天然次生林植物叶片生态化学计量特征及光合特性对长期N沉降的响应
|
摘要
2006年5月于吉林省抚松县露水河林业局实验林场布设了人工模拟氮沉降控制试验,共设置3个氮(N)添加梯度,分别为对照(CK 0 g·N·m~(-2)·a~(-1))、低N(LN 2.5 g·N·m~(-2)·a~(-1))和高N(HN 5.0 g·N·m~(-2)·a~(-1)),旨在探讨N沉降对天然次生林先锋树种白桦(Betula platyphylla)和山杨(Populus davidiana)鲜叶、凋落叶化学计量特征、养分重吸收的影响,以及鲜叶光合特性的变化和各性状之间的相互关系。结果表明:(1)模拟N沉降处理下白桦、山杨鲜叶的C含量较对照均无显著影响,LN处理显著降低了山杨鲜叶N、P含量(P<0.05),显著增加了C∶N、C∶P和N∶P(P<0.05);HN处理显著增加了白桦鲜叶N含量和N∶P,显著降低了C∶N(P<0.05)。(2)白桦、山杨鲜叶N、P重吸收率在两个梯度N添加下均显著下降(P<0.05),且均为负值。山杨鲜叶N重吸收率与P重吸收率呈显著正相关关系(P<0.05),与鲜叶C∶N呈显著负相关关系(P<0.05)。(3)N添加可以提高2种树木叶片氮素光合利用效率(PNUE)(P<0.05)、净光合速率(P_n)(P<0.05)。白桦鲜叶N含量与P_n、PNUE呈显著正相关(P<0.05);白桦、山杨鲜叶比叶重(LMA)与N含量呈显著负相关(P<0.05);P_n与PNUE呈显著正相关(P<0.05)。本试验研究表明:在生长季,白桦、山杨鲜叶中N、P均表现为富集状态,土壤养分及外源N可供林木较快吸收并促进其生长,无需从凋落叶中吸收养分。N添加可以增强白桦、山杨鲜叶的光合性能,进而促进植物养分吸收和叶片发育。HN对长白山天然次生林的生长有促进作用。
Three nitrogen(N)concentration gradients were set up in Experimental Forest Farm of Lushuihe Forestry Bureau in Fusong County,Jilin Province,including control(CK 0 g·N·m~(-2)·a~(-1)),low nitrogen(LN 2.5 g·N·m~(-2)·a~(-1))and high nitrogen(HN 5.0 g·N·m~(-2)·a~(-1)),to study the effects of N deposition on the stoichiometric characteristics,nutrient reuptake and changes of photosynthetic characteristics of leaves of birch(Betula platyphylla)and poplar(Populus davidiana),a pioneer tree in natural secondary forest,and the relationship between the characters.The results showed that:(1)Under the simulated N deposition,the C content of the fresh leaves of birch and poplar showed no significant influence compared with that of the control group,and the N and P contents of the fresh leaves of poplar were significantly reduced by LN treatment(P<0.05),and C∶N,C∶P and N∶P were significantly increased(P<0.05).HN treatment significantly increased N content and N∶P in birch leaf and significantly decreased C∶N(P<0.05).(2)The resorption efficiency of leaves N and P of birch and poplar decreased significantly in the treatment of LN(P<0.05)and were all negative.There was a significant positive correlation between N and P resorption efficiency(P<0.05),and a significant negative correlation between C∶N and N resorption efficiency(P<0.05).(3)N addition can improve the nitrogen photosynthetic efficiency(PNUE)and net photosynthetic rate(P_n)of two kinds of trees(P<0.05),leaf dry matter to leaf area(LMA)of birch and poplar showed a significant negative correlation with N content(P<0.05),and there was significant positive correlation between P_n and PNUE(P<0.05).The N content of birch leaf was significantly positively correlated with P_n and PNUE(P<0.05).The results showed that N and P in birch and poplar leaves were enriched in the growing season,and soil nutrient and exogenous N could be used to absorb and promote the growth of trees quickly,without the need to absorb nutrients from litters leaves.N addition can enhance the photosynthetic performance of the leaves of birch and poplar,thus promoting the plant nutrient absorption and leaf development.HN deposition can promote the growth of natural secondary forest in Changbai Mountains.
Three nitrogen(N)concentration gradients were set up in Experimental Forest Farm of Lushuihe Forestry Bureau in Fusong County,Jilin Province,including control(CK 0 g·N·m~(-2)·a~(-1)),low nitrogen(LN 2.5 g·N·m~(-2)·a~(-1))and high nitrogen(HN 5.0 g·N·m~(-2)·a~(-1)),to study the effects of N deposition on the stoichiometric characteristics,nutrient reuptake and changes of photosynthetic characteristics of leaves of birch(Betula platyphylla)and poplar(Populus davidiana),a pioneer tree in natural secondary forest,and the relationship between the characters.The results showed that:(1)Under the simulated N deposition,the C content of the fresh leaves of birch and poplar showed no significant influence compared with that of the control group,and the N and P contents of the fresh leaves of poplar were significantly reduced by LN treatment(P<0.05),and C∶N,C∶P and N∶P were significantly increased(P<0.05).HN treatment significantly increased N content and N∶P in birch leaf and significantly decreased C∶N(P<0.05).(2)The resorption efficiency of leaves N and P of birch and poplar decreased significantly in the treatment of LN(P<0.05)and were all negative.There was a significant positive correlation between N and P resorption efficiency(P<0.05),and a significant negative correlation between C∶N and N resorption efficiency(P<0.05).(3)N addition can improve the nitrogen photosynthetic efficiency(PNUE)and net photosynthetic rate(P_n)of two kinds of trees(P<0.05),leaf dry matter to leaf area(LMA)of birch and poplar showed a significant negative correlation with N content(P<0.05),and there was significant positive correlation between P_n and PNUE(P<0.05).The N content of birch leaf was significantly positively correlated with P_n and PNUE(P<0.05).The results showed that N and P in birch and poplar leaves were enriched in the growing season,and soil nutrient and exogenous N could be used to absorb and promote the growth of trees quickly,without the need to absorb nutrients from litters leaves.N addition can enhance the photosynthetic performance of the leaves of birch and poplar,thus promoting the plant nutrient absorption and leaf development.HN deposition can promote the growth of natural secondary forest in Changbai Mountains.
引文
1.Galloway J N,Townsend A R,Erisman J W,et al.Transformation of the nitrogen cycle:recent trends,questions,and potential solutions[J].Science,2008,320(5878):889-892.
2.Liu X J,Zhang Y,Han W X,et al.Enhanced nitrogen deposition over China[J].Nature,2013,494(7438):459-462.
3.Galloway J N,Cowling E B.Reactive nitrogen and the world:200 years of change[J].AMBIO:A Journal of the Human Environment,2002,31(2):64-71.
4.Lü C Q,Tian H Q.Spatial and temporal patterns of nitrogen deposition in China:synthesis of observational data[J].Journal of Geophysical Research:Atmospheres,2007,112(D22):D22S05.
5.邓小文,韩士杰.氮沉降对森林生态系统土壤碳库的影响[J].生态学杂志,2007,26(10):1622-1627.Deng X W,Han S J.Impact of nitrogen deposition on forest soil carbon pool[J].Chinese Journal of Ecology,2007,26(10):1622-1627.
6.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.Wang J Y,Wang S Q,Li R L,et al.C∶N∶P stoichiometric characteristics of four forest types’dominant tree species in China[J].Chinese Journal of Plant Ecology,2011,35(6):587-595.
7.Magill A H,Aber J D,Currie W S,et al.Ecosystem response to 15 years of chronic nitrogen additions at the Harvard Forest LTER,Massachusetts,USA[J].Forest Ecology and Management,2004,196(1):7-28.
8.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.
9.Elser J J,Andersen T,Baron J S,et al.Shifts in lake N∶Pstoichiometry and nutrient limitation driven by atmospheric nitrogen deposition[J].Science,2009,326(5954):835-837.
10.Pe1uelas J,Sardans J,Rivas-Ubach A,et al.The human‐induced imbalance between C,N and P in Earth’s life system[J].Global Change Biology,2012,18(1):3-6.
11.Turner M M,Henry H A L.Interactive effects of warming and increased nitrogen deposition on15N tracer retention in a temperate old field:seasonal trends[J].Global Change Biology,2009,15(12):2885-2893.
12.Passarinho J A P,Lamosa P,Baeta J P,et al.Annual changes in the concentration of minerals and organic compounds of Quercus suber leaves[J].Physiologia Plantarum,2006,127(1):100-110.
13.Fife D N,Nambiar E K,Saur E.Retranslocation of foliar nutrients in evergreen tree species planted in a Mediterranean environment[J].Tree Physiology,2008,28(2):187-196.
14.Phoenix G K,Booth R E,Leake J R,et al.Simulated pollutant nitrogen deposition increases P demand and enhances root-surface phosphatase activities of three plant functional types in a calcareous grassland[J].New Phytologist,2004,161(1):279-290.
15.Fujita Y,De Ruiter P C,Wassen M J,et al.Time-dependent,species-specific effects of N∶P stoichiometry on grassland plant growth[J].Plant and Soil,2010,334(1-2):99-112.
16.Chapin III S F,Matson P A,Mooney H A.Principles of terrestrial ecosystem ecology[M].New York,NY:Springer,2002.
17.Koojiman S A L M.The stoichiometry of animal energetics[J].Journal of Theoretical Biology,1995,177(2):139-149.
18.Aerts R.Nutrient resorption from senescing leaves of perennials:are there general patterns?[J].Journal of Ecology,1996,84(4):597-608.
19.Cernusak L A,Aranda J,Marshall J D,et al.Large variation in whole-plant water-use efficiency among tropical tree species[J].New Phytologist,2007,173(2):294-305.
20.郭淑青,李文金,张仁懿,等.氮磷添加对金露梅叶片化学计量及光合特征的影响[J].广西植物,2014,34(5):629-634,718.Guo S Q,Li W J,Zhang R Y,et al.Effects of N and P additions on foliar stoichiometry and photosynthetic characteristics of Potentilla fruiticosa[J].Guihaia,2014,34(5):629-634,718.
21.安慧.子午岭林区典型植物生长的氮素调控机理[D].杨凌:西北农林科技大学,2008.An H.Regulating mechanism of nitrogen of typical plants growth in Ziwuling forest region[D].Yangling:Northwest A&F University,2008.
22.Warren C R,ADAMS M A.Evergreen trees do not maximize instantaneous photosynthesis[J].Trends in Plant Science,2004,9(6):270-274.
23.王振南,杨惠敏.植物碳氮磷生态化学计量对非生物因子的响应[J].草业科学,2013,30(6):927-934.Wang Z N,Yang H M.Response of ecological stoichiometry of carbon,nitrogen and phosphorus in plants to abiotic environmental factors[J].Pratacultural Science,2013,30(6):927-934.
24.吕丽华,赵明,赵久然,等.不同施氮量下夏玉米冠层结构及光合特性的变化[J].中国农业科学,2008,41(9):2624-2632.LüL H,Zhao M,Zhao J R,et al.Canopy structure and photosynthesis of summer maize under different nitrogen fertilizer application rates[J].Scientia Agricultura Sinica,2008,41(9):2624-2632.
25.杨浩,罗亚晨.糙隐子草功能性状对氮添加和干旱的响应[J].植物生态学报,2015,39(1):32-42.Yang H,Luo Y C.Responses of the functional traits in Cleistogenes squarrosa to nitrogen addition and drough[J].Chinese Journal of Plant Ecology,2015,39(1):32-42.
26.谌贤,刘洋,邓静,等.川西亚高山森林凋落物不同分解阶段碳氮磷化学计量特征及种间差异[J].植物研究,2017,37(2):216-226.Shen X,Liu Y,Deng J,et al.C,N and P stoichiometry at different stages of litter decomposition in subalpine forest of western Sichuan province and inter species comparison[J].Bulletin of Botanical Research,2017,37(2):216-226.
27.刘荣杰,吴亚丛,张英,等.中国北亚热带天然次生林与杉木人工林土壤活性有机碳库的比较[J].植物生态学报,2012,36(5):431-437.Liu R J,Wu Y C,Zhang Y,et al.Comparison of soil labile organic carbon in Chinese fir plantations and natural secondary forests in north subtropical areas of China[J].Chinese Journal of Plant Ecology,2012,36(5):431-437.
28.胡艳玲,韩士杰,李雪峰,等.长白山原始林和次生林土壤有效氮含量对模拟氮沉降的响应[J].东北林业大学学报,2009,37(5):36-38,42.Hu Y L,Han S J,Li X F,et al.Responses of soil available nitrogen of natural forest and secondary forest to simulated N deposition in Changbai Mountain[J].Journal of Northeast Forestry University,2009,37(5):36-38,42.
29.唐艳,王传宽.东北主要树种光合作用可行的离体测定方法[J].植物生态学报,2011,35(4):452-462.Tang Y,Wang C K.A feasible method for measuring photosynthesis in vitro for major tree species in northeastern China[J].Chinese Journal of Plant Ecology,2011,35(4):452-462.
30.李朝英,郑路,卢立华,等.测定植物全氮的H2SO4-H2O2消煮法改进[J].中国农学通报,2014,30(6):159-162.Li Z Y,Zheng L,LU L H,et al.Improvement in the H2SO4-H2O2digestion method for determining plant total nitrogen[J].Chinese Agricultural Science Bulletin,2014,30(6):159-162.
31.卢超.两种测定湿地植物总磷方法的比较研究[J].江西农业学报,2009,21(8):142-144.Lu C.Comparative study on two methods for determination of total phosphorus in wetland plants[J].Acta Agriculturae Jiangxi,2009,21(8):142-144.
32.宾振钧,王静静,张文鹏,等.氮肥添加对青藏高原高寒草甸6个群落优势种生态化学计量学特征的影响[J].植物生态学报,2014,38(3):231-237.Bin Z J,Wang J J,Zhang W P,et al.Effects of N addition on ecological stoichiometric characteristics in six dominant plant species of alpine meadow on the Qinghai-Xizang Plateau,China[J].Chinese Journal of Plant Ecology,2014,38(3):231-237.
33.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.
34.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):11001-11006.
35.任书杰,于贵瑞,陶波,等.中国东部南北样带654种植物叶片氮和磷的化学计量学特征研究[J].环境科学,2007,28(12):2665-2673.Ren S J,Yu G R,Tao B,et al.Leaf nitrogen and phosphorus stoichiometry across 654 terrestrial plant species in NSTEC[J].Environmental Science,2007,28(12):2665-2673.
36.Hoorens B,Aerts R,Stroetenga M.Does initial litter chemistry explain litter mixture effects on decomposition?[J].Oecologia,2003,137(4):578-586.
37.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 and Ecology,2012,44:255-263.
38.白小芳.施肥对华北落叶松养分含量及生态化学计量特征影响研究[D].杨凌:中国科学院研究生院(教育部水土保持与生态环境研究中心),2016.Bai X F.Effcets of fertilization on nutrient content and ecoological stoichiometry characteristics in Larix principisrupprechtii Mayr plantation[D].Yangling:Graduate School of Chinese Academy of Sciences(Center for Soil and Water Conservation and Ecological Environment Research,Ministry of Education),2016.
39.Graciano C,Goya J F,Frangi J L,et al.Fertilization with phosphorus increases soil nitrogen absorption in young plants of Eucalyptus grandis[J].Forest Ecology and Management,2006,236(2-3):202-210.
40.Wang Y F,Wang S P,Xin X R,et al.Competition strategies of resources between Stipa grandis and Cleistogenes squarrosa.Ⅰ.Morphological response of shoot and root on sulfur supply[J].Journal of Integrative Plant Biology,2004,46(1):35-45.
41.刘洋,张健,陈亚梅,等.氮磷添加对巨桉幼苗生物量分配和C∶N∶P化学计量特征的影响[J].植物生态学报,2013,37(10):933-941.Liu Y,Zhang J,Chen Y M,et al.Effect of nitrogen and phosphorus fertilization on biomass allocation and C∶N∶Pstoichiometric characteristics of Eucalyptus grandis seedlings[J].Chinese Journal of Plant Ecology,2013,37(10):933-941.
42.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 Z,Niu D C,Wen H Y,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[J].Chinese Journal of Plant Ecology,2011,35(8):801-807.
43.王方超,邹丽群,唐静,等.氮沉降对杉木和枫香土壤氮磷转化及碳矿化的影响[J].生态学报,2016,36(11):3226-3234.Wang F C,Zou L Q,Tang J,et al.Influence of nitrogen deposition on soil nutrient supply and organic carbon mineralization in Cunninghamia lanceolata and Liquidambar formosana plantations[J].Acta Ecologica Sinica,2016,36(11):3226-3234.
44.Vitousek P M,Howarth R W.Nitrogen limitation on land and in the sea:how can it occur?[J].Biogeochemistry,1991,13(2):87-115.
45.Matzek V,Vitousek P M.N∶P stoichiometry and protein:RNA ratios in vascular plants:an evaluation of the growthrate hypothesis[J].Ecology Letters,2009,12(8):765-771.
46.gren G I.Stoichiometry and nutrition of plant growth in natural communities[J].Annual Review of Ecology,Evolution,and Systematics,2008,39:153-170.
47.赵琼,刘兴宇,胡亚林,等.氮添加对兴安落叶松养分分配和再吸收效率的影响[J].林业科学,2010,46(5):14-19.Zhao Q,Liu X Y,Hu Y L,et al.Effects of nitrogen addition on nutrient allocation and nutrient resorption efficiency in Larix gmelinii[J].Scientia Silvae Sinicae,2010,46(5):14-19.
48.樊后保,刘文飞,徐雷,等.氮沉降下杉木(Cunninghamia lanceolata)人工林凋落叶分解过程中C、N元素动态变化[J].生态学报,2008,28(6):2546-2553.Fan H B,Liu W F,Xu L,et al.Carbon and nitrogen dynamics of decomposing foliar litter in a Chinese fir(Cunninghamia lanceolata)plantation exposed to simulated nitrogen deposition[J].Acta Ecologica Sinica,2008,28(6):2546-2553.
49.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.Wu T G,Chen B F,Xiao Y H,et al.Leaf stoichiometry of trees in three forest types in Pearl River Delta,South China[J].Chinese Journal of Plant Ecology,2010,34(1):58-63.
50.陆姣云,段兵红,杨梅,等.植物叶片氮磷养分重吸收规律及其调控机制研究进展[J].草业学报,2018,27(4):178-188.Lu J Y,Duan B H,Yang M,et al.Research progress in nitrogen and phosphorus resorption from senesced leaves and the influence of ontogenetic and environmental factors[J].Acta Prataculturae Sinica,2018,27(4):178-188.
51.LüX T,Freschet G T,Flynn D F B,et al.Plasticity in leaf and stem nutrient resorption proficiency potentially reinforces plant-soil feedbacks and microscale heterogeneity in a semi-arid grassland[J].Journal of Ecology,2012,100(1):144-150.
52.Herbert D A,Williams M,Rastetter E B.A model analysis of N and P limitation on carbon accumulation in Amazonian secondary forest after alternate land-use abandonment[J].Biogeochemistry,2003,65(1):121-150.
53.刘万德,苏建荣,李帅锋,等.云南普洱季风常绿阔叶林演替系列植物和土壤C、N、P化学计量特征[J].生态学报,2010,30(23):6581-6590.Liu W D,Su J R,Li S F,et al.Stoichiometry study of C,Nand P in plant and soil at different successional stages of monsoon evergreen broad-leaved forest in Pu’er,Yunnan Province[J].Acta Ecologica Sinica,2010,30(23):6581-6590.
54.Sardans J,Pe1uelas J.Trees increase their P∶N ratio with size[J].Global Ecology and Biogeography,2015,24(2):147-156.
55.Nakaji T,Fukami M,Dokiya Y,et al.Effects of high nitrogen load on growth,photosynthesis and nutrient status of Cryptomeria japonica and Pinus densiflora seedlings[J].Trees,2001,15(8):453-461.
56.Nakaji T,Takenaga S,Kuroha M,et al.Photosynthetic response of Pinus densiflora seedlings to high nitrogen load[J].Environmental Sciences,2002,9(4):269-282.
57.Hikosaka K,Hanba Y T,Hirose T,et al.Photosynthetic nitrogen-use efficiency in leaves of woody and herbaceous species[J].Functional Ecology,1998,12(6):896-905.
58.Wright I J,Reich P B,Cornelissen J H C,et al.Modulation of leaf economic traits and trait relationships by climate[J].Global Ecology and Biogeography,2005,14(5):411-421.
59.Bowman W D.Accumulation and use of nitrogen and phosphorus following fertilization in two alpine tundra communities[J].Oikos,1994,70(2):261-270.
60.郑淑霞,上官周平.不同功能型植物光合特性及其与叶氮含量、比叶重的关系[J].生态学报,2007,27(1):171-181.Zheng S X,Shangguan Z P.Photosynthetic characteristics and their relationships with leaf nitrogen content and leaf mass per area in different plant functional types[J].Acta Ecologica Sinica,2007,27(1):171-181.
61.孙金伟,吴家兵,任亮,等.氮添加对长白山阔叶红松林2种树木幼苗光合生理生态特征的影响[J].生态学报,2016,36(21):6777-6785.Sun J W,Wu J B,Ren L,et al.Response of photosynthetic physiological characteristics to nitrogen addition by seedlings of two dominant tree species in a broadleaved-Korean pine mixed forest on Changbai Mountain[J].Acta Ecologica Sinica,2016,36(21):6777-6785.
62.Takashima T,Hikosaka K,Hirose T.Photosynthesis or persistence:nitrogen allocation in leaves of evergreen and deciduous Quercus species[J].Plant,Cell&Environment,2004,27(8):1047-1054.
63.Galloway J N,Dentener F J,Capone D G,et al.Nitrogen cycles:past,present,and future[J].Biogeochemistry,2004,70(2):153-226.
64.Ericsson A,Nordén L G,Nsholm T,et al.Mineral nutrient imbalances and arginine concentrations in needles of Picea abies(L.)karst.From two areas with different levels of airborne deposition[J].Trees,1993,8(2):67-74.
2.Liu X J,Zhang Y,Han W X,et al.Enhanced nitrogen deposition over China[J].Nature,2013,494(7438):459-462.
3.Galloway J N,Cowling E B.Reactive nitrogen and the world:200 years of change[J].AMBIO:A Journal of the Human Environment,2002,31(2):64-71.
4.Lü C Q,Tian H Q.Spatial and temporal patterns of nitrogen deposition in China:synthesis of observational data[J].Journal of Geophysical Research:Atmospheres,2007,112(D22):D22S05.
5.邓小文,韩士杰.氮沉降对森林生态系统土壤碳库的影响[J].生态学杂志,2007,26(10):1622-1627.Deng X W,Han S J.Impact of nitrogen deposition on forest soil carbon pool[J].Chinese Journal of Ecology,2007,26(10):1622-1627.
6.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.Wang J Y,Wang S Q,Li R L,et al.C∶N∶P stoichiometric characteristics of four forest types’dominant tree species in China[J].Chinese Journal of Plant Ecology,2011,35(6):587-595.
7.Magill A H,Aber J D,Currie W S,et al.Ecosystem response to 15 years of chronic nitrogen additions at the Harvard Forest LTER,Massachusetts,USA[J].Forest Ecology and Management,2004,196(1):7-28.
8.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.
9.Elser J J,Andersen T,Baron J S,et al.Shifts in lake N∶Pstoichiometry and nutrient limitation driven by atmospheric nitrogen deposition[J].Science,2009,326(5954):835-837.
10.Pe1uelas J,Sardans J,Rivas-Ubach A,et al.The human‐induced imbalance between C,N and P in Earth’s life system[J].Global Change Biology,2012,18(1):3-6.
11.Turner M M,Henry H A L.Interactive effects of warming and increased nitrogen deposition on15N tracer retention in a temperate old field:seasonal trends[J].Global Change Biology,2009,15(12):2885-2893.
12.Passarinho J A P,Lamosa P,Baeta J P,et al.Annual changes in the concentration of minerals and organic compounds of Quercus suber leaves[J].Physiologia Plantarum,2006,127(1):100-110.
13.Fife D N,Nambiar E K,Saur E.Retranslocation of foliar nutrients in evergreen tree species planted in a Mediterranean environment[J].Tree Physiology,2008,28(2):187-196.
14.Phoenix G K,Booth R E,Leake J R,et al.Simulated pollutant nitrogen deposition increases P demand and enhances root-surface phosphatase activities of three plant functional types in a calcareous grassland[J].New Phytologist,2004,161(1):279-290.
15.Fujita Y,De Ruiter P C,Wassen M J,et al.Time-dependent,species-specific effects of N∶P stoichiometry on grassland plant growth[J].Plant and Soil,2010,334(1-2):99-112.
16.Chapin III S F,Matson P A,Mooney H A.Principles of terrestrial ecosystem ecology[M].New York,NY:Springer,2002.
17.Koojiman S A L M.The stoichiometry of animal energetics[J].Journal of Theoretical Biology,1995,177(2):139-149.
18.Aerts R.Nutrient resorption from senescing leaves of perennials:are there general patterns?[J].Journal of Ecology,1996,84(4):597-608.
19.Cernusak L A,Aranda J,Marshall J D,et al.Large variation in whole-plant water-use efficiency among tropical tree species[J].New Phytologist,2007,173(2):294-305.
20.郭淑青,李文金,张仁懿,等.氮磷添加对金露梅叶片化学计量及光合特征的影响[J].广西植物,2014,34(5):629-634,718.Guo S Q,Li W J,Zhang R Y,et al.Effects of N and P additions on foliar stoichiometry and photosynthetic characteristics of Potentilla fruiticosa[J].Guihaia,2014,34(5):629-634,718.
21.安慧.子午岭林区典型植物生长的氮素调控机理[D].杨凌:西北农林科技大学,2008.An H.Regulating mechanism of nitrogen of typical plants growth in Ziwuling forest region[D].Yangling:Northwest A&F University,2008.
22.Warren C R,ADAMS M A.Evergreen trees do not maximize instantaneous photosynthesis[J].Trends in Plant Science,2004,9(6):270-274.
23.王振南,杨惠敏.植物碳氮磷生态化学计量对非生物因子的响应[J].草业科学,2013,30(6):927-934.Wang Z N,Yang H M.Response of ecological stoichiometry of carbon,nitrogen and phosphorus in plants to abiotic environmental factors[J].Pratacultural Science,2013,30(6):927-934.
24.吕丽华,赵明,赵久然,等.不同施氮量下夏玉米冠层结构及光合特性的变化[J].中国农业科学,2008,41(9):2624-2632.LüL H,Zhao M,Zhao J R,et al.Canopy structure and photosynthesis of summer maize under different nitrogen fertilizer application rates[J].Scientia Agricultura Sinica,2008,41(9):2624-2632.
25.杨浩,罗亚晨.糙隐子草功能性状对氮添加和干旱的响应[J].植物生态学报,2015,39(1):32-42.Yang H,Luo Y C.Responses of the functional traits in Cleistogenes squarrosa to nitrogen addition and drough[J].Chinese Journal of Plant Ecology,2015,39(1):32-42.
26.谌贤,刘洋,邓静,等.川西亚高山森林凋落物不同分解阶段碳氮磷化学计量特征及种间差异[J].植物研究,2017,37(2):216-226.Shen X,Liu Y,Deng J,et al.C,N and P stoichiometry at different stages of litter decomposition in subalpine forest of western Sichuan province and inter species comparison[J].Bulletin of Botanical Research,2017,37(2):216-226.
27.刘荣杰,吴亚丛,张英,等.中国北亚热带天然次生林与杉木人工林土壤活性有机碳库的比较[J].植物生态学报,2012,36(5):431-437.Liu R J,Wu Y C,Zhang Y,et al.Comparison of soil labile organic carbon in Chinese fir plantations and natural secondary forests in north subtropical areas of China[J].Chinese Journal of Plant Ecology,2012,36(5):431-437.
28.胡艳玲,韩士杰,李雪峰,等.长白山原始林和次生林土壤有效氮含量对模拟氮沉降的响应[J].东北林业大学学报,2009,37(5):36-38,42.Hu Y L,Han S J,Li X F,et al.Responses of soil available nitrogen of natural forest and secondary forest to simulated N deposition in Changbai Mountain[J].Journal of Northeast Forestry University,2009,37(5):36-38,42.
29.唐艳,王传宽.东北主要树种光合作用可行的离体测定方法[J].植物生态学报,2011,35(4):452-462.Tang Y,Wang C K.A feasible method for measuring photosynthesis in vitro for major tree species in northeastern China[J].Chinese Journal of Plant Ecology,2011,35(4):452-462.
30.李朝英,郑路,卢立华,等.测定植物全氮的H2SO4-H2O2消煮法改进[J].中国农学通报,2014,30(6):159-162.Li Z Y,Zheng L,LU L H,et al.Improvement in the H2SO4-H2O2digestion method for determining plant total nitrogen[J].Chinese Agricultural Science Bulletin,2014,30(6):159-162.
31.卢超.两种测定湿地植物总磷方法的比较研究[J].江西农业学报,2009,21(8):142-144.Lu C.Comparative study on two methods for determination of total phosphorus in wetland plants[J].Acta Agriculturae Jiangxi,2009,21(8):142-144.
32.宾振钧,王静静,张文鹏,等.氮肥添加对青藏高原高寒草甸6个群落优势种生态化学计量学特征的影响[J].植物生态学报,2014,38(3):231-237.Bin Z J,Wang J J,Zhang W P,et al.Effects of N addition on ecological stoichiometric characteristics in six dominant plant species of alpine meadow on the Qinghai-Xizang Plateau,China[J].Chinese Journal of Plant Ecology,2014,38(3):231-237.
33.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.
34.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):11001-11006.
35.任书杰,于贵瑞,陶波,等.中国东部南北样带654种植物叶片氮和磷的化学计量学特征研究[J].环境科学,2007,28(12):2665-2673.Ren S J,Yu G R,Tao B,et al.Leaf nitrogen and phosphorus stoichiometry across 654 terrestrial plant species in NSTEC[J].Environmental Science,2007,28(12):2665-2673.
36.Hoorens B,Aerts R,Stroetenga M.Does initial litter chemistry explain litter mixture effects on decomposition?[J].Oecologia,2003,137(4):578-586.
37.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 and Ecology,2012,44:255-263.
38.白小芳.施肥对华北落叶松养分含量及生态化学计量特征影响研究[D].杨凌:中国科学院研究生院(教育部水土保持与生态环境研究中心),2016.Bai X F.Effcets of fertilization on nutrient content and ecoological stoichiometry characteristics in Larix principisrupprechtii Mayr plantation[D].Yangling:Graduate School of Chinese Academy of Sciences(Center for Soil and Water Conservation and Ecological Environment Research,Ministry of Education),2016.
39.Graciano C,Goya J F,Frangi J L,et al.Fertilization with phosphorus increases soil nitrogen absorption in young plants of Eucalyptus grandis[J].Forest Ecology and Management,2006,236(2-3):202-210.
40.Wang Y F,Wang S P,Xin X R,et al.Competition strategies of resources between Stipa grandis and Cleistogenes squarrosa.Ⅰ.Morphological response of shoot and root on sulfur supply[J].Journal of Integrative Plant Biology,2004,46(1):35-45.
41.刘洋,张健,陈亚梅,等.氮磷添加对巨桉幼苗生物量分配和C∶N∶P化学计量特征的影响[J].植物生态学报,2013,37(10):933-941.Liu Y,Zhang J,Chen Y M,et al.Effect of nitrogen and phosphorus fertilization on biomass allocation and C∶N∶Pstoichiometric characteristics of Eucalyptus grandis seedlings[J].Chinese Journal of Plant Ecology,2013,37(10):933-941.
42.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 Z,Niu D C,Wen H Y,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[J].Chinese Journal of Plant Ecology,2011,35(8):801-807.
43.王方超,邹丽群,唐静,等.氮沉降对杉木和枫香土壤氮磷转化及碳矿化的影响[J].生态学报,2016,36(11):3226-3234.Wang F C,Zou L Q,Tang J,et al.Influence of nitrogen deposition on soil nutrient supply and organic carbon mineralization in Cunninghamia lanceolata and Liquidambar formosana plantations[J].Acta Ecologica Sinica,2016,36(11):3226-3234.
44.Vitousek P M,Howarth R W.Nitrogen limitation on land and in the sea:how can it occur?[J].Biogeochemistry,1991,13(2):87-115.
45.Matzek V,Vitousek P M.N∶P stoichiometry and protein:RNA ratios in vascular plants:an evaluation of the growthrate hypothesis[J].Ecology Letters,2009,12(8):765-771.
46.gren G I.Stoichiometry and nutrition of plant growth in natural communities[J].Annual Review of Ecology,Evolution,and Systematics,2008,39:153-170.
47.赵琼,刘兴宇,胡亚林,等.氮添加对兴安落叶松养分分配和再吸收效率的影响[J].林业科学,2010,46(5):14-19.Zhao Q,Liu X Y,Hu Y L,et al.Effects of nitrogen addition on nutrient allocation and nutrient resorption efficiency in Larix gmelinii[J].Scientia Silvae Sinicae,2010,46(5):14-19.
48.樊后保,刘文飞,徐雷,等.氮沉降下杉木(Cunninghamia lanceolata)人工林凋落叶分解过程中C、N元素动态变化[J].生态学报,2008,28(6):2546-2553.Fan H B,Liu W F,Xu L,et al.Carbon and nitrogen dynamics of decomposing foliar litter in a Chinese fir(Cunninghamia lanceolata)plantation exposed to simulated nitrogen deposition[J].Acta Ecologica Sinica,2008,28(6):2546-2553.
49.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.Wu T G,Chen B F,Xiao Y H,et al.Leaf stoichiometry of trees in three forest types in Pearl River Delta,South China[J].Chinese Journal of Plant Ecology,2010,34(1):58-63.
50.陆姣云,段兵红,杨梅,等.植物叶片氮磷养分重吸收规律及其调控机制研究进展[J].草业学报,2018,27(4):178-188.Lu J Y,Duan B H,Yang M,et al.Research progress in nitrogen and phosphorus resorption from senesced leaves and the influence of ontogenetic and environmental factors[J].Acta Prataculturae Sinica,2018,27(4):178-188.
51.LüX T,Freschet G T,Flynn D F B,et al.Plasticity in leaf and stem nutrient resorption proficiency potentially reinforces plant-soil feedbacks and microscale heterogeneity in a semi-arid grassland[J].Journal of Ecology,2012,100(1):144-150.
52.Herbert D A,Williams M,Rastetter E B.A model analysis of N and P limitation on carbon accumulation in Amazonian secondary forest after alternate land-use abandonment[J].Biogeochemistry,2003,65(1):121-150.
53.刘万德,苏建荣,李帅锋,等.云南普洱季风常绿阔叶林演替系列植物和土壤C、N、P化学计量特征[J].生态学报,2010,30(23):6581-6590.Liu W D,Su J R,Li S F,et al.Stoichiometry study of C,Nand P in plant and soil at different successional stages of monsoon evergreen broad-leaved forest in Pu’er,Yunnan Province[J].Acta Ecologica Sinica,2010,30(23):6581-6590.
54.Sardans J,Pe1uelas J.Trees increase their P∶N ratio with size[J].Global Ecology and Biogeography,2015,24(2):147-156.
55.Nakaji T,Fukami M,Dokiya Y,et al.Effects of high nitrogen load on growth,photosynthesis and nutrient status of Cryptomeria japonica and Pinus densiflora seedlings[J].Trees,2001,15(8):453-461.
56.Nakaji T,Takenaga S,Kuroha M,et al.Photosynthetic response of Pinus densiflora seedlings to high nitrogen load[J].Environmental Sciences,2002,9(4):269-282.
57.Hikosaka K,Hanba Y T,Hirose T,et al.Photosynthetic nitrogen-use efficiency in leaves of woody and herbaceous species[J].Functional Ecology,1998,12(6):896-905.
58.Wright I J,Reich P B,Cornelissen J H C,et al.Modulation of leaf economic traits and trait relationships by climate[J].Global Ecology and Biogeography,2005,14(5):411-421.
59.Bowman W D.Accumulation and use of nitrogen and phosphorus following fertilization in two alpine tundra communities[J].Oikos,1994,70(2):261-270.
60.郑淑霞,上官周平.不同功能型植物光合特性及其与叶氮含量、比叶重的关系[J].生态学报,2007,27(1):171-181.Zheng S X,Shangguan Z P.Photosynthetic characteristics and their relationships with leaf nitrogen content and leaf mass per area in different plant functional types[J].Acta Ecologica Sinica,2007,27(1):171-181.
61.孙金伟,吴家兵,任亮,等.氮添加对长白山阔叶红松林2种树木幼苗光合生理生态特征的影响[J].生态学报,2016,36(21):6777-6785.Sun J W,Wu J B,Ren L,et al.Response of photosynthetic physiological characteristics to nitrogen addition by seedlings of two dominant tree species in a broadleaved-Korean pine mixed forest on Changbai Mountain[J].Acta Ecologica Sinica,2016,36(21):6777-6785.
62.Takashima T,Hikosaka K,Hirose T.Photosynthesis or persistence:nitrogen allocation in leaves of evergreen and deciduous Quercus species[J].Plant,Cell&Environment,2004,27(8):1047-1054.
63.Galloway J N,Dentener F J,Capone D G,et al.Nitrogen cycles:past,present,and future[J].Biogeochemistry,2004,70(2):153-226.
64.Ericsson A,Nordén L G,Nsholm T,et al.Mineral nutrient imbalances and arginine concentrations in needles of Picea abies(L.)karst.From two areas with different levels of airborne deposition[J].Trees,1993,8(2):67-74.