亚热带湿地松叶片多元素化学计量与养分回收对氮添加的短期响应
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摘要
在我国南方亚热带湿地松人工林设置了3个水平的野外氮添加控制试验(0、40、120kg N·hm~(-2)a(-1)),于2014和2015年生长季高峰期(7月底)和末期(10月底)采集湿地松成熟绿叶和落叶,分析外源氮添加对湿地松叶片碳(C)、氮(N)、磷(P)、钾(K)、钙(Ca)、镁(Mg)、铝(Al)、铁(Fe)、锰(Mn)9种元素浓度及其养分回收的影响.结果表明:N添加显著增加了湿地松绿叶中N、Al、Mn浓度,降低了P和2014年的Ca浓度,而对C、K、Mg、Fe浓度无显著影响.N添加显著提高了绿叶N/P,且该比值及绿叶养分浓度(N、P、Mn)对N添加的响应依赖于N的剂量(高N条件下响应更强).N添加显著降低了2015年N的回收效率,提高了2014年K的回收效率.相比于养分回收效率,回收能力对增加的可利用氮响应更强.N添加显著降低了N的回收能力,提高了P、K的回收能力,降低了枯叶中的Fe浓度,而对枯叶中Ca、Mg、Al、Mn浓度无显著影响.这表明,N添加对叶片化学计量的影响因不同元素而异,植物会通过调整自身的养分内循环(养分回收)来应对环境变化.N添加提高了绿叶N/P和K/P,说明氮添加条件下植物生长可能由N、P共同限制转变为P限制.氮添加增加了绿叶中Al、Mn浓度,表明N添加下湿地松面临潜在的金属离子毒性风险升高.
We conducted a field experiment with three levels of N addition( 0,40 and 120 kg N·hm~(-2)·a~(-1)) in a Pinus elliottii plantation in subtropical China and collected green and senesced needles of P. elliottii at the peak( July) and the end( October) of each growing season in 2014 and 2015 for clarifying effects of nitrogen additions on concentrations of nine elements( C,N,P,K,Ca,Mg,Al,Fe and Mn) in the green and senesced needles and their corresponding resorption efficiency and resorption proficiency. Our results showed that N addition had positive effects on concentrations of N,Al and Mn,negative effects on the P concentration and the Ca concentration in 2014,and neutral effects on concentrations of C,K,Mg and Fe in green needles. N addition significantly increased foliar N/P. These stoichiometric responses were N level-dependent( stronger at high N rate). N addition significantly decreased N resorption efficiency in 2015 and increased that of K in 2014. Compared with the resorption efficiency,resorption proficiency responded more strongly to increased available N. N addition significantly decreased resorption proficiency of N,and increased that of P,K,and the concentration of Fe in senesced needles,however,there were no significant effects on the concentrations of Ca,Mg,Al and Mn in senesced needles. We concluded that responses of foliar stoichiometry to N addition were element-specific,and plants might cope with changing environments via adjusting internal nutrient cycle( resorption). The elevated foliar N/P and K/P suggested a shift from N and P co-limitation to P limitation with N additions,and increased concentrations of Al and Mn might imply potential toxicity of metal ions to P. elliottii.
We conducted a field experiment with three levels of N addition( 0,40 and 120 kg N·hm~(-2)·a~(-1)) in a Pinus elliottii plantation in subtropical China and collected green and senesced needles of P. elliottii at the peak( July) and the end( October) of each growing season in 2014 and 2015 for clarifying effects of nitrogen additions on concentrations of nine elements( C,N,P,K,Ca,Mg,Al,Fe and Mn) in the green and senesced needles and their corresponding resorption efficiency and resorption proficiency. Our results showed that N addition had positive effects on concentrations of N,Al and Mn,negative effects on the P concentration and the Ca concentration in 2014,and neutral effects on concentrations of C,K,Mg and Fe in green needles. N addition significantly increased foliar N/P. These stoichiometric responses were N level-dependent( stronger at high N rate). N addition significantly decreased N resorption efficiency in 2015 and increased that of K in 2014. Compared with the resorption efficiency,resorption proficiency responded more strongly to increased available N. N addition significantly decreased resorption proficiency of N,and increased that of P,K,and the concentration of Fe in senesced needles,however,there were no significant effects on the concentrations of Ca,Mg,Al and Mn in senesced needles. We concluded that responses of foliar stoichiometry to N addition were element-specific,and plants might cope with changing environments via adjusting internal nutrient cycle( resorption). The elevated foliar N/P and K/P suggested a shift from N and P co-limitation to P limitation with N additions,and increased concentrations of Al and Mn might imply potential toxicity of metal ions to P. elliottii.
引文
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[8]Yan K,Duan C,Fu D,et al.Leaf nitrogen and phosphorus stoichiometry of plant communities in geochemically phosphorus-enriched soils in a subtropical mountainous region,SW China.Environmental Earth Sciences,2015,74:3867-3876
[9]Zhang J,Li H,Shen H,et al.Effects of nitrogen addition on nitrogen resorption in temperate shrub lands in Northern China.PLo S One,2015,10(6):e0130434
[10]Liu J,Huang W,Zhou G,et al.Nitrogen to phosphorus ratios of tree species in response to elevated carbon dioxide and nitrogen addition in subtropical forests.Global Change Biology,2013,19:208-216
[11]Agüero ML,Puntieri J,Mazzarino MJ,et al.Seedling response of Nothofagus species to N and P:Linking plant architecture to N/P ratio and resorption proficiency.Trees,2014,28:1185-1195
[12]Sardans J,Pe1uelas J.Potassium:A neglected nutrient in global change.Global Ecology and Biogeography,2015,24:261-275
[13]Sardans J,Janssens IA,Alonso R,et al.Foliar elemental composition of European forest tree species associated with evolutionary traits and present environmental and competitive conditions.Global Ecology and Biogeography,2015,24:240-255
[14]Yuan ZY,Chen HYH.Negative effects of fertilization on plant nutrient resorption.Ecology,2015,96:373-380
[15]Han W,Tang L,Chen Y,et al.Relationship between the relative limitation and resorption efficiency of nitrogen vs.phosphorus in woody plants.PLo S One,2013,8(12):e83366
[16]See CR,Yanai RD,Fisk MC,et al.Soil nitrogen affects phosphorus recycling:Foliar resorption and plantsoil feedbacks in a northern hardwood forest.Ecology,2015,96:2488-2498
[17]Wang M,Murphy MT,Moore TR.Nutrient resorption of two evergreen shrubs in response to long-term fertilization in a bog.Oecologia,2014,174:365-377
[18]Townsend AR,Cleveland CC,Houlton BZ,et al.Multielement regulation of the tropical forest carbon cycle.Frontiers in Ecology and the Environment,2011,9:9-17
[19]Marschner H.Marschner’s Mineral Nutrition of Higher Plants.London:Academic Press,1995
[20]Takagi S,Nagai R.Several aspects of current research into the role of calcium in plant physiology.Journal of Plant Research,1992,105:687-697
[21]Fenn LB,Feagley S.Review of beneficial uses of calcium and ammonium salts for stimulating plant growth and metabolite translocation.Communications in Soil Science&Plant Analysis,1999,30:2627-2641
[22]Hu Y,Schmidhalter U.Drought and salinity:A comparison of their effects on mineral nutrition of plants.Journal of Plant Nutrition and Soil Science,2005,168:541-549
[23]Poschenrieder C,GunséB,Corrales I,et al.A glance into aluminum toxicity and resistance in plants.Science of the Total Environment,2008,400:356-368
[24]He J-S(贺金生),Han X-G(韩兴国).Ecological stoichiometry:Searching for unifying principles from individuals to ecosystems.Chinese Journal of Plant Ecology(植物生态学报),2010,34(1):2-6(in Chinese)
[25]Huang W,Zhou G,Liu J,et al.Mineral elements of subtropical tree seedlings in response to elevated carbon dioxide and nitrogen addition.PLo S One,2015,10(3):e0120190
[26]Ma Z-Q(马泽清),Liu Q-J(刘琪璟),Wang H-M(王辉民),et al.The growth pattern of Pinus elliottii plantation in central subtropical China.Acta Ecologica Sinica(生态学报),2011,31(6):1525-1537(in Chinese)
[27]Zhang T-P(张太平),Ren H(任海),Peng S-L(彭少麟),et al.The ecological and biological characteristics of Pinus elliottii.Ecologic Science(生态科学),1999,18(2):10-14(in Chinese)
[28]Zhang WJ,Wang HM,Yang FT,et al.Underestimated effects of low temperature during early growing season on carbon sequestration of a subtropical coniferous plantation.Biogeosciences Discussions,2011,8:1667-1678
[29]Shao Q-Q(邵全琴),Yang H-J(杨海军),Liu J-Y(刘纪远),et al.Dynamic analysis on carbon accumulation of a plantation in Qianyanzhou based on tree ring data.Acta Geographica Sinica(地理学报),2009,64(1):69-83(in Chinese)
[30]Zhan XY,Yu GR,He NP,et al.Nitrogen deposition and its spatial pattern in main forest ecosystems along north-south transect of eastern China.Chinese Geographical Science,2014,24:137-146
[31]Killingbeck KT.Nutrients in senesced leaves:Keys to the search for potential resorption and resorption proficiency.Ecology,1996,77:1716-1727
[32]Schreeg LA,Santiago LS,Wright SJ,et al.Stem,root,and older leaf N:P ratios are more responsive indicators of soil nutrient availability than new foliage.Ecology,2014,95:2062-2068
[33]Elvir JA,Wiersma GB,Day ME,et al.Effects of enhanced nitrogen deposition on foliar chemistry and physiological processes of forest trees at the Bear Brook Watershed in Maine.Forest Ecology and Management,2006,221:207-214
[34]Shao J-F(邵继锋),Chen R-F(陈荣府),Dong X-Y(董晓英),et al.Effects of different phosphorus rates on variations of Mn,Al,Mg and Ca concentrations in soil solution and wheat growth in acid red soil.Soils(土壤),2016,48(1):36-41(in Chinese)
[35]He L-Y(何利元),Hu Z-M(胡中民),Guo Q(郭群),et al.Influence of nitrogen and phosphorus addition on the aboveground biomass in Inner Mongolia temperate steppe,China.Chinese Journal of Applied Ecology(应用生态学报),2016,27(11):3427-3434(in Chinese)
[36]Olde Venterink H,Wassen MJ,Verkroost AWM,et al.Species richness-productivity patterns differ between N-,P-,and K-limited wetlands.Ecology,2003,84:2191-2199
[37]White PJ,Brown PH.Plant nutrition for sustainable development and global health.Annals of Botany,2010,105:1073-1080
[38]Hauck M,Paul A,Gross S,et al.Manganese toxicity in epiphytic lichens:Chlorophyll degradation and interaction with iron and phosphorus.Environmental and Experimental Botany,2003,49:181-191
[39]Ren L-M(任立民),Liu P(刘鹏).Review of manganese toxicity&the mechanisms of plant tolerance.Acta Ecologica Sinica(生态学报),2007,27(1):357-367(in Chinese)
[40]Shi L,Zhang H,Liu T,et al.Consistent effects of canopy vs.understory nitrogen addition on the soil exchangeable cations and microbial community in two contrasting forests.Science of the Total Environment,2016,553:349-357
[41]Hong J-T(洪江涛),Wu J-B(吴建波),Wang X-D(王小丹).Effects of global climate change on the C,N,and P stoichiometry of terrestrial plants.Chinese Journal of Applied Ecology(应用生态学报),2013,24(9):2658-2665(in Chinese)
[42]Wright IJ,Westoby M.Nutrient concentration,resorption and lifespan:Leaf traits of Australian sclerophyll species.Functional Ecology,2003,17:10-19
[43]Kou L,Wang H,Gao W,et al.Nitrogen addition regulates tradeoff between root capture and foliar resorption of nitrogen and phosphorus in a subtropical pine plantation.Trees,2016,doi:10.1007/s00468-016-1457-7
[44]Lucas RW,Klaminder J,Futter MN,et al.A metaanalysis of the effects of nitrogen additions on base cations:Implications for plants,soils,and streams.Forest Ecology and Management,2011,262:95-104
[45]Zhang C(张闯),Zou H-T(邹洪涛),Zhang X-Y(张心昱),et al.Effects of nitrogen additions on soil hydrolase and oxidase activities in Pinus elliottii plantations.Chinese Journal of Applied Ecology(应用生态学报),2016,27(11):3427-3434(in Chinese)
[46]Kozovits AR,Bustamante MMC,Garofalo CR,et al.Nutrient resorption and patterns of litter production and decomposition in a Neotropical Savanna.Functional Ecology,2007,21:1034-1043
[47]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.Plant Biology,2015,17:9-15
[48]Chapin FS,Kedrowski RA.Seasonal changes in nitrogen and phosphorus fractions and autumn retranslocation in evergreen and deciduous taiga trees.Ecology,1983,64:376-391
[49]Chapin FS,Moilanen L.Nutritional controls over nitrogen and phosphorus resorption from Alaskan birch leaves.Ecology,1991,72:709-715
[50]Tian Q,Liu N,Bai W,et al.A novel soil manganese mechanism drives plant species loss with increased nitrogen deposition in a temperate steppe.Ecology,2016,97:65-74
[51]Kou L,Guo D,Yang H,et al.Growth,morphological traits and mycorrhizal colonization of fine roots respond differently to nitrogen addition in a slash pine plantation in subtropical China.Plant and Soil,2015,391:207-218
[2]Yuan ZY,Chen HYH.Global-scale patterns of nutrient resorption associated with latitude,temperature and precipitation.Global Ecology and Biogeography,2009,18:11-18
[3]Wright SJ,Yavitt JB,Wurzburger N,et al.Potassium,phosphorus,or nitrogen limit root allocation,tree growth,or litter production in a lowland tropical forest.Ecology,2011,92:1616-1625
[4]Sardans J,Rivas-Ubach A,Pe1uelas J.The C:N:P stoichiometry of organisms and ecosystems in a changing world:A review and perspectives.Perspectives in Plant Ecology,Evolution and Systematics,2012,14:33-47
[5]Güsewell S.N:P ratios in terrestrial plants:Variation and functional significance.New Phytologist,2004,164:243-266
[6]Sterner RW,Elser JJ.Ecological Stoichiometry:The Biology of Elements from Molecules to the Biosphere.Princeton,NJ:Princeton University Press,2002
[7]Yuan ZY,Chen HYH.Decoupling of nitrogen and phosphorus in terrestrial plants associated with global changes.Nature Climate Change,2015,5:465-469
[8]Yan K,Duan C,Fu D,et al.Leaf nitrogen and phosphorus stoichiometry of plant communities in geochemically phosphorus-enriched soils in a subtropical mountainous region,SW China.Environmental Earth Sciences,2015,74:3867-3876
[9]Zhang J,Li H,Shen H,et al.Effects of nitrogen addition on nitrogen resorption in temperate shrub lands in Northern China.PLo S One,2015,10(6):e0130434
[10]Liu J,Huang W,Zhou G,et al.Nitrogen to phosphorus ratios of tree species in response to elevated carbon dioxide and nitrogen addition in subtropical forests.Global Change Biology,2013,19:208-216
[11]Agüero ML,Puntieri J,Mazzarino MJ,et al.Seedling response of Nothofagus species to N and P:Linking plant architecture to N/P ratio and resorption proficiency.Trees,2014,28:1185-1195
[12]Sardans J,Pe1uelas J.Potassium:A neglected nutrient in global change.Global Ecology and Biogeography,2015,24:261-275
[13]Sardans J,Janssens IA,Alonso R,et al.Foliar elemental composition of European forest tree species associated with evolutionary traits and present environmental and competitive conditions.Global Ecology and Biogeography,2015,24:240-255
[14]Yuan ZY,Chen HYH.Negative effects of fertilization on plant nutrient resorption.Ecology,2015,96:373-380
[15]Han W,Tang L,Chen Y,et al.Relationship between the relative limitation and resorption efficiency of nitrogen vs.phosphorus in woody plants.PLo S One,2013,8(12):e83366
[16]See CR,Yanai RD,Fisk MC,et al.Soil nitrogen affects phosphorus recycling:Foliar resorption and plantsoil feedbacks in a northern hardwood forest.Ecology,2015,96:2488-2498
[17]Wang M,Murphy MT,Moore TR.Nutrient resorption of two evergreen shrubs in response to long-term fertilization in a bog.Oecologia,2014,174:365-377
[18]Townsend AR,Cleveland CC,Houlton BZ,et al.Multielement regulation of the tropical forest carbon cycle.Frontiers in Ecology and the Environment,2011,9:9-17
[19]Marschner H.Marschner’s Mineral Nutrition of Higher Plants.London:Academic Press,1995
[20]Takagi S,Nagai R.Several aspects of current research into the role of calcium in plant physiology.Journal of Plant Research,1992,105:687-697
[21]Fenn LB,Feagley S.Review of beneficial uses of calcium and ammonium salts for stimulating plant growth and metabolite translocation.Communications in Soil Science&Plant Analysis,1999,30:2627-2641
[22]Hu Y,Schmidhalter U.Drought and salinity:A comparison of their effects on mineral nutrition of plants.Journal of Plant Nutrition and Soil Science,2005,168:541-549
[23]Poschenrieder C,GunséB,Corrales I,et al.A glance into aluminum toxicity and resistance in plants.Science of the Total Environment,2008,400:356-368
[24]He J-S(贺金生),Han X-G(韩兴国).Ecological stoichiometry:Searching for unifying principles from individuals to ecosystems.Chinese Journal of Plant Ecology(植物生态学报),2010,34(1):2-6(in Chinese)
[25]Huang W,Zhou G,Liu J,et al.Mineral elements of subtropical tree seedlings in response to elevated carbon dioxide and nitrogen addition.PLo S One,2015,10(3):e0120190
[26]Ma Z-Q(马泽清),Liu Q-J(刘琪璟),Wang H-M(王辉民),et al.The growth pattern of Pinus elliottii plantation in central subtropical China.Acta Ecologica Sinica(生态学报),2011,31(6):1525-1537(in Chinese)
[27]Zhang T-P(张太平),Ren H(任海),Peng S-L(彭少麟),et al.The ecological and biological characteristics of Pinus elliottii.Ecologic Science(生态科学),1999,18(2):10-14(in Chinese)
[28]Zhang WJ,Wang HM,Yang FT,et al.Underestimated effects of low temperature during early growing season on carbon sequestration of a subtropical coniferous plantation.Biogeosciences Discussions,2011,8:1667-1678
[29]Shao Q-Q(邵全琴),Yang H-J(杨海军),Liu J-Y(刘纪远),et al.Dynamic analysis on carbon accumulation of a plantation in Qianyanzhou based on tree ring data.Acta Geographica Sinica(地理学报),2009,64(1):69-83(in Chinese)
[30]Zhan XY,Yu GR,He NP,et al.Nitrogen deposition and its spatial pattern in main forest ecosystems along north-south transect of eastern China.Chinese Geographical Science,2014,24:137-146
[31]Killingbeck KT.Nutrients in senesced leaves:Keys to the search for potential resorption and resorption proficiency.Ecology,1996,77:1716-1727
[32]Schreeg LA,Santiago LS,Wright SJ,et al.Stem,root,and older leaf N:P ratios are more responsive indicators of soil nutrient availability than new foliage.Ecology,2014,95:2062-2068
[33]Elvir JA,Wiersma GB,Day ME,et al.Effects of enhanced nitrogen deposition on foliar chemistry and physiological processes of forest trees at the Bear Brook Watershed in Maine.Forest Ecology and Management,2006,221:207-214
[34]Shao J-F(邵继锋),Chen R-F(陈荣府),Dong X-Y(董晓英),et al.Effects of different phosphorus rates on variations of Mn,Al,Mg and Ca concentrations in soil solution and wheat growth in acid red soil.Soils(土壤),2016,48(1):36-41(in Chinese)
[35]He L-Y(何利元),Hu Z-M(胡中民),Guo Q(郭群),et al.Influence of nitrogen and phosphorus addition on the aboveground biomass in Inner Mongolia temperate steppe,China.Chinese Journal of Applied Ecology(应用生态学报),2016,27(11):3427-3434(in Chinese)
[36]Olde Venterink H,Wassen MJ,Verkroost AWM,et al.Species richness-productivity patterns differ between N-,P-,and K-limited wetlands.Ecology,2003,84:2191-2199
[37]White PJ,Brown PH.Plant nutrition for sustainable development and global health.Annals of Botany,2010,105:1073-1080
[38]Hauck M,Paul A,Gross S,et al.Manganese toxicity in epiphytic lichens:Chlorophyll degradation and interaction with iron and phosphorus.Environmental and Experimental Botany,2003,49:181-191
[39]Ren L-M(任立民),Liu P(刘鹏).Review of manganese toxicity&the mechanisms of plant tolerance.Acta Ecologica Sinica(生态学报),2007,27(1):357-367(in Chinese)
[40]Shi L,Zhang H,Liu T,et al.Consistent effects of canopy vs.understory nitrogen addition on the soil exchangeable cations and microbial community in two contrasting forests.Science of the Total Environment,2016,553:349-357
[41]Hong J-T(洪江涛),Wu J-B(吴建波),Wang X-D(王小丹).Effects of global climate change on the C,N,and P stoichiometry of terrestrial plants.Chinese Journal of Applied Ecology(应用生态学报),2013,24(9):2658-2665(in Chinese)
[42]Wright IJ,Westoby M.Nutrient concentration,resorption and lifespan:Leaf traits of Australian sclerophyll species.Functional Ecology,2003,17:10-19
[43]Kou L,Wang H,Gao W,et al.Nitrogen addition regulates tradeoff between root capture and foliar resorption of nitrogen and phosphorus in a subtropical pine plantation.Trees,2016,doi:10.1007/s00468-016-1457-7
[44]Lucas RW,Klaminder J,Futter MN,et al.A metaanalysis of the effects of nitrogen additions on base cations:Implications for plants,soils,and streams.Forest Ecology and Management,2011,262:95-104
[45]Zhang C(张闯),Zou H-T(邹洪涛),Zhang X-Y(张心昱),et al.Effects of nitrogen additions on soil hydrolase and oxidase activities in Pinus elliottii plantations.Chinese Journal of Applied Ecology(应用生态学报),2016,27(11):3427-3434(in Chinese)
[46]Kozovits AR,Bustamante MMC,Garofalo CR,et al.Nutrient resorption and patterns of litter production and decomposition in a Neotropical Savanna.Functional Ecology,2007,21:1034-1043
[47]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.Plant Biology,2015,17:9-15
[48]Chapin FS,Kedrowski RA.Seasonal changes in nitrogen and phosphorus fractions and autumn retranslocation in evergreen and deciduous taiga trees.Ecology,1983,64:376-391
[49]Chapin FS,Moilanen L.Nutritional controls over nitrogen and phosphorus resorption from Alaskan birch leaves.Ecology,1991,72:709-715
[50]Tian Q,Liu N,Bai W,et al.A novel soil manganese mechanism drives plant species loss with increased nitrogen deposition in a temperate steppe.Ecology,2016,97:65-74
[51]Kou L,Guo D,Yang H,et al.Growth,morphological traits and mycorrhizal colonization of fine roots respond differently to nitrogen addition in a slash pine plantation in subtropical China.Plant and Soil,2015,391:207-218