油菜素甾醇对杂交粳稻灌浆期光合物质、群体氮素积累和利用的影响

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田晓雅, 刘欣, 王强盛, 蒋琪, 冯金侠, 张慧, 丁艳锋. . 油菜素甾醇对杂交粳稻灌浆期光合物质、群体氮素积累和利用的影响. 作物学报, 2015, 41(12): 1844-1857
TIAN Xiao-Ya, LIU Xin, WANG Qiang-Sheng, JIANG Qi, FENG Jin-Xia, ZHANG Hui, DING Yan-Feng. . Effects of Brassinosteroids on Photosynthetic Matter, Nitrogen Accumulation and Use Efficiency during Grain Filling Stage of Hybrid Japonica . ACTA AGRONOMICA SINICA, 2015, 41(12): 1844-1857 复制到剪切板

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油菜素甾醇对杂交粳稻灌浆期光合物质、群体氮素积累和利用的影响

田晓雅, 刘欣, 王强盛^*, 蒋琪, 冯金侠, 张慧, 丁艳锋

南京农业大学农业部作物生理生态与生产管理重点实验室, 江苏南京210095

^*通讯作者(Corresponding author): 王强盛, E-mail: qswang@njau.edu.cn, Tel: 025-84395313

第一作者联系方式: E-mail: aefslab@163.com, Tel: 025-84395313

收稿日期:2015-04-13 接受日期:2015-09-06 网络出版日期:2015-09-14

基金:本研究由中央财政农业技术推广项目[TG(14)027, TG(14)013], 江苏省科技支撑重大项目(BE2013355), 江苏省农业科技自主创新资金项目[CX(14)4072]和江苏省农业三新工程项目(SXGC[2015]112, SXGC[2014]128, SXGC[2014]251, SXGC[2013]335)资助

摘要

以常规杂交粳稻常优3号和籼粳杂交超级稻甬优12为材料, 通过叶面施肥分析了油菜素甾醇(BR)对2个杂交粳稻灌浆期剑叶光合特性、群体干物质积累、氮素吸收利用、非结构性碳水化合物(NSC)积累和植株不同器官NSC/N的影响。结果表明, BR对2个水稻叶片光合生理指标的影响表现为 g_s> T_r> P_n> C_i。喷施2,4-表油菜素内酯(eBL)后能够增强灌浆期叶片净光合速率和蒸腾速率, 维持叶绿素含量和胞间CO₂浓度, 从而提升叶片光合能力; 以及增加群体干物质、氮素和NSC积累量以及干物质积累比例和积累速率、氮素吸收比例和吸收速率、NSC转运比例和转运速率, 但降低了氮素干物质生产效率; 喷施油菜素吡咯(Brz)后的上述生理效应相反。eBL提升了2个水稻品种花后干物质、NSC和氮素的转移量及氮素对籽粒贡献率, 降低了干物质和NSC对籽粒贡献率; Brz则降低了干物质、NSC及氮素的转移量和转移率, 提升了干物质和NSC对籽粒贡献率。2个杂交粳稻整个灌浆期地上部不同器官NSC、氮素转运量均表现为eBL>CK>Brz, 籽粒灌浆前期>籽粒灌浆后期。2个水稻品种地上部不同器官NSC/N随着籽粒的不断充实呈现上升趋势, 其中茎鞘、穗部NSC/N变化最为显著; eBL减小了叶片和茎鞘中的NSC/N, Brz则反之, 但喷施BRs后穗部变化规律不明显。籼粳杂交超级稻甬优12比常规杂交粳稻常优3号具有更强的剑叶光合速率, 更高的干物质和氮素及NSC积累量, 更大的灌浆前期NSC和氮素转运量, 最终籽粒实际产量较常优3号增加5.03%~ 9.32%; eBL对2个水稻品种干物质和氮素积累及NSC转运作用规律不一, Brz对甬优12干物质、氮素积累及NSC转运抑制效应大于常优3号。

关键词: 杂交粳稻; 油菜素甾醇; 光合特性; 干物质转运; 氮素利用; NSC积累

Effects of Brassinosteroids on Photosynthetic Matter, Nitrogen Accumulation and Use Efficiency during Grain Filling Stage of Hybrid Japonica

TIAN Xiao-Ya, LIU Xin, WANG Qiang-Sheng^*, JIANG Qi, FENG Jin-Xia, ZHANG Hui, DING Yan-Feng

Key Laboratory of Crop Physiology, Ecology and Production Management, Ministry of Agriculture / Nanjing Agricultural University, Nanjing 210095, China

Fund:This study was supported by the Central Finance for Agricultural Innovative Technology Extension [TG(14)027, TG(14)013], the Key Technology R&D Program of Jiangsu Province (BE2013355), the Independent Innovation Fund for Agriculture Science and Technology of Jiangsu Province [CX(14)4072] and Agrotechnical Innovation Projects of Jiangsu Province (SXGC[2015]112, SXGC[2014]128, SXGC[2014]251, SXGC[2013]335)

Abstract

Taking the conventional japonica hybrid rice Changyou 3 and indica-japonica super hybrid rice Yongyou 12 as materials, the experiment analyzed the effects of brassinosteroids (BRs) on photosynthetic characteristics, dry matter accumulation, nitrogen absorption and utilization, non-structural carbohydrates (NSC) accumulation and the NSC/N ratio of different plant organs during grain filling stage. Results showed that the effect of BRs on photosynthetic indexes presented as g_s> T_r> P_n> C_i. Spraying 2,4-epibrassinolide (eBL) could enhance net photosynthetic rate ( P_n) and transpiration rate ( T_r), maintain chlorophyll content and intercellular CO₂ concentration ( C_i), and eventually improve the leaves’ photosynthetic capacity during grain filling stage. It could also increase the amounts of dry matter, nitrogen and NSC accumulation, the proportions and rates of dry matter accumulation, nitrogen absorption and NSC transportation, but reduce the dry matter production efficiency of nitrogen nutrition. The effect of Brz was on the contrary. Spraying eBL improved amounts of the dry matter, NSC, nitrogen and the contribution rate to grains made by nitrogen of the two rice varieties, reduced the contribution rate to grains made by dry matters and NSC after flowering. However, brassinazole (Brz) decreased transfer amounts and rates of the dry matter, NSC and nitrogen, promoted the contribution rate to grains made by dry matters and NSC. The transfer amounts of NSC and nitrogen of different organs in shoot of the two hybrid rice varieties were shown as eBL>CK>Brz, and early stage of grain filling (EGF) > late stage of grain filling (LGF). The NSC/N of different organs in shoot of the two rice varieties showed an increasing tendency with filling, among which the changes in stem-sheath and panicle were the most significant. Spraying eBL decreased the NSC/N of stem-sheath and leaf, Spraying Brz promoted it, while the effects of BRs on panicle was not clear. Yongyou 12 had a stronger net photosynthetic rate in flag leaves, greater accumulations of dry matter, NSC and nitrogen, larger transfer amounts of NSC and nitrogen in early stage of grain filling, and finally its actual yield was 5.03%-9.32% higher than that of Changyou 3. Spraying eBL showed a diverse rule on the accumulations of dry matter, nitrogen and NSC transportation of the two rice varieties. The inhibiting effect of Brz on dry matter and nitrogen accumulations and NSC transferring was stronger in Yongyou 12 than that of Changyou 3.

Keyword: Hybrid rice; Brassinosteroids; Photosynthetic characteristics; Dry matter transportation; N use efficiency; NSC accumulation

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油菜素甾醇(BR)类物质作为一种新型的调控植物生长发育的重要激素^[1], 参与了植物的营养生长^[2]与生殖生长^[3], 能够增强水稻叶片的光能捕获能力^[4], 因而在植物生长调控中起重要作用。目前, 关于油菜素甾醇类激素对农作物生长发育的研究主要集中在提高产量^{[4, 5]}、促进植株伸长^[6]、BR合成途径^[7]及信号传导^[8]等方面。李勇等^{[9, 10]}研究指出, 油菜素内酯能够显著提升冬小麦的光合作用参数、总干物质积累量和植株氮积累量; 李晓玲等^[11]研究发现, 高油菜素内酯能够改善不同施肥水平下小麦幼苗的生长、生理生化指标及产量构成因子, 使低肥区增产5.8%; 刘大永等^[12]发现表油菜素内酯能提高水稻吸收肥料和土壤的P₂O₅量; Wu等^[4]也指出BRs能够改变叶片构型促进叶片及籽粒中的糖代谢, 从而促进粒重增加和增产。

提高单产是增加稻谷产量的永恒主题, 种植杂交粳稻是提高产量的一种有效途径, 大穗型杂交水稻品种叶片净光合速率高, 物质积累能力强^[13], 但由于源库矛盾问题^[14]难以充分发挥高产潜力, 因而目前杂交水稻品种普遍存在结实率低、产量不稳定等问题^[15], 严重阻碍了其增产潜力的发挥^[16]。纪洪亭等^[17]指出超级杂交水稻干物质和养分的优势在于快速增长期持续时间较长, 中后期干物质和养分积累速率较快, 而不同生育阶段干物质积累量的协调能够促进水稻增产^[18]。殷春渊等^[19]认为高产基因型水稻在各个生育阶段均具有较高氮素积累量, 茎叶部非结构性物质积累量和含量较高的品种具有稳产和高产潜力^[20]。

至今未见有关油菜素甾醇对杂交粳稻干物质积累、氮素营养吸收及非结构性碳水化合物转运的报道。本试验通过表油菜素内酯(促进型激素)和油菜素吡咯(抑制型激素)两种油菜素甾醇的喷施, 揭示油菜素甾醇对大穗型杂交粳稻后期光合特性、物质积累、氮素利用、非结构碳水化合物运转及植株器官碳氮分配的生理调控效应, 为现行大穗型杂交粳稻高产高效生产的激素调控提供依据。

1 材料与方法

1.1 试验设计

供试品种为2个大穗型杂交粳稻品种, 即三系杂交粳稻常优3号和三系籼粳杂交超级稻甬优12。

试验于2012— 2013年在南京农业大学试验农场(31° 54'N, 119° 28'E)进行, 供试土壤为黄壤土, pH 6.85, 0~20 cm耕层土壤含全氮1.26 g kg^-1、速效氮102.86 mg kg^-1、速效磷13.15 mg kg^-1、速效钾109.14 mg kg^-1、有机质20.76 g kg^-1, 前茬为小麦。5月28日播种育秧, 6月20日用插秧机移栽。采用完全随机区组设计, 株行距平均为13.3 cm× 30.0 cm, 每穴2苗, 小区面积66 m², 重复3次, 小区间筑埂隔离, 薄膜包埂, 单独灌排。基蘖肥为纯氮165 kg hm^-2, 穗肥为纯氮120 kg hm^-2, 分促花肥(倒四叶)和保花肥(倒二叶)等量施用, 磷肥P₂O₅ 135 kg hm^-2全部基施, 钾肥K₂O 150 kg hm^-2分基肥和促花肥等量施入。田间管理按当地高产栽培技术进行, 并于抽穗期选择同日抽穗、生长整齐单茎挂牌标记。本试验中, 抽穗期至开花期间隔2~3 d。

试验施用两种油菜素甾醇(brassinosteroid, BR), 促进型BR为2, 4-表油菜素内酯(2, 4-epibrassinolide, eBL), 抑制型BR为油菜素吡咯(brassinazole, Brz)。在挂牌当日分别叶面喷施预先试验明确的适宜浓度0.1 mg L^-1 eBL、2 mg L^-1 Brz, 溶液中加入0.1% (v/v)的Tween-20作为展布剂, 使溶液能够均匀附着, 并以喷施0.1% (v/v) Tween-20的蒸馏水溶液为对照, 每平米喷施30 mL, 喷施时间为16:00— 18:00, 连续喷施3日。

1.2 分析方法

1.2.1 标记方法

依据常优3号和甬优12籽粒灌浆特性, 以单穗籽粒灌浆最早达到最大平均粒重(earliest reach the high weight of grain, EHW)为时间节点, 常优3号该节点为花后35 d, 甬优12为花后30 d。依据该时间点将灌浆期分为籽粒灌浆前期(early stage of grain filling, EGF)和灌浆后期(late stage of grain filling, LGF); EGF为花后0 d~EHW, LGF为EHW~成熟期, 灌浆期为花后0 d~成熟期, 成熟期表示籽粒成熟时间点。2个品种各处理灌浆期历时天数如表1所示。

表1 油菜素甾醇各处理灌浆期历时天数 Table 1 Duration of grain filling stage in different BRs treatments (d)

1.2.2 SPAD测定

花后每隔5 d用日产SPAD- 502型叶绿素计测定剑叶SPAD值, 从各处理选20片叶, 测定每片叶上、中、下3点, 取平均值。

1.2.3 产量测定

成熟期从各处理连续取150样株考种, 测定穗数、每穗粒数、结实率和千粒重, 各小区的1/3用于测定实产。

1.2.4 氮素、NSC测定

于花后0 d、常优3号花后35 d、甬优12花后30 d和成熟期, 取各处理具有代表性植株3穴, 分为茎、叶、穗后105℃杀青30 min, 80℃烘至恒重, 磨粉过筛, 待测干物质重、全氮及NSC (non-structural carbohydrates, 非结构性碳水化合物, 为可溶性糖和可溶性淀粉之和)含量。采用硫酸消煮, 半微量凯氏定氮法^[21]测定全氮含量。

称取0.100 g样品, 置10 mL离心管, 在冰水浴中加入2 mL 9.2 mol L^-1高氯酸, 用涡旋仪振荡提取15 min再加6 mL蒸馏水, 混匀后离心15 min, 转移上清液至50 mL容量瓶。重复提取2次, 离心2次。合并3次离心所得上清液, 用水洗沉淀并离心2次, 合并所有上清液并用蒸馏水定容至50 mL, 采用蒽酮法^[22]测定NSC含量。

1.2.5 数据分析

2年试验重复性较好, 处理间各指标值变化趋势一致, 本研究主要采用2013年的数据。使用SPSS 19.0统计分析, Microsoft Excel 2010、Origin 8.0软件计算数据、绘制图形。

1.3 计算方法

SPAD衰减率 = (花后0 d SPAD值-花后35 d SPAD值)× 100%/花后35 d SPAD值

干物质积累速率=2次采样单位面积地上部干物质积累量差值/2次采样间隔时间

营养器官干物质(氮素或NSC)表观转移量=花后0 d地上部营养器官干物质(氮素或NSC)积累量-成熟期地上部营养器官干物质(氮素或NSC)积累量

营养器官干物质(氮素或NSC)表观转移率=花后0 d营养器官干物质(氮素或NSC)表观转移量× 100%/花后0 d地上部营养器官干物质(氮素或NSC)积累量

营养器官转移的干物质(氮素或NSC)对籽粒的贡献率=花后0 d营养器官干物质(氮素或NSC)表观转移量× 100%/成熟期籽粒干物质(氮素或NSC)积累量

氮素干物质生产效率=成熟期单位面积全株地上部干物重/氮素积累总量

单位产量吸氮量=群体氮素积累总量/稻谷实际产量

2 结果与分析

2.1 花后叶片光合特性

花后0~35 d植株剑叶SPAD值(叶绿素)呈现下降趋势, 常优3号SPAD值在花后0~20 d、甬优12在花后0~25 d下降较为缓慢, 其后2个品种均出现快速下降趋势(图1)。

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	图1 油菜素甾醇对杂交粳稻叶片SPAD值的影响 CK: 对照; eBL: 2, 4-表油菜素内酯; Brz: 油菜素吡咯。Fig. 1 Effects of BRs on leaf SPAD value of hybrid japonica rice CK: control; eBL: 2, 4-epibrassinolide; Brz: brassinazole.

花后0 d, 2个品种各处理间SPAD值差异不明显, 但常优3号各处理在花后10 d出现明显差异, 甬优12则在花后5 d各处理出现明显差异。相较CK, eBL能减缓叶片SPAD值的下降趋势, 而Brz作用相反。花后0~35 d, 常优3号CK、eBL、Brz处理的叶片SPAD值衰减率分别为23.45%、20.32%、25.38%, 甬优12分别为19.94%、18.76%、21.67%, 表明eBL减小了叶片SPAD值衰减率, 而Brz增大了叶片SPAD值衰减率。

整个灌浆期, 甬优12的叶片SPAD值均高于常优3号, 且下降幅度较常优3号更小(图1)。说明超级杂交水稻较常规杂交水稻具有更高的叶绿素浓度, 且能够在灌浆前期将叶绿素浓度维持在较高水平。以BRs处理后2个品种变化幅度最大时间点为例, 常优3号eBL处理在花后35 d叶片SPAD值上升5.27%, Brz处理在花后30 d下降8.45%; 而甬优12 eBL处理在花后15 d叶片SPAD值升高2.19%, Brz处理在花后30 d下降2.39%。说明超级杂交水稻在灌浆前期具有更高的光合能力, 而eBL能够维持叶片叶绿素含量, 进而促进光合作用。

BR对杂交粳稻花后叶片光合速率(P_n)、蒸腾速率(T_r)、气孔导度(g_s)和胞间CO₂浓度(C_i)有明显影响(表2和表3)。2个水稻品种叶片P_n 表现为eBL> CK> Brz, 随着灌浆进程呈下降趋势。花后10 d处理间P_n出现显著差异, 但花后40 d显著差异消失, 说明BR对杂交水稻叶片光合生理作用有一定的持续时间, 其中eBL处理减缓了籽粒灌浆前期P_n的下降趋势, Brz的作用则相反。T_r和g_s变化趋势与P_n相同, 花后10 d处理间T_r出现显著差异, 而花后0 d g_s即出现显著差异, 因为BRs能够直接调控植株气孔形成和分布^[23], 所以相较于T_r和P_n, BR对G_s的作用时间最快, 喷施后1~2 d即产生生理效应且变化幅度较大。C_i的变化趋势则与P_n、T_r和g_s相反, 且处理间差异并不显著。

表2 油菜素甾醇对杂交粳稻剑叶净光合速率和蒸腾速率的影响(2013) Table 2 Net photosynthetic rate (P_n) and transpiration rate (T_r) in hybrid japonica rice under different BRs treatments in 2013

相同处理条件下, 甬优12的P_n和T_r均高于常优3号, 而C_i相反, 但g_s数值相似。因此, 甬优12在灌浆期的光合能力优于常优3号。BR处理后, 相同处理2个品种间P_n、T_r、g_s和C_i变化程度不同, 可见BRs对于叶片光合作用的各项指标影响程度不同, eBL总体表现为增强叶片净光合速率和蒸腾速率, 维持胞间CO₂浓度和气孔导度, 最终提高叶片光合能力, 而Brz作用相反。

表3 油菜素甾醇对杂交粳稻剑叶气孔导度和胞间CO₂浓度的影响(2013) Table 3 Stomatal conductance (g_s) and intercellular CO₂ concentration (C_i) in hybrid japonica rice at different BRs treatments in 2013

2.2 成熟期产量及其构成

由表4可知, 2年间杂交粳稻3个处理实际产量均表现为eBL> CK> Brz, 其中eBL处理后产量较对照提高3.41%~5.69%, Brz处理水稻产量较对照降低2.34%~6.19%。甬优12产量高于常优3号, eBL对常优3号增产效应为4.69%~5.69%, 高于甬优12为 3.41%~4.33%; 而Brz对常优3号的减产效应为2.34%~2.36%, 低于甬优12的5.45%~6.19%。因而不同杂交粳稻产量对eBL和Brz的生理响应有差别。BRs处理后穗数和穗粒数没有显著差异, 而结实率和千粒重出现明显差异。eBL使结实率较CK提高0.69~3.59个百分点, 常优3号提高幅度大于甬优12; Brz使结实率降低0.99~2.46个百分点, 常优3号下降幅度小于甬优12。eBL处理后千粒重增大0.56~1.64 g, Brz令其减少0.33~0.65 g, 2012年常优3号千粒重增减数量大于甬优12, 2013年反之。

表4 油菜素甾醇对杂交粳稻成熟期产量构成因素的影响 Table 4 Effects of BRs on rice yield components of hybrid rice at maturity stage

2.3 植株干物质积累转运

2个品种籽粒灌浆前期干物质积累量、积累比率和积累速率均高于灌浆后期, BR处理后各指标均出现明显变化(表5)。相较于CK, eBL显著提升了籽粒灌浆前期、后期和成熟期植株干物质积累量、积累比率和积累速率, 使表观转移量和转移率分别提高了1.21%~1.39%和0.09~0.11个百分点, 但对籽粒贡献率下降了0.22~0.47个百分点, 这主要是eBL的显著增产效应所致。Brz显著降低了植株干物质积累量、积累比率和积累速率, 使表观转移量和转移率分别下降1.42%~1.50%和0.11~0.12个百分点, 但对籽粒贡献率升高0.10~0.54个百分点, 这主要是因为Brz降低了杂交水稻的实际产量。BR处理后, 籽粒灌浆后期的干物质积累量、积累比率和积累速率明显高于前期, 且甬优12的Brz处理高于常优3号而eBL处理低于常优3号, 因此籼粳杂交超级稻干物质变化对Brz更敏感, 常规杂交粳稻则对eBL更敏感。

甬优12的干物质积累量、积累比例、积累速率明显高于常优3号, 因此籼粳杂交超级稻在整个灌浆期较常规杂交粳稻具有更强的物质积累能力, 这为其成熟期高产形成提供了保证。甬优12积累速率在籽粒灌浆前期、后期和灌浆期比常优3号分别高24.93%~29.56%、1.43%~5.82%和13.62%~17.12%, 可见籽粒灌浆前期2个品种干物质积累速率差异较大, 因而在籽粒灌浆前期干物质的快速积累能够保证最终高产。虽然甬优12 表观转移量比常优3号高1.82%~2.09%, 表观转移率高0.05~0.08个百分点, 但其对籽粒贡献率比常优3号低0.44~0.88个百分点, 这主要是由于籼粳杂交超级稻在花后0 d干物质积累量大且明显高于常规杂交粳稻。

表 5 Table 5

2.4 植株NSC积累转运

eBL显著增大了植株NSC积累量、转运比率和转运速率, Brz则显著降低这些生理指标(表6)。eBL处理后2个品种花后营养器官NSC 表观转移量较CK增加0.39%~2.07%, 而对籽粒贡献率下降0.95~ 0.96个百分点, 但常优3号表观转移率上升0.19个百分点, 甬优12则降低0.01个百分点, 说明eBL有利于杂交水稻灌浆期的NSC转运积累, 但由于eBL处理一方面增大了杂交粳稻NSC含量, 另一方面提高了水稻实际产量, 最终导致NSC对籽粒贡献率下降。而Brz处理后上述指标变化规律相反, 说明Brz削弱了杂交粳稻NSC积累和转运量, 但对常规杂交粳稻和籼粳杂交超级稻作用规律不完全一致。

籽粒灌浆前期、后期和成熟期, 甬优12 NSC转运量、转运比率和转运速率均比常优3号高。甬优12花后营养器官NSC 表观转移量比常优3号高2.96%~ 6.34%, 因而籼粳杂交超级稻比常规杂交粳稻具有更强的NSC积累能力, 这可能是甬优12高产的关键因素。另外, 甬优12 NSC对籽粒的贡献率比常优3号低0.59~1.72个百分点, 而表观转移率除Brz处理甬优12比常优3号高0.29个百分点外, CK和eBL处理均比常优3号低0.05~0.25个百分点。以上指标中, 甬优12的Brz处理除花后营养器官NSC转运量和转运速率比常优3号低外, 其余指标均高于常优3号, 但eBL对2个品种的影响程度规律并不明显。可能是因为籼粳杂交超级稻库容较大造成其贡献率比常规杂交粳稻低, 造成籽粒充实度低; 且甬优12 NSC积累转运比常优3号对Brz处理更敏感。

表 6 Table 6

2.5 植株氮素吸收利用

BR对杂交粳稻灌浆期氮素营养积累转运的影响见表7, BR处理后2个品种籽粒灌浆前期、后期和成熟期(或灌浆期)氮素积累量、积累比例、积累速率及花后营养器官氮素表观转移量变化与植株干物质和NSC积累转运相似。

eBL增加了植株氮素积累量、氮素积累比率、积累速率、营养器官氮素表观转移量和对籽粒贡献率, 而Brz作用反之。但2个品种氮素表观转移率对eBL响应不一致, 甬优12下降了0.39个百分点, 但常优3号上升了0.86个百分点, 因此eBL有利于杂交粳稻灌浆期的氮素积累转运, 而甬优12氮素转移率下降是由于群体在花后0 d具有较高的氮素积累量。另外, eBL使甬优12单位产量吸氮量增加0.41%, 而常优3号下降0.42%, 因此eBL能够促进籼粳杂交超级稻籽粒对氮素的吸收并最终增产。Brz处理相较于CK, 常优3号吸氮量降低了2.92%, 而甬优12提高0.41%, 但差异不显著, 可见Brz对常规杂交粳稻吸氮量作用更显著。eBL使氮素干物质生产效率减少2.21%~2.54%, 可能是由于eBL增加了杂交粳稻产量形成过程中所需要的氮素消耗量, 使氮素积累量增大且升高幅度大于干物质积累增加幅度。Brz增加了氮素干物质生产效率, 因此Brz能够减少杂交粳稻的氮素消耗量, 提升地上部氮素干物质生产效率。

表 7 Table 7

甬优12氮素积累量、积累比例和积累速率均大于常优3号。除充实后期氮素吸收速率外, 常优3号eBL处理的各项指标较甬优12上升幅度更大, 而甬优12 Brz处理降低幅度均大于常优3号。由此可知常规杂交粳稻的氮素吸收对eBL响应更明显, 而籼粳杂交超级稻对Brz响应更明显。甬优12氮素表观转移量比常优3号高27.26%~33.81%, 因而籼粳杂交超级稻比常规杂交水稻具有更强的氮素营养吸收转运能力。甬优12的表观转移率和对籽粒贡献率比常优3号分别高4.14~5.59和2.17~3.75个百分点, 表明籼粳杂交超级稻植株氮素营养积累量和氮素表观转移能力是其高产的主要原因。成熟期2个水稻品种单位产量吸氮量为0.0233~0.0244 kg, 甬优12比常优3号高1.25%~4.72%, 因此籼粳杂交超级稻较常规杂交水稻单位产量需求更多氮素。eBL增强了籼粳杂交超级稻氮素吸收, 因而高产需要结合BR喷施并注重氮素营养的合理施用。甬优12氮素干物质生产效率较常优3号降低了4.30%~5.78%, 可见籼粳杂交超级稻单位干物质形成过程中籼粳杂交超级稻要比常规超级稻消耗更多氮素, 可以配合不同种类BR的施用增强氮素干物质生产效率, 使杂交粳稻生产获得较适宜的氮素, 发挥其增产潜力。

2.6 不同器官NSC、氮素转运量及NSC/N比例

2个水稻品种灌浆期地上部NSC转运量和氮素吸收量均表现为籽粒灌浆前期> 籽粒灌浆后期, eBL> CK> Brz (图2)。籽粒中的氮素来自于叶片输出多于茎鞘, 2个营养器官中NSC输出则相当。

籽粒灌浆前期, 叶片、茎鞘中NSC和氮素转运量均表现为eBL> CK> Brz, 籽粒中也呈现相同趋势, 甬优12 NSC和氮素转运量> 常优3号NSC和氮素转运量。籽粒灌浆后期, 2个水稻品种各处理叶片和茎鞘均表现为NSC集聚, eBL< CK, 但甬优12 Brz处理积累量最小, 常优3号则相反。因此, 甬优12氮素和NSC对籽粒的输出主要发生在前期, 而后期营养器官NSC聚集导致供应量不足可能是造成其结实率低的主要原因。而Brz能够削弱超级杂交粳稻籽粒灌浆后期NSC滞留于茎叶中, eBL可以减少杂交粳稻茎叶中NSC残留。

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	图2 油菜素甾醇对杂交粳稻灌浆期不同部位NSC和氮素含量的影响缩写同表2和表5。Abbreviations are the same as those given in Tables 2 and 5.Fig. 2 Effects of BRs on NSC and N accumulation in different organs of hybrid japonica rice during grain filling stage

2个品种叶片在籽粒灌浆后期均向外输出氮素。常优3号3个处理茎鞘均表现氮素滞留, 其中茎鞘eBL处理氮素积累量最小, Brz最大。因此BR将导致常规杂交粳稻灌浆后期氮素在茎鞘中集聚。甬优12营养器官则均表现向外输出氮素, 叶片中氮素输出量> 茎鞘中氮素输出量, 可见叶片是籼粳杂交超级稻氮素输出主要器官。因此, 利用eBL提升籽粒灌浆前期穗部的氮素和NSC积累, Brz减弱超级杂交粳稻籽粒灌浆后期营养器官的NSC残留, 在实际生产中可以适当提高籼粳杂交超级稻的产量。

杂交粳稻地上部不同器官NSC/N随籽粒的灌浆充实呈现上升趋势(表8)。eBL处理后, 叶片NSC/N与CK相比变化不显著, 只有甬优12成熟期eBL处理显著减小。整个灌浆期叶片NSC/N均表现为Brz> CK> eBL, 这可能是因为eBL促进叶片氮素积累的程度高于NSC积累。穗部和茎鞘NSC/N均出现较为显著的变化, 茎鞘eBL处理NSC/N较CK显著降低, 而Brz则显著升高; 穗部对BR的响应并不完全一致且规律不明显。

表8 油菜素甾醇对杂交粳稻不同器官NSC/N的影响 Table 8 Effects of BRs on NSC/N of different organs of hybrid japonica rice (kg kg^-1)

3 讨论

3.1 BRs对杂交粳稻灌浆期植株光合效应的影响

有研究指出保证超级稻高产超高产需要提升后期叶片光合效率^[24]并保持叶片光合功能期^[25], 本试验结果表明, eBL能够增强叶片净光合速率和蒸腾速率, 维持气孔导度和胞间CO₂浓度, 促进杂交粳稻叶片净光合速率, 这与李勇等在冬小麦上研究结果相一致^{[9, 10]}。相较于CK, 在花后0~35 d, eBL处理2个水稻叶片SPAD值变化了-0.46%~5.27%, Brz处理变化了-8.45%~0.62%, 并且常优3号变化幅度大于甬优12 (图1), 其叶片SPAD值对BR敏感性高于常优3号, 这与曹云英等^[25]关于eBL喷施增加水稻叶片叶绿素含量的研究结果基本相似。但本试验还表明, 常优3号叶片净光合速率受BR影响小于甬优12, 叶片净光合速率低于甬优12, 常优3号叶片SPAD值变大意味着叶绿素含量增加, 但常规杂交粳稻常优3号净光合速率增加幅度并没有超过籼粳杂交超级稻甬优12, 正如孙旭生等^[26]提出, 叶绿素含量仅是光合速率的一个“ 必要条件” 。也就是eBL 能够通过提升一定叶绿素含量来促进光合效率, 但这种促进作用并不能超越品种间差异。而Brz喷施后的光合参数变化相反。

水稻群体干物质积累对产量形成有着重要的促进作用^{[17, 18]}, 进一步提高产量主要依靠生物学产量的提高, 特别是籽粒充实期的物质生产是高产水稻关键所在^[28], 本试验结果表明, CK、eBL和Brz处理后灌浆期常规杂交粳稻常优3号积累的干物质分别为10.19、10.77和9.95 t hm^-2, 分别占最终产量的76.84%、77.16%和76.68%, 籼粳杂交超级稻甬优12积累干物质分别为11.14、11.52和10.45 t hm^-2, 分别占实际产量的78.37%、78.56%和77.89%, 再次揭示提高灌浆期干物质积累量是水稻实现高产超高产的重要途径, 并且eBL能够使籽粒灌浆前期和后期的干物质积累量升高, 为高产水稻奠定了生物学基础, 但本试验结果显示eBL对甬优12干物质积累的促进作用比常优3号小, 这主要是因为籼粳杂交超级稻甬优12比常规杂交粳稻常优3号具有较高的群体干物质积累量, 姜元华等^[29]也有相同试验结果; 而Brz喷施后对干物质积累效应相反, 并且籼粳杂交超级稻甬优12抑制效果更为明显; 但常规杂交粳稻干物质积累则比籼粳杂交超级稻对eBL响应更大。

3.2 BR对杂交粳稻灌浆期植株碳氮代谢的影响

水稻产量的高低与氮素吸收利用关系密切^{[30, 31]}, Norman等^[32]指出开花后水稻植株营养器官转运的氮素是籽粒氮素积累量的主要组成部分, 因此适当提高抽穗后氮素积累量和吸收比例有利于水稻的产量提高^[33]。本试验结果表明, eBL处理后2个杂交粳稻灌浆期氮素积累量、吸收比例和吸收速率均得到显著增加(表7), 产量也获得显著提升; Brz处理从反面也进一步说明了这一结果。

Cock等^[34]认为每个颖花分配的贮存碳水化合物多, 能够提高结实率和千粒重, 因此提高千粒重的关键是茎鞘转移到每个颖花中的非结构性碳水化合物总量高^[35]。本试验表明甬优12 NSC转运量明显高于常优3号, 但甬优12结实率和千粒重均低于常优3号, 其原因是籼粳杂交超级稻具有更多的颖花数, 导致其NSC平均分配量小, 而eBL处理能够增加杂交粳稻NSC转运量, 因此能够提高千粒重和结实率, 形成增产效应; Brz处理具有相反效应。

本试验结果还表明, 杂交粳稻叶片为主要氮素输出器官, 而叶片和茎鞘是NSC的主要输出器官(图2), 2个水稻品种叶片和茎鞘在充实前期均表现为氮素、NSC输出; 充实后期叶片、茎鞘积累NSC, 而茎鞘氮素为甬优12转运输出、常优3号吸收累积, 赵全志等^[36]研究认为, 高产水稻在灌浆前中期茎鞘物质输出多容易导致灌浆后期回升多, 而后期茎鞘物质一直向外输出是低产的表现。因此, 籼粳杂交超级稻充实后期茎鞘氮素向外输出可能是造成其充实度低的主要原因, 而eBL降低了籼粳杂交超级稻茎鞘氮素输出量, 是其增产效应的一个原因, Brz则反之。Yang等^[37]发现水稻灌浆期叶片茎鞘积累的非结构性物质通过酶分解为蔗糖等可溶性糖后向穗部转移, Hirose等^[38]研究认为, 叶片茎鞘可溶性糖的含量升高可以增加库源之间可溶性糖的浓度差, 促进非结构性物质向穗部转移。本试验结果显示, 甬优12籽粒充实后期虽然茎叶非结构性碳水化合物转运量高于常优3号, 但籽粒的NSC积累量小于常优3号, 这可能是甬优12后期茎叶中氮素的不断转运, 籽粒积累氮并消耗碳的影响使籽粒NSC积累量减少, 而eBL喷施后能够提升籽粒充实后期获得的NSC积累量, 因此eBL能够调节杂交粳稻植株碳氮代谢的平衡, 改善籽粒灌浆。Brz喷施后甬优12籽粒灌浆后期NSC转运量显著降低, 导致籽粒充实度差和产量降低。

植株C/N改变能够影响植株的生理代谢^[39], 本研究显示植株不同器官的NSC/N随着籽粒充实进程升高, BR喷施后显著改变了茎鞘和穗部的NSC/N (表8), 因而BR能够调控水稻灌浆期植株氮素吸收量和光合物质转运量, 增强茎鞘物质输送强度和穗部灌浆物质积累能力。另外, 本研究表明BRs喷施后对叶片NSC/N没有显著差异, 这主要是叶片存在NSC和氮素的协同转运变化, 这与阮新民等^[40]研究认为水稻叶片碳氮比差异主要在于叶片氮含量差异的结果略有不同。因而利用BRs合理调节水稻植株C/N, 能够发挥激素对杂交粳稻产量形成的调控效应。

4 结论

eBL处理提高了杂交粳稻叶片净光合速率和蒸腾速率, 促进了籽粒灌浆期植株干物质、氮素和NSC的积累量、积累比例和积累速率, 增强了干物质、NSC和氮素转移量, 但降低了干物质和NSC对籽粒贡献率以及叶片、茎鞘的NSC/N; Brz处理效应相反。植株地上部不同器官NSC、氮素转运量为eBL> CK> Brz, 籽粒灌浆前期> 籽粒灌浆后期。籼粳杂交超级稻甬优12比常规杂交粳稻常优3号具有更大的叶片光合速率以及干物质、氮素和NSC积累量, 但甬优12较常优3号对Brz更敏感。喷施eBL能够调节杂交粳稻植株碳氮代谢, 促进杂交粳稻灌浆过程籽粒充实, 具增产效应。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。The authors have declared that no competing interests exist.

参考文献

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[1]	Nakamura A, Fujioka S, Sunohara H, Kamiya N, Hong Z, Inukai Y, Miura K, Takatsuto S, Yoshida S, Ueguchi T M, Hasegawa Y, Kitano H, Matsuoka M. The role of OsBRI1 and its homologous genes, OsBRL1 and OsBRL3, in rice. Plant Physiol, 2006, 140: 580-590 [本文引用:1]
[2]	Zurek D M, Clouse S D. Molecular cloning and characterization of a brassinosteroid-regulated gene from elongating soybean (Glycine max L. ) epicotyls. Plant Physiol, 1994, 104: 161-170 [本文引用:1]
[3]	Khripach V A, Zhabinskii V N, Groot A D. Twenty years of brassinosteroids: steroidal plant hormones warrant better crops for the XXI century. Ann Bot, 2000, 86: 441-447 [本文引用:1]
[4]	Chuanyin W, Anthony T, Parthiban R, Shing F K, SamHarris, Ke Z, Jiulin W, Jianmin W, Huqu Z, Suguru T, Shogo M, Shozo F, Kenneth A F, Roger I P. Brassinosteroids regulate grain filling in rice. Plant Cell, 2008, 20: 2130-2145 [本文引用:3]
[5]	肖琳, 庞瑞华, 蔡荣先, 于萍, 黄新华, 王磊. 水稻初花期喷施油菜素内酯的生理效应及增产作用. 安徽农业科学, 2007, 35: 3317-3330 Xiao L, Pang R H, Cai R X, Yu P, Huang X H, Wang L. Physiological effect and yield increase action after spraying BR in rice early blooming stage. J Anhui Agric Sci, 2007, 35: 3317-3330 (in Chinese with English abstract) [本文引用:1]
[6]	张海丽, 高静, 张昊, 李生辉, 邢继红, 王凤茹, 董金皋. 油菜素内酯对水稻细胞伸长和分裂的调控. 农业生物技术学报, 2015, 23: 71-79 Zhang H L, Gao J, Zhang H, Li S H, Xing J H, Wang F R, Dong J G. The regulation of brassinosteroid (BR) on elongation and division of rice (Oryza sativa L. ) cells. J Agric Biotechnol, 2015, 23: 71-79 (in Chinese with English abstract) [本文引用:1]
[7]	Fujioka S, Yokota T. Biosynthesis and metabolism of brassinosteroids. Annu Rev Plant Biol, 2003, 54: 137-164 [本文引用:1]
[8]	王梦姣, 邓百万, 杨国鹏. 拟南芥油菜素内酯的合成、修饰、信号转导及其在农作物育种中的应用研究进展. 河南农业科学, 2015, 44(2): 1-6 Wang M J, Deng B W, Yang G P. Research progress of brassinosteroid in Arabidopsis thaliana and its application in crop breeding. J Henan Agric Sci, 2015, 44(2): 1-6 (in Chinese with English abstract) [本文引用:1]
[9]	李勇, 王红光, 李瑞奇, 李雁鸣. 表油菜素内酯对冬小麦产量及氮素吸收、积累和分配的影响. 麦类作物学报, 2015, 35: 239-250 Li Y, Wang H G, Li R Q, Li Y M. Effect of epibrassinoilide on grain yield, and absorption, accumulation and distribution of nitrogen in winter wheat. J Triticeae Crops, 2015, 35: 239-250 (in Chinese with English abstract) [本文引用:2]
[10]	李勇, 王红光, 李瑞奇, 李雁鸣. 表油菜素内酯对冬小麦开花后光合特性和物质生产的影响. 河北农业大学学报, 2015, 38(1): 1-7 Li Y, Wang H G, Li R Q, Li Y M. Effect of epibrassinolide on photosynthetic performance and dry matter production after anthesis of winter wheat. J Agric Univ Hebei, 2015, 38(1): 1-7 (in Chinese with English abstract) [本文引用:2]
[11]	李晓玲, 骆炳山, 屈映兰. 不同施肥水平下高油菜素内酯对小麦生长发育的影响. 华中农业大学学报, 2000, 19: 417-420 Li X L, Luo B S, Qu Y L. Effects of homobrassinolide on the growth of wheat on assorted fertility levels. J Huazhong Agric Univ, 2000, 19: 417-420 (in Chinese with English abstract) [本文引用:1]
[12]	刘大永, 万兆良. 油菜素内酯对水稻营养效应的研究. 核农学报, 1992, 13: 132-134 Liu D Y, Wan Z L. Research to the effect of brassinolide in rice nutritional absorption. J Nucl Agric Sci, 1992, 13: 132-134 (in Chinese) [本文引用:1]
[13]	严建明, 翟虎渠, 万建民, 焦德茂, 张荣铣. 亚种间重穗型杂交稻光合产物的运转特性及其生理机制. 中国农业科学, 2003, 36: 502-507 Yan J M, Zhai H Q, Wan J M, Jiao D M, Zhang R X. Transportation characteristics of assimilate and physiologic mechanisms in subspecific heavy ear hybrid rice (Oryza sativa L. ). Sci Agric Sin, 2003, 36: 502-507 (in Chinese with English abstract) [本文引用:1]
[14]	王元辉, 白朴, 谢长晓, 金瑜雪, 赖士膺. 重穗型杂交稻的籽粒灌浆特点及库源关系的协调. 浙江农业学报, 2004, 16: 400-404 Wang Y H, Bai P, Xie C X, Jin Y X, Lai S Y. Characteristics of grain filling and coordination of source-sink for heavy panicle hybrid rice. Acta Agric Zhejiangensis, 2004, 16: 400-404 (in Chinese with English abstract) [本文引用:1]
[15]	敖和军, 王淑红, 邹应斌, 彭少兵, 唐启源, 方远祥, 肖安民, 陈玉梅, 熊昌明. 超级杂交稻干物质生产特点与产量稳定性研究. 中国农业科学, 2008, 41: 1927-1936 Ao H J, Wang S H, Zou Y B, Peng S B, Tang Q Y, Fang Y X, Xiao A M, Chen Y M, Xiong C M. Study on yield stability and dry matter characteristics of super hybrid rice. Sci Agric Sin, 2008, 41: 1927-1936 (in Chinese with English abstract). [本文引用:1]
[16]	Yang J C, Zhang J H. Grain-filling problem in ‘super’ rice. J Exp Bot, 2010, 61: 1-5 [本文引用:1]
[17]	纪洪亭, 冯跃华, 何腾兵, 潘剑, 范乐乐, 李云, 武彪, 肖铭, 梁显林. 超级杂交稻群体干物质和养分积累动态模型与特征分析. 中国农业科学, 2012, 45: 3709-3720 Ji H T, Feng Y H, He T B, Pan J, Fan L L, Li Y, Wu B, Xiao M, Liang X L. A dynamic model of dry matter and nutrient accumulation in super hybrid rice and analysis of its characteristics. Sci Agric Sin, 2012, 45: 3709-3720 (in Chinese with English abstract) [本文引用:2]
[18]	邓飞, 王丽, 刘利, 刘代银, 任万军, 杨文钰. 不同生态条件下栽培方式对水稻干物质生产和产量的影响. 作物学报, 2012, 38: 1930-1942 Deng F, Wang L, Liu L, Liu D Y, Ren W J, Yang W Y. Effects of cultivation methods on dry matter production and yield of rice under different ecological conditions. Acta Agron Sin, 2012, 38: 1930-1942 (in Chinese with English abstract) [本文引用:2]
[19]	殷春渊, 张庆, 魏海燕, 张洪程, 戴其根, 霍中洋, 许轲, 马群, 杭杰, 张胜飞. 不同产量类型水稻基因型氮素吸收、利用效率的差异. 中国农业科学, 2010, 43: 39-50 Yin C Y, Zhang Q, Wei H Y, Zhang H C, Dai Q G, Huo Z Y, Xu K, Ma Q, Hang J, Zhang S F. Differences in nitrogen absorption and use efficiency in rice genotypes with different yield performance. Sci Agric Sin, 2010, 43: 39-50 (in Chinese with English abstract) [本文引用:1]
[20]	辛阳, 杨重法, 陈慧娟, 王清峰. 水稻结实期茎叶部贮藏物质再转移能力的简易评价方法. 热带作物学报, 2013, 34: 1128-1132 Xin Y, Yang Z F, Chen H J, Wang Q F. Studies on accumulation and translocation rate of the non-structural matter in seven different rice varieties. Chin J Tropical Crops, 2013, 34: 1128-1132 (in Chinese with English abstract) [本文引用:1]
[21]	中国科学院上海植物生理研究所, 上海市植物生理学会. 现代植物生理学实验指南. 北京: 科学出版社, 1999. pp 133-134 Shanghai Institute of Plant Physiology of Chinese Academy of Sciences, Shanghai Association of Plant Physiology ed. Modern Laboratory Manual of Plant Physiology. Beijing: Sciences Press, 1999. pp 133-134(in Chinese) [本文引用:1]
[22]	蔡武城, 袁厚积. 生物物质常用化学分析方法. 北京: 科学出版社, 1982. pp 15-16 Cai W C, Yuan H J. Chemical Analysis Method of Biological Substances Commonly Used. Beijing: Sciences Press, 1982. pp 15-16(in Chinese) [本文引用:1]
[23]	Serna L. The role of brassinosteroids and abscisic acid in stomatal development. Plant Sci, 2014, 225: 95-101 [本文引用:1]
[24]	马均, 朱庆森, 马文波, 田彦华, 杨建昌, 周开达. 重穗型水稻光合作用、物质积累与运转的研究. 中国农业科学, 2003, 36: 375-381 Ma J, Zhu Q S, Ma W B, Tian Y H, Yang J C, Zhou K D. Studies on the photosynthetic characteristics and accumulation and transformation of assimilation product in heavy panicle type of rice. Sci Agric Sin, 2003, 36: 375-381 (in Chinese with English abstract) [本文引用:1]
[25]	于光辉, 陈国祥, 江玉珍, 苑中原, 吕川根. 生殖生长期两优培九功能叶光反应特性. 作物学报, 2010, 36: 1959-1966 Yu G H, Chen G X, Jiang Y Z, Yuan Z Y, Lü C G. Light reaction characteristics in functional leaves of Liangyoupeijiu in the reproductive growth stage. Acta Agron Sin, 2010, 36: 1959-1966 (in Chinese with English abstract) [本文引用:2]
[26]	曹云英, 赵华. 高温胁迫下油菜素内酯对水稻幼苗的保护作用. 中国水稻科学, 2007, 21: 525-529 Cao Y Y, Zhao H. Protective roles of brassinolide in rice seedlings under heat stress. Chin J Rice Sci, 2007, 21: 525-529 (in Chinese with English abstract) [本文引用:1]
[27]	孙旭生, 林琪, 李玲燕, 姜雯, 翟延举. 氮素对超高产小麦生育后期光合特性及产量的影响. 植物营养与肥料学报, 2008, 14: 840-844 Sun X S, Lin Q, Li L Y, Jiang W, Zhai Y J. Effect of nitrogen on photosynthetic characteristics and yield of super-high-yielding wheat at late growth impact. Plant Nutr Fert Sci, 2008, 14: 840-844 (in Chinese with English abstract) [本文引用:1]
[28]	Ying J F, Peng S B, He Q R, Yang H, Yang C D, Visperas R M, Cassman K G. Comparison of high-yield in tropical and subtropical environments: I. Determinants of grain and dry matter yields. Field Crops Res, 1998, 57: 71-84 [本文引用:1]
[29]	姜元华, 许轲, 赵可, 孙建军, 韦还和, 许俊伟, 魏海燕, 郭保卫, 霍中洋, 戴其根, 张洪程. 甬优系列籼粳杂交稻的冠层结构与光合特性. 作物学报, 2015, 41: 286-296 Jiang Y H, Xu K, Zhao K, Sun J J, Wei H H, Xu J W, Wei H Y, Guo B W, Huo Z Y, Dai Q G, Zhang H C. Canopy structure and photosynthetic characteristics of Yongyou series of indica- japonica hybrid rice under high-yielding cultivation condition. Acta Agron Sin, 2015, 41: 286-296 (in Chinese with English abstract) [本文引用:1]
[30]	Li G H, Xue L H, Gu W, Yang C D, Wang S H, Ling Q H, Qin X, Ding Y F. Comparison of yield components and plant type characteristics of high-yield rice between Taoyuan, a “special eco-site” and Nanjing, China. Field Crops Res, 2009, 112: 214-221 [本文引用:1]
[31]	Camargo F A O, Gianello C, Tedesco M J. Soil nitrogen availability evaluated by kinetic mineralization parameters. Commun Soil Sci Plant Anal, 2004, 35: 1293-1307 [本文引用:1]
[32]	Norman R J, Guindo O, Wells B R, Wilson J C E. Seasonal accumulation and partitioning of nitrogen-15 in rice. ·Soil Sci Soc Am J, 1992, 56: 1521-1527 [本文引用:1]
[33]	王强盛, 甄若宏, 丁艳锋, 朱艳, 王绍华, 曹卫星. 钾对不同类型水稻氮素吸收利用的影响. 作物学报, 2009, 35: 704-710 Wang Q S, Zhen R H, Ding Y F, Zhu Y, Wang S H, Cao W X. Effect of potassium application rates on nitrogen absorption and utilization of different types of rice. Acta Agron Sin, 2009, 35: 704-710 (in Chinese with English abstract). [本文引用:1]
[34]	Cock J H, Yoshida S. Accumulation of ¹⁴C-labelled carbohydrate before flowering and the subsequent redistribution and respiration in the rice plant. Proc Crop Sci Soc Jpn, 1972, 41: 226-234 (in Japanese with English abstract) [本文引用:1]
[35]	林贤青, 王雅芬, 朱德峰, 罗玉坤. 水稻茎鞘非结构性碳水化合物与穗部性状关系的研究. 中国水稻科学, 2001, 15: 155-157 Lin X Q, Wang Y F, Zhu D F, Luo Y K. The non-structural carbohydrate of the stem and sheath in relation to the panicle characteristics in rice. Chin J Rice Sci, 2001, 15: 155-157 (in Chinese with English abstract) [本文引用:1]
[36]	赵全志, 黄丕生, 凌启鸿. 水稻群体光合速率和茎鞘贮藏物质与产量关系的研究. 中国农业科学, 2001, 34: 304-310 Zhao Q Z, Huang P S, Ling Q H. Relations between canopy apparent photosynthesis and store matter in stem and sheath between and yield and nitrogen regulations in rice. Sci Agric Sin, 2001, 34: 304-310 (in Chinese with English abstract) [本文引用:1]
[37]	Yang Z F, Inoue N, Fujita K, Masakazu K. Analysis of dry-matter translocation during grain filling stage of rice. Jpn J Crop Sci, 2004, 73: 416-423 (in Japanese with English abstract) [本文引用:1]
[38]	Hirose T, Ohsugi R. Sucrose transport pathways in source and sink tissues of plants and crops. Jpn J Crop Sci, 1998, 67: 129-135 (in Japanese with English abstract) [本文引用:1]
[39]	林忠成, 李土明, 吴福观, 张洪程, 戴其根, 叶世超, 郭宏文. 基蘖肥与穗肥氮比例对双季稻产量和碳氮比的影响. 植物营养与肥料学报, 2011, 17: 269-275 Lin Z C, Li T M, Wu F G, Zhang H C, Dai Q G, Ye S C, Guo H W. Effects of nitrogen application on yield and C/N of double-cropping rice. Plant Nutr Fert Sci, 2011, 17: 269-275 (in Chinese with English abstract) [本文引用:1]
[40]	阮新民, 施伏芝, 罗志祥. 施氮对高产杂交粳稻生育后期叶碳氮比与氮素吸收利用的影响, 中国土壤与肥料, 2011, (2): 35-38 Ruan X M, Shi F Z, Luo Z X. Effects of nitrogen application on the leaf of C/N and nitrogen uptake and utilization at later developmental stages in different high yield hybrid rice varieties. Soil Fert Sci China, 2011, (2): 35-38 (in Chinese with English abstract) [本文引用:1]

2006

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... 油菜素甾醇(BR)类物质作为一种新型的调控植物生长发育的重要激素^[1], 参与了植物的营养生长^[2]与生殖生长^[3], 能够增强水稻叶片的光能捕获能力^[4], 因而在植物生长调控中起重要作用 ...

1994

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2000

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2008

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... 目前, 关于油菜素甾醇类激素对农作物生长发育的研究主要集中在提高产量^[4,5]、促进植株伸长^[6]、BR合成途径^[7]及信号传导^[8]等方面 ...

... Wu等^[4]也指出BRs能够改变叶片构型促进叶片及籽粒中的糖代谢, 从而促进粒重增加和增产 ...

2007

0.0

... 目前, 关于油菜素甾醇类激素对农作物生长发育的研究主要集中在提高产量^[4,5]、促进植株伸长^[6]、BR合成途径^[7]及信号传导^[8]等方面 ...

2015

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张海丽, 高静, 张昊, 李生辉, 邢继红, 王凤茹, 董金皋. 油菜素内酯对水稻细胞伸长和分裂的调控. 农业生物技术学报, 2015, 23: 71-79

Zhang H

, Gao

, Zhang

, Li S

, Xing J

, Wang F

, Dong J

. The regulation of brassinosteroid (BR) on elongation and division of rice (Oryza sativa L. ) cells. J Agric Biotechnol, 2015, 23: 71-79 (in Chinese with English abstract)

Brassinosteroid (BR) is one of the most important hormones which regulate plant growth and development. To clarify the regulatory function of BR in rice (Oryza sativa) cell development, rice calli and suspension cells treated with different concentrations of exogenous BR and the mutant OsBZR1-OE in which the endogenous BR signal enhanced was used to analyze the effect of BR on the volume and cell distribution in rice callus, the shape and size of the single cell suspension, and we observed the cells in OsBZR1-OE transgenic rice seedlings, analyzed expression level of the key genes in regulating cell division and the accumulation of cytoskeletal protein (F-actin) which plays an important role in cell division. We found that low concentration (1×10-10 mol/L) BR could increase the calli volume but the high concentration (1×10-6 mol/L) BR blocked the growth of rice calli. Consistent with calli size, the rice calli treated with high concentrations of BR showed a high density nuclei distribution pattern (cell volume was small) whereas the calli treated with low concentrations of BR showed a low density nuclei distribution pattern (cell volume was large). In addition, BR treatments of single suspension cells showed similar results, low concentration of BRpromoted the cell elongation but the high concentration blocked the cell elongation. Moreover, the length of hypocotyl cells in OsBZR1-OE transgenic rice plants (in which the BR signaling was enhanced) was shorter than that in wild type rice. Propidium iodide(PI) and fluorescein isothiocyanate isomerⅠ(FITC)-phalloidin staining showed that BR treatment could promote the division of suspension and cytoskeleton, and the promoted effect of the high concentration BR treatment was more obvious. Two dimensional electrophoresis results indicated that BR could promote the accumulation of actin cytoskeleton protein. RT-PCR results showed that BR could increase the expression of CDC48 (cell division cycle protein 48) gene which can promote mitosis and CYCD2 (cyclin D2) gene which can shorten the cell cycle. Taken together, these results suggested that low concentration of BR could promote the elongation of rice cells but high concentration inhibited the elongation of rice cells via affecting the expression of cytoskeleton protein F-actin and the cell cycle regulatory genes CDC48 which can promote mitosis and CYCD2 which can shorten the cell cycle. These results provide a theoretical basis for the application of BR in control of the rice height and increasing the lodging resistance.

油菜素内酯(brassinosteroid, BR)是调控植物生长发育的重要的激素之一。为明确BR在调控水稻(Oryza sativa)细胞发育过程中的作用，本研究以不同浓度外源BR处理的水稻愈伤组织、悬浮系和内源BR信号增强的BR突变体OsBZR1-OE为材料，对BR处理后水稻愈伤组织的体积和细胞分布状况、单个悬浮细胞的形状和大小、OsBZR1-OE转基因水稻的细胞进行了观察，并对细胞分裂关键基因的表达量、对细胞分裂具有重要作用的细胞骨架蛋白(F-actin)的积累进行了分析。结果表明，低浓度(1×10-10 mol/L)BR处理的愈伤组织块体积(1.03 cm3)大于对照(0.73 cm3)，但高浓度BR(1×10-6 mol/L)处理的愈伤组织块的体积(0.27 cm3)明显小于对照；从细胞核的分布也能印证上述结果，低浓度BR处理后的愈伤组织细胞核排列疏松，表明细胞体积较大，即1×10-10 mol/L BR促进了水稻愈伤组织细胞体积的增加，高浓度BR处理后的愈伤组织细胞核密度大，表明细胞体积较小，即1×10-6 mol/L BR抑制了水稻愈伤组织细胞体积的增加；低浓度BR促进水稻悬浮系单个细胞的伸长，而高浓度的BR抑制悬浮系细胞的伸长；内源BR信号增强的突变体OsBZR1-OE下胚轴细胞明显短于野生型水稻下胚轴细胞；本研究通过碘化丙碇(propidium iodide, PI)和异硫氰酸荧光素(fluorescein isothiocyanate isomerⅠ, FITC)-鬼笔环肽染色对悬浮系细胞的形态和微丝细胞骨架进行观察，发现BR处理可促进水稻悬浮系细胞和微丝骨架的分裂，高浓度BR对细胞分裂的促进效果更明显；双向电泳结果表明，BR处理可促进微丝骨架蛋白的表达；RT-PCR结果表明，BR可增加促进有丝分裂的CDC48(cell division cycle protein 48)和能缩短细胞周期的CYCD2(cyclin D2)基因的表达。上述结果表明，低浓度BR促进水稻细胞的伸长，高浓度BR抑制水稻细胞的伸长，BR促进微丝骨架F-actin的积累，通过促进有丝分裂基因CDC48和缩短细胞周期的因CYCD2的表达而调控水稻细胞的分裂。本研究为BR在作物控高抗倒伏方面的应用提供了理论依据。

... 目前, 关于油菜素甾醇类激素对农作物生长发育的研究主要集中在提高产量^[4,5]、促进植株伸长^[6]、BR合成途径^[7]及信号传导^[8]等方面 ...

2003

0.0

... 目前, 关于油菜素甾醇类激素对农作物生长发育的研究主要集中在提高产量^[4,5]、促进植株伸长^[6]、BR合成途径^[7]及信号传导^[8]等方面 ...

2015

0.0

王梦姣, 邓百万, 杨国鹏. 拟南芥油菜素内酯的合成、修饰、信号转导及其在农作物育种中的应用研究进展. 河南农业科学, 2015, 44(2): 1-6

Wang M

, Deng B

, Yang G

. Research progress of brassinosteroid in Arabidopsis thaliana and its application in crop breeding. J Henan Agric Sci, 2015, 44(2): 1-6 (in Chinese with English abstract)

... 目前, 关于油菜素甾醇类激素对农作物生长发育的研究主要集中在提高产量^[4,5]、促进植株伸长^[6]、BR合成途径^[7]及信号传导^[8]等方面 ...

2015

0.0

... 李勇等^[9,10]研究指出, 油菜素内酯能够显著提升冬小麦的光合作用参数、总干物质积累量和植株氮积累量 ...

... 1 BRs对杂交粳稻灌浆期植株光合效应的影响有研究指出保证超级稻高产超高产需要提升后期叶片光合效率^[24]并保持叶片光合功能期^[25], 本试验结果表明, eBL能够增强叶片净光合速率和蒸腾速率, 维持气孔导度和胞间CO₂浓度, 促进杂交粳稻叶片净光合速率, 这与李勇等在冬小麦上研究结果相一致^[9,10] ...

2015

0.0

... 李勇等^[9,10]研究指出, 油菜素内酯能够显著提升冬小麦的光合作用参数、总干物质积累量和植株氮积累量 ...

2000

0.0

... 李晓玲等^[11]研究发现, 高油菜素内酯能够改善不同施肥水平下小麦幼苗的生长、生理生化指标及产量构成因子, 使低肥区增产5 ...

1992

0.0

... 刘大永等^[12]发现表油菜素内酯能提高水稻吸收肥料和土壤的P₂O₅量 ...

2003

0.0

严建明, 翟虎渠, 万建民, 焦德茂, 张荣铣. 亚种间重穗型杂交稻光合产物的运转特性及其生理机制. 中国农业科学, 2003, 36: 502-507

Yan J

, Zhai H

, Wan J

, Jiao D

, Zhang R

. Transportation characteristics of assimilate and physiologic mechanisms in subspecific heavy ear hybrid rice (Oryza sativa L. ). Sci Agric Sin, 2003, 36: 502-507 (in Chinese with English abstract)

南京农业大学作物遗传与种质创新国家重点实验室

利用14 C、3 2 P同位素示踪技术 ,研究了籼粳亚种间重穗型杂交稻Ⅱ优 16 2、两优培九的光合产物运转特性及其生理机制。结果表明 ,两组合在籽粒形成期的各个阶段 ,其14 C 光合产物的单株生产总量明显高于汕优 6 3,但光合产物运输至穗中的速度皆不如汕优 6 3快 ,最终分配至穗中的比率也比汕优 6 3低 ,且14 C 光合产物在这两个组合的茎鞘中有明显的积累和滞留现象发生。随稻穗发育进程 ,供试亚种间重穗型杂交稻叶片蔗糖磷酸合成酶活性和根系活力由齐穗期、乳熟期、蜡熟期逐步大幅度下降 ,这种蔗糖磷酸合成酶活性和根系活力下降的特性与光合产物运转分配效率较差有一定的内在联系。

... 提高单产是增加稻谷产量的永恒主题, 种植杂交粳稻是提高产量的一种有效途径, 大穗型杂交水稻品种叶片净光合速率高, 物质积累能力强^[13], 但由于源库矛盾问题^[14]难以充分发挥高产潜力, 因而目前杂交水稻品种普遍存在结实率低、产量不稳定等问题^[15], 严重阻碍了其增产潜力的发挥^[16] ...

2004

0.0

2008

0.0

敖和军, 王淑红, 邹应斌, 彭少兵, 唐启源, 方远祥, 肖安民, 陈玉梅, 熊昌明. 超级杂交稻干物质生产特点与产量稳定性研究. 中国农业科学, 2008, 41: 1927-1936

Ao H

, Wang S

, Zou Y

, Peng S

, Tang Q

, Fang Y

, Xiao A

, Chen Y

, Xiong C

. Study on yield stability and dry matter characteristics of super hybrid rice. Sci Agric Sin, 2008, 41: 1927-1936 (in Chinese with English abstract).

[Objective] The yield performance, the growth characteristics and the ecological adaptability of super hybrid rice were ascertained under different fertilizer application conditions in different tested sites. [Method] Field experiments with three fertilizer rate treatments (135, 180, 225 kg N/hm2) and two combinations (Zhunliangyou-527 and Liangyou-293) were conducted in ChangSha, GuiDong, HengYang, NanXian and YongZhou of Hunan province in 2004 and 2005. [Results] The significant differences of the dry matter accumulation, the harvest yield and its components were founded among the five tested sites and between two years, but no significant differences were founded among three fertilizer rate treatments. The yield components of super hybrid rice performed multifarious properties, of which the harvest yield showed the significantly positive correlations with the productive panicles, the grain filling percentage and 1000 grains weight, and the significantly negative correlation with grain number per panicle and no correlation with spikelet number per unit area. [Conclusion] The harvest yield of super hybrid rice was mainly influenced by the temperature and sunshine conditions of the tested sites and years. The fertilizer rate in the range from 135 kg N/hm2 to 225 kg N/hm2 was not the factor of harvest yield limited. And to increase panicle number was the possible path to increase harvest yield.

湖南农业大学农学院

【目的】探明超级杂交稻在不同种植地点和不同施肥量条件下的产量表现及干物质生产特点。【方法】于2004～2005年在湖南省桂东、长沙、衡阳、南县和永州5个地点进行大田试验，按照N﹕P2O5﹕K2O为1﹕0.5﹕1的比例，设置3种施肥量处理（135、180、225 kg N•ha-1），田间采用随机区组排列，4次重复，以超级杂交稻组合准两优527和两优293为试验材料。【结果】超级杂交稻收获产量以桂东点产量最高，地点间差异显著，其中准两优527平均为7 492.3～12 209.2 kg•ha-1，两优293为6 984.0～11 679.5 kg•ha-1。产量构成因子和干物质生产量的地点间变化与收获产量一致，但在同一地点的不同施肥量处理间收获产量和干物质生产量差异均不显著。收获产量与单位面积穗数、结实率和千粒重表现为正相关，而与每穗粒数表现为负相关。【结论】超级杂交稻存在适宜的种植区域，且在施肥量为135～225 kg N•ha-1的范围内，施肥量不是超高产栽培的限制因子。超级杂交稻的库容量大，提高结实率和粒重是获得超高产的可能途径。

2010

0.0

2012

0.0

纪洪亭, 冯跃华, 何腾兵, 潘剑, 范乐乐, 李云, 武彪, 肖铭, 梁显林. 超级杂交稻群体干物质和养分积累动态模型与特征分析. 中国农业科学, 2012, 45: 3709-3720

Ji H

, Feng Y

, He T

, Pan

, Fan L

, Li

, Wu

, Xiao

, Liang X

. A dynamic model of dry matter and nutrient accumulation in super hybrid rice and analysis of its characteristics. Sci Agric Sin, 2012, 45: 3709-3720 (in Chinese with English abstract)

【Objective】 Modeling dynamics of dry matter and nutrient accumulation of super hybrid rice is an important task for further improvement of yield and rational application of fertilizer.【Method】A field experiment with two super hybrid cultivars (Zhunliangyou527, Qyou 6) and CK(Ⅱyou838) was conducted in 2011, and dry matter accumulation (DMA) and nitrogen, phosphorus and potassium accumulation（NA, PA, KA）were measured. The dynamic equations of relative dry matter accumulation (DMAR) and nitrogen, phosphorus and potassium accumulation （NAR, PAR, KAR） were established with the normalized DMA, normalized NA, PK and KA and normalized growth time, and the dynamic characteristics of DMA and NA, PA, KA changes were analyzed based on the relative equations.【Result】Gompertz simulated better than the other equations for the DMA and nutrient dynamic accumulation characteristics of super hybrid rice. The equations were validated with field experimental data of 2009 and 2011.The NRMSE of DMA simulation were relatively smaller, and R2 values were above 0.9820. The accuracies were about 1, and the NS values were above 0.9530. Validation for the nutrient accumulation equation showed that the model simulated well. The R2, k and NS were all about 1. The analysis of growth characteristics of super hybrid rice showed that the rate of DMA in super hybrid rice was slightly higher than CK at earlier stage and significantly at last stage. The maximum rate of DMA of super hybrid rice was from boot stage to heading stage. The initial of DMA rapid growth stage for super hybrid rice were lower than the CK. The duration of rapid growth period of DMA of super hybrid rice was 71-75 d, which was 16-19 d longer than the CK and the ratio of DMA of rapid growth period in total DMA of super hybrid rice was 4.47%-11.25% higher than that of the CK. The maximum NA rate of super hybrid rice was in 10 d before boot stage. The NA (nitrogen accumulation) rapid growth stage was from 12 d before jointing stage to heading stage, and the NA in rapid growth period was 65.60% of total NA. The maximum PA rate of super hybrid rice was in 8 d before boot stage. The PA（phosphorus accumulation）rapid growth stage was from after jointing stage to 7 d before heading stage, and the PA in rapid growth period was 68.36% of total PA. The maximum KA rate of super hybrid rice was in 3-4 d after jointing stage. The PA(potassium accumulation) rapid growth stage was from 12-16 d after jointing stage to 1-5 d before heading stage, and the KA in rapid growth period was 60.10%-61.71% of total KA. 【Conclusion】The study used the Gompertz equation to simulate the DMA and nutrient dynamic accumulation characteristics of super hybrid rice. The advantage of DMA and nutrient of super hybrid rice were that the duration of rapid growth was longer than that of conventional rice, and the accumulation rate at last stage was faster relatively.

1.贵州大学农学院，贵阳550025

【目的】定量描述超级杂交稻干物质与养分积累的动态变化，为进一步提高超级杂交稻干物质积累的潜力以及肥料的合理施用提供依据。【方法】于2011年在大田条件下，以超级杂交稻准两优527、Q优6号和对照品种Ⅱ优838为供试材料，通过连续测定干物质和氮磷钾含量，并对干物质和氮磷钾积累量及生长时间进行归一化处理，建立超级杂交稻相对干物质和相对氮磷钾积累动态模型，进而分析超级杂交稻干物质和氮磷钾积累的动态特征。【结果】Gompertz方程对超级杂交稻干物质和养分的动态模拟效果较好。利用2009年和2011年试验数据对模型进行验证，干物质积累模型模拟的NRMSE较小，R2值均达到0.9820以上，模拟的准确度（以k表示）约为1，NS均在0.9530以上；对养分积累模型检验表明，模型模拟效果较好，不同模型NRMSE较小，R2和k值以及NS值趋近于1。对超级杂交稻生长特性分析表明，超级杂交稻干物质积累速率在前期略高于对照，后期尤为明显，最大干物质积累速率出现在孕穗至抽穗期；超级杂交稻干物质积累快速增长期持续的时间为71—75 d，比对照持续时间长15—19 d；快速增长期干物质积累量占总积累量的比例，超级杂稻比对照品种高4.47%—11.25%。超级杂交稻氮积累的最大速率出现在孕穗期前10 d；氮积累的快速增长期出现在拔节期前12 d至抽穗期，在快速增长期氮的积累量占氮总积累量的65.60%。超级杂交稻磷积累的最大速率出现在孕穗期前8 d；磷积累的快速增长期出现在拔节期至抽穗期前7 d，在快速增长期的磷积累量占磷总积累量的68.36%；超级杂交稻钾积累的最大速率出现在拔节期后3—4 d；钾积累的快速增长期出现在拔节期前12—16 d至孕穗期前1—5 d，在快速增长期的钾积累量占钾总积累量的60.10%—61.71%。【结论】采用Gompertz模型y=ae-exp(b-cx)模拟了超级杂交稻干物质和养分的积累动态。超级杂交稻干物质和养分的优势在于快速增长期持续时间较长，中后期干物质和养分积累速率较快。

... 纪洪亭等^[17]指出超级杂交水稻干物质和养分的优势在于快速增长期持续时间较长, 中后期干物质和养分积累速率较快, 而不同生育阶段干物质积累量的协调能够促进水稻增产^[18] ...

... 水稻群体干物质积累对产量形成有着重要的促进作用^[17,18], 进一步提高产量主要依靠生物学产量的提高, 特别是籽粒充实期的物质生产是高产水稻关键所在^[28], 本试验结果表明, CK、eBL和Brz处理后灌浆期常规杂交粳稻常优3号积累的干物质分别为10 ...

2012

0.0

邓飞, 王丽, 刘利, 刘代银, 任万军, 杨文钰. 不同生态条件下栽培方式对水稻干物质生产和产量的影响. 作物学报, 2012, 38: 1930-1942

Deng

, Wang

, Liu

, Liu D

, Ren W

, Yang W

. Effects of cultivation methods on dry matter production and yield of rice under different ecological conditions. Acta Agron Sin, 2012, 38: 1930-1942 (in Chinese with English abstract)

To explore the effects of cultivation methods on dry matter production and grain yield of rice under different ecological conditions, we measured dry matter accumulation, distribution, translocation, grain yield and its components in a field experiment with different seedling ages and transplanting methods in Renshou, Pixian and Ya’an using a randomized block experimental design. The results showed as follows. (1) The dry matter accumulation of rice such as dry matter weight per stem or in a population was clearly influenced by ecological condition, seedling age, transplanting method, and there were significant interactions of ecological condition and seedling age on dry matter weight of population from tillering to heading stages. Moreover, the interactions of ecological condition and transplanting method, seedling age and transplanting method, and all of the three factors mainly affected dry matter weight per stem and in a population after heading stage. (2) The diversification of dry matter production characteristics was due to the ecological condition changes. The grain yield was primarily come from the accumulation of photosynthate after heading stage in Renshou, but not significantly correlated with the dry matter translocation from stem-sheath. In Pixian, the dry matter translocation from stem-sheath affected grain yield intensely, rice yield was positively correlated with dry matter translocation ratio from stem-sheath at booting stage ( r = 0.775 * ), but negatively with that at maturity ( r = –0.757 * ), significantly. In addition, the dry matter redistribution from stem-sheath after heading was positively correlated with rice yield in Ya’an. (3) The yield in Renshou increased by 5.52% and 17.65% respectively, compared with that in Pixian and Ya’an. Seedling age and transplanting method had an effect on the yield, that was significantly different under the different cultivation methods. Cultivation methods influenced the yield through changing effective panicles per unit area, seed-setting rate, and 1000-grain weight in Renshou, while by changing effective panicles per unit area and spikelets per panicle in Pixian, and by changing total spikelets and 1000-grain weight in Ya’an. (4) It was suggested that, with the effective regulation of cultivation methods on rice dry matter production, a higher yield would be reached. Overall, the 50 d single seedling with optimized-broadcasting treatment could coordinate the dry matter accumulation at different growth stages to promote the grain yield in Renshou and Pixian. Furthermore, the 50 d single seedlings with hand-transplanting treatment enhanced the dry matter accumulation before heading and the dry matter translocation from stem-sheath after heading, resulting in the extraordinarily increased rice yield inYa’an.

1 Sichuan Agricultural University, Key Laboratory of Crop Ecophysiology and Farming System in Southwest China, Ministry of P.R. China, Wenjiang 611130, China; 2 Sichuan General Popularization Centre of Agricultural Technique, Chengdu 610041, China

为明确不同生态条件下栽培方式对水稻物质生产及产量的影响, 采用随机区组多点试验设计, 研究了不同秧龄和移栽方式对仁寿、郫县和雅安生态点水稻干物质积累与分配、茎鞘干物质输出与转化、产量及构成因素的影响。结果表明: (1)水稻的干物质积累特性为生态条件、秧龄、移栽方式及其互作效应共同作用的结果。水稻单茎和群体干物重均受三者显著主效作用; 生态条件与秧龄互作效应极显著影响分蘖盛期至抽穗期群体干物重, 而生态条件与移栽方式、秧龄与移栽方式及三者互作则主要影响抽穗后水稻单茎和群体干物重。(2 ) 高产水稻干物质生产特性因生态条件的变化而改变。仁寿的产量主要来自抽穗后光合产物在籽粒中的积累, 与茎鞘物质的输出和转化相关不显著; 在郫县, 茎鞘物质输出和转化对产量贡献大于在仁寿, 产量与孕穗期茎鞘干物质分配比例( r = 0.775 * )显著正相关, 与成熟期茎鞘干物质分配比例( r = -0.757 * )则呈显著负相关; 在雅安, 抽穗后茎鞘干物质的输出和转化与产量正相关。(3)水稻产量以仁寿最高, 较郫县和雅安高5.52%和17.65%; 秧龄和移栽方式均能影响水稻最终产量, 不同栽培方式间产量差异显著; 仁寿的栽培方式主要通过单位面积有效穗数、结实率和千粒重来影响产量, 在郫县则通过影响单位面积有效穗数和每穗颖花数实现对产量的调控, 雅安的栽培方式主要通过群体颖花量和千粒重调控产量。(4)适宜的栽培方式能有效调控水稻干物质生产, 促进产量的提高。在仁寿和郫县, 50 d秧龄单苗优化定抛有效地协调了不同生育阶段干物质积累量, 促进水稻增产; 在雅安, 50 d秧龄单苗手插具有较高穗前干物质积累量和抽穗后茎鞘干物质输出与转化效率, 增产显著。

2010

0.0

殷春渊, 张庆, 魏海燕, 张洪程, 戴其根, 霍中洋, 许轲, 马群, 杭杰, 张胜飞. 不同产量类型水稻基因型氮素吸收、利用效率的差异. 中国农业科学, 2010, 43: 39-50

Yin C

, Zhang

, Wei H

, Zhang H

, Dai Q

, Huo Z

, Xu

, Ma

, Hang

, Zhang S

. Differences in nitrogen absorption and use efficiency in rice genotypes with different yield performance. Sci Agric Sin, 2010, 43: 39-50 (in Chinese with English abstract)

【Objective】 The objective of this study was to identify the differences in N absorption and use efficiency in rice genotypes with different yield performances. 【Method】 A field experiment was conducted with 20 rice genotypes(10 high-yielding and 10 low-yielding performance) selected from 108 rice genotypes including early (125-135 d, growth duration), medium (136-145 d) and late-maturing (146-155 d) medium japonica and early (156-165 d) and medium-maturing (166-175 d) late japonica rice under 225 kg?hm-2 N fertilizer application in 2006 and 2007 on the farm of Yangzhou University, Jiangsu province, China. Yield, N accumulation, N uptake rate, N use efficiency and their relationship were analyzed. 【Result】 The results showed that the mean yields of high-yielding rice genotypes were 31.6%, 31.94%, 39.47%, 26.21%,and 21.82% higher than those of low-yielding rice genotypes, respectively, with the extension of the growth duration. N accumulation and use efficiency increased with the extension of the growth duration, and which of high-yielding genotypes was significantly higher than those of low-yielding genotypes. N accumulation and uptake rate of high-yielding genotypes were higher than those of low-yielding during the growth phases from transplanting to elongation and from elongation to heading, while it showed a reversed trend during the growth phase from heading to maturing for some genotypes. Correlation analysis indicated that there existed significant or highly significant positive correlations between yield and N accumulation, N use efficiency, N uptake rate during the growth phase from transplanting to elongation.【Conclusion】 Compared with the low-yielding rice genotypes, N accumulation in various growing stages, N uptake rate before heading and N use efficiency of high-yielding rice genotypes were higher. It was obvious that high yield and increasing N accumulation and use efficiency could be coordinated development on improving genotype.

（扬州大学农学院/江苏省作物遗传生理重点实验室/农业部长江中下游作物生理生态与栽培重点开放实验室）

【目的】明确不同产量基因型水稻氮素营养吸收、利用差异。【方法】采用大田试验,以长江中下游地区5种生育期类型中相对高产和低产的20个粳稻品种为材料,研究施氮量为225 kg?hm-2水平下两种产量类型水稻的产量、氮素积累量、氮素吸收速率和氮素利用效率的差异及其之间的相互关系。【结果】随着生育期的延长,高产水稻类型的平均产量分别比低产类型高31.62%、31.94%、39.47%、26.21%和21.82%;氮素生育期累积量和氮素利用率随着生育期的延长呈逐渐增加的趋势,高产类型明显高于低产类型;氮素阶段性积累量和阶段性吸收速率的变化为：移栽至拔节和拔节至抽穗阶段为高产类型高于低产类型;抽穗至成熟阶段部分基因型表现为低产类型高于高产类型,但差异不大。相关分析表明,水稻产量与氮素积累量、氮素利用率呈显著或极显著正相关,与各阶段的氮素吸收速率亦呈正相关关系,其中移栽至拔节阶段达极显著水平,其余阶段未达显著水平。【结论】相对于低产基因型水稻,高产基因型在各个生育阶段的氮素积累量、抽穗前的氮素吸收速率及氮素利用效率均较高。可见,高产与提高氮素的积累和利用在品种本身的改良上可得到统一。

... 殷春渊等^[19]认为高产基因型水稻在各个生育阶段均具有较高氮素积累量, 茎叶部非结构性物质积累量和含量较高的品种具有稳产和高产潜力^[20] ...

2013

0.0

... 殷春渊等^[19]认为高产基因型水稻在各个生育阶段均具有较高氮素积累量, 茎叶部非结构性物质积累量和含量较高的品种具有稳产和高产潜力^[20] ...

1999

0.0

... 采用硫酸消煮, 半微量凯氏定氮法^[21]测定全氮含量 ...

1982

0.0

... 合并3次离心所得上清液, 用水洗沉淀并离心2次, 合并所有上清液并用蒸馏水定容至50 mL, 采用蒽酮法^[22]测定NSC含量 ...

2014

0.0

... T_r和g_s变化趋势与P_n相同, 花后10 d处理间T_r出现显著差异, 而花后0 d g_s即出现显著差异, 因为BRs能够直接调控植株气孔形成和分布^[23], 所以相较于T_r和P_n, BR对G_s的作用时间最快, 喷施后1~2 d即产生生理效应且变化幅度较大 ...

2003

0.0

马均, 朱庆森, 马文波, 田彦华, 杨建昌, 周开达. 重穗型水稻光合作用、物质积累与运转的研究. 中国农业科学, 2003, 36: 375-381

, Zhu Q

, Ma W

, Tian Y

, Yang J

, Zhou K

. Studies on the photosynthetic characteristics and accumulation and transformation of assimilation product in heavy panicle type of rice. Sci Agric Sin, 2003, 36: 375-381 (in Chinese with English abstract)

四川农业大学水稻研究所

以 18个不同穗重型籼稻品种为材料 ,研究了重穗型水稻的光合作用及物质运转。结果表明 ,重穗型水稻在齐穗后的剑叶净光合速率明显高于中、轻穗型品种。其机理是剑叶的Rubisco活性提高及叶绿素含量增加 ;有在高、低光强、高温和低CO2 浓度下对CO2 较强的同化能力的优势 ,光合“午休”现象较轻 ,能保持较高的净光合速率 ,显示其对环境有较强的适应性 ;气孔性状优良、气孔开度积 (单位面积气孔密度×气孔开度 )大 ,也是重穗型水稻净光合速率高的重要气孔因素。重穗型水稻抽穗后物质积累的优势明显 ,且茎鞘物质向穗部的转运量、转运率和转化率也远高于中、轻穗型品种。

2010

0.0

于光辉, 陈国祥, 江玉珍, 苑中原, 吕川根. 生殖生长期两优培九功能叶光反应特性. 作物学报, 2010, 36: 1959-1966

Yu G

, Chen G

, Jiang Y

, Yuan Z

, Lü C

. Light reaction characteristics in functional leaves of Liangyoupeijiu in the reproductive growth stage. Acta Agron Sin, 2010, 36: 1959-1966 (in Chinese with English abstract)

The chloroplast light reaction characteristics were compared between super-high-yield hybrid rice Liangyoupeijiu and traditional hybrid rice Shanyou 63 intending to provide theoretical insights into physiological basis for high yield. Using fluorescence dynamics analysis and physiological and biochemical research techniques, in the field we systematically studied the primary response, electron transport chain and photophosphorylation during the reproductive period. The results showed that: (1) as compared to Shanyou 63, chlorophyll content in functional leaves of Liangyoupeijiu was 22.90% higher, it had a relatively longer photosynthetic function duration and chla/chlb ratio was 36.78% higher; (2) there was no significant difference in light absorption per unit leaf area of functional leaves, but Liangyoupeijiu maintained a high light energy absorption capacity and long stability period , the number of active reaction centers was more, the energy of heat dissipation was relatively lower, the energy transferred into the electron transport chain was higher; (3) fluorescence analysis showed that structures and status of the body side, the receptor side and the reaction center in PSII performed better than those of Shanyou 63, indicating that Liangyoupeijiu had a higher efficiency in the transforming light energy into electric energy; (4) in addition, oxygen evolution of the chloroplast, activities of electron transport chain and photophosphorylation were significantly higher, indicating that Liangyoupeijiu possessed some advantages in the energy convertsion from electric energy to active chemical energy. With higher light energy absorption, transmission and conversion efficiency, Liangyoupeijiu established a physicological basis of super-high-yield.

1 College of Life Sciences, Nanjing Normal University, Nanjing 210046, China; 2 Institute of food & Crops, Jiangsu Academy of Agricultural Sciences, Nanjing 210014, China

在野外条件下，利用荧光动力学分析和生理生化研究技术，对生殖生长期超高产杂交稻两优培九和大面积推广稻汕优63功能叶原初反应、电子传递和光合磷酸化水平进行系统研究，比较光反应特性。结果表明: (1) 与汕优63相比，两优培九功能叶叶绿素含量高22.90%，光合功能期长约50%，叶绿素a/b比值高36.78%；(2) 光能吸收与分配方面，两优培九功能叶单位叶面积吸收的光能不占优势，但保持高吸收的稳定期长，有活性的反应中心数量多，热耗散的能量比例较少，进入电子传递链的能量高；(3) 荧光参数分析发现两优培九PSII供体侧、受体侧和反应中心性能优良，光能吸收、传递和转化为电能效率较高；(4) 叶绿体放氧活性、电子传递链和光合磷酸化活性均显著高于汕优63，表明两优培九电能转化为活跃化学能能力强。

... 62%, 并且常优3号变化幅度大于甬优12 (图1), 其叶片SPAD值对BR敏感性高于常优3号, 这与曹云英等^[25]关于eBL喷施增加水稻叶片叶绿素含量的研究结果基本相似 ...

2007

0.0

曹云英, 赵华. 高温胁迫下油菜素内酯对水稻幼苗的保护作用. 中国水稻科学, 2007, 21: 525-529

Cao Y

, Zhao

. Protective roles of brassinolide in rice seedlings under heat stress. Chin J Rice Sci, 2007, 21: 525-529 (in Chinese with English abstract)

Two indica rice(Oryza sativa L.) materials,Xieqingzao B(heat-sensitive) and 082(heat-tolerant),were used to study the role of brassinolide(BR) in protection of rice seedlings from heat stress.Young seedlings were subjected to higher temperature and sprayed with 0.005 mg/L of BR.Analysis was conducted on the contents of chlorophyll,protein,malondialdehyde(MDA),electrolyte leakage,the activities of peroxidase(POD) and superoxide dismutase(SOD) and their isozymes expression levels in leaves.Under the heat stress treatment,application of BR significantly increased the contents of chlorophyll and protein,the activities of POD and SOD,and reduced the level of MDA and the leakage of electrolyte in the leaves of the heat-sensitive material Xieqingzao B,whereas BR had less effect on those of the heat-tolerant material 082,relatively.The BR treatment enhanced the expression level of POD isozymes in the leaves of both materials.Under the heat stress and BR treatment,the expression of four SOD isozymes were reduced in 082,but the expression of two SOD isozymes A and B were increased in Xieqingzao B.This suggested that BR plays an important role in protection rice seedlings from heat stress through enhancing the activities or expression levels of protective enzymes in the leaves.The materials with various heat-tolerance might differ in the mechanism of response to heat stress with BR application.

以两个籼稻材料082（耐热型）和协青早B（热敏型）为研究对象探讨高温胁迫下油菜素内酯（BR）对水稻秧苗的保护作用。在苗期进行高温和喷施BR处理，观察高温胁迫下BR对叶绿素、蛋白质和MDA含量、电解质外渗率和保护酶（SOD、POD）活性及其同工酶表达等的影响。结果表明，高温胁迫下，喷施BR能明显提高热敏型材料协青早B的叶片叶绿素和蛋白质含量，增加SOD和POD活性,降低叶片MDA含量和电解质外渗率，但对耐热型材料082的影响相对较小。经BR处理后两个材料的POD同工酶谱带表达均明显增强。在高温胁迫下，BR处理减弱了耐热材料082的4条SOD同工酶谱带的表达，但明显增强了热敏感材料协青早B的A、B两同工酶谱带的表达，表明BR能够增强稻苗内活性氧保护酶系统的活性或诱导其表达，能有效减轻高温对秧苗的伤害。BR处理后稻苗对高温的应答机制，在温敏性不同的材料间存在差异。

... 但本试验还表明, 常优3号叶片净光合速率受BR影响小于甬优12, 叶片净光合速率低于甬优12, 常优3号叶片SPAD值变大意味着叶绿素含量增加, 但常规杂交粳稻常优3号净光合速率增加幅度并没有超过籼粳杂交超级稻甬优12, 正如孙旭生等^[26]提出, 叶绿素含量仅是光合速率的一个#cod#x0201c ...

2008

0.0

孙旭生, 林琪, 李玲燕, 姜雯, 翟延举. 氮素对超高产小麦生育后期光合特性及产量的影响. 植物营养与肥料学报, 2008, 14: 840-844

Sun X

, Lin

, Li L

, Jiang

, Zhai Y

. Effect of nitrogen on photosynthetic characteristics and yield of super-high-yielding wheat at late growth impact. Plant Nutr Fert Sci, 2008, 14: 840-844 (in Chinese with English abstract)

The effects of nitrogen supply on photosynthetic characteristics at late developing stage in superhigh-yield winter wheat were studied under field conditions. The net photosynthesis rate（Pn）, intercellular CO 2 concentration（Ci） and stomatal conductance（Gs） in superhigh-yield winter wheat were measured using Li-6400 Portable Photosynthesis System with an open flow gas exchange system. The results showed that nitrogen supply improved the photosynthetic characteristics at late developing stages of superhigh-yield winter wheat. The net photosynthesis rate was enhanced with the increasing of nitrogen supply. However, surplus nitrogen supply (N 375 kg/hm 2 ) accelerated the leaf senescence. The highest grain yield was observed in the treatment with N 300 kg/hm 2 applied.

1 College of Plant Science and Technology, Qingdao Agricultural University , Qingdao, Shandong 266109,China; 2 Laizhou Jinhai Seed Limited Company, Laizhou, Shandong 261400, China

本试验在大田条件下研究了施氮量对超高产小麦生育后期光合特性的影响。利用LI-6400便携式光合测定仪，采用开放式气路测定了超高产麦田旗叶的净光合速率、胞间CO 2 浓度、气孔导度等相关指标。结果表明，氮素对超高产小麦生育后期的光合特性有较大的调节作用，随着施氮量增加，小麦的净光合速率增强，但过高的施氮量（N 375 kg/hm 2 ）导致灌浆后期叶片衰老快，净光合速率下降迅速，叶面积指数降低，千粒重下降明显，最终导致产量的减少。在本试验条件下，超高产麦田的适宜施氮量为N 300 kg/hm 2 。

1998

0.0

2015

0.0

姜元华, 许轲, 赵可, 孙建军, 韦还和, 许俊伟, 魏海燕, 郭保卫, 霍中洋, 戴其根, 张洪程. 甬优系列籼粳杂交稻的冠层结构与光合特性. 作物学报, 2015, 41: 286-296

Jiang Y

, Xu

, Zhao

, Sun J

, Wei H

, Xu J

, Wei H

, Guo B

, Huo Z

, Dai Q

, Zhang H

. Canopy structure and photosynthetic characteristics of Yongyou series of indica- japonica hybrid rice under high-yielding cultivation condition. Acta Agron Sin, 2015, 41: 286-296 (in Chinese with English abstract)

Compared with japonica hybrid rice cultivars(B), conventional japonica rice cultivars (C) and indica hybrid rice cultivars (D), Yongyou series of indica-japonica hybrid rice cultivars (A) were adopted to analyze the lightened posture of leave, light distribution of canopy and photosynthetic characteristics of plant systematically. Results were as follows: (1) leaf length, leaf width of top three leaves showed the trend of A>B>D>C, canopy extinction coefficient showed the trend of C>D>B>A, leaf basic angle of top three leaves and relative height of the largest leaf area density were D>A,>B>C, lower canopy leaf area density was C>B>A>D, plant height and largest leaf area density showed the trend of A>D>B>C, leaf drop angle of top three leaves showed the trend of D>B>C>A, upper canopy leaf area density observed D>A>B>C, lower canopy relative light showed the trend of A>B>C>D, lower canopy relative light and canopy average relative light showed the trend of B>A>C>D; (2) population LAI and high valid leaf area rate showed the trend of D>A>B>C at heading, valid leaf area rate, spikelets per cm2 leaf area, filled grains per cm2 leaf area, total biomass, economic yield, Chlorophyll (a+b) content, carotenoid content, PSII actual photochemical efficiency, net photosynthetic rate, enzyme activities of SOD, POD and CAT after heading showed the trend of A>B>C>D, harvest index and MDA content showed the trend of D>C>B>A. Therefore, compared with other three types of rice, Yongyou Series of indica-japonica hybrid rice had significant advantages in the canopy structure and photosynthetic characteristics, which are both normal ecological and physiological bases of high yield, and maybe an important way to improve the population productivity of inter-subspecific hybrid rice.

Innovation Center of Rice Cultivation Technology in the Yangtze Valley, Ministry of Agriculture / Key Laboratory of Crop Genetics and Physiology of Jiangsu Province, Yangzhou University, Yangzhou 225009, China

旨在阐明甬优系列籼粳杂交稻的冠层结构与光合特性。以甬优籼粳杂交稻(A)为研究对象，以三系杂交粳稻(B)、超级常规粳稻(C)和超级杂交籼稻(D)为对照，在机插高产栽培条件下，对不同类型水稻的叶片受光姿态和群体冠层光分布、光合特性系统比较研究。结果表明: (1)上3张叶片的长度、宽度表现为A>B>D>C，叶基角表现为D>A>B>C，披垂度表现为D>B>C>A。冠层上部叶面积密度表现为D>A>B>C，冠层下部叶面积密度表现为C>B>A>D，最大叶面积密度表现为A>D>B>C，最大叶面积密度出现的相对高度表现为D>A>B>C。冠层上部相对光照表现为A>B>C>D，冠层下部相对光照表现为B>A>C>D，冠层平均相对光照表现为B>A>C>D，冠层消光系数表现为C>D>B>A。(2)抽穗期群体叶面积指数、高效叶面积率均呈D>A>B>C趋势，有效叶面积率呈A>B>C>D趋势；颖花/叶和实粒/叶均表现为A>B>C>D。经济产量、生物产量均表现为A>B>C>D，经济系数呈D>C>B>A趋势；蜡熟期和成熟期剑叶的叶绿素含量、类胡萝卜素含量、PSII的光化学效率及净光合速率呈A>B>C>D趋势；抽穗至成熟期剑叶的MAD含量呈D>C>B>A趋势，SOD、POD、CAT酶活性呈A>B>C>D趋势。与其他3种类型水稻相比，甬优系列籼粳杂交稻的冠层结构与光合特性具有显著优势，这是甬优系列杂交稻产量潜力正常发挥的生态生理基础，也是进一步提高亚种间杂交稻群体生产力的重要途径。

... 89%, 再次揭示提高灌浆期干物质积累量是水稻实现高产超高产的重要途径, 并且eBL能够使籽粒灌浆前期和后期的干物质积累量升高, 为高产水稻奠定了生物学基础, 但本试验结果显示eBL对甬优12干物质积累的促进作用比常优3号小, 这主要是因为籼粳杂交超级稻甬优12比常规杂交粳稻常优3号具有较高的群体干物质积累量, 姜元华等^[29]也有相同试验结果 ...

2009

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... 2 BR对杂交粳稻灌浆期植株碳氮代谢的影响水稻产量的高低与氮素吸收利用关系密切^[30,31], Norman等^[32]指出开花后水稻植株营养器官转运的氮素是籽粒氮素积累量的主要组成部分, 因此适当提高抽穗后氮素积累量和吸收比例有利于水稻的产量提高^[33] ...

2004

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1992

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2009

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王强盛, 甄若宏, 丁艳锋, 朱艳, 王绍华, 曹卫星. 钾对不同类型水稻氮素吸收利用的影响. 作物学报, 2009, 35: 704-710

Wang Q

, Zhen R

, Ding Y

, Zhu

, Wang S

, Cao W

. Effect of potassium application rates on nitrogen absorption and utilization of different types of rice. Acta Agron Sin, 2009, 35: 704-710 (in Chinese with English abstract).

Potassium, nitrogen and their interaction play important roles in plant growth, development and yield of rice. Appropriate potassium can stimulate nitrogen accumulation and translocation in rice. In order to identify plant nitrogen absorption, distribution and utilization characteristics of rice under potassium supply, a field experiment was conducted with four potassium rates (0, 90, 180, 270 kg h a - 1 K 2 O) using conventional japonica cultivars (Wuyunjing 7 and Wuxiangjing 14) and hybrid japonica cultivars (86 you 8, Siyou 422) from 2004 to 2005 on Experimental Farm of Nanjing Agricultural University, Jiangsu province, China. Potassium application increased rice yield (4.56 - 14.77%) and nitrogen accumulation during growth stages, with the maximum from elongation stage to heading stage, but nitrogen uptake ratio decreased. Potassium application enhanced nitrogen transport amount and percentage after anthesis, as well as nitrogen accumulation in different rice organs, nitrogen distribution percentage in leaves and panicles, whereas declined nitrogen distribution percentage in culm and sheath. Nitrogen absorption from fertilizer, basic nitrogen recovery efficiency before elongation stage, nitrogen recovery efficiency during the whole growing stage, and nitrogen harvest index were raised significantly, but plant nitrogen productivity was opposite. Grain yield, nitrogen accumulation, nitrogen translocation amount and percentage after anthesis, and nitrogen distribution in different rice organs amounted to the maximum at K 2 O application rate of 180 kg ha -1 , as well as nitrogen absorption from fertilizer, basic nitrogen recovery efficiency before elongation stage, nitrogen recovery efficiency during the whole growing stage. However, nitrogen absorption percentage from elongation stage to heading stage, nitrogen distribution percentage in culm and sheath, plant nitrogen productivity decreased to the minimum. The yield and nitrogen recovery efficiency of conventional japonica cultivars were higher than those of hybrid japonica cultivars.

1College of Agronomy,Nanjing Zgricultural University,Nanjing 210095,China;2Jiangsu Academy of Agricultural Sciences,Nanjing 210014,China

以常规粳稻武运粳 7 号、武香粳 14 和杂交粳稻 86 优 8 号、泗优 422 为材料，研究了钾对水稻氮素吸收、分配和利用的影响。结果表明，钾对水稻有明显增产效应，增产率 4.56%~14.77% ；钾提高了水稻不同生育阶段植株吸氮量，以拔节期到抽穗期氮素积累增量最大，但吸氮比例下降；钾促进了抽穗后氮素转运量和转运率，提高了氮素在不同器官分配量及叶片、穗分配比例，但降低了茎鞘分配比例；钾增强了植株对肥料氮吸收，基肥氮素利用率、全生育期氮素利用率和氮素收获指数显著增长，但降低了植株氮生产效率；以 180 kg h m - 2 K 2 O 处理产量最高，氮素积累量、转运量和转运率以及不同器官分配量最大，基肥氮素利用效率、植株对肥料氮吸收量和全生育期氮素利用率最大，但拔节期到抽穗期氮素吸收比例、茎鞘氮素分配比例和植株氮生产效率最低；常规粳稻产量高于杂交粳稻，其氮素利用率相对较大。

1972

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... Cock等^[34]认为每个颖花分配的贮存碳水化合物多, 能够提高结实率和千粒重, 因此提高千粒重的关键是茎鞘转移到每个颖花中的非结构性碳水化合物总量高^[35] ...

2001

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林贤青, 王雅芬, 朱德峰, 罗玉坤. 水稻茎鞘非结构性碳水化合物与穗部性状关系的研究. 中国水稻科学, 2001, 15: 155-157

Lin X

, Wang Y

, Zhu D

, Luo Y

. The non-structural carbohydrate of the stem and sheath in relation to the panicle characteristics in rice. Chin J Rice Sci, 2001, 15: 155-157 (in Chinese with English abstract)

The field experiments were conducted using three rice combinations with different panicle traits. The results showed that the higher the non structural carbohydrate (NSC) of the stem and sheath at flowering stage and the lower at mature stage, the higher the seed setting rate was. The higher the average of each spikelet owning NSC at flowering stage, the higher 1000-grain weight was.

( 1 China National Rice Research Institute; Hangzhou 310006; China; 2 College of Food Science; Biology & Environmental Engineering; Hangzhou 310035; China)

以3个不同穗部性状类型的组合广超1号、特优明86和65396为材料进行大田试验，结果表明开花期茎鞘非结构性碳水化合物含量高，成熟期含量低的品种，结实率高，反之，结实率低。每个颖花平均从茎鞘获得非结构性碳水化合物高的品种，千粒重高。反之，千粒重低。

2001

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赵全志, 黄丕生, 凌启鸿. 水稻群体光合速率和茎鞘贮藏物质与产量关系的研究. 中国农业科学, 2001, 34: 304-310

Zhao Q

, Huang P

, Ling Q

. Relations between canopy apparent photosynthesis and store matter in stem and sheath between and yield and nitrogen regulations in rice. Sci Agric Sin, 2001, 34: 304-310 (in Chinese with English abstract)

南京农业大学农学院

采用汕优６３和盐粳１８７品种（组合），通过设置前后期不同氮素配比，对水稻群体光合速率（ＣＡＰ）和茎鞘贮藏物质与产量的关系进行了研究。结果表明，在抽穗后ＣＡＰ随生育进程而逐渐降低。汕优６３以６：４配比ＣＡＰ较高。上３茎鞘物质干重变化呈“Ｖ”字型，在穗后２３ｄ左右达最低值，其中以顶三茎鞘物质输出最多。ＣＡＰ与群体质量指标、穗后物质生产和产量的关系除穗后２３ｄ左右外比较密切，穗后２３ｄ茎鞘贮藏物质大量输出。

... 充实后期叶片、茎鞘积累NSC, 而茎鞘氮素为甬优12转运输出、常优3号吸收累积, 赵全志等^[36]研究认为, 高产水稻在灌浆前中期茎鞘物质输出多容易导致灌浆后期回升多, 而后期茎鞘物质一直向外输出是低产的表现 ...

2004

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... Yang等^[37]发现水稻灌浆期叶片茎鞘积累的非结构性物质通过酶分解为蔗糖等可溶性糖后向穗部转移, Hirose等^[38]研究认为, 叶片茎鞘可溶性糖的含量升高可以增加库源之间可溶性糖的浓度差, 促进非结构性物质向穗部转移 ...

1998

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2011

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林忠成, 李土明, 吴福观, 张洪程, 戴其根, 叶世超, 郭宏文. 基蘖肥与穗肥氮比例对双季稻产量和碳氮比的影响. 植物营养与肥料学报, 2011, 17: 269-275

Lin Z

, Li T

, Wu F

, Zhang H

, Dai Q

, Ye S

, Guo H

. Effects of nitrogen application on yield and C/N of double-cropping rice. Plant Nutr Fert Sci, 2011, 17: 269-275 (in Chinese with English abstract)

The experiment was conducted to study effects of different ratios of basic and tillering, and panicle nitrogen fertilization on yield, dry matter accumulation, nitrogen accumulation and the ratio of carbon and nitrogen of doublecropping rice. The results show that when the total nitrogen rate is N 225 kg/ha, the yield is the highest under the treatment which the ratio of basic and tillering nitrogen amount to panicle nitrogen amount is about 7∶3, the followings are the treatments of ratios of 6∶4 and 8∶2, which are all higher than that of the conventional nitrogen application pattern (10∶0). When the ratio of basic and tillering nitrogen application to total nitrogen application amount is from 60% to 70% of doublecropping rice, the dry matter accumulation, nitrogen accumulation, nitrogen utilization efficiency, N-agronomy efficiency are all increased under the ratio of 7∶3, the carbon metabolism and nitrogen metabolism are also in a good coordination. The usable carbohydrate including soluble sugars and starch of early rice leaf at booting stage is 17%-18%, C/N is 50-55, and the usable carbohydrate of late rice is 19%-21% and C/N is 50-60These would be the physiological base of getting higher rice yield when the ratios are 7∶3 and 6∶4In all, synthesizing the yield, nitrogen utilization efficiency and C/N of doublecropping rice, the ratio of nitrogen application at the panicle stage of rice to total nitrogen application amount is from 30% to 40%.

1 Yangzhou University/Key Laboratory of Crop Genetics and Physiology of Jiangsu Province/Innovation Center of Rice Cultivation Technology in Yangtze Valley, Ministry of Agriculture, Yangzhou, Jiangsu 225009, China; 2 Agricultural Committee of Wujiang County, Wujiang, Jiangsu 215200, China; 3 Ganzhou Institute of Agricultural Sciences, Ganzhou, Jiangxi 341000, China

试验研究了基蘖肥与穗肥氮比例对双季早、晚稻产量,干物质积累量,氮素积累量和碳氮比的影响。结果表明，当总施氮量为N 225 kg/hm2时，基蘖肥与穗肥氮比例7∶3处理的产量最高，其次为6∶4、8∶2处理，均比当地习惯施肥法（10∶0）高产。同时，当基蘖肥占总施氮量60%70%时，双季早、晚稻具有较高的干物质积累量、氮素积累量、氮素当季利用率、氮素农艺效率，群体的碳氮代谢也比较协调。早稻孕穗期叶片可用性糖（可溶性总糖＋淀粉）含量17%18%，碳氮比5055，晚稻孕穗期叶片可用性糖含量19%21%，碳氮比5060，这可能是基蘖肥与穗肥氮比例为7∶3和6∶4时双季水稻高产的生理基础。综合双季早、晚稻产量、氮素利用率及碳氮比值，穗肥施氮量占总施氮量的适宜比例为30%-40%。

... 植株C/N改变能够影响植株的生理代谢^[39], 本研究显示植株不同器官的NSC/N随着籽粒充实进程升高, BR喷施后显著改变了茎鞘和穗部的NSC/N (表8), 因而BR能够调控水稻灌浆期植株氮素吸收量和光合物质转运量, 增强茎鞘物质输送强度和穗部灌浆物质积累能力 ...

2011

0.0

阮新民, 施伏芝, 罗志祥. 施氮对高产杂交粳稻生育后期叶碳氮比与氮素吸收利用的影响, 中国土壤与肥料, 2011, (2): 35-38

Ruan X

, Shi F

, Luo Z

. Effects of nitrogen application on the leaf of C/N and nitrogen uptake and utilization at later developmental stages in different high yield hybrid rice varieties. Soil Fert Sci China, 2011, (2): 35-38 (in Chinese with English abstract)

大田试验条件下,研究了施肥对水稻生育后期高产杂交水稻吸氮特性、叶碳氮比及氮肥利用率的影响.结果表明,吸氮量主要集中在生育前期,抽穗期前吸氮量占成熟期的70%以上.皖稻125在适量施氮及氮胁迫条件下氮素积累能力均较强,促进了干物质积累和产量形成.叶碳含量基因型变异较小,且在生育后期总体呈下降趋势.叶碳氮比变异较大,抽穗期、灌浆期及成熟期在无氮区分别较施氮区高28.6%、18.3%、36.4%.适宜的叶碳氮比有利于氮素运转和产量的形成,在适量施氮下,皖稻125的氮肥农学利用率和生理利用效率均最高,分别为13.97和56.5 kg/kg,较汕优63高10.2%和24.4%.

... 另外, 本研究表明BRs喷施后对叶片NSC/N没有显著差异, 这主要是叶片存在NSC和氮素的协同转运变化, 这与阮新民等^[40]研究认为水稻叶片碳氮比差异主要在于叶片氮含量差异的结果略有不同 ...