自然条件下中熟籼稻初穗期剑叶光合的气孔和非气孔限制特征
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摘要
为探究自然条件下中熟籼稻初穗期剑叶光合作用的气孔与非气孔限制特征,利用LI-6400光合仪测定了4个水稻品种(恢复系R7-35,不育系井大3S,杂交F_1代井大3S/R7-35和对照品种9311)的气体交换数据,获得了它们的净光合速率、气孔导度、胞间CO_2浓度和气孔限制值对光的响应曲线。结果表明:与最大净光合速率相对应的饱和光强(I_(sat))分别为1500、1800、1200和1200μmol·m~(-2)·s~(-1);除井大3S/R7-35外,其他3个水稻品种的气孔导度随光强的增加而非线性增加,由此可知,在较高光强处限制这4个水稻叶片光合作用的因素是非气孔限制。与最小胞间CO_2浓度(C_i)相对应的光强分别为1600、1600、1400和1600μmol·m~(-2)·s~(-1),与最大气孔限制值(L_s)相对应的饱和光强分别为1400、1200、1400和600μmol·m~(-2)·s~(-1)。根据Ci降低和Ls升高作为判断叶片光合速率降低的主要原因是气孔因素的判据标准,品种9311则在光强小于1600μmol·m~(-2)·s~(-1)时,Ci随光强的增加而降低,在光强小于600μmol·m~(-2)·s~(-1)时,Ls随光强的增加而升高,但该品种在光强小于1200μmol·m~(-2)·s~(-1)时,其光合能力随光强的增加而非线性增加。这个结果与以Ci降低和Ls升高作为判断准则相互矛盾。为此,提出用植物叶片是否存在I_(sat)作为判断光合速率下降的非气孔限制因素准则,用气孔导度是否存在最大值作为植物叶片的气孔限制因素准则。
This study aims to explore the characteristics of stomatal and non-stomatal limitations on photosynthesis of flag leaves of medium mature indica rice(Oryza sativa) under natural conditions in Jiangxi.Leaf gas exchange of four rice cultivars(i.e.restorer line R7-35,sterile line Jingda 3S,F_1 hybrid of Jingda 3S/R7-35,and CK 9311)at the early earring stage were examined using a LI-6400 portable photosynthesis analyzer.The light-response curves of net assimilation rate(A_n),stomatal conductance(g_s),intercellular CO_2 concentration(C_i)and stomatal limitation value(L_s)were measured.The results showed that the saturated light intensity(I_(sat))values,corresponding to the maximum A_n,for the four cultivars were 1500,1800,1200,and1200μmol·m~(-2)·s~(-1),respectively.The g_sof R7-35,Jingda 3S and 9311 increased with theincreases of light intensity(I).At higher I levels,non-stomatal limitation played a major role in limiting A_nof the four cultivars.The I values corresponding to the minimum C_iwere 1600,1600,1400,and 1600μmol·m~(-2)·s~(-1),respectively.The saturation I values corresponding to the maximum L_swere 1400,1200,1400,and 600μmol·m~(-2)·s~(-1),respectively.If the decrease of C_iand the increase of L_swere the main reasons for the decreases of photosynthetic rate,the criterion of stomatal factor would be biased.Cultivar 9311 showed a decrease of C_iwith the increases of I until reaching 1600μmol·m~(-2)·s~(-1),an increase of L_swith the increases of I until reaching600μmol·m~(-2)·s~(-1),and a nonlinear increase of A_nwith the increases of I until reaching 1200μmol·m~(-2)·s~(-1).These results indicated that a combination of C_idecrease and of L_sincrease would not necessarily lead to a decrease in A_n.Accordingly,we propose a new criterion for determining non-stomatal limitation whether it had I_(sat)and for determining stomatal limitation whether it had the maximum g_s.
This study aims to explore the characteristics of stomatal and non-stomatal limitations on photosynthesis of flag leaves of medium mature indica rice(Oryza sativa) under natural conditions in Jiangxi.Leaf gas exchange of four rice cultivars(i.e.restorer line R7-35,sterile line Jingda 3S,F_1 hybrid of Jingda 3S/R7-35,and CK 9311)at the early earring stage were examined using a LI-6400 portable photosynthesis analyzer.The light-response curves of net assimilation rate(A_n),stomatal conductance(g_s),intercellular CO_2 concentration(C_i)and stomatal limitation value(L_s)were measured.The results showed that the saturated light intensity(I_(sat))values,corresponding to the maximum A_n,for the four cultivars were 1500,1800,1200,and1200μmol·m~(-2)·s~(-1),respectively.The g_sof R7-35,Jingda 3S and 9311 increased with theincreases of light intensity(I).At higher I levels,non-stomatal limitation played a major role in limiting A_nof the four cultivars.The I values corresponding to the minimum C_iwere 1600,1600,1400,and 1600μmol·m~(-2)·s~(-1),respectively.The saturation I values corresponding to the maximum L_swere 1400,1200,1400,and 600μmol·m~(-2)·s~(-1),respectively.If the decrease of C_iand the increase of L_swere the main reasons for the decreases of photosynthetic rate,the criterion of stomatal factor would be biased.Cultivar 9311 showed a decrease of C_iwith the increases of I until reaching 1600μmol·m~(-2)·s~(-1),an increase of L_swith the increases of I until reaching600μmol·m~(-2)·s~(-1),and a nonlinear increase of A_nwith the increases of I until reaching 1200μmol·m~(-2)·s~(-1).These results indicated that a combination of C_idecrease and of L_sincrease would not necessarily lead to a decrease in A_n.Accordingly,we propose a new criterion for determining non-stomatal limitation whether it had I_(sat)and for determining stomatal limitation whether it had the maximum g_s.
引文
丁雷,李英瑞,李勇,等.2014.梯度干旱胁迫对水稻叶片光合和水分状况的影响.中国水稻科学,28(1):65-70.
高冠龙,冯起,张小由,等.2018.植物叶片光合作用的气孔与非气孔限制研究综述.干旱区研究,35(4):929-937.
贺燕燕,王朝英,袁中勋,等.2018.三峡库区消落带不同水淹强度下池杉与落羽杉的光合生理特性.生态学报,38(8):2722-2731.
蒋英,刘素慧.2018.不同光质对蕹菜光合特性及品质的影响.江苏农业科学,46(2):92-95.
李俊周,乔江方,李梦琪,等.2017.短时水分胁迫对水稻叶片光合作用的影响.干旱地区农业研究,35(3):126-129,277.
李征珍,杨琼,石莎,等.2017.蒙古沙冬青光合作用特征及其影响因素.生态学杂志,36(9):2481-2488.
李周,赵雅洁,宋海燕,等.2018.不同水分处理下喀斯特土层厚度异质性对两种草本叶片解剖结构和光合特性的影响.生态学报,38(2):721-732.
吕晓菡,傅鸿妃,吴根良,等.2018.低温弱光下杭椒类辣椒不同世代间光合特性的比较.浙江农业科学,59(2):190-193.
麻雪艳,周广胜.2018.夏玉米叶片气体交换参数对干旱过程的响应.生态学报,38(7):2372-2383.
任博,李俊,同小娟,等.2018.太行山南麓栓皮栎和刺槐光合作用-CO2响应模拟.应用生态学报,29(1):1-10.
王海珍,韩路,徐雅丽,等.2017.干旱胁迫下胡杨光合光响应过程模拟与模型比较.生态学报,37(7):2315-2324.
徐斌,朱报著,潘文,等.2017.广东含笑的光响应特性及其最适模型研究.林业科学研究,30(4):604-609.
许大全.1997.光合作用气孔限制分析中的一些问题.植物生理学通讯,33(4):241-244.
徐俊增,彭世彰,魏征,等.2010.节水灌溉水稻叶片胞间CO2浓度及气孔与非气孔限制.农业工程学报,26(7):76-80.
杨泽粟,张强,郝小翠.2015.自然条件下半干旱雨养春小麦生育后期旗叶光合的气孔和非气孔限制.中国生态农业学报,23(2):174-182.
杨世琼,杨再强,王琳,等.2018.高温高湿交互对设施番茄叶片光合特性的影响.生态学杂志,37(1):57-63.
俞芹,王倩颖,刘志高,等.2018.光强与水分处理下景宁木兰光合光响应模型拟合比较.生态学杂志,37(3):898-905.
张效星,樊毅,贾悦,等.2018.水分亏缺对滴灌柑橘光合和产量及水分利用效率的影响.农业工程学报,34(3):143-150.
周宁,沈士博,景立权,等.2016.自由空气中CO2浓度和温度增高对粳稻叶片光合作用日变化的影响.生态学杂志,35(9):2404-2416.
周玉霞,巨天珍,王引弟,等.2018.4种光响应曲线模型对3种高寒草甸植物的实用性分析.草地学报,26(2):488-496.
朱大恒,张闪闪,莫骏坚,等.2018.光强对湘南烟草生育后期光合及生理特性的影响.江苏农业科学,46(8):81-84.
Anev S,Ivanova A,Tzvetkova N,et al.2016.Stomatal control on photosynthesis in drought-treated subalpine pine saplings.Genetics and Plant Physiology,6:43-53.
Ball JT,Woodrow IE,Berry JA.1987.A model predicting stomatal conductance and its contribution to the control of photosynthesis under different environmental conditions.In Progress in Photosynthesis Research(ed.J.Biggens),Martinus-Nijhoff Publishers,Dordrecht,the Netherlands:221-224.
Bellasio C,Quirk J,Beerling DJ.2018.Stomatal and non-stomatal limitations in savanna trees and C4grasses grown at low,ambient and high atmospheric CO2.Plant Science,274:181-192.
Farquhar GD,Sharkey TD.1982.Stomatal conductance and photosynthesis.Annual Review of Plant Physiology,33:317-345.
Flexas J,Medrano H.2002.Drought-inhibition of photosynthesis in C3plants:Stomatal and non-stomatal limitations revisited.Annals of Botany,89:183-189.
Franks PJ,Doheny-Adams TW,Britton-Harper ZJ,et al.2015.Increasing water-use efficiency directly through genetic manipulation of stomatal density.New Phytologist,207:188-195.
Khler IH,Macdonald AJ,Schnyder H.2016.Last-century increases in intrinsic water-use ef ficiency of grassland communities have occurred over a wide range of vegetation composition,nutrient inputs,and soil p H.Plant Physiology,170:881-890.
Mantlana K,Arneth A,Veenendaal E,et al.2008.Photosynthetic properties of C4plants growing in an African savanna/wetland mosaic.Journal of Experimental Botany,59:3941-3952.
Murata N,Nishiyama Y.2018.ATP is a driving force in the repair of photosystem II during photoinhibition.Plant,Cell&Environment,41:285-299.
Nakamura S,Izumi M.2018.Regulation of chlorophagy during photoinhibition and senescence:Lessons from mitophagy.Plant&Cell Physiology,59:1135-1143.
Niyogi KK,Truong TB.2013.Evolution of flexible non-photochemical quenching mechanisms that regulate light harvesting in oxygenic photosynthesis.Current Opinion in Plant Biology,16:307-314.
Urban O,Hlavcov M,Klem K,et al.2018.Combined effects of drought and high temperature on photosynthetic characteristics in four winter wheat genotypes.Field Crops Research,223:137-149.
Ye ZP,Suggett JD,Robakowski P,et al.2013.A mechanistic model for the photosynthesis-light response based on the photosynthetic electron transport of PS II in C3 and C4 species.New Phytologist,152:1251-1262.
Ye ZP,Yu Q.2008.A coupled model of stomatal conductance and photosynthesis for winter wheat.Photosynthetica,46:637-640.
高冠龙,冯起,张小由,等.2018.植物叶片光合作用的气孔与非气孔限制研究综述.干旱区研究,35(4):929-937.
贺燕燕,王朝英,袁中勋,等.2018.三峡库区消落带不同水淹强度下池杉与落羽杉的光合生理特性.生态学报,38(8):2722-2731.
蒋英,刘素慧.2018.不同光质对蕹菜光合特性及品质的影响.江苏农业科学,46(2):92-95.
李俊周,乔江方,李梦琪,等.2017.短时水分胁迫对水稻叶片光合作用的影响.干旱地区农业研究,35(3):126-129,277.
李征珍,杨琼,石莎,等.2017.蒙古沙冬青光合作用特征及其影响因素.生态学杂志,36(9):2481-2488.
李周,赵雅洁,宋海燕,等.2018.不同水分处理下喀斯特土层厚度异质性对两种草本叶片解剖结构和光合特性的影响.生态学报,38(2):721-732.
吕晓菡,傅鸿妃,吴根良,等.2018.低温弱光下杭椒类辣椒不同世代间光合特性的比较.浙江农业科学,59(2):190-193.
麻雪艳,周广胜.2018.夏玉米叶片气体交换参数对干旱过程的响应.生态学报,38(7):2372-2383.
任博,李俊,同小娟,等.2018.太行山南麓栓皮栎和刺槐光合作用-CO2响应模拟.应用生态学报,29(1):1-10.
王海珍,韩路,徐雅丽,等.2017.干旱胁迫下胡杨光合光响应过程模拟与模型比较.生态学报,37(7):2315-2324.
徐斌,朱报著,潘文,等.2017.广东含笑的光响应特性及其最适模型研究.林业科学研究,30(4):604-609.
许大全.1997.光合作用气孔限制分析中的一些问题.植物生理学通讯,33(4):241-244.
徐俊增,彭世彰,魏征,等.2010.节水灌溉水稻叶片胞间CO2浓度及气孔与非气孔限制.农业工程学报,26(7):76-80.
杨泽粟,张强,郝小翠.2015.自然条件下半干旱雨养春小麦生育后期旗叶光合的气孔和非气孔限制.中国生态农业学报,23(2):174-182.
杨世琼,杨再强,王琳,等.2018.高温高湿交互对设施番茄叶片光合特性的影响.生态学杂志,37(1):57-63.
俞芹,王倩颖,刘志高,等.2018.光强与水分处理下景宁木兰光合光响应模型拟合比较.生态学杂志,37(3):898-905.
张效星,樊毅,贾悦,等.2018.水分亏缺对滴灌柑橘光合和产量及水分利用效率的影响.农业工程学报,34(3):143-150.
周宁,沈士博,景立权,等.2016.自由空气中CO2浓度和温度增高对粳稻叶片光合作用日变化的影响.生态学杂志,35(9):2404-2416.
周玉霞,巨天珍,王引弟,等.2018.4种光响应曲线模型对3种高寒草甸植物的实用性分析.草地学报,26(2):488-496.
朱大恒,张闪闪,莫骏坚,等.2018.光强对湘南烟草生育后期光合及生理特性的影响.江苏农业科学,46(8):81-84.
Anev S,Ivanova A,Tzvetkova N,et al.2016.Stomatal control on photosynthesis in drought-treated subalpine pine saplings.Genetics and Plant Physiology,6:43-53.
Ball JT,Woodrow IE,Berry JA.1987.A model predicting stomatal conductance and its contribution to the control of photosynthesis under different environmental conditions.In Progress in Photosynthesis Research(ed.J.Biggens),Martinus-Nijhoff Publishers,Dordrecht,the Netherlands:221-224.
Bellasio C,Quirk J,Beerling DJ.2018.Stomatal and non-stomatal limitations in savanna trees and C4grasses grown at low,ambient and high atmospheric CO2.Plant Science,274:181-192.
Farquhar GD,Sharkey TD.1982.Stomatal conductance and photosynthesis.Annual Review of Plant Physiology,33:317-345.
Flexas J,Medrano H.2002.Drought-inhibition of photosynthesis in C3plants:Stomatal and non-stomatal limitations revisited.Annals of Botany,89:183-189.
Franks PJ,Doheny-Adams TW,Britton-Harper ZJ,et al.2015.Increasing water-use efficiency directly through genetic manipulation of stomatal density.New Phytologist,207:188-195.
Khler IH,Macdonald AJ,Schnyder H.2016.Last-century increases in intrinsic water-use ef ficiency of grassland communities have occurred over a wide range of vegetation composition,nutrient inputs,and soil p H.Plant Physiology,170:881-890.
Mantlana K,Arneth A,Veenendaal E,et al.2008.Photosynthetic properties of C4plants growing in an African savanna/wetland mosaic.Journal of Experimental Botany,59:3941-3952.
Murata N,Nishiyama Y.2018.ATP is a driving force in the repair of photosystem II during photoinhibition.Plant,Cell&Environment,41:285-299.
Nakamura S,Izumi M.2018.Regulation of chlorophagy during photoinhibition and senescence:Lessons from mitophagy.Plant&Cell Physiology,59:1135-1143.
Niyogi KK,Truong TB.2013.Evolution of flexible non-photochemical quenching mechanisms that regulate light harvesting in oxygenic photosynthesis.Current Opinion in Plant Biology,16:307-314.
Urban O,Hlavcov M,Klem K,et al.2018.Combined effects of drought and high temperature on photosynthetic characteristics in four winter wheat genotypes.Field Crops Research,223:137-149.
Ye ZP,Suggett JD,Robakowski P,et al.2013.A mechanistic model for the photosynthesis-light response based on the photosynthetic electron transport of PS II in C3 and C4 species.New Phytologist,152:1251-1262.
Ye ZP,Yu Q.2008.A coupled model of stomatal conductance and photosynthesis for winter wheat.Photosynthetica,46:637-640.
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