玉米叶片水分利用效率的保守性
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摘要
水分利用效率是植物个体或生态系统水分利用过程的重要特征参数,可表征不同时空尺度的植物碳-水耦合关系,对植物适应气候变化研究具有重要意义。以玉米为例,利用中国气象局固城农业气象野外科学试验基地2013—2014年玉米不同灌溉方案模拟试验资料,对不同叶位叶片的水分利用效率特征及其影响因素进行分析。结果表明:植株顶部第1片叶片水分利用效率在拔节期和乳熟期呈现明显的峰值,反映出明显的周期变化规律及其与叶片生理生态特征的紧密相关。在相同环境条件下,不同叶位叶片的水分利用效率不存在显著性差异,即玉米叶片水分利用效率具有空间稳定性与叶龄保守性。同时,研究指出叶片光合速率和蒸腾速率在叶位之间的协调变化是导致空间稳定性和叶龄保守性的主要原因。研究结果可为植物水分关系研究提供参考,也可为水分利用效率的尺度化研究提供依据。
Water use efficiency(WUE), which is the ratio of CO_2 assimilation to water loss, has been recognized as an important parameter in water use of plant individuals or ecosystems. As a classic index of carbon-water coupling at various temporal and spatial scales, from leaves to ecosystems, it is useful to determine the adaptation of plants to climate change. The North China Plain is the main maize(Zea mays L.) production region in China, where water is the key resource for optimum crop production. Leaf water use efficiency is defined as the ratio of P_n to T_r, which is mostly affected by biological and environmental factors. Although many findings about leaf water use efficiency have been published previously, limited attention has been focused on the water use efficiency relationship between leaves from the same individual plant under water stress during the whole growth period. Therefore, to understand changes of water use efficiency among different leaf positions at the leaf scale and investigate the main factors controlling leaf water use efficiency under water stress, an experiment with different water treatments(only one time irrigation, water controlled at seven and three leaf stages in 2013 and 2014, respectively) for maize was designed and conducted at the Gucheng Agro-meteorological Field Scientific Experiment Base, China Meteorological Administration(115°40′E, 39°08′N) in northern China. The amount of water irrigated in the experiment was in accordance with the total average precipitation during late July and July during 1980—2010 separately in 2013 and 2014. The results showed that there was an obvious pattern for the first opened leaf′s water use efficiency during progressive water stress, as it had two peaks at the jolting and milking stage. Then, leaf water use efficiency in the later growth stage was higher than that in the earlier stage; moderate water deficit improves water use efficiency, but the quantitative relationship between soil water availability and plant water use efficiency needs further investigation. Moreover, leaf water use efficiency was closely connected to its physiological and ecological characteristics, such as leaf water content and specific leaf area. Furthermore, leaf water use efficiency among different leaf positions along the maize plant showed little variation under nearly identical environments, such as soil water content and atmospheric conditions, which means leaf water use efficiency had spatial stability or conservation regardless of leaf ages to some degree. Meanwhile, the results also showed the coordinative changes between the photosynthetic and transpiration rates had mostly contributed to the leaf water use efficiency stability at different leaf positions. The findings provide information for plant-water relationship research and facilitate a mechanistic understanding of the carbon-water relationships at a leaf scale under water stress conditions, and lay a foundation for and up-scaled study of water use efficiency.
Water use efficiency(WUE), which is the ratio of CO_2 assimilation to water loss, has been recognized as an important parameter in water use of plant individuals or ecosystems. As a classic index of carbon-water coupling at various temporal and spatial scales, from leaves to ecosystems, it is useful to determine the adaptation of plants to climate change. The North China Plain is the main maize(Zea mays L.) production region in China, where water is the key resource for optimum crop production. Leaf water use efficiency is defined as the ratio of P_n to T_r, which is mostly affected by biological and environmental factors. Although many findings about leaf water use efficiency have been published previously, limited attention has been focused on the water use efficiency relationship between leaves from the same individual plant under water stress during the whole growth period. Therefore, to understand changes of water use efficiency among different leaf positions at the leaf scale and investigate the main factors controlling leaf water use efficiency under water stress, an experiment with different water treatments(only one time irrigation, water controlled at seven and three leaf stages in 2013 and 2014, respectively) for maize was designed and conducted at the Gucheng Agro-meteorological Field Scientific Experiment Base, China Meteorological Administration(115°40′E, 39°08′N) in northern China. The amount of water irrigated in the experiment was in accordance with the total average precipitation during late July and July during 1980—2010 separately in 2013 and 2014. The results showed that there was an obvious pattern for the first opened leaf′s water use efficiency during progressive water stress, as it had two peaks at the jolting and milking stage. Then, leaf water use efficiency in the later growth stage was higher than that in the earlier stage; moderate water deficit improves water use efficiency, but the quantitative relationship between soil water availability and plant water use efficiency needs further investigation. Moreover, leaf water use efficiency was closely connected to its physiological and ecological characteristics, such as leaf water content and specific leaf area. Furthermore, leaf water use efficiency among different leaf positions along the maize plant showed little variation under nearly identical environments, such as soil water content and atmospheric conditions, which means leaf water use efficiency had spatial stability or conservation regardless of leaf ages to some degree. Meanwhile, the results also showed the coordinative changes between the photosynthetic and transpiration rates had mostly contributed to the leaf water use efficiency stability at different leaf positions. The findings provide information for plant-water relationship research and facilitate a mechanistic understanding of the carbon-water relationships at a leaf scale under water stress conditions, and lay a foundation for and up-scaled study of water use efficiency.
引文
[1] Cowan I R,Farquhar G D.Stomatal function in relation to leaf metabolism and environment.Symposia of the Society for Experimental Biology,1997,31:471- 505.
[2] Farquhar G D,Richards R A.Isotopic composition of plant carbon correlates with water-use efficiency of wheat genotypes.Australian Journal of Plant Physiology,1984,11(6):539- 552.
[3] Sinclair T R,Tanner C B,Bennett J M.Water-use efficiency in crop production.BioScience,1984,34(1):36- 40.
[4] Farquhar G D,Hubick K T,Condon A G,Richards R A.Carbon isotope fractionation and plant water-use efficiency//Rundel P W,Ehleringer J R,Nagy K A,eds.Stable Isotopes in Ecological Research.New York:Springer,1989:21- 40.
[5] Beer C,Ciais P,Reichstein M,Baldocchi D,Law B E,Papale D,Soussana J F,Ammann C,Buchmann N,Frank D,Gianelle D,Janssens I A,Knohl A,K?stner B,Moors E,Roupsard O,Verbeeck H,Vesala T,Williams C A,Wohlfahrt G.Temporal and among-site variability of inherent water use efficiency at the ecosystem level.Global Biogeochemical Cycles,2009,23(2):GB2018.
[6] Sun Y,Piao S L,Huang M T,Ciais P,Zeng Z Z,Cheng L,Li X R,Zhang X P,Mao J F,Peng S S,Poulter B,Shi X Y,Wang X H,Wang Y P,Zeng H.Global patterns and climate drivers of water-use efficiency in terrestrial ecosystems deduced from satellite-based datasets and carbon cycle models.Global Ecology and Biogeography,2016,25(3):311- 323.
[7] Niu S L,Xing X R,Zhang Z,Xia J Y,Zhou X H,Song B,Li L H,Wan S Q.Water-use efficiency in response to climate change:from leaf to ecosystem in a temperate steppe.Global Change Biology,2011,17(2):1073- 1082.
[8] Hsiao T C,Acevedo E.Plant responses to water deficits,water-use efficiency,and drought resistance.Agricultural Meteorology,1974,14(1/2):59- 84.
[9] Liu C C,He N P,Zhang J H,Li Y,Wang Q F,Sack L,Yu G R.Variation of stomatal traits from cold temperate to tropical forests and association with water use efficiency.Functional Ecology,2017,32(1):20- 28.
[10] Tolk J A,Evett S R,Xu W W,Schwartz R C.Constraints on water use efficiency of drought tolerant maize grown in a semi-arid environment.Field Crops Research,2016,186:66- 77.
[11] Boese S,Jung M,Carvalhais N,Reichstein M.The importance of radiation for semiempirical water-use efficiency models.Biogeosciences,2017,14(12):3015- 3026.
[12] Drake B G,Gonzàlez-Meler M A,Long S P.More efficient plants:a consequence of rising atmospheric CO2?Annual Review of Plant Physiology and Plant Molecular Biology,1997,48:609- 639.
[13] Leakey A D B,Bernacchi C J,Dohleman F G,Ort D R,Long S P.Will photosynthesis of maize (Zea mays) in the US corn belt increase in future [CO2] rich atmospheres?An analysis of diurnal courses of CO2 uptake under free-air concentration enrichment (FACE).Global Change Biology,2004,10(6):951- 962.
[14] Eamus D.The interaction of rising CO2 and temperatures with water use efficiency.Plant Cell & Environment,1991,14(8):843- 852.
[15] Schymanski S J,Or D.Wind increases leaf water use efficiency.Plant,Cell & Environment,2016,39(7):1448- 1459.
[16] Ben-Asher J,Garcia y Garcia A,Hoogenboom G.Effect of high temperature on photosynthesis and transpiration of sweet corn (Zea mays L.var.rugosa).Photosynthetica,2008,46(4):595- 603.
[17] Kang S Z,Zhang F C,Hu X T,Zhang J H.Benefits of CO2 enrichment on crop plants are modified by soil water status.Plant and Soil,2002,238(1):69- 77.
[18] Way D A,Katul G G,Manzoni S,Vico G.Increasing water use efficiency along the C3 to C4 evolutionary pathway:a stomatal optimization perspective.Journal of Experimental Botany,2014,65(13):3683- 3693.
[19] 于晓芳,高聚林,宋国栋,冯勇,赵瑞霞,孙峰成,王志刚,李丽君.玉米叶片水分利用效率及其相关性状的研究.玉米科学,2008,16(3):64- 69.
[20] Field C,Merino J,Mooney H A.Compromises between water-use efficiency and nitrogen-use efficiency in five species of California evergreens.Oecologia,1983,60(3):384- 389.
[21] Wieser G,Oberhuber W,Gruber A,Leo M,Matyssek R,Grams T E E.Stable water use efficiency under climate change of three sympatric conifer species at the alpine treeline.Frontiers in Plant Science,2016,7:799.
[22] Wellstein C,Poschlod P,Gohlke A,Chelli S,Campetella G,Rosbakh S,Canullo R,Kreyling J,Jentsch A,Beierkuhnlein C.Effects of extreme drought on specific leaf area of grassland species:a meta-analysis of experimental studies in temperate and sub-Mediterranean systems.Global Change Biology,2017,23(6):2473- 2481.
[23] Zhao W S,Sun Y L,Kjelgren R,Liu X P.Response of stomatal density and bound gas exchange in leaves of maize to soil water deficit.Acta Physiologiae Plantarum,2015,37(1):1704.
[24] Ainsworth E A,Rogers A.The response of photosynthesis and stomatal conductance to rising [CO2]:mechanisms and environmental interactions.Plant,Cell & Environment,2007,30(3):258- 270.
[25] Lawson T,Blatt M R.Stomatal size,speed,and responsiveness impact on photosynthesis and water use efficiency.Plant Physiology,2014,164(4):1556- 1570.
[26] Fang S B,Su H,Liu W,Tan K Y,Ren S X.Infrared warming reduced winter wheat yields and some physiological parameters,which were mitigated by irrigation and worsened by delayed sowing.PLoS One,2013,8(7):e67518.
[27] Francis C A,Rutger J N,Palmer A F E.A rapid method for plant leaf area estimation in maize (Zea mays L.).Crop Science,1969,9(5):537- 539.
[28] 于文颖,纪瑞鹏,冯锐,赵先丽,张玉书.不同生育期玉米叶片光合特性及水分利用效率对水分胁迫的响应.生态学报,2015,35(9):2902- 2909.
[29] Tuzet A,Perrier A,Leuning R.A coupled model of stomatal conductance,photosynthesis and transpiration.Plant,Cell & Environment,2003,26(7):1097- 1116.
[30] 严昌荣,韩兴国,陈灵芝.六种木本植物水分利用效率和其小生境关系研究.生态学报,2011,21(11):1952- 1956.
[31] 赵明,李少昆,王美云.影响玉米单叶水分利用效率内在因素的研究.中国农业大学学报,1997,2(1):89- 94.
[32] Wang R H,Chang J C,Li K T,Lin T S,Chang L S.Leaf age and light intensity affect gas exchange parameters and photosynthesis within the developing canopy of field net-house-grown papaya trees.Scientia Horticulturae,2014,165:365- 373.
[33] Constable G A,Rawson H M.Effect of leaf position,expansion and age on photosynthesis,transpiration and water use efficiency of cotton.Australian Journal of Plant Physiology,1980,7(1):89- 100.
[34] 刘莹,李鹏,沈冰,冯朝红,刘琦,张祎.采用稳定碳同位素法分析白羊草在不同干旱胁迫下的水分利用效率.生态学报,2017,37(9):3055- 3064.
[35] Kang S Z,Shi W J,Zhang J H.An improved water-use efficiency for maize grown under regulated deficit irrigation.Field Crops Research,2000,67(3):207- 214.
[36] Wang Y Q,Zhang Y H,Zhang R,Li J P,Zhang M,Zhou S L,Wang Z M.Reduced irrigation increases the water use efficiency and productivity of winter wheat-summer maize rotation on the North China Plain.Science of the Total Environment,2018,618:112- 120.
[37] Peňuelas J,Canadell J G,Ogaya R.Increased water-use efficiency during the 20th century did not translate into enhanced tree growth.Global Ecology and Biogeography,2011,20(4):597- 608.
[38] Bernacchi C J,VanLoocke A.Terrestrial ecosystems in a changing environment:a dominant role for water.Annual Review of Plant Biology,2015,66:599- 622.
[39] Flanagan L B,Farquhar G D.Variation in the carbon and oxygen isotope composition of plant biomass and its relationship to water-use efficiency at the leaf-and ecosystem-scales in a northern Great Plains grassland.Plant,Cell & Environment,2014,37(2):425- 438.
[40] Medlyn B E,De Kauwe M G,Lin Y S,Knauer J,Duursma R A,Williams C A,Arneth A,Clement R,Isaac P,Limousin J M,Linderson M L,Meir P,Martin-StPaul N,Wingate L.How do leaf and ecosystem measures of water-use efficiency compare?New Phytologist,2017,216(3):758- 770.
[2] Farquhar G D,Richards R A.Isotopic composition of plant carbon correlates with water-use efficiency of wheat genotypes.Australian Journal of Plant Physiology,1984,11(6):539- 552.
[3] Sinclair T R,Tanner C B,Bennett J M.Water-use efficiency in crop production.BioScience,1984,34(1):36- 40.
[4] Farquhar G D,Hubick K T,Condon A G,Richards R A.Carbon isotope fractionation and plant water-use efficiency//Rundel P W,Ehleringer J R,Nagy K A,eds.Stable Isotopes in Ecological Research.New York:Springer,1989:21- 40.
[5] Beer C,Ciais P,Reichstein M,Baldocchi D,Law B E,Papale D,Soussana J F,Ammann C,Buchmann N,Frank D,Gianelle D,Janssens I A,Knohl A,K?stner B,Moors E,Roupsard O,Verbeeck H,Vesala T,Williams C A,Wohlfahrt G.Temporal and among-site variability of inherent water use efficiency at the ecosystem level.Global Biogeochemical Cycles,2009,23(2):GB2018.
[6] Sun Y,Piao S L,Huang M T,Ciais P,Zeng Z Z,Cheng L,Li X R,Zhang X P,Mao J F,Peng S S,Poulter B,Shi X Y,Wang X H,Wang Y P,Zeng H.Global patterns and climate drivers of water-use efficiency in terrestrial ecosystems deduced from satellite-based datasets and carbon cycle models.Global Ecology and Biogeography,2016,25(3):311- 323.
[7] Niu S L,Xing X R,Zhang Z,Xia J Y,Zhou X H,Song B,Li L H,Wan S Q.Water-use efficiency in response to climate change:from leaf to ecosystem in a temperate steppe.Global Change Biology,2011,17(2):1073- 1082.
[8] Hsiao T C,Acevedo E.Plant responses to water deficits,water-use efficiency,and drought resistance.Agricultural Meteorology,1974,14(1/2):59- 84.
[9] Liu C C,He N P,Zhang J H,Li Y,Wang Q F,Sack L,Yu G R.Variation of stomatal traits from cold temperate to tropical forests and association with water use efficiency.Functional Ecology,2017,32(1):20- 28.
[10] Tolk J A,Evett S R,Xu W W,Schwartz R C.Constraints on water use efficiency of drought tolerant maize grown in a semi-arid environment.Field Crops Research,2016,186:66- 77.
[11] Boese S,Jung M,Carvalhais N,Reichstein M.The importance of radiation for semiempirical water-use efficiency models.Biogeosciences,2017,14(12):3015- 3026.
[12] Drake B G,Gonzàlez-Meler M A,Long S P.More efficient plants:a consequence of rising atmospheric CO2?Annual Review of Plant Physiology and Plant Molecular Biology,1997,48:609- 639.
[13] Leakey A D B,Bernacchi C J,Dohleman F G,Ort D R,Long S P.Will photosynthesis of maize (Zea mays) in the US corn belt increase in future [CO2] rich atmospheres?An analysis of diurnal courses of CO2 uptake under free-air concentration enrichment (FACE).Global Change Biology,2004,10(6):951- 962.
[14] Eamus D.The interaction of rising CO2 and temperatures with water use efficiency.Plant Cell & Environment,1991,14(8):843- 852.
[15] Schymanski S J,Or D.Wind increases leaf water use efficiency.Plant,Cell & Environment,2016,39(7):1448- 1459.
[16] Ben-Asher J,Garcia y Garcia A,Hoogenboom G.Effect of high temperature on photosynthesis and transpiration of sweet corn (Zea mays L.var.rugosa).Photosynthetica,2008,46(4):595- 603.
[17] Kang S Z,Zhang F C,Hu X T,Zhang J H.Benefits of CO2 enrichment on crop plants are modified by soil water status.Plant and Soil,2002,238(1):69- 77.
[18] Way D A,Katul G G,Manzoni S,Vico G.Increasing water use efficiency along the C3 to C4 evolutionary pathway:a stomatal optimization perspective.Journal of Experimental Botany,2014,65(13):3683- 3693.
[19] 于晓芳,高聚林,宋国栋,冯勇,赵瑞霞,孙峰成,王志刚,李丽君.玉米叶片水分利用效率及其相关性状的研究.玉米科学,2008,16(3):64- 69.
[20] Field C,Merino J,Mooney H A.Compromises between water-use efficiency and nitrogen-use efficiency in five species of California evergreens.Oecologia,1983,60(3):384- 389.
[21] Wieser G,Oberhuber W,Gruber A,Leo M,Matyssek R,Grams T E E.Stable water use efficiency under climate change of three sympatric conifer species at the alpine treeline.Frontiers in Plant Science,2016,7:799.
[22] Wellstein C,Poschlod P,Gohlke A,Chelli S,Campetella G,Rosbakh S,Canullo R,Kreyling J,Jentsch A,Beierkuhnlein C.Effects of extreme drought on specific leaf area of grassland species:a meta-analysis of experimental studies in temperate and sub-Mediterranean systems.Global Change Biology,2017,23(6):2473- 2481.
[23] Zhao W S,Sun Y L,Kjelgren R,Liu X P.Response of stomatal density and bound gas exchange in leaves of maize to soil water deficit.Acta Physiologiae Plantarum,2015,37(1):1704.
[24] Ainsworth E A,Rogers A.The response of photosynthesis and stomatal conductance to rising [CO2]:mechanisms and environmental interactions.Plant,Cell & Environment,2007,30(3):258- 270.
[25] Lawson T,Blatt M R.Stomatal size,speed,and responsiveness impact on photosynthesis and water use efficiency.Plant Physiology,2014,164(4):1556- 1570.
[26] Fang S B,Su H,Liu W,Tan K Y,Ren S X.Infrared warming reduced winter wheat yields and some physiological parameters,which were mitigated by irrigation and worsened by delayed sowing.PLoS One,2013,8(7):e67518.
[27] Francis C A,Rutger J N,Palmer A F E.A rapid method for plant leaf area estimation in maize (Zea mays L.).Crop Science,1969,9(5):537- 539.
[28] 于文颖,纪瑞鹏,冯锐,赵先丽,张玉书.不同生育期玉米叶片光合特性及水分利用效率对水分胁迫的响应.生态学报,2015,35(9):2902- 2909.
[29] Tuzet A,Perrier A,Leuning R.A coupled model of stomatal conductance,photosynthesis and transpiration.Plant,Cell & Environment,2003,26(7):1097- 1116.
[30] 严昌荣,韩兴国,陈灵芝.六种木本植物水分利用效率和其小生境关系研究.生态学报,2011,21(11):1952- 1956.
[31] 赵明,李少昆,王美云.影响玉米单叶水分利用效率内在因素的研究.中国农业大学学报,1997,2(1):89- 94.
[32] Wang R H,Chang J C,Li K T,Lin T S,Chang L S.Leaf age and light intensity affect gas exchange parameters and photosynthesis within the developing canopy of field net-house-grown papaya trees.Scientia Horticulturae,2014,165:365- 373.
[33] Constable G A,Rawson H M.Effect of leaf position,expansion and age on photosynthesis,transpiration and water use efficiency of cotton.Australian Journal of Plant Physiology,1980,7(1):89- 100.
[34] 刘莹,李鹏,沈冰,冯朝红,刘琦,张祎.采用稳定碳同位素法分析白羊草在不同干旱胁迫下的水分利用效率.生态学报,2017,37(9):3055- 3064.
[35] Kang S Z,Shi W J,Zhang J H.An improved water-use efficiency for maize grown under regulated deficit irrigation.Field Crops Research,2000,67(3):207- 214.
[36] Wang Y Q,Zhang Y H,Zhang R,Li J P,Zhang M,Zhou S L,Wang Z M.Reduced irrigation increases the water use efficiency and productivity of winter wheat-summer maize rotation on the North China Plain.Science of the Total Environment,2018,618:112- 120.
[37] Peňuelas J,Canadell J G,Ogaya R.Increased water-use efficiency during the 20th century did not translate into enhanced tree growth.Global Ecology and Biogeography,2011,20(4):597- 608.
[38] Bernacchi C J,VanLoocke A.Terrestrial ecosystems in a changing environment:a dominant role for water.Annual Review of Plant Biology,2015,66:599- 622.
[39] Flanagan L B,Farquhar G D.Variation in the carbon and oxygen isotope composition of plant biomass and its relationship to water-use efficiency at the leaf-and ecosystem-scales in a northern Great Plains grassland.Plant,Cell & Environment,2014,37(2):425- 438.
[40] Medlyn B E,De Kauwe M G,Lin Y S,Knauer J,Duursma R A,Williams C A,Arneth A,Clement R,Isaac P,Limousin J M,Linderson M L,Meir P,Martin-StPaul N,Wingate L.How do leaf and ecosystem measures of water-use efficiency compare?New Phytologist,2017,216(3):758- 770.