高CO_2浓度对杂交水稻光合作用日变化的影响——FACE研究
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摘要
大气二氧化碳(CO_2)浓度增高导致全球变暖,但作为光合作用底物促进绿色作物的光合作用。为了明确高CO_2浓度对杂交水稻结实期光合日变化的影响,2014年利用稻田FACE(Free Air CO_2Enrichment)平台,以生产上曾创高产纪录的两个杂交稻新组合甬优2640和Y两优2号为供试材料,设置环境CO_2和高CO_2浓度(增200μmol/mol)两个水平,测定杂交稻抽穗期和灌浆中期光合作用日变化和成熟期生物量。结果表明,高CO_2浓度环境下两组合抽穗期叶片净光合速率均大幅增加(全天平均52%),但灌浆中期的平均增幅减半,其中Y两优2号这种光合下调表现更为明显。大气CO_2浓度升高使两杂交稻组合抽穗和灌浆中期叶片气孔导度均大幅下降,导致蒸腾速率下降而水分利用效率大幅增加,Y两优2号气孔导度和蒸腾速率对CO_2的响应上午大于下午,而甬优2640表现相反。尽管大气CO_2浓度升高使杂交稻结实期不同时刻胞间CO_2浓度均大幅增加,但对气孔限制值特别是胞间CO_2与空气CO_2浓度之比多无显著影响,两品种趋势一致。大气CO_2浓度升高对甬优2640地上部生物量及其组分的影响明显大于Y两优2号,CO_2与品种间多存在互作效应。以上结果表明,与甬优2640相比,Y两优2号最终生产力从高CO_2浓度环境中获益较少可能与该品种生长后期存在明显的光合适应有关,但这种光合适应似乎不是由气孔限制造成的。
The rising of atmospheric carbon dioxide( CO_2) concentration has been blamed for global warming,but it promotes the leaf photosynthesis of crops because CO_2 is the main substrate for photosynthesis. In order to understand the effect of elevated CO_2 concentration on diurnal courses of CO_2 uptake of hybrid rice,a paddy field experiment utilizing freeair CO_2enrichment( FACE) technology was undertaken to determine diurnal courses of leaf photosynthesis at heading and middle grain filling stages,and its association with the final productivity of rice at maturity. Two hybrid rice variety Yongyou2640( YY 2640) and YLiangyou No.2( YLY 2) were grown in the ambient CO_2 and the elevated CO_2concentration( 200μmol/mol above ambient) from plant transplanting until grain maturity. Elevated CO_2 concentration significantly increased net photosynthetic CO_2 assimilation of flag leaves of two hybrids by 52% on average at heading,but the enhancement was reduced to half at the middle grain filling stage. This photosynthesis acclimation was more pronounced in YLY 2. Elevated CO_2 concentration significantly decreased stomatal conductance of two hybrids at both heading and grain filling stages,resulting in lower transpiration and higher water use efficiency. Greater CO_2 responses of leaf transpiration and stomatal conductance were observed in the morning for YLY 2,but YY 2640 showed higher CO_2 effects in the afternoon. Growth at elevated CO_2 concentration significantly increased intercellular CO_2 concentration,but had no clear effects on the ratio of intercellular to air CO_2 concentration and stomata limitation value,and the same trend was observed for the two varieties.The CO_2 effects on the above-ground biomass and its components were greater in YY 2640 than YLY 2,and it was reflected in the significant CO_2 by variety interactions. The results indicated that compared with YY 2640,the lower CO_2 gain on final productivity of YLY 2 might result from photosynthesis acclimation at the late growth stage,and this down-regulation in leaf photosynthesis was not caused by stomatal limitation.
The rising of atmospheric carbon dioxide( CO_2) concentration has been blamed for global warming,but it promotes the leaf photosynthesis of crops because CO_2 is the main substrate for photosynthesis. In order to understand the effect of elevated CO_2 concentration on diurnal courses of CO_2 uptake of hybrid rice,a paddy field experiment utilizing freeair CO_2enrichment( FACE) technology was undertaken to determine diurnal courses of leaf photosynthesis at heading and middle grain filling stages,and its association with the final productivity of rice at maturity. Two hybrid rice variety Yongyou2640( YY 2640) and YLiangyou No.2( YLY 2) were grown in the ambient CO_2 and the elevated CO_2concentration( 200μmol/mol above ambient) from plant transplanting until grain maturity. Elevated CO_2 concentration significantly increased net photosynthetic CO_2 assimilation of flag leaves of two hybrids by 52% on average at heading,but the enhancement was reduced to half at the middle grain filling stage. This photosynthesis acclimation was more pronounced in YLY 2. Elevated CO_2 concentration significantly decreased stomatal conductance of two hybrids at both heading and grain filling stages,resulting in lower transpiration and higher water use efficiency. Greater CO_2 responses of leaf transpiration and stomatal conductance were observed in the morning for YLY 2,but YY 2640 showed higher CO_2 effects in the afternoon. Growth at elevated CO_2 concentration significantly increased intercellular CO_2 concentration,but had no clear effects on the ratio of intercellular to air CO_2 concentration and stomata limitation value,and the same trend was observed for the two varieties.The CO_2 effects on the above-ground biomass and its components were greater in YY 2640 than YLY 2,and it was reflected in the significant CO_2 by variety interactions. The results indicated that compared with YY 2640,the lower CO_2 gain on final productivity of YLY 2 might result from photosynthesis acclimation at the late growth stage,and this down-regulation in leaf photosynthesis was not caused by stomatal limitation.
引文
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[13]杨连新,王余龙,黄建晔,杨洪建,刘红江.开放式空气CO2浓度增高对水稻生长发育影响的研究进展.应用生态学报,2006,17(7):1331-1337.
[14]景立权,赖上坤,王云霞,杨连新,王余龙.大气CO2浓度和温度互作对水稻生长发育影响的研究进展.生态学报,2016,DOI:10.5846/stxb201412272588.
[15]邵在胜,赵轶鹏,宋琪玲,贾一磊,王云霞,杨连新,王余龙.大气CO2和O3浓度升高对汕优63叶片光合作用的影响.中国生态农业学报,2014,22(4):422-529.
[16]Zhu C W,Zhu J G,Cao J,Jiang Q,Liu G,Ziska L H.Biochemical and molecular characteristics of leaf photosynthesis and relative seed yield of two contrasting rice cultivars in response to elevated[CO2].Journal of Experimental Botany,2014,65(20):6049-6056
[17]Long S P,Ainsworth E A,Leakey A D B,Nosberger J,Ort D R.Food for Thought:Lower-than-expected crop yield stimulation with rising CO2concentrations.Science,2006,312:1918-1921.
[18]Sun J D,Yang L X,Wang Y L,Ort D R.FACE-ing the global change:Opportunities for improvement in photosynthetic radiation use efficiency and crop yield(A research review).Plant Science,2009,177:511-522.
[19]Kobayashi K,Okada M,Kim H Y,Lieffering M,Miura S,Hasegawa T.Paddy rice responses to free-air[CO2]enrichment.//Nosberger J,Long S P,Norby R J,Stitt M,Hendrey G R,Blum H Eds,Ecological Studies Series,Springer-Verlag Berlin Heidelberg,2006,187:87-104.
[20]刘钢,韩勇,朱建国,冈田益己,中村浩史,吉本真由美.稻麦轮作FACE系统平台I.系统结构与控制.应用生态学报,2002,13(10):1253-1258.
[21]彭玉林,李鸿,何森林,姜国泉,吴朝晖,闻尉宏,袁隆平.Y两优2号在安徽舒城低海拔地区“百亩方”单产突破12.5 t/hm2栽培技术.杂交水稻,2013,28(6):50-52.
[22]赖上坤,庄时腾,吴艳珍,王云霞,朱建国,杨连新,王余龙.大气CO2浓度和温度升高对超级稻生长发育的影响.生态学杂志,2015,34(5):1253-1262.
[23]沈陈华.气象因子对江苏省水稻单产的影响.生态学报,2015,35(12):4155-5167.
[24]彭少兵.对转型时期水稻生产的战略思考.中国科学:生命科学,2014,44(8):845-850.
[25]朱德峰,张玉屏,陈惠哲,向镜,张义凯.中国水稻高产栽培技术创新与实践.中国农业科学,2015,48(17):3404-3414
[26]Cure J D,Acock B.Crop responses to carbon dioxide doubling:A literature survey.Agricultural and Forest Meteorology,1986,38:127-145.
[27]Moore B D,Cheng S H,Sims D and Seemann J R.The biochemical and molecular basis for photosynthetic acclimation to elevated atmospheric CO2.Plant Cell Environment,1999,22:567-582.
[28]Rogers A,Humphries S W.A mechanistic evaluation of photosynthetic acclimation at elevated CO2.Global Change Biology.2000,6:1005-1011.
[29]Sakai H,Hasegawa T,Kobayashi K.Enhancement of rice canopy carbon gain by elevated CO2is sensitive to growth stage and leaf nitrogen concentration.New Phytologist,2006,170:321-332.
[30]Ziska L H,Manalo P A,Ordonez R A.Intraspecific variation in the response of rice(Oryza sativa L.)to increased CO2and temperature:growth and yield response of 17 cultivars.Journal of Experimental Botany,1996,47(9):1353-1359.
[31]Moya T B,Ziska L H,Namuco O S,Olszyk D.Growth dynamics and genotypic variation in tropical,field-grown paddy rice(Oryza sativa L.)in response to increasing carbon dioxide and temperature.Global Change Biology,1998,4(6):645-656.
[32]Hasegawa T,Sakai H,Tokida T,Nakamura H,Zhu C,Usui Y,Yoshimoto M,Fukuoka M,Wakatsuki H,Katayanagi N,Matsunami T,Kaneta Y,Sato T,Takakai F,Sameshima R,Okada M,Mae T,Makino A.Rice cultivar responses to elevated CO2at two free-air CO2enrichment(FACE)sites in Japan.Functional Plant Biology,2013,40:148-159.
[2]Fisher B,Nakicenovic N,Alfsen K,Corfee Morlot J,de la Chesnaye F,Hourcade J C et al.Issues related to mitigation in the long term context.//In Climate Change 2007:Mitigation.Contribution of Working Group III to the Fourth Assessment Report of the Inter-governmental Panel on Climate Change.Edited by Metz L A M B,Davidson O R,Bosch P R and Dave R.Cambridge,UK.2007:169-250.
[3]IPCC(Intergovernmental Panel on Climate Change).The Physical Science Basis//Lisa V A,Simon K A,Nathaniel L B,eds.Contribution of Working GroupⅠto the Fifth Annual Assessment Report of the IPCC.Cambridge UK:Cambridge University Press,2013:29-29.
[4]Kimball B A,Kobayashi K,Bindi M.Responses of agricultural crops to free-air CO2enrichment.Advances in Agronomy,2002,77:293-368.
[5]王云霞,杨连新,Remy Manderscheid,王余龙.C4作物FACE(free air CO2enrichment)研究进展.生态学报,2011,31(5):1450-1459.
[6]杨连新,王云霞,朱建国,王余龙.十年水稻FACE研究的产量响应.生态学报,2009,29(3):1486-1497.
[7]杨连新,王云霞,朱建国,Toshihiro Hasegawa,王余龙.开放空气中CO2浓度增高(FACE)对水稻生长和发育的影响.生态学报,2010,30(6):1573-1585.
[8]Wang J Y,Wang C,Chen N N,Xiong Z Q,Wolfe D,Zou J W.Response of rice production to elevated[CO2]and its interaction with rising temperature or nitrogen supply:a meta-analysis.Climatic Change,2015,130:529-543.
[9]Liu H J,Yang L X,Wang Y L,Huang J Y,Zhu J G,Wang Y X,Dong G C,Liu G.Yield formation of CO2-enriched hybrid rice cultivar Shanyou63 under fully open-air field conditions.Field Crops Research,2008,108:93-100.
[10]Yang L X,Liu H J,Wang Y X,Zhu J G,Huang J Y,Liu G,Dong G C,Wang Y L.Yield formation of CO2-enriched inter-subspecific hybrid rice cultivar Liangyoupeijiu under fully open-air field condition in a warm sub-tropical climate.Agriculture,Ecosystems and Environment,2009,129:193-200.
[11]Yang L X,Liu H J,Wang Y X,Zhu J G,Huang J Y,Liu G,Dong G C,Wang Y L.Impact of elevated CO2concentration on inter-subspecific hybrid rice cultivar Liangyoupeijiu under fully open air field conditions.Field Crops Research,2009,112:7-15.
[12]赖上坤,周三妮,顾伟锋,庄时腾,周娟,朱建国,杨连新,王余龙.二氧化碳、施氮量和移栽密度对汕优63产量形成的影响——FACE研究.农业环境科学学报,2014,33(5):836-843.
[13]杨连新,王余龙,黄建晔,杨洪建,刘红江.开放式空气CO2浓度增高对水稻生长发育影响的研究进展.应用生态学报,2006,17(7):1331-1337.
[14]景立权,赖上坤,王云霞,杨连新,王余龙.大气CO2浓度和温度互作对水稻生长发育影响的研究进展.生态学报,2016,DOI:10.5846/stxb201412272588.
[15]邵在胜,赵轶鹏,宋琪玲,贾一磊,王云霞,杨连新,王余龙.大气CO2和O3浓度升高对汕优63叶片光合作用的影响.中国生态农业学报,2014,22(4):422-529.
[16]Zhu C W,Zhu J G,Cao J,Jiang Q,Liu G,Ziska L H.Biochemical and molecular characteristics of leaf photosynthesis and relative seed yield of two contrasting rice cultivars in response to elevated[CO2].Journal of Experimental Botany,2014,65(20):6049-6056
[17]Long S P,Ainsworth E A,Leakey A D B,Nosberger J,Ort D R.Food for Thought:Lower-than-expected crop yield stimulation with rising CO2concentrations.Science,2006,312:1918-1921.
[18]Sun J D,Yang L X,Wang Y L,Ort D R.FACE-ing the global change:Opportunities for improvement in photosynthetic radiation use efficiency and crop yield(A research review).Plant Science,2009,177:511-522.
[19]Kobayashi K,Okada M,Kim H Y,Lieffering M,Miura S,Hasegawa T.Paddy rice responses to free-air[CO2]enrichment.//Nosberger J,Long S P,Norby R J,Stitt M,Hendrey G R,Blum H Eds,Ecological Studies Series,Springer-Verlag Berlin Heidelberg,2006,187:87-104.
[20]刘钢,韩勇,朱建国,冈田益己,中村浩史,吉本真由美.稻麦轮作FACE系统平台I.系统结构与控制.应用生态学报,2002,13(10):1253-1258.
[21]彭玉林,李鸿,何森林,姜国泉,吴朝晖,闻尉宏,袁隆平.Y两优2号在安徽舒城低海拔地区“百亩方”单产突破12.5 t/hm2栽培技术.杂交水稻,2013,28(6):50-52.
[22]赖上坤,庄时腾,吴艳珍,王云霞,朱建国,杨连新,王余龙.大气CO2浓度和温度升高对超级稻生长发育的影响.生态学杂志,2015,34(5):1253-1262.
[23]沈陈华.气象因子对江苏省水稻单产的影响.生态学报,2015,35(12):4155-5167.
[24]彭少兵.对转型时期水稻生产的战略思考.中国科学:生命科学,2014,44(8):845-850.
[25]朱德峰,张玉屏,陈惠哲,向镜,张义凯.中国水稻高产栽培技术创新与实践.中国农业科学,2015,48(17):3404-3414
[26]Cure J D,Acock B.Crop responses to carbon dioxide doubling:A literature survey.Agricultural and Forest Meteorology,1986,38:127-145.
[27]Moore B D,Cheng S H,Sims D and Seemann J R.The biochemical and molecular basis for photosynthetic acclimation to elevated atmospheric CO2.Plant Cell Environment,1999,22:567-582.
[28]Rogers A,Humphries S W.A mechanistic evaluation of photosynthetic acclimation at elevated CO2.Global Change Biology.2000,6:1005-1011.
[29]Sakai H,Hasegawa T,Kobayashi K.Enhancement of rice canopy carbon gain by elevated CO2is sensitive to growth stage and leaf nitrogen concentration.New Phytologist,2006,170:321-332.
[30]Ziska L H,Manalo P A,Ordonez R A.Intraspecific variation in the response of rice(Oryza sativa L.)to increased CO2and temperature:growth and yield response of 17 cultivars.Journal of Experimental Botany,1996,47(9):1353-1359.
[31]Moya T B,Ziska L H,Namuco O S,Olszyk D.Growth dynamics and genotypic variation in tropical,field-grown paddy rice(Oryza sativa L.)in response to increasing carbon dioxide and temperature.Global Change Biology,1998,4(6):645-656.
[32]Hasegawa T,Sakai H,Tokida T,Nakamura H,Zhu C,Usui Y,Yoshimoto M,Fukuoka M,Wakatsuki H,Katayanagi N,Matsunami T,Kaneta Y,Sato T,Takakai F,Sameshima R,Okada M,Mae T,Makino A.Rice cultivar responses to elevated CO2at two free-air CO2enrichment(FACE)sites in Japan.Functional Plant Biology,2013,40:148-159.