glucose pyrophosphorylase), ADPGPPase (ADP-glucose pyrophosphorylase), SSS (soluble starch synthase) and GBSS (starch granule bound starch synthase) and subsequently produced much higher single grain weight than inferior grains. Nitrogen increased enzyme activities and starch accumulation rates, and thus improved individual grain weight, especially for inferior grains. The SS, ADPGPPase and SSS were significantly correlated to amylopectin accumulation, while SS, ADPGPPase, SSS and GBSS were significantly correlated to amylose accumulation. This infers that SS, ADPGPPase and starch synthase play key roles in regulating starch accumulation and grain weight in superior and inferior grains on a wheat spike.     

水稻谷蛋白GluA-2基因5′上游序列控制下的UidA基因在转基因水稻胚乳中的表达模式

为了研究谷蛋白胚乳特异性表达启动子在我国栽培稻品种中的表达模式,将UidA基因分别置于水稻谷蛋白GluA—2基因750bp和2．3kb上游序列下游,利用农杆菌转化法导人栽培稻品种中花8号并获得转基因植株。Southern blot检测表明,UidA基因已经整合到水稻基因组当中并以单拷贝存在。Northern blot检测表明,开花后13～15d和11～13d,UidA基因和水稻内源的GluA—2基因的表达量分别达到最高,随后逐渐降低。对转基因植株种子的GUS染色表明,UidA基因仅在胚乳中表达,在糊粉层中GUS表达量最高。测定了2．3kb和750bp转基因植株种子的GUS活性,结果表明前者的GUS活性是后者的2～3倍。序列分析表明,位于GluA—2基因转录启始位点上游2170bD的G-box可能是一个与表达量相关的顺式调控元件。     

Factors affecting dry weight accumulation in developing barley endosperm

Some factors that may be concerned in determining final grain weight in barley ( Hordeum vulgare L. var. distichum ) have been investigated. Variation in endosperm fresh and dry weight, volume and starch content have been recorded at different stages of grain development between anthesis and harvest-ripeness for two barleys, cvs Kym and Golden Promise, differing in final grain weight. Results were recorded under both field and glasshouse conditions. The results suggest that the higher final dry weight of Kym, in comparison with Golden Promise, is a function of both rate and duration of grain filling. Only at later stages of endosperm development did the differences in volume become significant and the Kym endosperms continued to increased rapidly in volume for two to three days after endosperm volume had reached a maximum in Golden Promise. The rates of starch accumulation in both cultivars were very similar but starch deposition continued in Kym endosperms for four to five days after deposition in Golden Promise endosperms had slowed down.     

IAA level and dry matter accumulation at different positions within a wheat ear

Dry matter accumulation and IAA level of grains in various positions within the wheat ( Triticum aestivum L. cvs Solo and Kolibri) ear were investigated during the grain filling period. Main shoot ears were partitioned into basal, central and apical regions and further into proximal and distal grains. Results from a two-year experiment with two wheat cultivars of different grain size are presented. The IAA level increased rapidly from about 8 until 20 to 30 days after anthesis, and then decreased dependent upon year and cultivar. The differences in IAA levels, both among spikelets in different regions of the ear (i.e. basal, central and apical) and also among florets within a spikelet (i.e. proximal and distal), were positively correlated with the differences in dry matter accumulation. Although the correlations held true within cultivars in a particular year, differences in dry matter between cvs Solo and Kolibri or differences from year to year were not well correlated with the corresponding changes in IAA levels. Possible interactions between dry matter accumulation and IAA levels are briefly discussed.