天山高寒草原植物叶片氮磷化学计量特征对氮沉降的响应
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摘要
人类活动和气候变化导致的大气氮沉降急剧升高,将会影响高寒草原植物养分循环。为了研究氮沉降对高寒草原植物养分含量及生态化学计量学特征的影响。依托2009年在新疆天山巴音布鲁克高寒草原设置的长期氮沉降研究平台,于2017年7月和10月采集溚草(Koeleria cristata)、二裂委陵菜(Potentilla bifurca)和天山赖草(Leymus tianschanicus) 3种植物的成熟叶和老叶,分析3种植物叶片N、P元素生态化学计量对不同施氮水平的响应规律。结果表明:①3种植物养分浓度及其化学计量比在成熟叶和老叶之间存在较大差异,成熟叶和老叶的N、P含量分别为12. 78 mg·g~(-1)、0. 88 mg·g~(-1)和6. 88 mg·g~(-1)、0. 46 mg·g~(-1)。3种植物成熟叶和老叶的N元素含量均为:二裂委陵菜>天山赖草>溚草,P元素含量同N元素含量趋势一致;②氮添加增加了成熟叶和老叶N浓度,分别达12%和48%;成熟叶与老叶P浓度对氮添加的响应规律不一致,成熟叶P浓度对氮添加的响应与物种有关,而老叶P浓度呈一致的下降趋势;氮添加对成熟叶和老叶N∶P的比值影响亦不同,其中相对于成熟叶,老叶的N∶P呈显著增加趋势。③植物叶片的N元素、P元素和N∶P比三者之间的相关分析表明:N元素和N∶P之间呈显著正相关关系(P <0. 05),在生长旺盛期和枯黄期,溚草和天山赖草的N与P元素之间没有发现显著相关性(P>0. 05),而二裂委陵菜呈现一定的差异性,在生长旺盛期二裂委陵菜叶片N与P元素呈显著的正相关、枯黄期呈显著的负相关(P <0. 05)。表明长期氮的添加显著影响了3种植物叶片的养分含量和生态化学计量,不同物种间存在一定差异,未来氮沉降升高将会影响天山山地草原的植物养分循环。
Atmospheric nitrogen( N) deposition is rapidly increased due to the increase of human activities and climate change,which enhances the N availability and affects the nutrient cycling of plants in alpine grasslands.However,there is still much debate about how the change of nutrient availability affects P content in plants,therefore,the effects of N addition on N and P contents and N∶ P stoichiometry in mature and senesced leaves at plant organ level in the Bayinbuluk alpine grasslands in the northwest Tianshanin Mountains in China were examined based on a 10-year field experiment. The nitrogen and phosphorus contents in mature and senesced leaves of Potentilla bifurca,Leymus tianschanicus and Koeleria cristata were measured,and the values of foliar stoichiometry under N addition treatments at levels of 0,3,9 g·m~(-2)·a~(-1)) were estimated. Results showed that there were the significant differences of the values of nutrient concentration and foliar stoichiometry between the mature and senesced leaves.The N contents in mature and senesced leaves were 12. 78 mg·g~(-1) and 6. 88 mg·g~(-1),and the P contents were0. 88 mg·g~(-1) and 0. 46 mg·g~(-1),respectively. N and P concentrations in mature and senesced leaves were in an order of P. bifurca > L. tiansecalinus > K. cristata. After the N treatments for 10 years,the N contents in mature and senesced leaves were increased by 12% and 48% respectively. It was found that there was a divergent response of P content in leaves,the effect of N addition on foliar P content was different from different plant species,and the P content in senesced leaves was decreased by N addition. N∶ P in senesced leaves was increased significantly but not in mature leaves. In addition,the correlations between N and P elements and plant stoichiometry in mature and senesced leaves were different. There was a positive correlation between foliar N and P contents of P. bifurca in mature leaves( P < 0. 05),but a negative correlation in senesced leaves. Our finds revealed that nitrogen addition affected significantly the nutrient content and stoichiometry in leaves of the three species,but there was a significant difference among species. The increase of N deposition would affect the nutrient cycling of plants in grasslands in the Tianshan Mountains.
Atmospheric nitrogen( N) deposition is rapidly increased due to the increase of human activities and climate change,which enhances the N availability and affects the nutrient cycling of plants in alpine grasslands.However,there is still much debate about how the change of nutrient availability affects P content in plants,therefore,the effects of N addition on N and P contents and N∶ P stoichiometry in mature and senesced leaves at plant organ level in the Bayinbuluk alpine grasslands in the northwest Tianshanin Mountains in China were examined based on a 10-year field experiment. The nitrogen and phosphorus contents in mature and senesced leaves of Potentilla bifurca,Leymus tianschanicus and Koeleria cristata were measured,and the values of foliar stoichiometry under N addition treatments at levels of 0,3,9 g·m~(-2)·a~(-1)) were estimated. Results showed that there were the significant differences of the values of nutrient concentration and foliar stoichiometry between the mature and senesced leaves.The N contents in mature and senesced leaves were 12. 78 mg·g~(-1) and 6. 88 mg·g~(-1),and the P contents were0. 88 mg·g~(-1) and 0. 46 mg·g~(-1),respectively. N and P concentrations in mature and senesced leaves were in an order of P. bifurca > L. tiansecalinus > K. cristata. After the N treatments for 10 years,the N contents in mature and senesced leaves were increased by 12% and 48% respectively. It was found that there was a divergent response of P content in leaves,the effect of N addition on foliar P content was different from different plant species,and the P content in senesced leaves was decreased by N addition. N∶ P in senesced leaves was increased significantly but not in mature leaves. In addition,the correlations between N and P elements and plant stoichiometry in mature and senesced leaves were different. There was a positive correlation between foliar N and P contents of P. bifurca in mature leaves( P < 0. 05),but a negative correlation in senesced leaves. Our finds revealed that nitrogen addition affected significantly the nutrient content and stoichiometry in leaves of the three species,but there was a significant difference among species. The increase of N deposition would affect the nutrient cycling of plants in grasslands in the Tianshan Mountains.
引文
[1]Sterner R W,Elser J J.Ecological stoichiometry:The biology of elements from molecules to the biosphere[J].Journal of Plankton Research 2003,25(9):1 183.
[2]Venterink H O,Wassen M J,Verkroost A W M,et al.Species richness-productivity patterns differ between N-,P-,and K-limited wetlands[J].Ecology,2003,84(8):2 191-2 199.
[3]庄晓玲,李凯辉,王晓丽,等.新疆两种不同类型农田生态系统大气氮素干沉降分析[J].干旱区研究,2017,34(6):1 395-1 401.[Zhuang Xiaoling,Li Kaihui,Wang Xiaoli,et al.Atmospheric nitrogen dry deposition in two agroecosystems in Xinjiang[J].Arid Zone Research,2017,34(6):1 395-1 401.]
[4]Zhan S X,Wang Y,Zhu Z C,et al.Nitrogen enrichment alters plant N∶P stoichiometry and intensifies phosphorus limitation in a steppe ecosystem[J].Environmental and Experimental Botany,2017,134:21-32.
[5]Huang W J,Zhou G Y,Liu J X,et al.Effects of elevated carbon dioxide and nitrogen addition on foliar stoichiometry of nitrogen and phosphorus of five tree species in subtropical model forest ecosystems[J].Environmental Pollution,2012,168:113-120.
[6]LüX T,Kong D L,Pan Q M,et al.Nitrogen and water availability interact to affect leaf stoichiometry in a semi-arid grassland[J].Oecologia,2012,168:301-310.
[7]Li L,Gao X P,Li X Y,et al.Nitrogen(N)and phosphorus(P)resorption of two dominant alpine perennial grass species in response to contrasting N and P availability[J].Environmental and Experimental Botany,2016,127:37-44
[8]Yuan Z Y,Chen H Y.Decoupling of nitrogen and phosphorus in terrestrial plants associated with global changes[J].Nature Climate Change,2015,5(5):465-469.
[9]Rowe E C,Smart S M,Kennedy V H,et al.Nitrogen deposition increases the acquisition of phosphorus and potassium by heather Calluna vulgaris[J].Environmental Pollution,2008,155(2):201-207.
[10]Wang X G,LüX T,Han X G.Responses of nutrient concentrations and stoichiometry of senesced leaves in dominant plants to nitrogen addition and prescribed burning in a temperate steppe[J].Ecological Engineering,2014,70:154-161.
[11]Erschbamer B,Unterluggauer P,Winkler E,et al.Changes in plant species diversity revealed by long-term monitoring on mountain summits in the Dolomites(Northern Italy)[J].Preslia Praha,2011,83(3):387-401.
[12]岳平,宋韦,李凯辉,等.天山中部巴音布鲁克高寒草原大气无机氮沉降[J].应用生态学报,2014,25(6):1 592-1 598.[Yue Ping,Song Wei,Li Kaihui,et al.Inorganic N deposition in the Bayinbuluk alpine grassland of the central Tian shan Mountains[J].Chinese Journal of Applied Ecology,2014,25(6):1 592-1 598.]
[13]Liu Y Y,Taxipulati T,Gong Y M,et al.N-P fertilization inhibits growth of root hemiparasite Pedicularis kansuensis in Natural Grassland[J].Frontiers in Plant Science,2017,8:2 088.
[14]Li K H,Liu X J,Song L,et al.Response of alpine grassland to elevated nitrogen deposition and water supply in China[J].Oecologia,2015,177(1):65-72.
[15]张珂,陈永乐,高艳红,等.阿拉善荒漠典型植物功能群氮、磷化学计量特征[J].中国沙漠,2014,34(5):1 261-1 267.[Zhang Ke,Chen Yongle,Gao Yanhong,et al.Stoichiometric characteristics of leaf nitrogen and phosphorus of different plant functional groups in Alashan desert region[J].Journal of Desert Research,2014,34(5):1 261-1 267.]
[16]李玉霖,崔夺,陈静,等.科尔沁沙质草地优势多年生植物氮素回收效率的分异特征[J].中国沙漠,2013,33(3):688-695.[Li Yulin,Cui Duo,Chen Jing,et al.Divergence of nitrogen resorption efficiency of dominant perennial plants in Horqin sandy grassland[J].Journal of Desert Research,2013,33(3):688-695.]
[17]Carrera A L,Sain C L,Bertiller M B.Patterns of nitrogen conservation in shrubs and grasses in the Patagonian Monte,Argentina[J].Plant and Soil,2000,224(2):185-193.
[18]Gusewell S.N∶P ratios in terrestrial plants:Variation and functional significance[J].New Phytologist,2004,164(2):243-266.
[19]Yu Q,Chen Q S,Elser J J,et al.Linking stoichiometry homeostasis with ecosystem structure,functioning and stability[J].Ecology Letters,2010,13(11):1 390-1 399.
[20]Li X,Liu J,Fan J,et al.Combined effects of nitrogen addition and litter manipulation on nutrient resorption of Leymus chinensis in a semi-arid grassland of northern China[J].Plant Biology,2015,17(1):9-15.
[21]Cui Q,LüX T,Wang Q B,et al.Nitrogen fertilization and fire act independently on foliar stoichiometry in a temperate steppe[J].Plant and Soil,2010,334(1-2):209-219.
[22]LüX T,Cui Q,Wang Q B,et al.Nutrient resorption response to fire and nitrogen addition in a semiarid grassland[J].Ecological Engineering,2011,37(3):534-538
[23]Kobe R K,Lepczyk C A,Iyer M.Resorption efficiency decreases with increasing green leaf nutrients in a global data set[J].Ecology,2005,86(10):2 780-2 792.
[24]Menge D N L,Field C B.Simulated global changes alter phosphorus demand in annual grassland[J].Global Change Biology,2007,13(12):2 582-2 591.
[25]Phoenix G K,Booth R E,Leake J R,et al.Effects of enhanced nitrogen deposition and phosphorus limitation on nitrogen budgets of semi-natural grasslands[J].Global Change Biology,2003,9(9):1 309-1 321.
[26]LüX T,Reed S,Yu Q,et al.Convergent responses of nitrogen and phosphorus resorption to nitrogen inputs in a semi-arid grassland[J].Global Change Biology,2013,19(9):2 775-2 784
[27]LüX T,Han X G.Nutrient resorption responses to water and nitrogen amendment in semi-arid grassland of Inner Mongolia,China[J].Plant and Soil,2010,27(1-2):481-491
[28]Güsewell S,Gessner M O.N∶P ratios influence litter decomposition and colonization by fungi and bacteria in microcosms[J].Functional Ecology,2009,23(1):211-219.
[29]Liu X J,Zhang Y,Han W X,et al.Enhanced nitrogen deposition over China[J].Nature,2013,494(7 438):459-562.
[30]Elser J J,Sterner R W,Gorokhova E,et al.Biological stoichiometry from genes to ecosystems[J].Ecology Letters,2010,3(6):540-550.
[31]Rong Q Q,Liu J T,Cai Y P,et al.Leaf carbon,nitrogen and phosphorus stoichiometry of Tamarix chinensis Lour.in the Laizhou Bay coastal wetland,China[J].Ecological Engineering,2015,76:57-65.
[32]Wang Z N,Lu J Y,Yang M,et al.Stoichiometric characteristics of carbon,nitrogen,and phosphorus in leaves of differently aged lucerne(Medicago sativa)stands[J].Frontiers in Plant Science,2015,6:1 062.
[33]李征,韩琳,刘玉虹,等.滨海盐地碱蓬不同生长阶段叶片C、N、P化学计量特征[J].植物生态学报,2012,36(10):1 054-1 061.[Li Zheng,Han Lin,Liu Yuhong,et al.C,N and P stoichiometric characteristics in leaves of Suaeda salsa during different growthphase in coastal wetlands of China[J].Chinese Journal of Plant Ecology,2012,36(10):1 054-1 061.]
[34]罗艳,贡璐,朱美玲,等.塔里木河上游荒漠区4种灌木植物叶片与土壤生态化学计量特征[J].生态学报,2017,37(24):8 326-8 335.[Luo Yan,Gong Lu,Zhu Meiling,et al.Stoichiometry characteristics of leaves and soil of four shrubs in the upper reaches of the TarimRiver Desert[J].Acta Ecologica Sinica,2017,37(24):8 326-8 335.]
[35]石贤萌,杞金华,宋亮,等.哀牢山中山湿性常绿阔叶林两种优势幼苗C、N、P化学计量特征及其对N沉降增加的响应[J].植物生态学报,2015,39(10):962-970.[Shi Xianmeng,Qi Jinhua,Song Liang,et al.C,N and P stoichiometry of two dominant seedlings and their responses to nitrogen additionsin the montane moist evergreen broad-leaved forest in Ailao Mountains,Yunnan[J].Chinese Journal of Plant Ecology,2015,39(10):962-970.]
[36]曾德慧,陈广生.生态化学计量学:复杂生命系统奥秘的探索[J].植物生态学报,2005,29(6):1 007-1 019.[Ceng Dehui,Chen Guangsheng.Ecological stoichiometry:A science to explore of the complexity of living systems[J].Chinese Journal of Plant Ecology,2005,29(6):1 007-1 019.]
[37]安卓,牛得草,文海燕,等.氮素添加对黄土高原典型草原长芒草氮磷重吸收率及C∶N∶P化学计量特征的影响[J].植物生态学报,2011,35(8):801-807.[An Zhuo,Niu Decao,Wen Haiyan,et al.Effects of N addition on nutrient resorption efficiency and C∶N∶P stoichiometric characteristics in Stipa bungeana of steppe grasslands in the Loess Plateau,China[J].Chinese Journal of Plant Ecology,2011,35(8):801-807.]
[2]Venterink H O,Wassen M J,Verkroost A W M,et al.Species richness-productivity patterns differ between N-,P-,and K-limited wetlands[J].Ecology,2003,84(8):2 191-2 199.
[3]庄晓玲,李凯辉,王晓丽,等.新疆两种不同类型农田生态系统大气氮素干沉降分析[J].干旱区研究,2017,34(6):1 395-1 401.[Zhuang Xiaoling,Li Kaihui,Wang Xiaoli,et al.Atmospheric nitrogen dry deposition in two agroecosystems in Xinjiang[J].Arid Zone Research,2017,34(6):1 395-1 401.]
[4]Zhan S X,Wang Y,Zhu Z C,et al.Nitrogen enrichment alters plant N∶P stoichiometry and intensifies phosphorus limitation in a steppe ecosystem[J].Environmental and Experimental Botany,2017,134:21-32.
[5]Huang W J,Zhou G Y,Liu J X,et al.Effects of elevated carbon dioxide and nitrogen addition on foliar stoichiometry of nitrogen and phosphorus of five tree species in subtropical model forest ecosystems[J].Environmental Pollution,2012,168:113-120.
[6]LüX T,Kong D L,Pan Q M,et al.Nitrogen and water availability interact to affect leaf stoichiometry in a semi-arid grassland[J].Oecologia,2012,168:301-310.
[7]Li L,Gao X P,Li X Y,et al.Nitrogen(N)and phosphorus(P)resorption of two dominant alpine perennial grass species in response to contrasting N and P availability[J].Environmental and Experimental Botany,2016,127:37-44
[8]Yuan Z Y,Chen H Y.Decoupling of nitrogen and phosphorus in terrestrial plants associated with global changes[J].Nature Climate Change,2015,5(5):465-469.
[9]Rowe E C,Smart S M,Kennedy V H,et al.Nitrogen deposition increases the acquisition of phosphorus and potassium by heather Calluna vulgaris[J].Environmental Pollution,2008,155(2):201-207.
[10]Wang X G,LüX T,Han X G.Responses of nutrient concentrations and stoichiometry of senesced leaves in dominant plants to nitrogen addition and prescribed burning in a temperate steppe[J].Ecological Engineering,2014,70:154-161.
[11]Erschbamer B,Unterluggauer P,Winkler E,et al.Changes in plant species diversity revealed by long-term monitoring on mountain summits in the Dolomites(Northern Italy)[J].Preslia Praha,2011,83(3):387-401.
[12]岳平,宋韦,李凯辉,等.天山中部巴音布鲁克高寒草原大气无机氮沉降[J].应用生态学报,2014,25(6):1 592-1 598.[Yue Ping,Song Wei,Li Kaihui,et al.Inorganic N deposition in the Bayinbuluk alpine grassland of the central Tian shan Mountains[J].Chinese Journal of Applied Ecology,2014,25(6):1 592-1 598.]
[13]Liu Y Y,Taxipulati T,Gong Y M,et al.N-P fertilization inhibits growth of root hemiparasite Pedicularis kansuensis in Natural Grassland[J].Frontiers in Plant Science,2017,8:2 088.
[14]Li K H,Liu X J,Song L,et al.Response of alpine grassland to elevated nitrogen deposition and water supply in China[J].Oecologia,2015,177(1):65-72.
[15]张珂,陈永乐,高艳红,等.阿拉善荒漠典型植物功能群氮、磷化学计量特征[J].中国沙漠,2014,34(5):1 261-1 267.[Zhang Ke,Chen Yongle,Gao Yanhong,et al.Stoichiometric characteristics of leaf nitrogen and phosphorus of different plant functional groups in Alashan desert region[J].Journal of Desert Research,2014,34(5):1 261-1 267.]
[16]李玉霖,崔夺,陈静,等.科尔沁沙质草地优势多年生植物氮素回收效率的分异特征[J].中国沙漠,2013,33(3):688-695.[Li Yulin,Cui Duo,Chen Jing,et al.Divergence of nitrogen resorption efficiency of dominant perennial plants in Horqin sandy grassland[J].Journal of Desert Research,2013,33(3):688-695.]
[17]Carrera A L,Sain C L,Bertiller M B.Patterns of nitrogen conservation in shrubs and grasses in the Patagonian Monte,Argentina[J].Plant and Soil,2000,224(2):185-193.
[18]Gusewell S.N∶P ratios in terrestrial plants:Variation and functional significance[J].New Phytologist,2004,164(2):243-266.
[19]Yu Q,Chen Q S,Elser J J,et al.Linking stoichiometry homeostasis with ecosystem structure,functioning and stability[J].Ecology Letters,2010,13(11):1 390-1 399.
[20]Li X,Liu J,Fan J,et al.Combined effects of nitrogen addition and litter manipulation on nutrient resorption of Leymus chinensis in a semi-arid grassland of northern China[J].Plant Biology,2015,17(1):9-15.
[21]Cui Q,LüX T,Wang Q B,et al.Nitrogen fertilization and fire act independently on foliar stoichiometry in a temperate steppe[J].Plant and Soil,2010,334(1-2):209-219.
[22]LüX T,Cui Q,Wang Q B,et al.Nutrient resorption response to fire and nitrogen addition in a semiarid grassland[J].Ecological Engineering,2011,37(3):534-538
[23]Kobe R K,Lepczyk C A,Iyer M.Resorption efficiency decreases with increasing green leaf nutrients in a global data set[J].Ecology,2005,86(10):2 780-2 792.
[24]Menge D N L,Field C B.Simulated global changes alter phosphorus demand in annual grassland[J].Global Change Biology,2007,13(12):2 582-2 591.
[25]Phoenix G K,Booth R E,Leake J R,et al.Effects of enhanced nitrogen deposition and phosphorus limitation on nitrogen budgets of semi-natural grasslands[J].Global Change Biology,2003,9(9):1 309-1 321.
[26]LüX T,Reed S,Yu Q,et al.Convergent responses of nitrogen and phosphorus resorption to nitrogen inputs in a semi-arid grassland[J].Global Change Biology,2013,19(9):2 775-2 784
[27]LüX T,Han X G.Nutrient resorption responses to water and nitrogen amendment in semi-arid grassland of Inner Mongolia,China[J].Plant and Soil,2010,27(1-2):481-491
[28]Güsewell S,Gessner M O.N∶P ratios influence litter decomposition and colonization by fungi and bacteria in microcosms[J].Functional Ecology,2009,23(1):211-219.
[29]Liu X J,Zhang Y,Han W X,et al.Enhanced nitrogen deposition over China[J].Nature,2013,494(7 438):459-562.
[30]Elser J J,Sterner R W,Gorokhova E,et al.Biological stoichiometry from genes to ecosystems[J].Ecology Letters,2010,3(6):540-550.
[31]Rong Q Q,Liu J T,Cai Y P,et al.Leaf carbon,nitrogen and phosphorus stoichiometry of Tamarix chinensis Lour.in the Laizhou Bay coastal wetland,China[J].Ecological Engineering,2015,76:57-65.
[32]Wang Z N,Lu J Y,Yang M,et al.Stoichiometric characteristics of carbon,nitrogen,and phosphorus in leaves of differently aged lucerne(Medicago sativa)stands[J].Frontiers in Plant Science,2015,6:1 062.
[33]李征,韩琳,刘玉虹,等.滨海盐地碱蓬不同生长阶段叶片C、N、P化学计量特征[J].植物生态学报,2012,36(10):1 054-1 061.[Li Zheng,Han Lin,Liu Yuhong,et al.C,N and P stoichiometric characteristics in leaves of Suaeda salsa during different growthphase in coastal wetlands of China[J].Chinese Journal of Plant Ecology,2012,36(10):1 054-1 061.]
[34]罗艳,贡璐,朱美玲,等.塔里木河上游荒漠区4种灌木植物叶片与土壤生态化学计量特征[J].生态学报,2017,37(24):8 326-8 335.[Luo Yan,Gong Lu,Zhu Meiling,et al.Stoichiometry characteristics of leaves and soil of four shrubs in the upper reaches of the TarimRiver Desert[J].Acta Ecologica Sinica,2017,37(24):8 326-8 335.]
[35]石贤萌,杞金华,宋亮,等.哀牢山中山湿性常绿阔叶林两种优势幼苗C、N、P化学计量特征及其对N沉降增加的响应[J].植物生态学报,2015,39(10):962-970.[Shi Xianmeng,Qi Jinhua,Song Liang,et al.C,N and P stoichiometry of two dominant seedlings and their responses to nitrogen additionsin the montane moist evergreen broad-leaved forest in Ailao Mountains,Yunnan[J].Chinese Journal of Plant Ecology,2015,39(10):962-970.]
[36]曾德慧,陈广生.生态化学计量学:复杂生命系统奥秘的探索[J].植物生态学报,2005,29(6):1 007-1 019.[Ceng Dehui,Chen Guangsheng.Ecological stoichiometry:A science to explore of the complexity of living systems[J].Chinese Journal of Plant Ecology,2005,29(6):1 007-1 019.]
[37]安卓,牛得草,文海燕,等.氮素添加对黄土高原典型草原长芒草氮磷重吸收率及C∶N∶P化学计量特征的影响[J].植物生态学报,2011,35(8):801-807.[An Zhuo,Niu Decao,Wen Haiyan,et al.Effects of N addition on nutrient resorption efficiency and C∶N∶P stoichiometric characteristics in Stipa bungeana of steppe grasslands in the Loess Plateau,China[J].Chinese Journal of Plant Ecology,2011,35(8):801-807.]
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