落叶松人工林草本植物群落特征和生物量对氮添加的响应
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摘要
以华北落叶松(Larix principis-rupprechtii)人工林幼龄林为研究对象,设置3个梯度的氮添加水平(0,20和50kgN/(ha·a),即N0,N20和N50),研究氮添加对人工林草本植物群落特征及地上生产力的影响,得到以下结果。1)氮添加改变了土壤微环境,降低土壤温度、显著地增加土壤水分。2)氮添加在一定程度上促进落叶松人工林草本植物群落的生长(P>0.05)。与对照组(N0)相比,植物群落总盖度在N20和N50样方中分别增加6.83%和15.03%(2014年),或9.80%和12.94%(2016年),但草本植物地上生物量的累积无显著变化;群落总盖度和地上生物量与土壤含水量显著正相关(P<0.05)。3)氮添加对落叶松人工林草本植物群落的多样性和均匀度无显著影响,但是低浓度氮添加(即N20)显著地增加单位面积内物种的丰富度(2014年单位面积内物种丰富度在N20样方中增加23.33%,P<0.05)。4)氮添加对不同植物功能群的影响不同,其中氮添加促进蓼科和莎草科植物的生长,抑制禾本科和菊科植物的生长,对豆科植物无影响,说明温带落叶松人工林草本植物群落对氮沉降的响应敏感,氮添加在一定程度上促进草本植物的生长,同时氮添加改变了草本植物不同功能群间的竞争力。
The research was conducted in a sapling Larix principis-rupprechtii, plantation set three nitrogen addition levels(0, 20 and 50 kg N/(ha·a), called N0, N20 and N50), in order to assess the influence of nitrogen addition on plant communities characteristics and above-ground biomass. The results showed that nitrogen addition altered soil micro-environment, reduced soil temperature and significantly increased soil moisture. Nitrogen addition partly promotes the growth of herb community(increased by 6.83% and 6.83% in N20 and N50 in 2014,respectively, compared with the control; while increased by 9.80% and 6.83% in N20 and N50 in 2016,respectively, P>0.05). The community coverage and aboveground biomass were significantly and positively correlated with soil moisture(P<0.05). The diversity and evenness had no significant response to nitrogen addition,but N20 significantly increases the species richness per area(significantly increased by 23.33% in N20 samples in2014, P<0.05). Nitrogen addition exerted different influences on plant functional groups, i.e., nitrogen addition promoted the growth of polygonaceae and cyperaceae plants, restrained gramineae and compositae plants and had no influence of leguminous plants. The results suggested that herb communities are sensitive to nitrogen addition in temperate Larix principis-rupprechtii plantations, and nitrogen addition partly promotes the growth of herb plants and changes competition among different herbaceous plants functional groups.
The research was conducted in a sapling Larix principis-rupprechtii, plantation set three nitrogen addition levels(0, 20 and 50 kg N/(ha·a), called N0, N20 and N50), in order to assess the influence of nitrogen addition on plant communities characteristics and above-ground biomass. The results showed that nitrogen addition altered soil micro-environment, reduced soil temperature and significantly increased soil moisture. Nitrogen addition partly promotes the growth of herb community(increased by 6.83% and 6.83% in N20 and N50 in 2014,respectively, compared with the control; while increased by 9.80% and 6.83% in N20 and N50 in 2016,respectively, P>0.05). The community coverage and aboveground biomass were significantly and positively correlated with soil moisture(P<0.05). The diversity and evenness had no significant response to nitrogen addition,but N20 significantly increases the species richness per area(significantly increased by 23.33% in N20 samples in2014, P<0.05). Nitrogen addition exerted different influences on plant functional groups, i.e., nitrogen addition promoted the growth of polygonaceae and cyperaceae plants, restrained gramineae and compositae plants and had no influence of leguminous plants. The results suggested that herb communities are sensitive to nitrogen addition in temperate Larix principis-rupprechtii plantations, and nitrogen addition partly promotes the growth of herb plants and changes competition among different herbaceous plants functional groups.
引文
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[28]Hirao T,Murakami M,Kashizaki A.Importance of the understory stratum to entomofaunal diversity in a temperate deciduous forest.Ecological Research,2009,24(2):263-272
[29]Nilsson M C,Wardle D A.Understory vegetation as a forest ecosystem driver:evidence from the northern Swedish boreal forest.Frontiers in Ecology and the Environment,2005,3(8):421-428
[30]王大鹏,王文斌,郑亮,等.中国主要人工林土壤有机碳的比较.生态环境学报,2014,23(4):698-704
[31]Piao S L,Fang J Y,Ciais P,et al.The carbon balance of terrestrial ecosystems in China.Nature,2009,458:1009-1013
[32]李国雷,刘勇,于海群,等.油松(Pinus tabulaeformis)人工林林下植被发育对油松生长节律的响应.生态学报,2009,29(3):1264-1275
[33]于景金,谷建才,李校,等.塞罕坝落叶松人工林下植物多样性研究.林业资源管理,2008(5):90-94
[34]Neufeld H S,Young D R.Ecophysiology of the herbaceous layer in temperate deciduous forests//Gilliam F S.The herbaceous layer in forests of eastern North America.2nd ed.New York:Oxford University Press,2014:34-39
[35]Ma S,Verheyen K,Props R,et al.Plant and soil microbe responses to light,warming and nitrogen addition in a temperate forest.Functional Ecology,2018,32:1293-1303
[36]Strengbom J,N?sholm T,Ericson L.Light,not nitrogen,limits growth of the grass Deschampsia flexuosa in boreal forests.Canadian Journal of botany,2004,82(4):430-435
[37]鲁显楷,莫江明,董少峰.氮沉降对森林生物多样性的影响.生态学报,2008,28(11):5532-5548
[38]Valladares F,Laanisto L,Niinemets U,et al.Shedding light on shade:ecological perspectives of understory plant life.Transactions of the Botanical Society of Edinburgh,2016,9(3):237-251
[39]Beatty S W.Habitat heterogeneity and maintenance of species in understory communities//Gilliam F S.The herbaceous layer of forests in eastern North America,2nd ed.New York:Oxford University Press,2014:215-232
[40]Small C J,Mccarthy B C.Spatial and temporal variability of herbaceous vegetation in an eastern deciduous forest.Plant Ecology,2003,164(1):37-48
[41]Yang K,Zhu J,Gu J,et al.Changes in soil phosphorus fractions after 9 years of continuous nitrogen addition in a Larix gmelinii plantation.Annals of Forest Science,2015,72(4):435-442
[42]Hu Y L,Zeng D H,Liu Y X,et al.Responses of soil chemical and biological properties to nitrogen addition in a Dahurian larch plantation in northeast China.Plant&Soil,2010,333(1/2):81-92
[43]Li F,Chen B,Liang L,et al.Responses of tree and insect herbivores to elevated nitrogen inputs:a metaanalysis.Acta Oecologica,2016,77:160-167
[44]Lau J A,Bowling E J,Gentry L E,et al.Direct and interactive effects of light and nutrients on the legume-rhizobia mutualism.Acta Oecologica,2012,39(2):80-86
[45]Reef R,Slot M,Motro U,et al.The effects of CO2and nutrient fertilisation on the growth and temperature response of the mangrove avicennia germinans.Photosynthesis Research,2016,129(2):159-170
[2]Bobbink R,Hicks K,Galloway J,et al.Global assessment of nitrogen deposition effects on terrestrial plant diversity:a synthesis.Ecological Applications:A Publication of the Ecological Society of America,2010,20(1):30-59
[3]Manning P,Newington J E,Robson H R,et al.Decoupling the direct and indirect effects of nitrogen deposition on ecosystem function.Ecology Letters,2006,9(9):1015-1024
[4]毛晋花,邢亚娟,闫国永,等.陆生植物生物量分配对模拟氮沉降响应的Meta分析.生态学报,2018,38(9):1-11
[5]Walter C A,Raiff D T,Burnham M B,et al.Nitrogen fertilization interacts with light to increase Rubus spp.cover in a temperate forest.Plant Ecology,2016,217(4):1-10
[6]Wu J,Liu W,Fan H,et al.Asynchronous responses of soil microbial community and understory plant community to simulated nitrogen deposition in a subtropical forest.Ecology&Evolution,2013,3(11):3895-3905
[7]Lebauer D S,Treseder K K.Nitrogen limitation of net primary productivity in terrestrial ecosystems is globally distributed.Ecology,2008,89(2):371-379
[8]Neff J C,Townsend A R,Gleixner G,et al.Variable effects of nitrogen additions on the stability and turnover of soil carbon.Nature,2002,419:915-917
[9]Paoli G D,Curran L M.Soil nutrients limit fine litter production and tree growth in mature lowland forest of southwestern borneo.Ecosystems,2007,10(3):503-518
[10]Baribault T W,Kobe R K,Finley A O.Tropical tree growth is correlated with soil phosphorus,potassium,and calcium,though not for legumes.Ecological Monographs,2012,82(2):189-203
[11]李化山,汪金松,刘星,等.模拟N沉降对太岳山油松人工林和天然林草本群落的影响.生态学报,2015,35(11):3710-3721
[12]Talhelm A F,Burton A J,Pregitzer K S,et al.Simulated nitrogen deposition reduces the abundance of dominant forest understory and groundcover plants.Forest Ecology&Management,2013,293:39-48
[13]Lu X K,Mo J M,Glliam F S,et al.Effects of experimental nitrogen additions on plant diversity in an old-growth tropical forest.Global Change Biology,2010,16(10):2688-2700
[14]Gilliam F S.Response of the herbaceous layer of forest ecosystems to excess nitrogen deposition.Journal of Ecology,2006,94(6):1176-1191
[15]Concilio A L,Loik M E.Elevated nitrogen effects on Bromus tectorum dominance and native plant diversity in an arid montane ecosystem.Applied Vegetation Science,2013,16(4):598-609
[16]Thomas R Q,Canham C D,Weathers K C,et al.Increased tree carbon storage in response to nitrogen deposition in the US.Nature Geoscience,2010,3(1):229-244
[17]Eilts J A,Mittelbach G G,Reynolds H L,et al.Resource heterogeneity,soil fertility,and species diversity:effects of clonal species on plant communities.American Naturalist,2011,177(5):574-588
[18]Vitousek P M,Porder S,Houlton B Z,et al.Terrestrial phosphorus limitation:mechanisms,implications,and nitrogen-phosphorus interactions.Ecological Applications,2010,20(1):5-15
[19]Cusack D F,Silver W L,Torn M S,et al.Changes in microbial community characteristics and soil organic matter with nitrogen additions in two tropical forests.Ecology,2011,92(3):621-632
[20]Van der Heiiden M G A,Bardgett R D,Straalen N MV.The unseen majority:soil microbes as drivers of plant diversity and productivity in terrestrial ecosystems.Ecology Letters,2008,11(3):296-310
[21]Gilliam F S.Hockenberry A W,Adams M B.Effects of atmospheric nitrogen deposition on the herbaceous layer of a central Appalachian hardwood forest.Journal of the Torrey Botanical Society,2006,133(2):240-254
[22]Vitousek P M,Howarth R W.Nitrogen limitation on land and in the sea:how can it occur?.Biogeochemistry,1991,13(2):87-115
[23]Xia J Y,Wan S Q.Global response patterns of terrestrial plant species to nitrogen addition.New Phytologist,2008,179(2):428-439
[24]Galloway J N,Townsend A R,Erisman J W,et al.Transformation of the nitrogen cycle:recent trends,questions,and potential solutions.Science,2008,320:889-892
[25]Takafumi H,Hiura T.Effects of disturbance history and environmental factors on the diversity and productivity of understory vegetation in a cool-temperate forest in Japan.Forest Ecology&Management,2009,257(3):843-857
[26]Tchouto M G P,Boer W F D,Wilde J J F E D,et al.Diversity patterns in the flora of the Campo-Ma’an Rain Forest,Cameroon:do tree species tell it all?.Biodiversity&Conservation,2006,15(4):1353-1374
[27]Royo A A,Carson W P.On the formation of dense understory layers in forests worldwide:consequences and implications for forest dynamics,biodiversity,and succession.Canadian Journal of Forest Research,2006,36(6):1345-1362
[28]Hirao T,Murakami M,Kashizaki A.Importance of the understory stratum to entomofaunal diversity in a temperate deciduous forest.Ecological Research,2009,24(2):263-272
[29]Nilsson M C,Wardle D A.Understory vegetation as a forest ecosystem driver:evidence from the northern Swedish boreal forest.Frontiers in Ecology and the Environment,2005,3(8):421-428
[30]王大鹏,王文斌,郑亮,等.中国主要人工林土壤有机碳的比较.生态环境学报,2014,23(4):698-704
[31]Piao S L,Fang J Y,Ciais P,et al.The carbon balance of terrestrial ecosystems in China.Nature,2009,458:1009-1013
[32]李国雷,刘勇,于海群,等.油松(Pinus tabulaeformis)人工林林下植被发育对油松生长节律的响应.生态学报,2009,29(3):1264-1275
[33]于景金,谷建才,李校,等.塞罕坝落叶松人工林下植物多样性研究.林业资源管理,2008(5):90-94
[34]Neufeld H S,Young D R.Ecophysiology of the herbaceous layer in temperate deciduous forests//Gilliam F S.The herbaceous layer in forests of eastern North America.2nd ed.New York:Oxford University Press,2014:34-39
[35]Ma S,Verheyen K,Props R,et al.Plant and soil microbe responses to light,warming and nitrogen addition in a temperate forest.Functional Ecology,2018,32:1293-1303
[36]Strengbom J,N?sholm T,Ericson L.Light,not nitrogen,limits growth of the grass Deschampsia flexuosa in boreal forests.Canadian Journal of botany,2004,82(4):430-435
[37]鲁显楷,莫江明,董少峰.氮沉降对森林生物多样性的影响.生态学报,2008,28(11):5532-5548
[38]Valladares F,Laanisto L,Niinemets U,et al.Shedding light on shade:ecological perspectives of understory plant life.Transactions of the Botanical Society of Edinburgh,2016,9(3):237-251
[39]Beatty S W.Habitat heterogeneity and maintenance of species in understory communities//Gilliam F S.The herbaceous layer of forests in eastern North America,2nd ed.New York:Oxford University Press,2014:215-232
[40]Small C J,Mccarthy B C.Spatial and temporal variability of herbaceous vegetation in an eastern deciduous forest.Plant Ecology,2003,164(1):37-48
[41]Yang K,Zhu J,Gu J,et al.Changes in soil phosphorus fractions after 9 years of continuous nitrogen addition in a Larix gmelinii plantation.Annals of Forest Science,2015,72(4):435-442
[42]Hu Y L,Zeng D H,Liu Y X,et al.Responses of soil chemical and biological properties to nitrogen addition in a Dahurian larch plantation in northeast China.Plant&Soil,2010,333(1/2):81-92
[43]Li F,Chen B,Liang L,et al.Responses of tree and insect herbivores to elevated nitrogen inputs:a metaanalysis.Acta Oecologica,2016,77:160-167
[44]Lau J A,Bowling E J,Gentry L E,et al.Direct and interactive effects of light and nutrients on the legume-rhizobia mutualism.Acta Oecologica,2012,39(2):80-86
[45]Reef R,Slot M,Motro U,et al.The effects of CO2and nutrient fertilisation on the growth and temperature response of the mangrove avicennia germinans.Photosynthesis Research,2016,129(2):159-170