土壤动物对森林凋落物分解的影响:机制和模拟
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摘要
土壤动物是森林生态系统的重要组成部分。本文综述了土壤动物在不同森林生态系统中对凋落物分解过程的贡献大小及影响因素、土壤动物影响凋落物分解的作用机制以及凋落物分解模型的研究进展,以期为更好地理解森林生态系统中土壤动物在地上、地下生态过程中的作用。我们试图建立一个概念模型来模拟土壤动物在凋落物分解过程中的贡献。土壤动物可以通过直接作用于凋落物(包括移动、破碎、取食等),或间接作用于土壤(穿梭、掘穴等影响凋落物分解微环境)和微生物(影响定殖于凋落物中的微生物群落种类、数量和活性)影响凋落物的分解过程。温度和水分条件是影响土壤动物活动的重要因素,普遍认为热带森林中的土壤动物作用要大于亚热带森林、温带森林和高山/亚高山森林。未来该领域的研究应注重如何在凋落物分解模型中体现土壤动物的作用机制以及利用野外实验数据量化土壤动物对凋落物分解过程的贡献等。
Soil fauna is an important component of forest ecosystems. We reviewed the contribution of soil fauna on litter decomposition in different forest ecosystems and the influencing factors,the underlying mechanisms of soil fauna affecting litter decomposition,and research advances on litter decomposition models,aiming to better understand the effects of soil fauna on aboveground and belowground ecological processes in forest ecosystems. We proposed a novel framework for modeling litter decomposition by considering the contribution of soil fauna. Soil fauna can affect litter decomposition by acting directly on litter( including moving,breaking and feeding,etc.) or indirectly on the soil( affecting litter decomposition microenvironment through shuttling and burrowing) and microorganisms( influencing the composition,quantity and activities of microbial communities which colonized in litter). Temperature and moisture are important factors affecting activities of soil fauna. The effects of soil fauna on litter decomposition in tropical forests are stronger than in subtropical forests,temperate forests and alpine/subalpine forests. Future modeling studies should focus on how to mimic the effects of soil fauna on litter decomposition and validate the modeling results using field experiment data.
Soil fauna is an important component of forest ecosystems. We reviewed the contribution of soil fauna on litter decomposition in different forest ecosystems and the influencing factors,the underlying mechanisms of soil fauna affecting litter decomposition,and research advances on litter decomposition models,aiming to better understand the effects of soil fauna on aboveground and belowground ecological processes in forest ecosystems. We proposed a novel framework for modeling litter decomposition by considering the contribution of soil fauna. Soil fauna can affect litter decomposition by acting directly on litter( including moving,breaking and feeding,etc.) or indirectly on the soil( affecting litter decomposition microenvironment through shuttling and burrowing) and microorganisms( influencing the composition,quantity and activities of microbial communities which colonized in litter). Temperature and moisture are important factors affecting activities of soil fauna. The effects of soil fauna on litter decomposition in tropical forests are stronger than in subtropical forests,temperate forests and alpine/subalpine forests. Future modeling studies should focus on how to mimic the effects of soil fauna on litter decomposition and validate the modeling results using field experiment data.
引文
包剑利,殷秀琴,李晓强.2015.长白山牛皮杜鹃凋落物分解及土壤动物的作用.生态学报,35(10):3320-3328.
范换,王绍军,阮宏华,等.2014.苏北沿海不同土地利用类型土壤动物群落及其对凋落物分解的影响.南京林业大学学报:自然科学版,38(3):1-7.
黄玉梅,杨万勤,张健.2015.川西亚高山云杉叶凋落物质量损失过程及土壤生物的作用.长江流域资源与环境,24(4):676-683.
廖姝,杨万勤,彭艳,等.2016.土壤动物对四川盆地四种树种凋落叶在不同物候时期可溶性碳和氮动态的影响.生态学杂志,35(12):3172-3182.
刘瑞龙,李维民,杨万勤,等.2013.土壤动物对川西高山/亚高山森林凋落物分解的贡献.应用生态学报,24(12):3354-3360.
谭羽,杨万勤,廖姝,等.2017.川西高寒生态系统不同海拔土壤动物对冬季凋落叶腐殖化过程的影响.生态学报,37(5):1595-1602.
王利峰,和润莲,林杨,等.2016.川西高山林线交错带土壤动物对岷江冷杉和高山杜鹃凋落物分解的贡献.应用生态学报,27(11):3689-3697.
王文君,杨万勤,谭波,等.2013.四川盆地亚热带常绿阔叶林土壤动物对几种典型凋落物分解的影响.生态环境学报,22(9):1488-1495.
王文君,杨万勤,谭波,等.2015.四川盆地亚热带常绿阔叶林不同物候土壤动物对凋落物氮和磷释放的影响.林业科学,51(1):1-11.
王振海,殷秀琴,张成蒙.2016.土壤动物在长白山臭冷杉凋落物分解中的作用.林业科学,52(7):59-67.
夏磊,吴福忠,杨万勤,等.2012.川西亚高山森林凋落物分解初期土壤动物对红桦凋落叶质量损失的贡献.应用生态学报,23(2):301-306.
许湘琴,林植华,陈慧丽.2011.凋落物分解对土壤生物的影响.生态学杂志,30(6):1258-1264.
杨万勤,邓仁菊,张健.2007.森林凋落物分解及其对全球气候变化的响应.应用生态学报,18(12):2889-2895.
杨效东,邹晓明.2006.西双版纳热带季节雨林凋落叶分解与土壤动物群落:两种网孔分解袋的分解实验比较.植物生态学报,30(5):791-801.
殷秀琴,仲伟彦,王海霞,等.2002.小兴安岭森林落叶分解与土壤动物的作用.地理研究,21(6):689-699.
Allison SD,Wallenstein MD,Bradford MA.2010.Soil-carbon response to warming dependent on microbial physiology.Nature Geoscience,3:336-340.
Amosse J,Turberg P,Kohler-Milleret R,et al.2015.Effects of endogeic earthworms on the soil organic matter dynamics and the soil structure in urban and alluvial soil materials.Geoderma,243:50-57.
Anderson JM.1973.The breakdown and decomposition of sweet chestnut(Castanea sativa Mill.)and beech(Fagus sylvatica L.)leaf litter in two deciduous woodland soils.I.Breakdown,leaching and decomposition.Oecologia,12:251-274.
Aubert M,Margerie P,Trap J,et al.2010.Aboveground-belowground relationships in temperate forests:Plant litter composes and microbiota orchestrates.Forest Ecology and Management,259:563-572.
Barajas-Guzmn G,Alvarez-Snchez J.2003.The relationships between litter fauna and rates of litter decomposition in a tropical rain forest.Applied Soil Ecology,24:91-100.
Bonan GB.2008.Forests and climate change:Forcings,feedbacks,and the climate benefits of forests.Science,320:1444-1449.
Bonan GB,Hartman MD,Parton WJ,et al.2013.Evaluating litter decomposition in earth system models with long-term litterbag experiments:An example using the Community Land Model version 4(CLM4).Global Change Biology,19:957-974.
Bonanomi G,Capodilupo M,Incerti G,et al.2015.Litter quality and temperature modulate microbial diversity effects on decomposition in model experiments.Community Ecology,16:167-177.
Bottinelli N,Jouquet P,Capowiez Y,et al.2015.Why is the influence of soil macrofauna on soil structure only considered by soil ecologists?Soil&Tillage Research,146:118-124.
Campbell EE,Parton WJ,Soong JL,et al.2016.Using litter chemistry controls on microbial processes to partition litter carbon fluxes with the Litter Decomposition and Leaching(LIDEL)model.Soil Biology&Biochemistry,100:160-174.
Carcamo HA,Prescott CE,Chanway CP,et al.2001.Do soil fauna increase rates of litter breakdown and nitrogen release in forests of British Columbia,Canada?Canadian Journal of Forest Research,31:1195-1204.
Chertov O,Shaw C,Shashkov M,et al.2017.Romul_Hum model of soil organic matter formation coupled with soil biota activity.III.Parameterisation of earthworm activity.Ecological Modelling,345:140-149.
Coq S,Souquet JM,Meudec E,et al.2010.Interspecific variation in leaf litter tannins drives decomposition in a tropical rain forest of French Guiana.Ecology,91:2080-2091.
Coulis M,Hattenschwiler S,Coq S,et al.2016.Leaf litter consumption by macroarthropods and burial of their faeces enhance decomposition in a Mediterranean ecosystem.Ecosystems,19:1104-1115.
Davidson DA,Grieve IC.2006.Relationships between biodiversity and soil structure and function:Evidence from laboratory and field experiments.Applied Soil Ecology,33:176-185.
Diek9tter T,Wamser S,Wolters V.2010.Landscape and management ffects on structure and function of soil arthropod communities in winter wheat.Agriculture,Ecosystems and Environment,137:108-112.
Frouz J.2002.The effect of soil macrofauna on litter decomposition and soil organic matter accumulation during soil formation in spoil heaps after brown coal mining:A preliminary results.Ekologia Bratislava,21:363-369.
Frouz J.2018.Effects of soil macro-and mesofauna on litter decomposition and soil organic matter stabilization.Geoderma,332:161-172.
Frouz J,Roubickova A,Hedenec P,et al.2015.Do soil fauna really hasten litter decomposition?A meta-analysis of enclosure studies.European Journal of Soil Biology,68:18-24.
Fujii S,Makita N,Mori AS,et al.2016.Plant species control and soil faunal involvement in the processes of above-and below-ground litter decomposition.Oikos,125:883-892.
Garcia-Palacios P,Maestre FT,Kattge J,et al.2013.Climate and litter quality differently modulate the effects of soil fauna on litter decomposition across biomes.Ecology Letters,16:1045-1053.
Gonzalez G,Seastedt TR.2001.Soil fauna and plant litter decomposition in tropical and subalpine forests.Ecology,82:955-964.
Haettenschwiler S,Jorgensen HB.2010.Carbon quality rather than stoichiometry controls litter decomposition in a tropical rain forest.Journal of Ecology,98:754-763.
Handa IT,Aerts R,Berendse F,et al.2014.Consequences of biodiversity loss for litter decomposition across biomes.Nature,509:218-221.
Hou PCL,Zou XM,Huang CY,et al.2005.Plant litter decomposition influenced by soil animals and disturbance in a subtropical rainforest of Taiwan.Pedobiologia,49:539-547.
Illig J,Schatz H,Scheu S,et al.2008.Decomposition and colonization by micro-arthropods of two litter types in a tropical montane rain forest in southern Ecuador.Journal of Tropical Ecology,24:157-167.
Kars H,Eijkel L.2016.Soil in Criminal and Environmental Forensics.Switzerland:Springer International Publishing Switzerland.
Komarov A,Chertov O,Bykhovets S,et al.2017.Romul_Hum model of soil organic matter formation coupled with soil biota activity.I.Problem formulation,model description,and testing.Ecological Modelling,345:113-124.
Li C,Frolking S,Frolking TA.1992.A model of nitrous oxide evolution from soil driven by rainfall events:1.Model structure and sensitivity.Journal of Geophysical Research:Atmosphere,97:9759-9776.
Li X,Yin X,Wang Z,et al.2015.Litter mass loss and nutrient release influenced by soil fauna of Betula ermanii forest floor of the Changbai Mountains,China.Applied Soil Ecology,95:15-22.
Liao S,Ni X,Yang W,et al.2016.Water,rather than temperature,dominantly impacts how soil fauna affect dissolved carbon and nitrogen release from fresh litter during early litter decomposition.Forests,7:249-265.
Makkonen M,Berg MP,Handa IT,et al.2012.Highly consistent effects of plant litter identity and functional traits on decomposition across a latitudinal gradient.Ecology Letters,15:1033-1041.
Milcu A,Manning P.2011.All size classes of soil fauna and lit-ter quality control the acceleration of litter decay in its home environment.Oikos,120:1366-1370.
Parton W,Rasmussen P.1994.Long-term effects of crop management in wheat-fallow:Ⅱ.CENTURY model simulations.Soil Science Society of America Journal,58:530-536.
Porre RJ,van Groenigen JW,De Deyn GB,et al.2016.Exploring the relationship between soil mesofauna,soil structure and N2O emissions.Soil Biology&Biochemistry,96:55-64.
Schmidt P,Dickow K,Rocha AA,et al.2008.Soil macrofauna and decomposition rates in southern Brazilian Atlantic rainforests.Ecotropica,14:89-100.
Schon NL,Mackay AD,Gray RA,et al.2017.Influence of earthworm abundance and diversity on soil structure and the implications for soil services throughout the season.Pedobiologia,62:41-47.
Slade EM,Riutta T.2012.Interacting effects of leaf litter species and macrofauna on decomposition in different litter environments.Basic and Applied Ecology,13:423-431.
Soong JL,Nielsen UN.2016.The role of microarthropods in emerging models of soil organic matter.Soil Biology&Biochemistry,102:37-39.
Soong JL,Parton WJ,Calderon F,et al.2015.A new conceptual model on the fate and controls of fresh and pyrolized plant litter decomposition.Biogeochemistry,124:27-44.
Tan B,Wu F,Yang W,et al.2015.Soil fauna significantly contributes to litter decomposition at low temperatures inthe alpine/subalpine forests.Polish Journal of Ecology,63:377-386.
Wardle DA,Bardgett RD,Klironomos JN,et al.2004.Ecological linkages between aboveground and belowground biota.Science,304:1629-1633.
Xin WD,Yin XQ,Song B.2012.Contribution of soil fauna to litter decomposition in Songnen sandy lands in northeastern China.Journal of Arid Environments,77:90-95.
Yang X,Chen J.2009.Plant litter quality influences the contribution of soil fauna to litter decomposition in humid tropical forests,southwestern China.Soil Biology&Biochemistry,41:910-918.
Zhang W,Chao L,Yang Q,et al.2016.Litter quality mediated nitrogen effect on plant litter decomposition regardless of soil fauna presence.Ecology,97:2834-2843.
Zhang W,Yuan S,Hu N,et al.2015.Predicting soil fauna effect on plant litter decomposition by using boosted regression trees.Soil Biology&Biochemistry,82:81-86.
Zhao J,He X,Wang K.2015.A hypothetical model that explains differing net effects of inorganic fertilization on biomass and/or abundance of soil biota.Theoretical Ecology,8:505-512.
范换,王绍军,阮宏华,等.2014.苏北沿海不同土地利用类型土壤动物群落及其对凋落物分解的影响.南京林业大学学报:自然科学版,38(3):1-7.
黄玉梅,杨万勤,张健.2015.川西亚高山云杉叶凋落物质量损失过程及土壤生物的作用.长江流域资源与环境,24(4):676-683.
廖姝,杨万勤,彭艳,等.2016.土壤动物对四川盆地四种树种凋落叶在不同物候时期可溶性碳和氮动态的影响.生态学杂志,35(12):3172-3182.
刘瑞龙,李维民,杨万勤,等.2013.土壤动物对川西高山/亚高山森林凋落物分解的贡献.应用生态学报,24(12):3354-3360.
谭羽,杨万勤,廖姝,等.2017.川西高寒生态系统不同海拔土壤动物对冬季凋落叶腐殖化过程的影响.生态学报,37(5):1595-1602.
王利峰,和润莲,林杨,等.2016.川西高山林线交错带土壤动物对岷江冷杉和高山杜鹃凋落物分解的贡献.应用生态学报,27(11):3689-3697.
王文君,杨万勤,谭波,等.2013.四川盆地亚热带常绿阔叶林土壤动物对几种典型凋落物分解的影响.生态环境学报,22(9):1488-1495.
王文君,杨万勤,谭波,等.2015.四川盆地亚热带常绿阔叶林不同物候土壤动物对凋落物氮和磷释放的影响.林业科学,51(1):1-11.
王振海,殷秀琴,张成蒙.2016.土壤动物在长白山臭冷杉凋落物分解中的作用.林业科学,52(7):59-67.
夏磊,吴福忠,杨万勤,等.2012.川西亚高山森林凋落物分解初期土壤动物对红桦凋落叶质量损失的贡献.应用生态学报,23(2):301-306.
许湘琴,林植华,陈慧丽.2011.凋落物分解对土壤生物的影响.生态学杂志,30(6):1258-1264.
杨万勤,邓仁菊,张健.2007.森林凋落物分解及其对全球气候变化的响应.应用生态学报,18(12):2889-2895.
杨效东,邹晓明.2006.西双版纳热带季节雨林凋落叶分解与土壤动物群落:两种网孔分解袋的分解实验比较.植物生态学报,30(5):791-801.
殷秀琴,仲伟彦,王海霞,等.2002.小兴安岭森林落叶分解与土壤动物的作用.地理研究,21(6):689-699.
Allison SD,Wallenstein MD,Bradford MA.2010.Soil-carbon response to warming dependent on microbial physiology.Nature Geoscience,3:336-340.
Amosse J,Turberg P,Kohler-Milleret R,et al.2015.Effects of endogeic earthworms on the soil organic matter dynamics and the soil structure in urban and alluvial soil materials.Geoderma,243:50-57.
Anderson JM.1973.The breakdown and decomposition of sweet chestnut(Castanea sativa Mill.)and beech(Fagus sylvatica L.)leaf litter in two deciduous woodland soils.I.Breakdown,leaching and decomposition.Oecologia,12:251-274.
Aubert M,Margerie P,Trap J,et al.2010.Aboveground-belowground relationships in temperate forests:Plant litter composes and microbiota orchestrates.Forest Ecology and Management,259:563-572.
Barajas-Guzmn G,Alvarez-Snchez J.2003.The relationships between litter fauna and rates of litter decomposition in a tropical rain forest.Applied Soil Ecology,24:91-100.
Bonan GB.2008.Forests and climate change:Forcings,feedbacks,and the climate benefits of forests.Science,320:1444-1449.
Bonan GB,Hartman MD,Parton WJ,et al.2013.Evaluating litter decomposition in earth system models with long-term litterbag experiments:An example using the Community Land Model version 4(CLM4).Global Change Biology,19:957-974.
Bonanomi G,Capodilupo M,Incerti G,et al.2015.Litter quality and temperature modulate microbial diversity effects on decomposition in model experiments.Community Ecology,16:167-177.
Bottinelli N,Jouquet P,Capowiez Y,et al.2015.Why is the influence of soil macrofauna on soil structure only considered by soil ecologists?Soil&Tillage Research,146:118-124.
Campbell EE,Parton WJ,Soong JL,et al.2016.Using litter chemistry controls on microbial processes to partition litter carbon fluxes with the Litter Decomposition and Leaching(LIDEL)model.Soil Biology&Biochemistry,100:160-174.
Carcamo HA,Prescott CE,Chanway CP,et al.2001.Do soil fauna increase rates of litter breakdown and nitrogen release in forests of British Columbia,Canada?Canadian Journal of Forest Research,31:1195-1204.
Chertov O,Shaw C,Shashkov M,et al.2017.Romul_Hum model of soil organic matter formation coupled with soil biota activity.III.Parameterisation of earthworm activity.Ecological Modelling,345:140-149.
Coq S,Souquet JM,Meudec E,et al.2010.Interspecific variation in leaf litter tannins drives decomposition in a tropical rain forest of French Guiana.Ecology,91:2080-2091.
Coulis M,Hattenschwiler S,Coq S,et al.2016.Leaf litter consumption by macroarthropods and burial of their faeces enhance decomposition in a Mediterranean ecosystem.Ecosystems,19:1104-1115.
Davidson DA,Grieve IC.2006.Relationships between biodiversity and soil structure and function:Evidence from laboratory and field experiments.Applied Soil Ecology,33:176-185.
Diek9tter T,Wamser S,Wolters V.2010.Landscape and management ffects on structure and function of soil arthropod communities in winter wheat.Agriculture,Ecosystems and Environment,137:108-112.
Frouz J.2002.The effect of soil macrofauna on litter decomposition and soil organic matter accumulation during soil formation in spoil heaps after brown coal mining:A preliminary results.Ekologia Bratislava,21:363-369.
Frouz J.2018.Effects of soil macro-and mesofauna on litter decomposition and soil organic matter stabilization.Geoderma,332:161-172.
Frouz J,Roubickova A,Hedenec P,et al.2015.Do soil fauna really hasten litter decomposition?A meta-analysis of enclosure studies.European Journal of Soil Biology,68:18-24.
Fujii S,Makita N,Mori AS,et al.2016.Plant species control and soil faunal involvement in the processes of above-and below-ground litter decomposition.Oikos,125:883-892.
Garcia-Palacios P,Maestre FT,Kattge J,et al.2013.Climate and litter quality differently modulate the effects of soil fauna on litter decomposition across biomes.Ecology Letters,16:1045-1053.
Gonzalez G,Seastedt TR.2001.Soil fauna and plant litter decomposition in tropical and subalpine forests.Ecology,82:955-964.
Haettenschwiler S,Jorgensen HB.2010.Carbon quality rather than stoichiometry controls litter decomposition in a tropical rain forest.Journal of Ecology,98:754-763.
Handa IT,Aerts R,Berendse F,et al.2014.Consequences of biodiversity loss for litter decomposition across biomes.Nature,509:218-221.
Hou PCL,Zou XM,Huang CY,et al.2005.Plant litter decomposition influenced by soil animals and disturbance in a subtropical rainforest of Taiwan.Pedobiologia,49:539-547.
Illig J,Schatz H,Scheu S,et al.2008.Decomposition and colonization by micro-arthropods of two litter types in a tropical montane rain forest in southern Ecuador.Journal of Tropical Ecology,24:157-167.
Kars H,Eijkel L.2016.Soil in Criminal and Environmental Forensics.Switzerland:Springer International Publishing Switzerland.
Komarov A,Chertov O,Bykhovets S,et al.2017.Romul_Hum model of soil organic matter formation coupled with soil biota activity.I.Problem formulation,model description,and testing.Ecological Modelling,345:113-124.
Li C,Frolking S,Frolking TA.1992.A model of nitrous oxide evolution from soil driven by rainfall events:1.Model structure and sensitivity.Journal of Geophysical Research:Atmosphere,97:9759-9776.
Li X,Yin X,Wang Z,et al.2015.Litter mass loss and nutrient release influenced by soil fauna of Betula ermanii forest floor of the Changbai Mountains,China.Applied Soil Ecology,95:15-22.
Liao S,Ni X,Yang W,et al.2016.Water,rather than temperature,dominantly impacts how soil fauna affect dissolved carbon and nitrogen release from fresh litter during early litter decomposition.Forests,7:249-265.
Makkonen M,Berg MP,Handa IT,et al.2012.Highly consistent effects of plant litter identity and functional traits on decomposition across a latitudinal gradient.Ecology Letters,15:1033-1041.
Milcu A,Manning P.2011.All size classes of soil fauna and lit-ter quality control the acceleration of litter decay in its home environment.Oikos,120:1366-1370.
Parton W,Rasmussen P.1994.Long-term effects of crop management in wheat-fallow:Ⅱ.CENTURY model simulations.Soil Science Society of America Journal,58:530-536.
Porre RJ,van Groenigen JW,De Deyn GB,et al.2016.Exploring the relationship between soil mesofauna,soil structure and N2O emissions.Soil Biology&Biochemistry,96:55-64.
Schmidt P,Dickow K,Rocha AA,et al.2008.Soil macrofauna and decomposition rates in southern Brazilian Atlantic rainforests.Ecotropica,14:89-100.
Schon NL,Mackay AD,Gray RA,et al.2017.Influence of earthworm abundance and diversity on soil structure and the implications for soil services throughout the season.Pedobiologia,62:41-47.
Slade EM,Riutta T.2012.Interacting effects of leaf litter species and macrofauna on decomposition in different litter environments.Basic and Applied Ecology,13:423-431.
Soong JL,Nielsen UN.2016.The role of microarthropods in emerging models of soil organic matter.Soil Biology&Biochemistry,102:37-39.
Soong JL,Parton WJ,Calderon F,et al.2015.A new conceptual model on the fate and controls of fresh and pyrolized plant litter decomposition.Biogeochemistry,124:27-44.
Tan B,Wu F,Yang W,et al.2015.Soil fauna significantly contributes to litter decomposition at low temperatures inthe alpine/subalpine forests.Polish Journal of Ecology,63:377-386.
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