南亚热带海岸沙地不同树种叶片化学计量学季节动态研究
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摘要
养分贫瘠是南亚热带海岸沙地植物生长与分布的主要限制因子之一。本文以南亚热带海岸沙地5种树种的叶片和凋落叶为研究对象,通过对其碳、氮、磷含量及生态化学计量学特征的研究,从树种水平上分析了不同树种鲜叶与凋落叶养分浓度及重吸收率的动态变化,探讨了不同树种的叶片化学计量特征及与养分含量的关系,以科学评判不同树种对海岸沙地贫瘠立地的适应性,为沿海防护林的可持续经营提供理论依据。结果表明:海岸沙地上5个树种叶片碳含量普遍较高,而氮、磷含量在不同树种间差异较大,且均呈显著的季节变化。不同树种的平均氮、磷重吸收率变化范围分别为25.0%～54.4%和52.1%～78.1%,其中固氮树种的平均氮重吸收率低于非固氮树种,而其磷重吸收率高于非固氮树种。在不同季节,不同树种受不同的养分限制。尾巨桉在春秋季为氮限制,在夏冬季为磷限制;厚荚相思和木麻黄在不同季节均为磷限制;湿地松在春季为氮磷共同限制,夏季为氮限制,秋冬季为磷限制;而潺槁木姜子在春季为氮限制,夏季为氮磷共同限制,秋冬季节为磷限制。
Nutrient depletion is one of the main limiting factors for plant growth and distribution in coastal sand dunes in southeast China. In order to evaluate the adaptability of different tree species to the infertile environment of coastal sand dunes and provide a theoretical basis for the sustainable management of coastal protection forest, we investigated the seasonal dynamics of leaf stoichiometry for five tree species(including Casuarina equisetifolia, Acacia crassicarpa, Pinus elliottii, Eucalyptus grandis × E. urophylla, and Litsea glutinosa) in a coastal sand dune in subtropical China. Results showed that the leaf carbon(C) content of the five tree species was generally high, while the nitrogen(N) and phosphorus(P) contents differed among the different tree species and exhibited significant seasonal variations. The mean N and P resorption rates of the five tree species ranged from 25.0%–54.4% and 52.1%–78.1%, respectively, and the mean N resorption rate of N-fixing trees was lower than that of non-N-fixing trees, while its P resorption rate was higher. The nutrient limitation of the five tree species varied in different seasons, briefly, E. grandis × E. urophylla was under N limitation in spring and P limitation in autumn, while A. crassicarpa and C. equisetifolia were under P limitation across all seasons. P. elliottii was under N and P co-limitation in spring, and N limitation in summer, and P limitation in autumn and winter. L. glutinosa was under N limitation in spring, N and P limitation in summer, and P limitation in autumn and winter.
Nutrient depletion is one of the main limiting factors for plant growth and distribution in coastal sand dunes in southeast China. In order to evaluate the adaptability of different tree species to the infertile environment of coastal sand dunes and provide a theoretical basis for the sustainable management of coastal protection forest, we investigated the seasonal dynamics of leaf stoichiometry for five tree species(including Casuarina equisetifolia, Acacia crassicarpa, Pinus elliottii, Eucalyptus grandis × E. urophylla, and Litsea glutinosa) in a coastal sand dune in subtropical China. Results showed that the leaf carbon(C) content of the five tree species was generally high, while the nitrogen(N) and phosphorus(P) contents differed among the different tree species and exhibited significant seasonal variations. The mean N and P resorption rates of the five tree species ranged from 25.0%–54.4% and 52.1%–78.1%, respectively, and the mean N resorption rate of N-fixing trees was lower than that of non-N-fixing trees, while its P resorption rate was higher. The nutrient limitation of the five tree species varied in different seasons, briefly, E. grandis × E. urophylla was under N limitation in spring and P limitation in autumn, while A. crassicarpa and C. equisetifolia were under P limitation across all seasons. P. elliottii was under N and P co-limitation in spring, and N limitation in summer, and P limitation in autumn and winter. L. glutinosa was under N limitation in spring, N and P limitation in summer, and P limitation in autumn and winter.
引文
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[26]马明睿,杨洁,王强,等.河岸带加拿大一枝黄花化学计量学及入侵机理研究[J].中国环境科学,2014,34(6):1 531-1 539.
[27]牛得草,李茜,江世高,等.阿拉善荒漠区6种主要灌木植物叶片C:N:P化学计量比的季节变化[J].植物生态学报,2013,37(4):317-325.
[28]张立华,林益明,叶功富,等.不同林分类型叶片氮磷含量、氮磷比及其内吸收率[J].北京林业大学学报,2009,31(5):67-72.
[29]赵亚芳,徐福利,王渭玲,等.华北落叶松针叶碳、氮、磷含量及化学计量比的季节变化[J].植物营养与肥料学报,2015,21(5):1 328-1 335.
[30]郑艳明,尧波,吴琴,等.鄱阳湖湿地两种优势植物叶片C、N、P动态特征[J].生态学报,2013,33(20):6 488-6 496.
[31]Elser J J,Fagan W F,Denno R F,et al.Nutritional constraints in terrestrial and freshwater food webs[J].Nature,2000,408(6 812):578-80.
[32]Pan F J,Zhang W,Liu S J,et al.Leaf N:P stoichiometry across plant functional groups in the karst region of southwestern China[J].Trees,2015,29(3):883-892.
[33]Vitousek P M,Cassman K,Cleveland C,et al.Towards an ecological understanding of biological nitrogen fixation[J].Biogeochemistry,2002,57:1.
[34]吴统贵,吴明,刘丽,等.杭州湾滨海湿地3种草本植物叶片N、P化学计量学的季节变化[J].植物生态学报,2010,34(1):23-28.
[35]Zhang L H,Ye G F,Lin Y M,et al.Seasonal changes in tannin and nitrogen contents of Casuarina equisetifolia branchlets[J].Journal of Zhejiang University(Science B),2009,10(2):103-111.
[36]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.
[37]Killingbeck K T.Nutrients in senesced leaves:Keys to the search for potential resorption and resorption proficiency[J].Ecology,1996,77(6):1 716-1 727.
[38]任书杰,于贵瑞,姜春明,等.中国东部南北样带森林生态系统102个优势种叶片碳氮磷化学计量学统计特征[J].应用生态学报,2012,23(3):581-586.
[39]Wu T G,Yu M K,Wang G G,et al.Leaf nitrogen and phosphorus stoichiometry across forty-two woody species in Southeast China[J].Biochemical Systematics&Ecology,2012,44(10):255-263.
[2]Vitousek P M.Nutrient cycling and limitation:Hawaii as a model system[M].Princeton:Princeton University Press,2004.
[3]Richardson S J,Allen R B,Doherty J E.Shifts in leaf N:Pratio during resorption reflect soil P in temperate rainforest[J].Functional Ecology,2008,22(4):738-745.
[4]Von O G,Power S A,Falk K,et al.N:P ratio and the nature of nutrient limitation in calluna-dominated heathlands[J].Ecosystems,2010,13(2):317-327.
[5]Vitousek P.Nutrient cycling and nutrient use efficiency[J].The American Naturalist,1982,119(4):553-572.
[6]Koerselman W.The vegetation N:P ratio:a new tool to detect the nature of nutrient limitation[J].Journal of Applied Ecology,1996,33(6):1 441-1 450.
[7]Güsewell S,Freeman C.Nutrient limitation and enzyme activities during litter decomposition of nine wetland species in relation to litter N:P ratios[J].Functional Ecology,2005,19(4):582-593.
[8]Güsewell S,Koerselman W.Biomass N:P ratios as indicators of nutrient limitation for plant populations in wetlands[J].Ecological Applications,2008,13(2):372-384.
[9]Schreeg L A,Santiago L S,Wright S J,et al.Stem,root,and older leaf N:P ratios are more responsive indicators of soil nutrient availability than new foliage[J].Ecology,2014,95(8):2 062-2 068.
[10]Alvarezclare S,Mack M C.Do foliar,litter,and root nitrogen and phosphorus concentrations reflect nutrient limitation in a lowland tropical wet forest?[J].PLoS ONE,2015,10(4):e0123796.
[11]Güsewell S,Verhoeven J T.Litter N:P ratios indicate whether N or P limits the decomposability of graminoid leaf litter[J].Plant and Soil,2006,287(1):131-143.
[12]Sabine G,Marko G.N:P ratios influence litter decomposition and colonization by fungi and bacteria in microcosms[J].Functional Ecology,2009,23(1):211-219.
[13]Aerts R,Chapin F S I.The mineral nutrition of wild plants revisited:a re-evaluation of processes and patterns[J].Advances in Ecological Research,1999,30(8):1-67.
[14]Brant A N,Chen H Y H.Patterns and mechanisms of nutrient resorption in plants[J].Critical Reviews in Plant Sciences,2015,34(5):471-486.
[15]Vergutz L,Manzoni S,Porporato A,et al.Global resorption efficiencies and concentrations of carbon and nutrients in leaves of terrestrial plants[J].Ecological Monographs,2012,82(2):205-220.
[16]Townsend A R,Cleveland C C,Asner G P,et al.Controls over foliar N:P ratios in tropical rain forests[J].Ecology,2007,88(1):107.
[17]Kang H Z,Xin Z J,Berg B,et al.Global pattern of leaf litter nitrogen and phosphorus in woody plants[J].Annals of Forest Science,2010,67(8):811.
[18]Han W X,Fang J Y,Guo D L,et al.Leaf nitrogen and phosphorus stoichiometry across 753 terrestrial plant species in China[J].New Phytologist,2005,168(2):377-385.
[19]王晶苑,王绍强,李纫兰,等.中国四种森林类型主要优势植物的C:N:P化学计量学特征[J].植物生态学报,2011,35(6):587-595.
[20]吴统贵,陈步峰,肖以华,等.珠江三角洲3种典型森林类型乔木叶片生态化学计量学[J].植物生态学报,2010,34(1):58-63.
[21]阎恩荣,王希华,郭明,等.浙江天童常绿阔叶林、常绿针叶林与落叶阔叶林的C:N:P化学计量特征[J].植物生态学报,2010,34(1):48-57.
[22]Inagaki M,Kamo K,Miyamoto K,et al.Nitrogen and phosphorus retranslocation and N:P ratios of litterfall in three tropical plantations:luxurious N and efficient P use by Acacia mangium[J].Plant and Soil,2011,341(1):295-307.
[23]Li X W,Sun K,Li F Y.Variation in leaf nitrogen and phosphorus stoichiometry in the nitrogen-fixing Chinese sea-buckthorn(Hippophae rhamnoides L.subsp.sinensis Rousi)across northern China[J].Ecological Research,2014,29(4):723-731.
[24]Reich P B,Oleksyn J.Global patterns of plant leaf N and Pin relation to temperature and latitude[J].Proceedings of the National Academy of Sciences of the United States of America,2004,101(30):11 001-11 006.
[25]Yuan Z,Chen H Y H.Global trends in senesced-leaf nitrogen and phosphorus[J].Global Ecology&Biogeography,2009,18(5):532-542.
[26]马明睿,杨洁,王强,等.河岸带加拿大一枝黄花化学计量学及入侵机理研究[J].中国环境科学,2014,34(6):1 531-1 539.
[27]牛得草,李茜,江世高,等.阿拉善荒漠区6种主要灌木植物叶片C:N:P化学计量比的季节变化[J].植物生态学报,2013,37(4):317-325.
[28]张立华,林益明,叶功富,等.不同林分类型叶片氮磷含量、氮磷比及其内吸收率[J].北京林业大学学报,2009,31(5):67-72.
[29]赵亚芳,徐福利,王渭玲,等.华北落叶松针叶碳、氮、磷含量及化学计量比的季节变化[J].植物营养与肥料学报,2015,21(5):1 328-1 335.
[30]郑艳明,尧波,吴琴,等.鄱阳湖湿地两种优势植物叶片C、N、P动态特征[J].生态学报,2013,33(20):6 488-6 496.
[31]Elser J J,Fagan W F,Denno R F,et al.Nutritional constraints in terrestrial and freshwater food webs[J].Nature,2000,408(6 812):578-80.
[32]Pan F J,Zhang W,Liu S J,et al.Leaf N:P stoichiometry across plant functional groups in the karst region of southwestern China[J].Trees,2015,29(3):883-892.
[33]Vitousek P M,Cassman K,Cleveland C,et al.Towards an ecological understanding of biological nitrogen fixation[J].Biogeochemistry,2002,57:1.
[34]吴统贵,吴明,刘丽,等.杭州湾滨海湿地3种草本植物叶片N、P化学计量学的季节变化[J].植物生态学报,2010,34(1):23-28.
[35]Zhang L H,Ye G F,Lin Y M,et al.Seasonal changes in tannin and nitrogen contents of Casuarina equisetifolia branchlets[J].Journal of Zhejiang University(Science B),2009,10(2):103-111.
[36]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.
[37]Killingbeck K T.Nutrients in senesced leaves:Keys to the search for potential resorption and resorption proficiency[J].Ecology,1996,77(6):1 716-1 727.
[38]任书杰,于贵瑞,姜春明,等.中国东部南北样带森林生态系统102个优势种叶片碳氮磷化学计量学统计特征[J].应用生态学报,2012,23(3):581-586.
[39]Wu T G,Yu M K,Wang G G,et al.Leaf nitrogen and phosphorus stoichiometry across forty-two woody species in Southeast China[J].Biochemical Systematics&Ecology,2012,44(10):255-263.