粳稻生育后期剑叶光合日变化和光合色素对大气CO_2浓度和温度升高的响应——FACE研究
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摘要
为研究水稻叶片光合色素和光合日变化对大气CO_2浓度和气温升高的响应,我们采用在开放空气中控制升高CO_2浓度和温度的方法,以常规粳稻南粳9108为试验材料,设置了环境CO_2和高大气CO_2浓度(增加200μmol·mol~(-1))、环境温度和增温(增加1~2℃)交互的4个处理,测定了灌浆中期和后期水稻剑叶的光合日变化特征和光合色素含量.结果表明:水稻剑叶净光合速率(P_n)为双峰曲线,发生了光合"午休"现象;大气CO_2浓度升高提高了剑叶P_n,灌浆中期和后期平均分别增加了47.6%和39.1%;高温有降低P_n的趋势,但相关性未达到显著水平.大气CO_2浓度和温度升高导致水稻剑叶生育后期气孔导度(g_s)平均分别降低了17.0%和11.8%.高CO_2浓度水稻剑叶生育后期蒸腾速率(T_r)、叶绿素a、叶绿素b、类胡萝卜素、总叶绿素和叶绿素a/b值显著降低,平均降幅分别为5.9%、50.4%、21.3%、41.4%、39.4%和21.4%,明显增加了剑叶水分利用率(WUE),平均增幅达47.9%.与之相反,生育后期增温使水稻剑叶T_r增加了10.2%,使WUE平均降低了20.4%.综上所述,大气CO_2浓度升高对粳稻生育后期剑叶P_n、g_s和光合色素含量的影响明显大于增温效应.因此,应重视大气CO_2浓度和温度对水稻光合作用和光合色素的综合效应,减弱增温的负效应.
A local popular Japonica rice(Oryza sating L.) cultivar,Nanjing 9108,was tested with free air controlled enrichment(FACE) approach to study the responses of photosynthetic pigment content and diurnal variation of flag-leaf photosynthesis to elevated atmospheric CO_2 concentration and temperature. Four alternative treatments were designed with two CO_2 concentration levels(ambient and elevated 200 μmol·mol~(-1)) and two air temperature regimes(ambient and elevated1-2 ℃). Diurnal variation of flag-leaf photosynthesis was measured in the middle full stage and the late full stage,and photosynthetic pigment of the leaf was analyzed afterward. Results showed that diurnal variation of net photosynthetic rate(P_n) in each treatment followed a double-peak curve with midday depression feature during late growth stage. Compared to P_n under ambient condition,P_n under elevated CO_2 concentration increased by 47.6% and 39.1% on average at middle full stage and late full stage,respectively. There was a negative correlation between temperature and P_n with no significance. Both elevated CO_2 concentration and temperature had a significant negative effect on stomatal conductance(g_s),decreased by 17.0% and 11.8% on average,respectively. Elevated CO_2 concentration significantly reduced transpiration rate(T_r),chlorophyll a(Chl a),chlorophyll b(Chl b),carotene(Car),total chlorophyll(Chl t) and chlorophyll a/b ratio(Chl a/b) during late growth stage of rice by 5. 9%,50. 4%,21. 3%,41. 4%,39. 4% and 21. 4% on average,respectively,whereas water use efficiency(WUE) increased by 47. 9%. However,there were opposite effects on T_r,WUE and photosynthetic pigment content under elevated temperature,with Trincreased by 10. 2% and WUE decreased by 20. 4%. It could be concluded that elevated CO_2 concentration had a greater effect on P_n,g_s and photosynthetic pigment content of rice leaf than elevated temperature did during late growth stage. Therefore,it should be paid more attention to the colligate effects of elevated CO_2 concentration and high temperature on photosynthesis and photosynthetic pigment content to reduce negative effect of high air temperature.
A local popular Japonica rice(Oryza sating L.) cultivar,Nanjing 9108,was tested with free air controlled enrichment(FACE) approach to study the responses of photosynthetic pigment content and diurnal variation of flag-leaf photosynthesis to elevated atmospheric CO_2 concentration and temperature. Four alternative treatments were designed with two CO_2 concentration levels(ambient and elevated 200 μmol·mol~(-1)) and two air temperature regimes(ambient and elevated1-2 ℃). Diurnal variation of flag-leaf photosynthesis was measured in the middle full stage and the late full stage,and photosynthetic pigment of the leaf was analyzed afterward. Results showed that diurnal variation of net photosynthetic rate(P_n) in each treatment followed a double-peak curve with midday depression feature during late growth stage. Compared to P_n under ambient condition,P_n under elevated CO_2 concentration increased by 47.6% and 39.1% on average at middle full stage and late full stage,respectively. There was a negative correlation between temperature and P_n with no significance. Both elevated CO_2 concentration and temperature had a significant negative effect on stomatal conductance(g_s),decreased by 17.0% and 11.8% on average,respectively. Elevated CO_2 concentration significantly reduced transpiration rate(T_r),chlorophyll a(Chl a),chlorophyll b(Chl b),carotene(Car),total chlorophyll(Chl t) and chlorophyll a/b ratio(Chl a/b) during late growth stage of rice by 5. 9%,50. 4%,21. 3%,41. 4%,39. 4% and 21. 4% on average,respectively,whereas water use efficiency(WUE) increased by 47. 9%. However,there were opposite effects on T_r,WUE and photosynthetic pigment content under elevated temperature,with Trincreased by 10. 2% and WUE decreased by 20. 4%. It could be concluded that elevated CO_2 concentration had a greater effect on P_n,g_s and photosynthetic pigment content of rice leaf than elevated temperature did during late growth stage. Therefore,it should be paid more attention to the colligate effects of elevated CO_2 concentration and high temperature on photosynthesis and photosynthetic pigment content to reduce negative effect of high air temperature.
引文
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[10]Du Y-D(杜尧东),Li J-L(李键陵),Wang H(王华),et al.Effects of high temperature stress on the flag leaf photosynthesis and chlorophyll fluorescence parameters of rice.Chinese Journal of Ecology(生态学杂志),2012,31(10):2541-2548(in Chinese)
[11]Vu JCV,Allen LH,Boote KJ,et al.Effects of elevated CO2and temperature on photosynthesis and rubisco in rice and soybean.Plant,Cell and Environment,1997,20:68-76
[12]Lai S-K(赖上坤),Zhuang S-T(庄时腾),Wu Y-Z(吴艳珍),et al.Impact of elevated atmospheric CO2concentration and temperature on growth and development of super rice.Chinese Journal of Ecology(生态学杂志),2015,34(5):1253-1262(in Chinese)
[13]Zhou N(周宁),Shen S-B(沈士博),Jing L-Q(景立权),et al.Effects of elevated atmospheric CO2and temperature on diurnal courses of photosynthesis in leaves of Japonica rice.Chinese Journal of Ecology(生态学杂志),2016,35(9):2404-2416(in Chinese)
[14]Liu G(刘钢),Han Y(韩勇),Zhu J-G(朱建国),et al.Rice-wheat rotational FACE platform.Ⅰ.System structure and control.Chinese Journal of Applied Ecology(应用生态报),2002,13(10):1253-1258(in Chinese)
[15]Lawsor DW,Mitchell PAC.The effects of increasing CO2on crop photosynthesis and productivity:A review of field studies.Plant,Cell and Environment,1991,14:807-818
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[17]Xu D-Q(许大全),Xu B-J(徐宝基),Shen Y-G(沈允钢).Diurnal variation of photosynthetic efficiency in C3plants.Acta Phytophysiologica Sinica(植物生理学报),1990,16(1):1-5(in Chinese)
[18]Li X(李霞),Jiao D-M(焦德茂),Wang S-H(王守海),et al.Photosynthetic characteristics for rice hybrids with transgenic PEPC parent HPTER-01.Acta Agronomica Sinica(作物学报),2001,27(2):137-143(in Chinese)
[19]Zou Q(邹琦).A study on photosynthetic midday depression of wheat and mechanism under drought and well-watered conditions.Acta Agriculturae Boreali-Sinica(华北农学报),1997,12(4):48-51(in Chinese)
[20]Weng X-Y(翁晓燕),Lu Q(陆庆),Jiang D-A(蒋德安).Rubisco activase and its regulation on diurnal changes of photosynthetic rate and the activity of ribubose 1,5-bisphosphate carboxyase/oxygenase(rubisco).Chinese Journal of Rice Science(中国水稻科学),2001,15(1):36-41(in Chinese)
[21]Chen CP,Sakai H,Tokida T,et al.Do the rich always become Richer?Characterizing the leaf physiological response of the high-yielding rice cultivar Takanari to free-air CO2enrichment.Plant and Cell Physiology,2014,55:381-391
[22]Wang X-N(王小宁),Shen S-H(申双和),Wang ZM(王志明),et al.Effect of day-time and night-time warming on photosynthesis of rice.Jiangsu Journal of Agricultural Sciences(江苏农业学报),2008,24(3):237-240(in Chinese)
[23]Aoki N,Ono K,Sasaki H,et al.Effect of elevated CO2concentration on photosynthetic carbon metabolism in flag-leaf blades of rice before and after heading.Plant Production Science,2003,6:52-58
[24]Figueiredo N,Carranca C,Trindade H,et al.Elevated carbon dioxide and temperature effects on rice yield,leaf greenness,and phenological stages duration.Paddy Water Environment,2015,13:313-324
[25]Wang R-P(王荣平),Zhang F-T(张帆涛),Gao J-X(高家旭),et al.An overview of chlorophyll biosynthesis in higher plants.Acta Botanica Boreali-Occidentalia Sinica(西北植物学报),2009,29(3):629-636(in Chinese)
[2]Wang JY,Wang C,Chen NN,et al.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
[3]Chen GY,Yong ZH,Liao Y,et al.Photosynthetic acclimation in rice leaves to free air CO2enrichment related to both Ribulose-1,5-bisphosphate carboxylation limitation and Ribulose-1,5-bisphosphate regeneration limitation.Plant and Cell Physiology,2005,46:1036-1045
[4]Seneweara SP,Conroy JP,Ishimara K,et al.Changes in source-sink relations during development influence photosynthetic acclimation of rice to free air CO2enrichment(FACE).Functional Plant Biology,2002,29:945-953
[5]Hu J(胡健),Zhou J(周娟),Yang L-X(杨连新),et al.Effect of free air CO2enrichment(FACE)on dynamics of chlorophyll content and composition in flag leaves of rice during grain filling stage.Journal of Agro-Environment Science(农业环境科学学报),2007,26(4):1322-1326(in Chinese)
[6]Xie L-Y(谢立勇),Jiang L(姜乐),Feng Y-X(冯永祥),et al.Impacts of elevated CO2and increasing temperature on rice photosynthesis and yield in North China under FACE system.Journal of China Agricultural University(中国农业大学学报),2014,19(3):101-107(in Chinese)
[7]Hamid A,Khanam M,Biswas DK,et al.The effect of elevated CO2concentration on leaf chlorophyll and nitrogen contents in rice during post-flowering phases.Biologia Plantarum,2006,50:69-73
[8]Xie L-Y(谢立勇),Sun X(孙雪),Feng Y-X(冯永祥),et al.Responses of flag-leaf photosynthetic pigments at late growth stage and rice yield components to elevated CO2under FACE system.Chinese Journal of Eco-Agriculture(中国生态农业学报),2015,23(4):425-431(in Chinese)
[9]Zhang G-L(张桂莲),Chen L-Y(陈立云),Zhang ST(张顺堂),et al.Effects of high temperature on physiological and biochemical characteristics in flag leaf of rice during heading and flowering period.Scientia Agricultura Sinica(中国农业科学),2007,40(7):1345-1352(in Chinese)
[10]Du Y-D(杜尧东),Li J-L(李键陵),Wang H(王华),et al.Effects of high temperature stress on the flag leaf photosynthesis and chlorophyll fluorescence parameters of rice.Chinese Journal of Ecology(生态学杂志),2012,31(10):2541-2548(in Chinese)
[11]Vu JCV,Allen LH,Boote KJ,et al.Effects of elevated CO2and temperature on photosynthesis and rubisco in rice and soybean.Plant,Cell and Environment,1997,20:68-76
[12]Lai S-K(赖上坤),Zhuang S-T(庄时腾),Wu Y-Z(吴艳珍),et al.Impact of elevated atmospheric CO2concentration and temperature on growth and development of super rice.Chinese Journal of Ecology(生态学杂志),2015,34(5):1253-1262(in Chinese)
[13]Zhou N(周宁),Shen S-B(沈士博),Jing L-Q(景立权),et al.Effects of elevated atmospheric CO2and temperature on diurnal courses of photosynthesis in leaves of Japonica rice.Chinese Journal of Ecology(生态学杂志),2016,35(9):2404-2416(in Chinese)
[14]Liu G(刘钢),Han Y(韩勇),Zhu J-G(朱建国),et al.Rice-wheat rotational FACE platform.Ⅰ.System structure and control.Chinese Journal of Applied Ecology(应用生态报),2002,13(10):1253-1258(in Chinese)
[15]Lawsor DW,Mitchell PAC.The effects of increasing CO2on crop photosynthesis and productivity:A review of field studies.Plant,Cell and Environment,1991,14:807-818
[16]Zhang Z-L(张志良),Qu W-J(瞿伟菁),Li X-F(李小方).Experiment Guide to Plant Physiology.Beijing:Higher Education Press,2009(in Chinese)
[17]Xu D-Q(许大全),Xu B-J(徐宝基),Shen Y-G(沈允钢).Diurnal variation of photosynthetic efficiency in C3plants.Acta Phytophysiologica Sinica(植物生理学报),1990,16(1):1-5(in Chinese)
[18]Li X(李霞),Jiao D-M(焦德茂),Wang S-H(王守海),et al.Photosynthetic characteristics for rice hybrids with transgenic PEPC parent HPTER-01.Acta Agronomica Sinica(作物学报),2001,27(2):137-143(in Chinese)
[19]Zou Q(邹琦).A study on photosynthetic midday depression of wheat and mechanism under drought and well-watered conditions.Acta Agriculturae Boreali-Sinica(华北农学报),1997,12(4):48-51(in Chinese)
[20]Weng X-Y(翁晓燕),Lu Q(陆庆),Jiang D-A(蒋德安).Rubisco activase and its regulation on diurnal changes of photosynthetic rate and the activity of ribubose 1,5-bisphosphate carboxyase/oxygenase(rubisco).Chinese Journal of Rice Science(中国水稻科学),2001,15(1):36-41(in Chinese)
[21]Chen CP,Sakai H,Tokida T,et al.Do the rich always become Richer?Characterizing the leaf physiological response of the high-yielding rice cultivar Takanari to free-air CO2enrichment.Plant and Cell Physiology,2014,55:381-391
[22]Wang X-N(王小宁),Shen S-H(申双和),Wang ZM(王志明),et al.Effect of day-time and night-time warming on photosynthesis of rice.Jiangsu Journal of Agricultural Sciences(江苏农业学报),2008,24(3):237-240(in Chinese)
[23]Aoki N,Ono K,Sasaki H,et al.Effect of elevated CO2concentration on photosynthetic carbon metabolism in flag-leaf blades of rice before and after heading.Plant Production Science,2003,6:52-58
[24]Figueiredo N,Carranca C,Trindade H,et al.Elevated carbon dioxide and temperature effects on rice yield,leaf greenness,and phenological stages duration.Paddy Water Environment,2015,13:313-324
[25]Wang R-P(王荣平),Zhang F-T(张帆涛),Gao J-X(高家旭),et al.An overview of chlorophyll biosynthesis in higher plants.Acta Botanica Boreali-Occidentalia Sinica(西北植物学报),2009,29(3):629-636(in Chinese)