北方水稻生育后期剑叶可溶性物质含量及植株生产力对CO<sub>2</sub>浓度增高的响应

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王惠贞, 赵洪亮, 冯永祥, 姜乐, 宁大可, 谢立勇, 林而达. 北方水稻生育后期剑叶可溶性物质含量及植株生产力对CO₂浓度增高的响应 . 作物学报, 2014, 40(2): 320-328[WANG Hui-Zhen, ZHAO Hong-Liang, FENG Yong-Xiang, JIANG Le, NING Da-Ke, XIE Li-Yong, LIN Er-Da. Response of Soluble Substances Content in Flag Leaves during Late Growth Stage and Plant Productivity of Rice to Elevated CO₂ in North China . 作物学报, 2014, 40(2): 320-328] 复制到剪切板

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北方水稻生育后期剑叶可溶性物质含量及植株生产力对CO₂浓度增高的响应

王惠贞¹, 赵洪亮¹, 冯永祥², 姜乐¹, 宁大可¹, 谢立勇^1,^*, 林而达^3,^*

¹沈阳农业大学农学院, 辽宁沈阳 110866

² 黑龙江八一农垦大学农学院, 黑龙江大庆 163319

³ 中国农业科学院农业环境与可持续发展研究所, 北京 100081

^* 通讯作者(Corresponding authors): 谢立勇, E-mail:xly0910@163.com, Tel: 024-88487135; 林而达,lined@ami.ac.cn, Tel: 010-82105998

收稿日期:2013-08-31

基金:本研究由国家自然科学基金项目(41175097)和国家科技支撑计划项目(2013BAD11B03)资助。

摘要

以高产优质粳稻松粳9号和稻花香2号为试材, 利用中国北方FACE (free air CO₂ enrichment)实验平台研究北方水稻生理代谢对CO₂浓度增高的响应规律。在水稻抽穗期开始测定剑叶可溶性糖含量、蛋白质含量和总叶绿素含量, 收获后实测小区产量及产量构成因素, 比较处理间和品种间差异。结果表明, CO₂浓度增高显著提高水稻抽穗期、乳熟期和完熟期剑叶可溶性糖含量, 松粳9号和稻花香2号最高增幅分别达11.7%和47.5%。CO₂浓度增高显著降低抽穗期和完熟期剑叶可溶性蛋白含量, 松粳9号和稻花香2号最大降幅分别为16.2%和10.5%。CO₂浓度增高使抽穗期和乳熟期剑叶总叶绿素含量显著增加, 松粳9号和稻花香2号最大增幅达18.9%和22.5%, 之后便逐渐降低。CO₂浓度增高使松粳9号单株籽粒产量、生物学产量、经济系数分别提高6.82%、1.50%和12.64%, 稻花香2号平均升高2.56%、2.13%和26.05%。研究表明, CO₂浓度增高最终提高了水稻植株生产力, 对可溶性物质含量的影响在不同生育期存在差异。这可能由于CO₂浓度增高促进水稻生长发育, 导致水稻提早成熟, 叶片衰老促进了可溶性糖分解, 也加快了功能叶可溶性蛋白向籽粒运输速率。

关键词: FACE; 水稻; 生育后期; 可溶性糖; 可溶性蛋白质; 叶绿素; 植株生产力

Response of Soluble Substances Content in Flag Leaves during Late Growth Stage and Plant Productivity of Rice to Elevated CO₂ in North China

WANG Hui-Zhen¹, ZHAO Hong-Liang¹, FENG Yong-Xiang², JIANG Le¹, NING Da-Ke¹, XIE Li-Yong^1,^*, LIN Er-Da^3,^*

¹ College of Agronomy, Shenyang Agricultural University, Shenyang 110866, China

² College of Agronomy, Heilongjiang Bayi Agricultural University, Daqing 163319, China

³ Institute of Environment and Sustainable Development in Agriculture, Chinese Academy of Agricultural Sciences, Beijing 100081, China

第一作者联系方式: E-mail:whz0301@163.com; Tel: 18311485048

Abstract

The responses of high yield and good qualityjaponica varieties Songjing 9 and Daohuaxiang 2 to elevated CO₂ concentration in North China were studied by using FACE experimental system. The soluble sugar content, soluble protein content and total chlorophyll content in flag leaves of rice were measured at late growth stage, rice yield per plot and yield components were measured at harvest stage. The results showed that elevated CO₂ concentration increased soluble sugar content in flag leaf for Songjing 9 and Daohuaxiang 2 obviously during heading stage, milky stage and ripening stage, with the peak value of 11.7% and 47.5%, respectively. Elevated CO₂ concentration obviously reduced soluble protein content in flag leaf during heading stage and ripening stage for Songjing 9 and Daohuaxiang 2, which was 16.2% and 10.5% in the largest reduction, respectively. Elevated CO₂ concentration obviously increased total chlorophyll content in flag leaf for Songjing 9 and Daohuaxiang 2 from heading stage to milky stage, with the largest increase of 18.9% and 22.5%, and then decreased after milk stage. Elevated CO₂ concentration increased seed yield of individual plant, total biomass and economic coefficient with an average of 6.82%, 1.50%, 12.64% for Songjing 9, and 2.56%, 2.13%, 26.05% for Daohuaxiang 2. The results indicated that elevated atmospheric CO₂ concentration increases rice plant productivity, but with diffrence effects on soluble sugar, soluble protein and chlorophyll in flag leaf of different rice varieties at different growth stages. The reason is likely that elevated atmospheric CO₂ concentration promotes rice growth and development, leading to a early maturity, then senescent leaf promotes soluble sugar’s decomposition, and speeds up transportation of soluble protein from function leaf to seeds.

Keyword: FACE; Rice; Late growth stage; Soluble sugar; Soluble protein; Chlorophyll; Plant productivity

Show Figures

工业革命以来, 大气中二氧化碳(CO₂)含量明显增高。根据世界气象组织(WMO)的年度报告, 至2012年底全球大气CO₂浓度已经增加到400.9 µmol mol^-1, 比工业革命前的280 µmol mol^-1增加了约42.86%。预计到21世纪中期将达450~550 µmol mol^-1, 21世纪末将升至730~1020 µmol mol^{-1 [ 1, 2, 3]}。作为作物光合作用的主要底物, CO₂浓度增高必然对作物生理代谢与产量品质产生深刻影响^{[ 4, 5]}。水稻属于C₃作物, 对CO₂浓度增加反应更为敏感^{[ 6]}。水稻又是我国及东南亚等人口大国的主要粮食作物, 所以中国、日本以及菲律宾等东南亚国家都把水稻对高CO₂浓度的响应研究作为气候变化对农业生产影响的研究重点。如南京土壤研究所利用江都(32°35′N, 119°42′E) FACE实验平台在亚热带对小麦-水稻轮作开展研究^{[ 7, 8, 9, 10, 11, 12]}, 日本东北农业试验站利用岩手(39°38´N, 140°57´E) FACE实验平台在温带开展了水稻生态系统研究^{[ 13, 14, 15]}。研究表明CO₂浓度增高促进水稻生育进程, 多数品种表现增产, 但增幅差异较大, 区域间(纬度间)的差异也广泛存在。北方水稻特别是东北粳稻历来以高产优质且口感佳著称, 近年来国内外需求量大, 在发展高产优质高效农业中具有特殊意义, 对保证居民的粮食供应和出口创汇也具有重要意义。本研究对北方水稻剑叶生育后期可溶性物质含量及植株生产力对CO₂浓度增高的响应趋势开展试验研究, 探索其响应规律, 为全面揭示高CO₂浓度对北方水稻产量生理影响规律及比较与南方水稻反应差异提供理论依据。

1 材料与方法

1.1 试验地概况与土壤类型

FACE实验平台位于北京市昌平南部中国农业科学院昌平实验基地(40°08´N, 116°08´E), 京昌公路的西侧。土壤类型属褐潮土, 耕作方式为冬小麦-水稻轮作, 土壤含有机质18.27 g kg^-1、全氮 10.38 g kg^-1、全磷1.14 g kg^-1、全钾14.17 g kg^-1、碱解氮为94.83 mg kg^-1、速效磷49.1 mg kg^-1、速效钾104.28 mg kg^-1, pH 8.28。

1.2 供试材料

选用粳稻品种松粳9号(典型特征为高产稳产, 平均产量10.5 t hm^-2)和稻花香2号(典型特征为优质, 全部指标达到农业部优质米一级标准)。松粳9号从出苗到成熟需活动积温2650℃, 株高95~100 cm, 穗长20 cm左右; 稻花香2号需≥10℃活动积温2800℃左右, 株高100~105 cm左右, 穗长21.6 cm左右。

1.3 试验设计

中国北方FACE实验平台的系统构成、系统控制等见郝兴宇等^{[ 16]}报道。该试验设2个处理, CO₂为主处理, 为试验地实测大气CO₂浓度(389±40 μmol mol^-1)和FACE圈增高浓度(600±60 μmol mol^-1), 误差为10%, 各设3次重复(即3个FACE圈, 3个对照圈); 副处理为覆膜, 与对照相比, 土壤温度约提高0.5~1.0℃, 含水量约提高3%~6%。采用盆栽(盆长 55 cm, 宽40 cm, 深30 cm), 每圈种植60盆, 两品种各30盆, 每盆栽6穴, 每穴精选苗4株, 每品种半数盆覆地膜。

2012年5月16日育苗, 6月15日移栽, 同时开始对FACE圈内通CO₂气体并覆膜处理至收获。对各盆施尿素4 g、磷酸二铵4 g、氯化钾2 g, 7月6日追加氯化钾3 g。FACE圈与对照圈管理方式相同; 覆膜与对照水分管理一致, 移栽前盆土浇透水, 水分管理与大田相同。

1.4 生育日期确定

参照国家水稻研究中心水稻生育期的划分标准, 结合试验实际情况, 植株8月9日开始进入孕穗期(幼穗在叶鞘内伸长至叶鞘膨胀), 8月16日进入抽穗期(穗顶端从叶鞘中露出, 多数穗开花), 8月30日进入乳熟期(籽粒内容物呈白色乳浆状, 背部仍为绿色), 9月16日进入蜡熟期(籽粒内容物浓黏, 无乳状物出现, 手压穗中部籽粒有坚硬感, 米粒背部绿色逐渐消失, 谷壳微黄), 9月28日为完熟期(谷壳变黄, 不易破碎, 达收获期), 10月2日全部收获。

1.5 试验测定项目及方法

根据生育进程, 从水稻抽穗期开始共取样5次, 间隔7 d, 取样时间在上午8:00至9:00之间。选取各处理主茎剑叶10片, 分别放于自封袋, 标记后迅速放入冰箱中冷藏, 待测可溶性糖、可溶性蛋白含量及总叶绿素含量。收获期收割测产区植株测植株生产力相关指标。

1.5.1 可溶性糖含量测定方法采用蒽酮法^{[ 17]}。用牛血清蛋白制作标准曲线。称取剪碎混匀的新鲜叶片0.5 g, 放入盛有25 mL蒸馏水的三角瓶, 沸水浴煮30 min, 取出冷却, 过滤入100 mL容量瓶, 热水冲洗残渣数次, 定容至刻度。提取样品液0.5 mL加等量水, 加蒽酮试剂, 摇匀后用分光光度计于620 nm下比色测得光密度值, 计算可溶性糖含量。

1.5.2 可溶性蛋白含量测定方法采用考马斯亮蓝G-250法^{[ 17]}。用分析纯葡萄糖制作标准曲线。称取剪碎混匀的新鲜叶片0.5 g, 用5 mL蒸馏水研磨成匀浆, 于3000转 min^-1离心10 min, 提取上清液1 mL, 加5 mL考马斯亮蓝G-250溶液, 摇匀后放置2 min, 用分光光度计在595 nm波长下测定吸光度, 计算可溶性蛋白含量。

1.5.3 总叶绿素含量测定方法采用丙酮乙醇混合液法^{[ 17]}(略有改进)。用电子天秤称取0.5 g叶片, 剪碎后放入盛有10 mL的1∶1的乙醇-丙酮混合浸提液的试管中, 室温下黑暗浸提48 h, 待材料完全变白后摇匀, 稀释10倍倒入干净比色皿, 用分光光度法分别测定其在663 nm、645 nm下的光密度值, 计算单位质量叶片总叶绿素含量。

1.5.4 植株生产力测定方法在成熟期, 从各处理分别取4盆计算和分析植株生产力相关指标的差异。从各处理调查4盆植株的地上部干物质总量即生物学产量; 取代表性植株5株, 105℃杀青30 min,

80℃烘72 h称重, 根据单株平均穗粒数、结实率、千粒重计算单株籽粒产量。取测产区4盆植株, 机械收获称重得到实收经济产量, 经济产量与生物产量的比值即经济系数。

1.6 统计分析方法

以SPSS统计软件进行数据处理和方差分析, 检验CO₂处理和覆膜间的差异显著性, 用Microsoft Excel绘制图表。

2 结果与分析

2.1 CO₂浓度增高及覆膜对水稻生育后期剑叶可溶性物质含量的影响

2.1.1 可溶性糖含量可溶性糖是植物碳素营养主要形式, 为作物提供能源物质, 并在维持植物蛋白质稳定方面起重要作用。可溶性糖含量的变化与水稻功能叶片光合作用能力密切相关^{[ 18]}。松粳9号和稻花香2号剑叶可溶性糖含量在生育后期都呈先升高后降低的趋势, 在孕穗期, 二者剑叶可溶性糖含量分别升高13.9%和14.1%, 乳熟期达最大值(表1)。覆膜分别使其平均升高14.0%和18.2%, 增幅差异均达极显著水平, 但FACE条件下覆膜处理却分别降低3.4%和0.4%。

表1 FACE及覆膜处理对水稻生育后期剑叶可溶性糖含量的影响 Table 1 Effects of FACE and mulching on soluble sugar content in rice flag leaf during late growth stage

从抽穗至完熟期整个阶段, FACE条件下松粳9号剑叶可溶性糖含量平均升高11.7%、11.0%、11.3%和11.2%, 而稻花香2号平均升高15.9%、7.3%、3.5%和47.5%, 除蜡熟期, 增幅均达显著或极显著水平。覆膜处理使松粳9号平均增加17.9%、9.1%、4.0%和0.1%, 抽穗期和乳熟期增幅达显著或极显著水平; 稻花香2号平均增加13.0%、5.5%、1.3%和8.3%, 抽穗期和完熟期增幅达极显著水平。松粳9号在孕穗和乳熟期, FACE处理与覆膜处理间分别存在极显著( P<0.01)和显著( P<0.05)交互效应; 稻花香2号在孕穗和抽穗期, 分别存在极显著( P<0.01)和显著( P<0.05)交互效应。

FACE处理对剑叶可溶性糖含量的影响存在品种间差异。在乳熟和蜡熟期, 松粳9号增幅大于稻花香2号, 而孕穗、抽穗和完熟期, 松粳9号增幅小于稻花香2号。覆膜条件下抽穗至蜡熟期, 松粳9号增幅均大于稻花香2号; 完熟期松粳9号增幅小于稻花香2号。

2.1.2 可溶性蛋白含量可溶性蛋白是植物氮素存在的主要形式, 其含量常作为衡量叶片衰老程度重要指标, 尤其在灌浆期。剑叶可溶性蛋白含量升高有利于维持剑叶生长, 延长光合功能期, 为籽粒碳、氮化合物积累奠定物质基础, 同时反映叶片生

理生化作用状态和叶片氮向籽粒转运状况^{[ 19]}。松粳9号和稻花香2号剑叶可溶性蛋白含量在生育后期均呈下降趋势(表2)。与对照相比, 松粳9号从孕穗至完熟期剑叶可溶性蛋白含量平均降低5.6%、10.6%、9.8%、9.9%和16.2%, 增幅均达显著或极显著水平; 稻花香2号平均降低7.2%、10.5%、4.4%、9.8%和8.6%, 除乳熟期外增幅均达显著或极显著水平。覆膜处理使松粳9号平均提高7.8%、18.9%、15.3%、36.0%和49.7%, 而稻花香2号平均提高13.6%、18.8%、18.9%、38.7%和16.6%。除蜡熟期外, 松粳9号在FACE与覆膜处理间存在极显著( P<0.01)或显著( P<0.05)交互效应, 除乳熟和蜡熟期外, 稻花香2号在两处理间存在极显著( P<0.01)或显著( P<0.05)交互效应。

FACE处理下两品种剑叶可溶性蛋白含量均降低, 孕穗期松粳9号降幅小于稻花香2号, 在乳熟期和完熟期, 松粳9号降幅大于稻花香2号。覆膜条件下均升高, 孕穗、乳熟和蜡熟期, 松粳9号降幅小于稻花香2号, 完熟期松粳9号增幅大于稻花香2号。

2.1.3 总叶绿素含量功能叶片叶绿素含量与光合作用密切相关, 结实期功能叶片叶绿素含量的变化反映叶片光合能力的变化和叶片衰老进程^{[ 20, 21]}, 生育后期保持剑叶较高的叶绿素含量为叶片吸收更多光能提供了良好生理条件^{[ 22]}, 为提高产量提供有力保障。松粳9号和稻花香2号剑叶总叶绿素含量在生育后期均呈降低的趋势(表3)。孕穗至乳熟期, FACE处理使松粳9号剑叶总叶绿素含量平均升高24.9%、18.9%和17.7%, 增幅均达显著或极显著水平; 稻花香2号平均升高21.8%、22.5%和20.8%, 除孕穗期外增幅均达极显著水平。覆膜使松粳9号平均降低8.1%、12.4%、15.8%, 除孕穗期增幅均达显著水平; 使稻花香2号平均降低16.1%、10.7%和14.4%, 除孕穗期外增幅均达显著或极显著水平。蜡熟期FACE处理使松粳9号和稻花香2号剑叶总叶绿素含量降低6.0%和12.0%。覆膜使二者降低5.6%和5.8%, 但FACE条件下覆膜使二者剑叶总叶绿素含量升高11.2%和25.0%。松粳9号在FACE与覆膜处理间不存在显著交互效应; 稻花香2号在抽穗期两处理间存在显著( P<0.05)交互效应, 乳熟期两处理间存在极显著( P<0.01)交互效应。

表2 FACE及覆膜处理对水稻生育后期剑叶可溶性蛋白含量的影响 Table 2 Effects of FACE and mulching on soluble protein content in rice flag leaf during late growth stage

孕穗至乳熟期, FACE处理使两品种剑叶的总叶绿素含量都明显提高, 但孕穗期松粳9号增幅大于稻花香2号, 抽穗至乳熟期松粳9号增幅小于稻花香2号。蜡熟期FACE处理松粳9号降幅小于稻花香2号, 覆膜却使松粳9号增幅小于稻花香2号。覆膜降低两者总叶绿素含量, 品种间差异不显著; 但FACE条件下覆膜却使松粳9号增幅小于稻花香2号。

表3 FACE及覆膜处理对水稻生育后期剑叶总叶绿素含量的影响 Table 3 Effects of FACE and mulching on chlorophyll content in rice flag leaf during late growth stage

2.2 FACE及覆膜对水稻植株生产力的影响

2.2.1 单株籽粒产量如图1所示, 松粳9号单株籽粒产量低于稻花香2号, 品种间差异达极显著( P<0.01)水平。FACE使松粳9号比对照平均增加6.82%, 达极显著( P<0.01)水平。而稻花香2号比对照平均增加2.56%, 未达显著( P<0.05)水平。覆膜使松粳9号和稻花香2号两品种单株籽粒产量都极显著( P<0.01)提高了, 分别提高11.03%和8.07%。统计分析表明,两品种FACE×FILM之互作效应对水稻单株籽粒产量的影响均未达显著( P<0.05)水平。

2.2.2 生物学产量松粳9号生物学产量高于稻花香2号, 品种间差异达极显著( P<0.01)水平(图2)。两品种FACE处理都极显著( P<0.01)高于对照, 松粳9号和稻花香2号分别比对照平均增加1.50%和2.13%。覆膜同样使两品种生物学产量极显著( P<0.01)提高, 松粳9号和稻花香2号分别提高1.51%和1.94%。统计分析表明, 两品种FACE×FILM互作效应对水稻生物学产量的影响均未达显著水平。说明水稻生物学产量品种间差异显著, 并为FACE和FILM处理显著提高, 且增幅表现为稻花香2号大于松粳9号。

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图1 FACE及覆膜处理对水稻单株籽粒产量的影响FACE: 开放式空气CO₂浓度增加; AMB: 正常大气CO₂浓度; FILM: 对土壤做覆地膜处理; N: 土壤正常状况; Songjing 9: 松粳9号; Daohuaxiang 2: 稻花香2号。Fig. 1 Effects of FACE and mulching on seed yield of individual plant in riceFACE: free air CO₂ enrichment; AMB: ambient CO₂; FILM: plastic-film mulching; N: natural soil.

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图2 FACE及覆膜处理对水稻生物学产量的影响FACE: 开放式空气CO₂浓度增加; AMB: 正常大气CO₂浓度; FILM: 对土壤做覆地膜处理; N: 土壤正常状况; Songjing 9: 松粳9号; Daohuaxiang 2: 稻花香2号。Fig. 2 Effects of FACE and mulching on total biomass of riceFACE: free air CO₂ enrichment; AMB: ambient CO₂; FILM: plastic-film mulching; N: natural soil.

2.2.3 经济系数松粳9号经济系数高于稻花香2号, 品种间差异极显著( P<0.01)水平(图3)。两品种FACE处理均高于对照, 松粳9号和稻花香2号分别比对照平均增加12.64%和26.05%, 均达极显著( P<0.01)水平。覆膜使松粳9号极显著( P<0.01)降低, 平均降幅为2.54%; 而稻花香2号比对照平均增加12.84%, 达极显著( P<0.01)水平。

统计分析表明, 松粳9号FACE×FILM的互作效应对两品种经济系数影响达极显著水平, 而稻花香2号仅达显著水平(图4)。说明不同品种水稻经济系数对FACE和FILM处理的响应存在很大差异, 两处理对松粳9号影响的差异更明显, 但总体而言, 稻花香2号变幅大于松粳9号。

图4

FACE: free air CO₂ enrichment; AMB: ambient CO₂; FILM: plastic-film mulching; N: natural soil.
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	图3 FACE及覆膜处理对水稻经济系数的影响 Fig. 3 Effects of FACE and mulching on economic coefficient of riceFACE: 开放式空气CO₂浓度增加; AMB: 正常大气CO₂浓度; FILM: 对土壤做覆地膜处理; N: 土壤正常状况; Songjing 9: 松粳9号; Daohuaxiang 2: 稻花香2号。

3 讨论

一般认为, 从水稻孕穗期开始, 剑叶中可溶性糖开始积累, 并不断向籽粒运输, 乳熟期加速向籽粒输送, 因此生育后期剑叶可溶性糖含量呈先升高后降低的趋势, 乳熟期最高^{[ 23]}。生育后期功能叶片可溶性蛋白是籽粒重要氮源, 籽粒充实阶段剑叶向籽粒源源不断地输氮化合物, 所以剑叶可溶性蛋白含量呈下降趋势^{[ 24]}。土壤水分含量增加使抗旱性品种剑叶可溶性糖含量增加幅度较大, CO₂浓度增加使多数品种孕穗期到完熟期剑叶可溶性蛋白含量下降^{[ 25, 26]}。

孕穗期是碳氮代谢旺盛的时期, 可溶性糖是光合作用暗反应主要产物。本研究显示高浓度CO₂促进可溶性糖的积累; 而覆膜增加土壤温度, 超过幼穗分化最适温度则不利于糖的形成, 且增加呼吸对糖的消耗。CO₂浓度升高和覆膜都可能加快生育进程, 导致功能叶早衰, 增加糖积累量。抽穗至完熟期是决定结实率和粒重的关键时期, 土壤温度及叶片寿命等都可能影响最终产量^{[ 26]}。本研究显示高浓度CO₂和覆膜可能导致功能叶早衰, 衰老分解产生部分可溶性糖, 另外覆膜增加土壤含水量, 有利于根吸收水分运输到叶片中, 增强叶片蒸腾, 同时气孔开放增加CO₂的摄入, 抑制呼吸对部分糖的分解。蜡熟期对松粳9号影响大于稻花香2号, 原因可能是抗旱性品种剑叶可溶性糖含量水分含量增加时增加幅度较大。

呼吸代谢与氨基酸和蛋白质的生物合成、转化密切相关, 早在19世纪就有人发现呼吸作用随CO₂浓度升高而下降, 以后不少研究都证实了这一点^{[ 27]}。本研究显示高浓度CO₂和覆膜处理条件下, 水稻生育后期可溶性蛋白含量变化表现出一定规律性, 高浓度CO₂使剑叶中可溶性蛋白含量下降, 原因可能是环境中高浓度CO₂使叶片早衰, 加快功能叶片合成的蛋白向籽粒中运输, 且CO₂浓度升高有加快生育进程的可能, 导致叶片中蛋白提前运输到籽粒中。而覆膜却使两品种整个生育后期剑叶中可溶性蛋白含量都提高, 覆膜增加的土壤水分在一定程度上促进根系吸收更多水分, 延迟叶片衰老, 降低可溶性蛋白质的分解速率。有研究表明, 水分含量较高情况下, 植物体内正常的蛋白质合成受到抑制, 会被诱导出新蛋白, 使蛋白质的含量增加^{[ 24]}。本研究表明孕穗期对松粳9号的影响小于稻花香2号, 完熟期对松粳9号的影响大于稻花香2号, 说明不同时期两品种对高浓度CO₂和覆膜的响应程度有差异。

从孕穗期开始, 叶片开始进入功能衰退期, 叶绿素加速分解。本研究显示大气CO₂浓度增高使孕穗期至乳熟期剑叶叶绿素含量高于对照, 原因可能是CO₂浓度增加条件下, 植物充分利用环境资源, 增加对CO₂的同化作用, 需要通过增加叶片叶绿素的含量来提高对光能的捕获能力, 以满足碳同化时能量的需求; 同时CO₂浓度升高一定程度上加快生育进程, 减少叶绿素的分解。CO₂浓度升高使蜡熟期剑叶总叶绿素含量下降, 可能因为叶片早衰加快叶绿素分解速率, CO₂浓度升高条件下覆膜可提高剑叶总叶绿素含量, 抑制早衰。此外, 覆膜使土壤含水量增加, 叶片加速变黄, 影响叶绿素的合成, 与前人研究结果一致^{[ 28]}; 如果覆膜和高浓度CO₂同时作用则改善了这种状况, 提高剑叶总叶绿素含量, 有助于植物捕获更多光能供给光合作用。叶片失绿被认为是叶片衰老的最主要和最初症状, 本研究发现存在品种间差异, 松粳9号的抗性大于稻花香2号。蜡熟期至完熟期, 覆膜条件下CO₂浓度升高使两品种剑叶总叶绿素含量都升高了, 说明此时期CO₂影响更大。

水稻产量是反应植株生产力重要参数, 关于CO₂浓度增加对水稻产量影响的报道较多, 但研究条件各异, 多数结果表明, FACE使大部分品种产量极显著提高^{[ 13]}。生物学产量是作物茎叶光合产物贮存累积的结果, 体现该品种的总生产力。日本关于FACE对水稻生物学产量的影响有研究显示平均增幅为12%, FACE使武香粳14的生物学产量平均比对照增产16.2%^{[ 29, 30, 31]}, 经济系数比对照减少2.7%^{[ 16]}。有关CO₂浓度增加(FACE)对水稻产量及其构成因素影响的研究^{[ 13, 17, 19]}表明多数品种有增产趋势。而本研究显示高浓度CO₂使两水稻品种单株籽粒产量和总生物量大幅度提高, FACE处理和覆膜对两品种经济系数的影响存在互作效应, 说明FACE比对照显著增产的重要原因, 经济系数对FACE水稻产量的提高可能有明显影响。

总之, 大气CO₂浓度增高促进水稻生长发育, 抽穗后加速可溶性糖及叶绿素的积累, 孕穗开始促进可溶性蛋白向籽粒中运输, 且品种间响应程度在不同生育期有差异; 覆膜延长水稻功能叶片光合功能期, 促进可溶性糖和蛋白积累, 但加快生育后期叶绿素降解。在生育后期, 覆膜可一定程度抑制CO₂浓度增高对叶片早衰的影响。CO₂浓度增高和覆膜不同程度提高水稻品种植株生产力。由于FACE系统实验地点条件所限, 本试验对北方优质高产水稻的研究并非在北方水稻主产区, 以后的研究中将逐步加以改进。

4 结论

本研究发现, FACE和覆膜处理均显著提高两个品种抽穗至乳熟期剑叶的可溶性糖含量, FACE处理下松粳9号增幅小于稻花香2号, 覆膜条件下松粳9号增幅大于稻花香2号。FACE使松粳9号剑叶在整个生育后期的可溶性蛋白含量明显降低, 而稻花香2号除乳熟期外均显著增加。覆膜使两品种整个生育后期剑叶可溶性蛋白含量显著升高, 但孕穗期松粳9号变幅小于稻花香2号, 完熟期松粳9号变幅大于稻花香2号。孕穗至乳熟期, FACE使两品种剑叶总叶绿素含量显著升高, 两品种变幅各阶段有差异。FACE和覆膜都不同程度提高水稻单株籽粒产量, 稻花香2号增幅小于松粳9号, 总生物量稻花香2号增幅大于松粳9号; FACE和覆膜对水稻经济系数的影响有差异, 但总体上稻花香2号变幅大于松粳9号。

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.

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16	郝兴宇, 林而达, 杨锦忠, 居辉, 马占云, 韩雪, 王贺然, 杨万深. 自由大气CO₂浓度升高对夏大豆生长与产量的影响. 生态学报, 2009, 29: 4595-4603 Hao X Y, Lin E D, Yang J Z, Ju H, Ma Z Y, Han X, Wang H R, Yang W S. Effects of free air CO₂ enrichment (FACE) on growth and yield of summer soybean. Acta Ecol Sin, 2009, 29: 4595-4603 (in Chinese with English abstract) [本文引用:2]
17	张志良, 瞿伟菁. 植物生理学实验指导(第3版). 北京: 高等教育出版社, 2005. pp67-159 Zhang Z L, Zhai W J. The Guidance of Plant Physiology Experiments, 3rd edn. Beijing: Higher Education Press, 2005. pp67-159(in Chinese) [本文引用:4]
18	谢立勇, 林而达, 孙芳, 赵海燕. 全生育期二氧化碳与温度处理对水稻生理性状的影响初报. 中国农业大学学报, 2006, 11(1): 17-21 Xie L Y, Lin E D, Sun F, Zhao H Y. Effects of CO₂ enrichment and temperature increase during growth duration on physiological characteristics of rice. J China Agric Univ, 2006, 11(1): 17-21 (in Chinese with English abstract) [本文引用:1]
19	胡健, 王余龙, 杨连新, 周娟, 朱建国. FACE对武香粳14 各生育期功能叶片及根系可溶蛋白含量的影响. 农业环境科学学报, 2006, 25: 1117-1121 Hu J, Wang Y L, Yang L X, Zhou J, Zhu J G. Effect of free-air CO₂ enrichment (FACE) on concentrations of soluble protein in functional leaves and roots of japonica rice (Oryza sativa L. ) cultivar Wuxiangjing 14. J Agro-Environ Sci, 2006, 25: 1117-1121 (in Chinese with English abstract) [本文引用:2] [CJCR: 1.178]
20	刘贞琦, 刘振业, 马达鹏, 曾淑芬. 水稻叶绿素含量及其与光合速率关系的研究. 作物学报, 1984, 10: 57-61 Liu Z Q, Liu Z Y, Ma D P, Zeng S F. A study on the relation between chlorophyll content and photosynthetic rate of rice. Acta Agron Sin, 1984, 10: 57-61 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.667]
21	金松恒, 蒋德安, 王品美, 赵凯, 翁晓燕. 水稻孕穗期不同叶位叶片的气体交换与叶绿素荧光特性. 中国水稻科学, 2004, 18: 443-448 Jin S H, Jiang D A, Wang P M, Zhao K, Weng X Y. Characteristics of gas exchange and chlorophyll fluorescence in different position leaves at booting stage in rice plants. Chin J Rice Sci, 2004, 18: 443-448 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.494]
22	童汉华, 梅捍卫, 邢永忠, 曹一平, 余新桥, 章善庆, 罗利军. 水稻生育后期剑叶形态和生理特性的QTL定位. 中国水稻科学, 2007, 21: 493-499 Tong H H, Mei H W, Xing Y Z, Cao Y P, Yu X Q, Zhang S Q, Luo L J. QTL analysis for morphological and physiological characteristics of flag leaf at the late developmental stage in rice. Chin J Rice Sci, 2007, 21: 493-499 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.494]
23	Imai K, Mae T, Suzuki Y, Makino A, Mae T. Effects of nitrogen nutrition on the relationships between the levels of rbcS and rbcL mRNAs and the amount of ribulose 1, 5-bisphosphate carboxylase/oxygenase synthesized in the eighth leaves of rice from emergence through senescence. Plant Cell Environ, 2005, 28: 1589-1600 [本文引用:1] [JCR: 5.135]
24	胡健, 杨连新, 周娟, 王余龙, 朱建国. 开放式空气CO₂浓度增高和施氮量对水稻结实期叶片内肽酶活力的影响. 中国水稻科学, 2008, 22: 155-160 Hu J, Yang L X, Zhou J, Wang Y L, Zhu J G. Effect of free air CO₂ enrichment (FACE) and nitrogen level on endopeptidase activities in rice leaves during grain filling stage. Chin J Rice Sci, 2008, 22: 155-160 (in Chinese with English abstract) [本文引用:2] [CJCR: 1.494]
25	王贺正, 马均, 李旭毅, 李艳, 张荣萍, 汪仁全. 水分胁迫对水稻结实期一些生理性状的影响. 作物学报, 2006, 32: 1892-1897 Wang H Z, Ma J, Li X Y, Li Y, Zhang R P, Wang R Q. Effects of water stress on some physiological characteristics in rice during grain filling stage. Acta Agron Sin, 2006, 32: 1892-1897 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.667]
26	彭长连, 林植芳, 林桂珠. 加富CO₂条件下水稻叶片抗氧化能力的变化. 作物学报, 1999, 25: 39-43 Peng C L, Lin Z F, Lin G Z. Changes of antioxidative ability in leaves of rice cultivars grown under enriched CO₂. Acta Agron Sin, 1999, 25: 39-43 (in Chinese with English abstract) [本文引用:2] [CJCR: 1.667]
27	彭晓邦, 张硕新. 大气CO₂浓度升高对植物某些生理过程影响的研究进展. 西北林学院学报, 2006, 21(1): 68-71 Peng X B, Zhang S X. Research progress in effects of increased atmospheric CO₂ concentration on certain physiological process of plants. J Northwest For Univ, 2006, 21(1): 68-71 (in Chinese with English abstract) [本文引用:1]
28	刘少华, 陈国祥, 胡艳, 吕川根, 吴国荣, 杨艳华. 高产杂交稻“两优培九”功能叶抗氧化系统对水分胁迫的响应. 作物学报, 2004, 30: 1244-1249 Liu S H, Chen G X, Hu Y, Lü C G, Wu G R, Yang Y H. Effects of water stress on the antioxidant system in mature leaves of hybrid rice “Liangyoupeijiu”. Acta Agron Sin, 2004, 30: 1244-1249 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.667]
29	Kimball B A, 朱建国, 程磊, Kobayashi K, Bindi M. 开放系统中农作物对空气CO₂浓度增加的响应. 应用生态学报, 2002, 13: 1323-1338 Kimball B A, Zhu J G, Cheng L, Kobayashi K, Bindi M. Responses of agricultural crops to free air CO₂ enrichment. Chin J Appl Ecol, 2002, 13: 1323-1338 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.742]
30	Kim H Y, Lieffering M, Kobayashi K, Okada M, Miura S. Seasonal changes in the effects of elevated CO₂ on rice at three levels of nitrogen supply: a free air CO₂ enrichment (FACE) experiment. Global Change Biol, 2003, 9: 826-837 [本文引用:1] [JCR: 6.91]
31	黄建晔, 董桂春, 杨洪建, 王余龙, 朱建国, 杨连新, 单玉华. 开放式空气CO₂ 增高对水稻物质生产与分配的影响. 应用生态学报, 2003, 14: 253-257 Huang J Y, Dong G C, Yang H J, Wang Y L, Zhu J G, Yang L X, Shan Y H. Effects of free-air CO₂ enrichment on biomass accumulation and distribution in rice. Chin J Appl Ecol, 2003, 14: 253-257 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.742]

2011

2.578

0.0

... mol mol^{-1 [1,2,3]} ...

2011

0.0

Acta Ecol Sin. 2011, 31:277-

The impact of atmospheric CO₂ concentration enrichment on rice quality#cod#x02014;a research review

Wang

Y X

,Michael

,Song

Q L

,Yang

L X

.The impact of atmospheric CO₂ concentration enrichment on rice quality#cod#x02014;a research review.Acta Ecol Sin,2011,31:277-282

... mol mol^{-1 [1,2,3]} ...

2011

2.474

0.0

... mol mol^{-1 [1,2,3]} ...

2007

0.0

... 作为作物光合作用的主要底物, CO₂浓度增高必然对作物生理代谢与产量品质产生深刻影响^[4,5] ...

2010

0.0

0.795

Chin J Eco-Agric. 2010, 18(5):1116 - 1124

Effect of elevated CO₂ concentration and increased temperature on physiology and production of crops

大气[CO₂]和温度升高对农作物生理及生产的影响

(房世波),(沈斌),(谈凯炎),(高西宁).(大气CO_{2)和温度升高对农作物生理及生产的影响}.(中国生态农业学报),2010,18:1116-1124

房世波, 沈斌, 谭凯炎, 高西宁

Increasing concentration of global atmospheric CO₂ and temperature are the two most important factors in the national list of the impact of climate change on crop yield. This brief summary focused on research advances on the impact of increasing [CO₂] and temperature on crop photosynthesis, respiration, transpiration, yield, quality, etc. The review shows that increasing [CO₂] increases the rates of photosynthesis and transpiration, but decreases respiration rate and stomatal conductance. Though crop quality decreases with increasing yield, there is no general research consensus on this trend. C3 and C4 crops’ response to variations in CO₂ concentration is inconsistent, with short- and long-term effects on photosynthetic pathways. It is generally conceived that atmospheric temperature increase inhibits crop photosynthesis, reducing crop yield. Most of the existing reports on the response of crop yield to climate change focus on crop modeling or simulation experiment. Research also shows discrepant model/simulation results. There are also discrepant yield assessment results from different scholars. Some recent studies suggest that rising temperatures have a nonlinear effect on crop yield. Crop yield declines rapidly when temperatures exceed threshold values. As most of the simulation experiments are conducted in gas chambers (which are quite different from real-world conditions), their conclusions require further validation. The crop model results generally lack verifications by experiment.

全球大气[CO₂]和气温升高是全球气候变化对农作物产量影响最为重要的两个因子。本文着重介绍了[CO₂]和温度升高对农作物光合作用、呼吸作用、蒸腾作用、产量、品质等方面影响的研究进展。研究表明随着[CO₂]升高, 作物光合速率及蒸腾速率有上升趋势, 呼吸作用和气孔导度下降, 产量有所提高, 品质将会降低, 但研究仍有不确定性。随着[CO₂]变化, 不同光合途径(C3、C4)作物的响应不一致且存在短期和长期效应。普遍认为大气温度升高抑制作物光合作用, 作物产量下降。现有的研究多采用模型或模拟试验的方法研究气候变化对作物产量的影响, 但研究发现模型研究结果与模拟试验研究结果有差异, 不同学者对产量的评估结果也不一致。最新研究认为温度对作物产量影响成非线性, 当温度高于关键温度后产量会迅速下降。现阶段大部分模拟试验均在气室中研究, 与野外实际情况差异较大, 结论仍需进一步验证。目前尚缺乏对作物模型结果的实验验证。

... 作为作物光合作用的主要底物, CO₂浓度增高必然对作物生理代谢与产量品质产生深刻影响^[4,5] ...

2010

0.0

... 水稻属于C₃作物, 对CO₂浓度增加反应更为敏感^[6] ...

2004

0.0

1.889

Sci Agric Sin. 2004, 37(12):1824 - 1830

(开放式空气CO_{2)浓度增加(FACE)对水稻产量形成的影响及其与氮的互作效应}

FACE对水稻产量形成的影响及其原因分析

黄建晔,王余龙

扬州大学水稻研究所

... E) FACE实验平台在亚热带对小麦-水稻轮作开展研究^{[7,8,9,10,11,12]}, 日本东北农业试验站利用岩手(39#cod#x000b0 ...

2009

0.0

... E) FACE实验平台在亚热带对小麦-水稻轮作开展研究^{[7,8,9,10,11,12]}, 日本东北农业试验站利用岩手(39#cod#x000b0 ...

2007

0.0

1.667

Acta Agron Sin. 2007, 33(3):433 - 440

Response of Plant Height to Free Air CO₂ Enrichment in Rice (Oryza sativa L.)

水稻株高性状对大气CO₂浓度升高的响应

FAN Gui-Zhi^1,4;CAI Qing-Sheng^1,*;WANG Chun-Ming²;WAN Jian-Min²;ZHU Jian-Guo³

范桂枝^1,4;蔡庆生^1,*;王春明²;万建民²;朱建国³

¹ College of Life Sciences;² State Key Laboratory of Crop Genetics and Germplasm Enhancement, Nanjing Agricultural University, Nanjing 210095, Jiangsu;³ Institute of Soil Science, Chinese Academy of Sciences, Nanjing 210008, Jiangsu;⁴ College of Life Sciences, Northeast Forestry University, Harbin, 150040, Heilongjiang, China

Free air carbon dioxide enrichment (FACE) technique is capable of providing a means by which the environment around growing plants may be modified to realistically simulate future concentrations of atmospheric CO2. The plant growth and development, as well as its height, biomass, and yields can be accelerated and enhanced under FACE. Identifying genomic regions influencing the response of plant height and its components to elevated CO₂ will be useful for us to understand the genetic response to changed CO₂ environment and select materials or cultivars adapting to future elevated CO2 atmospheric environment in plant height characteristics. A mapping population of 65 indica (IR24) chromosome segment substitution lines (CSSLs) with japonica (Asominori) background was used to detect quantitative trait locus (QTL) for plant height and its components containing panicle length and various internode lengths under ambient atmosphere CO₂ concentration (Ambient, about 370 µmol CO2 mol^-1) and FACE (exceed ambient CO₂ concentration about 200 µmol mol^-1). The main results indicated that plant height and its components of Asominori and IR24 under FACE were significantly different from these of Ambient at 1% probability level. Both positive and negative responses to elevated CO₂ in plant height and its components of CSSLs were observed under FACE. Among them, eight CSSLs largely responded to elevated CO₂, they were AI63 (increased 14.2 cm) and AI44 (reduced 4.54 cm) in plant height, AI9 (increased 3.56 cm) and AI12 (reduced 2.39 cm) in panicle length, AI39 (increased 15.74 cm) and AI27 (reduced 1.49 cm) in the first internode, AI32 (increased 8.09 cm) and AI53 (reduced 3.00 cm) in the second internode. Fourteen QTLs and 15 QTLs were detected under FACE and Ambient, respectively. Among them, 5 QTLs, named qPH6^-4, qPH8^-4, qPL8^-4, qPL12^-4, and qLFN6^-4, were simultaneously detected under FACE and Ambient, the others were detected only under one CO₂ level. Three QTLs, named qPH6^-4QE, qPH8^-4QE and qLSN5^-4QE were found to have gene x environment interaction effects significantly. It is suggest that some new QTLs for plant height and its components can be detected under elevated CO₂ condition, the QTLs being detected under unitary CO₂condition, natural ambient atmosphere or FACE are sensitive to CO₂ concentration, plant height is controlled by the interaction effect between genotype and CO₂ concentration.

以粳稻品种Asominori与籼稻品种IR24的杂交组合所衍生的染色体片段置换系（CSSLs）为材料，田间试验分别在FACE（CO₂浓度约570 µmol mol^-1）和对照（CO₂浓度约370 µmol mol^-1）下，对水稻株高性状的数量性状位点（QTL）进行了分析。结果表明，Asominori和IR24的株高、穗长、上位第一节间长和上位第二节间长在FACE和对照下的差异达显著水平；供试株系的4个株高性状对CO₂浓度升高都呈正负两种响应，其变化最大的株系为AI7和AI44（株高分别增加14.2 cm和降低4.54 cm），AI9和AI12（穗长分别增加3.56 cm和降低2.39 cm），AI39和AI27（上位第一节间长分别增加15.74 cm和降低1.49 cm），AI32和AI53（上位第二节间长分别增加8.09 cm和降低3.00 cm）；FACE和对照下分别检测出14和15个QTL，分布在除第2、7、9和第10号染色体外的各染色体上，其中5个（qPH6^-4、qPH8^-4、qPL8^-4、qPL12^-4和qLFN6^-4）在FACE和对照条件下同时检测到，分布在第6、8和第12染色体上，而其余的只在FACE或对照下检测到。这29个QTLs中，3个（qPH6^-4QE、qPH8^-4QE和qLSN5^-4QE）具显著的基因型与环境互作。在不同的CO₂环境下，测试性状发生不同程度的表型变异。结果推论，对CO₂浓度增加敏感的QTL位点，可能受到CO₂浓度增加的诱导，可见控制水稻株高性状的QTL与CO₂增加的环境发生了互作效应。

... E) FACE实验平台在亚热带对小麦-水稻轮作开展研究^{[7,8,9,10,11,12]}, 日本东北农业试验站利用岩手(39#cod#x000b0 ...

2002

0.0

1.742

... E) FACE实验平台在亚热带对小麦-水稻轮作开展研究^{[7,8,9,10,11,12]}, 日本东北农业试验站利用岩手(39#cod#x000b0 ...

2005

0.0

1.667

Acta Agron Sin. 2005, 31(9):1221 - 1226

Effects of Free-air CO2 Enrichment on Root System and Yield in Rice (Oryza sativa L.)

FACE对水稻根系及产量的影响

YANG Hong-Jian; YANG Lian-Xin; LIU Hong-Jiang; HUANG Jian-Ye; DONG Gui-Chun; ZHU Jian-Guo; WANG Yu-Long

杨洪建; 杨连新; 刘红江; 黄建晔; 董桂春; 朱建国; 王余龙

Key Laboratory of Crop Cultivation & Physiology, Jiangsu Province, Yangzhou University, Yangzhou 225009, Jiangsu

Increasing atmospheric CO2 concentrations are generally expected to enhance photosynthesis and growth and as a result substantially increase yields in plants. However, little is known about the effect of elevated CO2 on root growth of rice. The experiment conducted at Anzhen town, Jiangsu, China, to

2002－2003年利用我国惟一的农田开放式空气CO2浓度增高(FACE)研究平台，以武香粳14为供试品种，设施Ｎ量150 kg·hm-2(LN)、250 kg·hm-2 (NN)、350 kg·hm-2 (HN) 3个处理，研究了大气CO2浓度比对照高200 μmol·mol-1的FACE处理对水稻根系形态性状、根系活性和产量的影响。结果表明，(1) FACE使LN、NN、HN处理水稻抽穗期的

... E) FACE实验平台在亚热带对小麦-水稻轮作开展研究^{[7,8,9,10,11,12]}, 日本东北农业试验站利用岩手(39#cod#x000b0 ...

2006

0.0

1.742

Chin J Appl Ecol. 2006, 17(7):1331 - 1337

Responses of rice growth and development to free-air CO₂ enrichment (FACE): A research review

开放式空气CO₂浓度增高对水稻生长发育影响的研究进展

YANG Lianxin^{1, 2}; WANG Yulong¹; HUANG Jianye¹; YANG Hongjian¹; LIU Hongjiang¹

杨连新^1，2;王余龙¹;黄建晔¹;杨洪建¹;刘红江¹

¹Key Laboratory of Crop Genetics & Physiology of Jiangsu Province, Yangzhou University, Yangzhou 225009, China;²Institute of Soil Science，Chinese Academy of Sciences, Nanjing 210008, China

The ongoing increasing of atmospheric CO₂ has become a hot topic in the last two decades due to its profound impact on crop growth and development.Oryza sativaL. (rice) is one of the most important crops in the world and the first food in China. This paper reviewed the results based on
chamberbased experiments, mainly focused on the effects of free-air carbon dioxide enrichment (FACE) on the photosynthesis, water relationship, growth stage, leaf and root growth, dry matter production and distribution, chemical composition, and grain yield and quality of rice crop. The further research directions in this field were discussed.

地球大气中CO₂浓度不断升高已是不争的事实.CO₂浓度升高势必对植物的生长发育过程产生深刻的影响.水稻是世界上最重要的作物之一，也是中国第一大作物.结合气室条件下的研究结果，从光合作用、水分关系、生育期、叶片和根系生长、物质生产与分配、化学组分以及产量和品质等方面，重点收集和整理了开放式空气中CO₂浓度增高 (FACE) 对水稻生长发育影响的研究进展，并讨论了该领域有待深入研究的方向.

... E) FACE实验平台在亚热带对小麦-水稻轮作开展研究^{[7,8,9,10,11,12]}, 日本东北农业试验站利用岩手(39#cod#x000b0 ...

2003

2.474

0.0

... E) FACE实验平台在温带开展了水稻生态系统研究^[13,14,15] ...

... 水稻产量是反应植株生产力重要参数, 关于CO₂浓度增加对水稻产量影响的报道较多, 但研究条件各异, 多数结果表明, FACE使大部分品种产量极显著提高^[13] ...

... 有关CO₂浓度增加(FACE)对水稻产量及其构成因素影响的研究^[13,17,19]表明多数品种有增产趋势 ...

2009

5.242

0.0

... E) FACE实验平台在温带开展了水稻生态系统研究^[13,14,15] ...

2001

6.736

0.0

... E) FACE实验平台在温带开展了水稻生态系统研究^[13,14,15] ...

2009

0.0

... 3 试验设计中国北方FACE实验平台的系统构成、系统控制等见郝兴宇等^[16]报道 ...

... 7%^[16] ...

2005

0.0

... 1 可溶性糖含量测定方法采用蒽酮法^[17] ...

... 2 可溶性蛋白含量测定方法采用考马斯亮蓝G-250法^[17] ...

... 3 总叶绿素含量测定方法采用丙酮乙醇混合液法^[17](略有改进) ...

... 有关CO₂浓度增加(FACE)对水稻产量及其构成因素影响的研究^[13,17,19]表明多数品种有增产趋势 ...

2006

0.0

... 可溶性糖含量的变化与水稻功能叶片光合作用能力密切相关^[18] ...

2006

0.0

1.178

... 理生化作用状态和叶片氮向籽粒转运状况^[19] ...

... 有关CO₂浓度增加(FACE)对水稻产量及其构成因素影响的研究^[13,17,19]表明多数品种有增产趋势 ...

1984

0.0

1.667

Acta Agron Sin. 1984, 10(1):57 - 62

A STUDY ON THE RELATION BETWEEN CHLOROPHYLL CONTENT AND PHOTOSYNTHETIC RATE OF RICE

水稻叶绿素含量及其与光合速率关系的研究

刘贞琦;

Guizhou College of Agriculture

Experiments on rice were conducted for 4 ycars.Differences of chl-orophyll(a+b)content and a/b ratio among varieties were significantwhen grown under the same condition.The changes in chlorophyll(a+b)content and a/b ratio resulted from the changes in chlorophyll a and/orchlorophyll b.The chlorophyll

水稻不同品种的叶绿素(a+b)含量及叶绿素 a/b 比值是不同的。叶绿素(a+b)含量生育初期较少,然后逐渐增多,到生育后期则下降。水稻各品种由苗期至乳熟期叶绿素(a+b)含量的变异系数为12～20%,而叶绿素 a/b比值的变异系数只有4～8%。剑叶的叶绿素 a/b 比值最大,越是下位叶越低。叶绿素(a+b)含量与净光合速率呈显著的正相关,其相

... 3 总叶绿素含量功能叶片叶绿素含量与光合作用密切相关, 结实期功能叶片叶绿素含量的变化反映叶片光合能力的变化和叶片衰老进程^[20,21], 生育后期保持剑叶较高的叶绿素含量为叶片吸收更多光能提供了良好生理条件^[22], 为提高产量提供有力保障 ...

2004

0.0

1.494

Chin J Rice Sci. 2004, 18(5):443 - 448

Characteristics of Gas Exchange and Chlorophyll Fluorescence in Different Position Leaves at Booting Stage in Rice Plants

水稻孕穗期不同叶位叶片的气体交换与叶绿素荧光特性

JIN Song-heng; JIANG De-an *; WANG Pin-mei; ZHAO Kai; WENG Xiao-yan

金松恒; 蒋德安*; 王品美; 赵凯; 翁晓燕

State Key Laboratory of Plant Physiology and Biochemistry; College of Life Sciences; Zhejiang University; Hangzhou 310029; China

To investigate the senescence among different position leaves at booting stage in rice plant, gas exchange and chlorophyll fluorescence properties for different leaf positions of six rice cultivars were examined. The net photosynthetic rate (P_n), stomatal conductance (g), transpiration rate (E) and water use efficiency (WUE) decreased significantly with lowering of leaf positions, while intercellular CO₂ concentration (C_i) had little change, which indicated that the decrease of photosynthetic rate did not result from the decrease ofg. The decrease of SPAD reading which had a close correlation with chlorophyll content was one of the apparent reasons resulted in decease ofP_n. Further evidence by chlorophyll fluorescence showed that the photochemical efficiency of PSⅡ and its electron transport rate (ETR) decreased substantially with lowering of the leaf positions but the variable-to-maximum fluorescence ratio (F_v/F_m) and efficiency of energy conversion of open PSⅡ (F′_v/F′_m) maintained rather stable, suggesting that primary limitation ofP_n was suppressed by electron transport of PSⅠ, and probably associated with carbon assimilation.

研究了6个水稻品种孕穗期不同叶位的气体交换和叶绿素荧光特性，结果表明净光合速率([i]P[/i][sub]n[/sub]）、气孔导度（[i]g[/i]）、蒸腾速率（[i]E[/i]）和水分利用效率（[i]WUE[/i]）均随叶位降低而明显下降；而胞间CO[sub]2[/sub]浓度（[i]C[/i][sub]i[/sub]）基本维持不变，说明叶片衰老时[i]P[/i][sub]n[/sub]的下降主要不是气孔因素引起的。叶绿素含量降低与[i]P[/i][sub]n[/sub]下降的表现一致，叶绿素荧光分析表明[i]F[/i][sub]v[/sub]/[i]F[/i][sub]m[/sub]和[i]F[/i]'[sub]v[/sub]/[i]F[/i]'[sub]m[/sub]没有随叶位发生变化，而PSⅡ的光化学效率[i]Ф[/i][sub]PSⅡ[/sub]和电子传递速率([i]ETR[/i]）随叶位降低明显下降，这表明可能是PSⅠ电子传递受阻导致CO[sub]2[/sub]同化下降。

2007

0.0

1.494

Chin J Rice Sci. 2007, 21(5):493 - 499

QTL Analysis for Morphological and Physiological Characteristics of Flag Leaf at the Late Developmental Stage in Rice

水稻生育后期剑叶形态和生理特性的QTL定位

TONG Han-hua , MEI Han-wei , XING Yong-zhong , CAO Yi-ping , YU Xin-qiao , ZHANG Shan-qing , LUO Li-jun

童汉华1,2,3 ;梅捍卫2;邢永忠3;曹一平1; 余新桥2 ;章善庆1,*; 罗利军2,3,*

A recombinant inbred line(RIL) population derived from a cross between two elite indica varieties,Zhenshan 97 and HR5,was used to investigate the morphological and physiological characteristics of the functional leaves(flag leaves) at the late developmental stage in rice.A total of 26 QTLs referring the flag leaf characteristics,including morphological characteristics(length,width,and the ratio of length to width),metabolism characteristics(fresh weight,dry weight,water content),and chlorophyll content(chlorophyll a,chlorophyll b,and total chlorophyll contents) were detected,and the phenotypic variation of each QTL ranged from 4.77%-31.67%.Among them,the QTLs for the flag leaf morphology were mainly located on chromosomes 1 and 4,and the QTLs underlying the flag leaf metabolite were on chromosomes 3,7 and 10,whereas those affecting the chlorophyll contents of flag leaf were on chromosomes 3,4 and 9.Interestingly,a main QTL at marker interval RM321-MR6444 on chromosome 9 was identified, which would be beneficial for breeding late-stage vigor super-high-yielding rice.

为了探讨水稻生育后期功能叶尤其是剑叶的形态性状及生理特性对水稻高产的影响，利用来源于籼稻珍汕97/HR5后代的重组自交系群体，对水稻生长后期剑叶形态（长、宽、长宽比）、剑叶代谢产物（鲜质量、干质量、含水量）和剑叶叶绿素含量（叶绿素a、叶绿素b、总叶绿素含量）进行了考查分析和QTL定位。共检测到26个QTL，单个QTL的表型贡献率为4.77%~31.67%。其中控制剑叶形态的QTL主要位于第1和第4染色体，控制剑叶代谢产物的QTL主要位于第3、7和10染色体，控制剑叶叶绿素含量的QTL主要位于第3、4和9染色体上。其中位于第9染色体上的QTL，在后期功能型超级稻育种上具有一定的应用价值。

2005

5.135

0.0

... 3 讨论一般认为, 从水稻孕穗期开始, 剑叶中可溶性糖开始积累, 并不断向籽粒运输, 乳熟期加速向籽粒输送, 因此生育后期剑叶可溶性糖含量呈先升高后降低的趋势, 乳熟期最高^[23] ...

2008

0.0

1.494

Chin J Rice Sci. 2008, 22(2):155 - 160

Effect of Free Air CO₂ Enrichment (FACE) and Nitrogen Level on Endopeptidase Activities in Rice Leaves During Grain Filling Stage

开放式空气CO₂浓度增高和施氮量对水稻结实期叶片内肽酶活力的影响

HU Jian , YANG Lian-xin , ZHOU Juan , WANG Yu-long , ZHU Jian-guo

胡健1,2; 杨连新1;周娟1;王余龙1,*; 朱建国3

A japonica rice variety Wuxiangjing 14 was grown under ambient(AMB) or elevated CO₂(FACE,AMB + 200 μmol/mol) conitions and supplied with three levels of nitrogen: low level(LN,15 g/m²),normal level(NN,25 g /m²,similar to the level recommended to local farmers) and high level(HN,35 g/m²) to investigate the effect of elevated CO₂ and N supply on endopeptidase(EP) activity and the specific activity in the flag and the second leaves from the top of rice.As compared with AMB,FACE significantly increased EP activity in the flag and the second leaves at 10 days after heading(DAH) and 20 DAH,as well as EP activity in the second leaves at heading.However,EP activity in the flag leaves at maturity was greatly reduced under FACE.FACE obviously increased the specific activity in the flag and the second leaves at 10,20 DAH and maturity,together with the specific activity in the second leaves at heading.In general,CO2 enrichment-induced changes in both EP activity and specific activity were larger at 10 and 20 DAH than at heading and maturity.EP activity at early filling stage decreased greatly with increasing N supply,while it showed adverse trends at late filling stages.The specific activity decreased with increasing N supply at all sampling stages,with CO2 enrichment-induced decreases being larger at early than at late filling stages.Compared with AMB,FACE significantly decreased the content of soluble protein in the flag and the second leaves at 10,20 DAH and maturity,and CO₂ enrichment-induced changes in the content of soluble protein significantly decreased with increasing N supply.Soluble protein contents were correlated positively with EP activity in the flag and the second leaves at the maturity stage,while it showed adverse trends at the other sampling stages.This suggested that the CO2 enrichment-induced reduction in soluble protein content in leaves at the filling stage closely linked with corresponding changes in EP activity at elevated CO₂.

利用农田开放式空气CO2浓度增高 (free air CO2 enrichment，FACE) 系统，CO2浓度设正常CO2 (ambient, AMB) 和高CO2 (FACE，AMB + 200 μmol/mol) 2个水平，施N量设低氮 (LN，15 g/m2，以纯氮计)、中氮 (NN，25 g/m2)和高氮 (HN，35 g / m2) 3个水平，对水稻品种武香粳14结实期剑叶和倒2叶的内肽酶活力变化情况进行了研究。结果表明: 1) 同AMB相比，FACE处理使剑叶抽穗后10 d、20 d以及倒2叶抽穗到抽穗后20 d内肽酶活力明显提高，但使剑叶成熟期内肽酶活力明显下降；使剑叶抽穗后10 d、20 d和成熟期以及倒2叶抽穗到成熟期内肽酶比活力明显提高；FACE对抽穗后10 d、20 d叶片内肽酶活力和比活力影响较大，而对抽穗期和成熟期的影响较小。同LN相比，HN使灌浆前期叶片内肽酶活力明显降低，灌浆中后期则呈增加趋势；HN降低了叶片各期内肽酶比活力，灌浆前期降幅大于后期。 2) 同AMB相比，FACE使叶片抽穗后10 d至成熟期可溶性蛋白含量明显降低；同LN相比，HN明显减缓FACE处理对抽穗后10 d至成熟期功能叶片可溶性蛋白含量的影响。 3）除成熟期叶片可溶性蛋白含量与内肽酶活力呈正相关外，结实期剑叶及倒2叶可溶性蛋白含量与对应时期内肽酶活力呈显著负相关。上述结果说明，FACE处理下水稻剑叶及倒2叶结实中后期可溶性蛋白含量明显下降与结实期叶片内肽酶活力的变化关系密切。

... 生育后期功能叶片可溶性蛋白是籽粒重要氮源, 籽粒充实阶段剑叶向籽粒源源不断地输氮化合物, 所以剑叶可溶性蛋白含量呈下降趋势^[24] ...

... 有研究表明, 水分含量较高情况下, 植物体内正常的蛋白质合成受到抑制, 会被诱导出新蛋白, 使蛋白质的含量增加^[24] ...

2006

0.0

1.667

Acta Agron Sin. 2006, 32(12):1892 - 1897

Effects of Water Stress on Some Physiological Characteristics in Rice during Grain Filling Stage

水分胁迫对水稻结实期一些生理性状的影响

WANG He-Zheng,MA Jun*,LI Xu-Yi,LI Yan,ZHANG Rong-Ping,WANG Ren-Quan

王贺正;马均*;李旭毅;李艳;张荣萍;汪仁全

Rice Research Institute of Sichuan Agricultural University, Wenjiang 611130, Sichuan, China

Along with the increasing scarcity of global water resource, water shortage is crucial for restricting agricultural development in china. Rice is one of the main staple food crops in the world, and may suffer serious water stress during its growth period. Water stress during grain filling results in decreasing grain yield, and its physiological mechanism is unclear under continuance of the water stress. So study on the relationship between water stress and drought resistance in rice have become increasingly important proposals of rice scientific researches. The experiments were carried out in growth chamber to investigate the effects of water stress on MDA，photosynthetic rate, stomatal conductance, osmotic regulators and antioxidative enzyme activities in three hybrid rice cultivars (combinations) with different drought resistant abilities under both normal and drought conditions during grain filling stage. The results indicated that water stress aggravated the membrane lipid peroxidation in rice leaves, and it was more severe in weaker drought resistant cultivars than in better drought resistant cultivars. During the period of water stress, decreases of photosynthetic rate, stomatal conductance and chlorophyll content in better drought resistant cultivars were obvious less than those in weaker drought resistant ones. Contents of soluble sugar and amino acid and activities of superoxide dismutase(SOD), catalase(CAT) and peroxidase(POD) in leaves increased obviously in 0–14 d after anthesis and subsequently decreased rapidly, and the increases were more in better drought resistant cultivars than in weaker drought resistant ones in 0–14 d after anthesis. These results suggest that water stress can induce physiological and biochemical changes in rice, and the changes indicate drought resistance of rice during grain filling stage. Changes of MDA content, photosynthesis, osmotic regulators and antioxidative enzyme activities are correlated significantly to drought resistance of rice varieties.

选用抗旱性不同的3个杂交水稻品种，在干旱棚内分别在正常供水和水分胁迫条件下，研究了水稻结实期丙二醛（MDA）含量、光合速率、气孔导度、渗透调节物质及保护性酶活性等生理性状变化。结果表明，结实期随土壤水分胁迫加剧和时间延长细胞膜脂过氧化加剧，抗旱性弱的品种膜脂过氧化更为严重；剑叶光合速率、气孔导度和叶绿素含量迅速下降，抗旱性弱的品种比抗旱性强的品种降速快、幅度大。抽穗后0～14 d随胁迫时间的延长叶片内可溶性糖、氨基酸含量增加，超氧化物歧化酶（SOD）、过氧化氢酶（CAT）和过氧化物酶（POD）3种酶活性升高，抗旱性强的品种增加或升高的幅度大。抽穗14 d后叶片内可溶性糖、氨基酸含量下降，SOD、CAT和POD酶活性降低。试验表明，水稻叶片MDA含量、光合作用、渗透调节物质及保护性酶活性的变化幅度与品种的抗旱性密切相关。

... 土壤水分含量增加使抗旱性品种剑叶可溶性糖含量增加幅度较大, CO₂浓度增加使多数品种孕穗期到完熟期剑叶可溶性蛋白含量下降^[25,26] ...

1999

0.0

1.667

Acta Agron Sin. 1999, 25(1):39 - 43

Changes of Antioxidative Ability in Leaves of Rice Cultivars Grown under Enriched C02

加富CO2条件下水稻叶片抗氧化能力的变化

Peng Changlian;Lin Zhifang;Lin Guizhu

彭长连;林植芳;林桂珠

South China Institute of Botany, Academiu Sinica, Guangzhou 510650

For understanding the changes of antioxidative ability in crop plants under CO2 enrichment，six rice cultivars (Oryza sativa L.)were grown at two chambers located outdoors and exposed to natural sunlight during the life time，The CO2 concentrations in these two chamber were 350μl/L(ambient) and 600

本文研究厂不同产地栽培稻品种(IR72-菲律宾，Amaroo-澳大利亚，绿黄占3号-广东，特三矮2号-广东)和广东产野生稻(药用野生稻，普通野生稻)在加富CO2条件下抗氧化能力的变化。结果表明在自然条件下其抗氧化酶活性因品种、产地的不同而存在一定的差异。与普通的空气CO2浓度(350μl/L)下生长的水稻相比，加富CO2下(600μl/L)下

... 土壤水分含量增加使抗旱性品种剑叶可溶性糖含量增加幅度较大, CO₂浓度增加使多数品种孕穗期到完熟期剑叶可溶性蛋白含量下降^[25,26] ...

... 抽穗至完熟期是决定结实率和粒重的关键时期, 土壤温度及叶片寿命等都可能影响最终产量^[26] ...

2006

0.0

... 呼吸代谢与氨基酸和蛋白质的生物合成、转化密切相关, 早在19世纪就有人发现呼吸作用随CO₂浓度升高而下降, 以后不少研究都证实了这一点^[27] ...

2004

0.0

1.667

Acta Agron Sin. 2004, 30(12):1244 - 1249

Effects of Water Stress on the Antioxidant System in Mature Leaves of Hybrid Rice “Liangyou-peijiu”

高产杂交稻“两优培九”功能叶抗氧化系统对水分胁迫的响应

LIU Shao-Hua;CHEN Guo-Xiang;HU Yan;LU Chuan-Gen;WU Guo-Rong;YANG Yan-Hua

刘少华;陈国祥;胡艳;吕川根;吴国荣;杨艳华

College of Life Science, Nanjing Normal University, Nanjing 210097, Jiangsu

The effect of water stress on the antioxidant system of the functional leaves were studied. The results showed that during the prophase of water stress, (1) the activity of SOD was descended slightly, but the activities of POD and CAT were enhanced; (2) the contents of AsA and GSH were reduced; (3)

主要研究了在不同程度的土壤水分胁迫下“两优培九”功能叶抗氧化系统的变化情况。结果显示，在水分胁迫的前期，抗氧化系统中各成分的反应并不完全一致，SOD活性略有下降，而POD和CAT的活性则快速上升；AsA和GSH的含量不同程度地下降；脯氨酸和可溶性蛋白的含量均大幅增加。O2．- 的产生速率及膜脂过氧化产物MDA的含量始终

... 此外, 覆膜使土壤含水量增加, 叶片加速变黄, 影响叶绿素的合成, 与前人研究结果一致^[28] ...

2002

0.0

1.742

... 2%^[29,30,31], 经济系数比对照减少2 ...

2003

6.91

0.0

... 2%^[29,30,31], 经济系数比对照减少2 ...

2003

0.0

1.742

... 2%^[29,30,31], 经济系数比对照减少2 ...