闽江河口湿地碳氮循环关键过程对氮输入的响应
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摘要
滨海湿地位于海陆相互作用地带，是响应全球变化和人类活动较为敏感的生态系统之一。滨海地区人口增长和工农业生产的迅速发展，使得越来越多的氮素营养通过大气干湿沉降、河流输入、地表径流、地下径流和污水排放等途径进入湿地，影响湿地碳氮生物地球化学过程。为了更深入的理解潮滩湿地碳氮循环过程对外源氮输入的响应特征，论文以闽江河口短叶茳芏和互花米草湿地为研究对象，通过野外原位观测、微区试验和实验室培养，研究了氮输入对湿地温室气体排放通量，湿地土壤碳氮化学转化过程、湿地植物生长与碳氮累积以及湿地植物残体分解及分解过程碳氮动态特征的影响，主要研究结论如下：
（1）闽江河口短叶茳芏和互花米草湿地在植物生长初期总体表现为CO_2的汇、CH_4的源和N_2O的弱源/弱汇，互花米草入侵增加了湿地CO_2吸收和CH_4释放，但对N_2O排放无显著影响。外源氮（N1，21g N m-2a-1；N_2，42g N m-2a-1）输入对湿地CO_2、CH_4和N_2O排放具有促进作用，N1和N_2处理使短叶茳芏湿地的CO_2、CH_4及N_2O排放通量分别增加了25.80%和29.72%、31.05%和123.50%及1604%和2706%，而互花米草湿地的CO_2、CH_4及N_2O排放通量则分别增加了30.38%和28.35%、63.88%和7.55%以及1384%和1443%。
（2）氮输入对湿地CO_2、CH_4和N_2O排放通量的影响具有明显的时间变异性，特别是N_2O排放通量，在施氮3小时后显著增加而8天后差异不显著，可见氮输入后的观测时间不同对研究结论影响较大，今后应注意加强该方面的研究。
（3）外源氮输入对两种湿地土壤电导率（EC）、pH和氧化还原电位（Eh）均未产生显著影响。两种湿地CO_2和CH_4的排放通量与气温、地温、EC、pH和Eh均存在一定程度的相关性，但氮输入影响下其相关性有所减弱，而不同处理下的N_2O排放通量与环境因子之间的相关性并不显著。
（4）在60％WHC和淹水条件下，不同浓度和形态的氮输入对短叶茳芏湿地土壤有机碳的累积矿化量多表现为抑制作用，而对互花米草湿地土壤的有机碳的累积矿化量则多表现为促进作用。水分条件是影响闽江河口潮汐湿地土壤有机碳矿化的重要因素，淹水条件下两种湿地土壤的有机碳矿化速率均显著低于好气条件。土壤累积矿化量（52d）、C0值、k值、C0k与土壤DOC含量和pH均呈极显著相关关系，外源氮输入引起的土壤DOC含量和pH的变化是导致土壤有机碳矿化差异的重要原因。
（5）不同处理下互花米草湿地土壤的CH_4产生潜力均显著高于短叶茳芏湿地。氮输入对两种湿地土壤的CH_4产生潜力具有显著影响，外源氮输入引起的土壤DOC和矿质氮含量以及EC的变化是导致CH_4产生潜力变化的重要原因。
（6）氮输入对两种湿地土壤CH_4氧化潜力的影响与外源氮的浓度和类型有关，且盐分和氮输入对湿地土壤CH_4氧化速率存在一定的交互影响，高盐输入对短叶茳芏和互花米草湿地土壤的CH_4氧化均具有抑制作用，高盐和高铵态氮输入对其抑制作用增强，而高盐和高硝态氮共同作用却对其产生了促进。
（7）硝化细菌反硝化作用是短叶茳芏和互花米草湿地土壤N_2O产生的关键过程，对湿地N_2O排放的贡献不容忽视。硝态氮输入主要通过促进反硝化作用和其他过程从而促进了两种湿地土壤的N_2O总生产。低浓度的铵态氮输入则通过促进反硝化作用、硝化细菌反硝化作用和其他过程从而增加了短叶茳芏湿地土壤N_2O的总产生量，而中氮和高氮输入则通过抑制硝化作用、硝化细菌反硝化作用以及其他过程从而降低了湿地土壤N_2O的总产生量。而对互花米草湿地土壤来说，三种铵态氮均通过促进硝化细菌反硝化作用和其他过程从而促进了其N_2O产生，但显著低于硝态氮的促进作用。
（8）不同形态的外源氮输入对短叶茳芏和互花米草各器官生物量、植物株高和基茎粗均具有一定程度的促进作用，且铵态氮的促进作用大于硝态氮。氮输入对两种植物不同器官的碳含量均没有显著影响，对氮含量则均具有促进作用。两种氮输入影响下短叶茳芏各器官的C/N均有所降低，且铵态氮输入影响下氮含量的增加和C/N的降低均大于硝态氮输入，但两种形态的氮对互花米草各器官C/N的影响却并不一致。
（9）氮输入对短叶茳芏、互花米草叶和叶鞘的相对分解速率均具有一定程度的促进，但对互花米草茎的分解速率影响不大。残体相对分解速率还受分解环境变化、残体基质质量、温度条件、土壤盐分和pH等的影响。四种残体在不同分解条件下均一直表现为碳的净释放，氮输入和环境条件变化对残体碳释放的影响呈波动变化，不同处理下残体碳绝对量的变化受物质残留量的影响程度明显。在各分解小区植物残体的氮在分解过程中多表现为不同程度的净释放，氮输入在分解初期抑制了短叶茳芏残体的氮净释放，而在分解后期则促进了其氮的净释放，但氮输入对互花米草植物残体多表现为抑制其氮净释放。
Coastal wetland, located in the interaction areas between land and sea, is a veryimportant ecosystem, which is sensitive to the global change and human activities.With the population growth and rapid development of industrial and agriculturalproduction in coastal zones, the estuarine wetlands often receive more and morenitrogen (N) nutrition originated from atmospheric deposition, river input, surface andunderground runoff and pollutant discharge, which may affect the carbon (C) and Nbiogeochemical process in the wetlands. In this paper, the native Cyperusmalaccensis and invaded Spartina alterniflora wetlands in the Min River estuarywere selected as study objects in order to understand the response of C and N cycle toexogenous N addition. The observations in situ, microcosm experiments andlaboratory control experiment were performed to study the effects of N addition ongreenhouse gas emission, transformation characteristics of C and N in wetland soils,the plant growth and C and N accumulation, the litter decomposition rules and C andN dynamics in decomposition process. The main results were drawn as follows:
(1) The C. malaccensis and S. alternifora wetlands acted as sinks of CO_2,sources of CH_4and weak sources or sinks of N_2O in the growing season. Theinvasion of S. alternifora into the Min River estuary stimulated CO_2uptake andCH_4emission, while no significant effects were found on N_2O emission. Exogenous N(N1,21g N m-2a-1; N_2,42g N m-2a-1) promoted CO_2, CH_4and N_2O emissions bothin native and in invaded tidal marsh. Compared to N0treatment, the mean gas fluxesof N1and N_2treatments increased by25.80%and29.72%for CO_2,31.05%and123.50%for CH_4, and1604%and2706%for N_2O in the C. malaccensis marsh.However, in the S. alternifora marsh, the mean gas fluxes of N1and N_2treatmentsincreased by30.38%and28.35%for CO_2,63.88%and7.55%for CH_4, and1384%and1443%for N_2O.
(2) Significant temporal variability of CO_2, CH_4and N_2O fuxes were observedafter the N was gradually added to the native and invaded marshes. Within3hours ofN addition, N_2O fuxes were signifcantly higher in plots receiving N additionsrelative to controls. After8days, when N concentration decreased, no significant differences were found between treatments. We conclude that the overall increase ofCO_2, CH_4and N_2O emissions affected by N addition in coastal regions may besignificantly underestimated due to the short-term temporal variations of gas fuxes. Inorder to better assess the global climatic role of salt marshes as affected by N addition,the short-term temporal variability of greenhouse gas emission should receive muchmore attention.
(3) EC, pH and Eh in different soil depths were not significantly affected by Naddition during the entire experiment. There were some significant correlationbetween CO_2(CH_4) fluxes and temperature (Ec, pH and Eh), while the correlationdecreased when N was added to the wetlands. The N_2O fluxes were not significantlycorrelated with temperature, Ec, pH and Eh.
(4) Under60%WHC and submerged conditions, the added N of differentconcentrations and forms restrained the mineralization of soil organic carbon (SOC)in the C. malaccensis marsh and promoted the SOC mineralization in the S.alternifora marsh. Water condition was an important factor affecting the SOCmineralization in the Min River estuarine wetlands, the SOC mineralization ratesunder submerged condition were significantly lower than those under60%WHCcondition. There were significant correlations between the amounts of cumulative C,C0, k or C0k and the DOC contents or pH, suggesting that the changes of DOC and pHaffected by N input might be two of the important reasons that led to the differencesof SOC mineralization.
(5) The CH_4production potential of different treatments in the S. alterniforamarsh were higher than those in the C. malaccensis marsh. N addition significantlyaffected the the CH_4production potential of the two marsh soils. The changes ofDOC and mineral N contents and EC in soils affected by N input might be theimportant reasons that led to the differences of CH_4production potential.
(6) The CH_4oxidation potential were affected by the concentrations and forms ofadded N. There were interaction effects of N and salinity addition. Salinity additiondecreased the CH_4oxidation rate both in C. malaccensis and in S. alterniforawetland soils. The interaction of salinity and NH_4+-N addition inhibited the CH_4oxidation, while the interaction of salinity and NO3--N addition increased the CH_4oxidation.
(7) The nitrifier denitrification is an important process of N_2O production inthe C. malaccensis and S. alternifora wetland soils, which can not be ignored when we study the N_2O production in wetland ecosystems. Nitrate N inputincreased the total N_2O production by promoting denitrification and other process.Low ammonium N addition increased the total N_2O production by promotingdenitrification, nitrifier denitrification and other process, however, the middle andhigh ammonium N addition decreased the total N_2O production by inhibitingnitrification, nitrifier denitrification and other proces in the C. malaccensiswetland soils. Three concentrations of ammonium N input increased the totalN_2O production by promoting nitrifier denitrification and other process in the S.alternifora wetland soils, but the increase were significantly lower than those ofnitrate N addition treatments.
(8) Different forms of exogenous N input promoted the biomass of different organs,plant height and stem diameter of C. malaccensis and S. alternifora, and thepromotions of ammonium N were greater than those of nitrate N. There were nosignificant difference between treatments in TC contents of different organs of plants,while the N addition increased the N content in different organs. The C/N of differentorgans of C. malaccensis decreased as affected by the N addition, and the decreaseas effected by the ammonium N input were greater than those of nitrate N. The effectsof two forms of added N on C/N of S. alternifora were not consistent.
(9) Nitrogen addition promoted the relative decomposition rate of C. malaccensislitters, leaf and leaf sheath litters of S. alternifora, while little effects were observedon the decomposition of stem litter of S. alternifora. In addition, relativedecomposition rate of litters were also affected by environment change, substratequality, temperature conditions, salinity and pH of soils. The C in litters in differentdecomposition sub-zones showed release at all times, the effects of N addition andchange of decompositon environment on C release in litters showed fluctuated change.The mass remaining rates significantly affected the C absolute contents. The N inlitters in different decomposition sub-zones showed release, the effects of N additionpromoted N release of C. malaccensis litter in the period of0~43d, while inhibited Nrelease in the periods of72~345d. However, N addition inhibited N release in S.alternifora litters.
（1）闽江河口短叶茳芏和互花米草湿地在植物生长初期总体表现为CO_2的汇、CH_4的源和N_2O的弱源/弱汇，互花米草入侵增加了湿地CO_2吸收和CH_4释放，但对N_2O排放无显著影响。外源氮（N1，21g N m-2a-1；N_2，42g N m-2a-1）输入对湿地CO_2、CH_4和N_2O排放具有促进作用，N1和N_2处理使短叶茳芏湿地的CO_2、CH_4及N_2O排放通量分别增加了25.80%和29.72%、31.05%和123.50%及1604%和2706%，而互花米草湿地的CO_2、CH_4及N_2O排放通量则分别增加了30.38%和28.35%、63.88%和7.55%以及1384%和1443%。
（2）氮输入对湿地CO_2、CH_4和N_2O排放通量的影响具有明显的时间变异性，特别是N_2O排放通量，在施氮3小时后显著增加而8天后差异不显著，可见氮输入后的观测时间不同对研究结论影响较大，今后应注意加强该方面的研究。
（3）外源氮输入对两种湿地土壤电导率（EC）、pH和氧化还原电位（Eh）均未产生显著影响。两种湿地CO_2和CH_4的排放通量与气温、地温、EC、pH和Eh均存在一定程度的相关性，但氮输入影响下其相关性有所减弱，而不同处理下的N_2O排放通量与环境因子之间的相关性并不显著。
（4）在60％WHC和淹水条件下，不同浓度和形态的氮输入对短叶茳芏湿地土壤有机碳的累积矿化量多表现为抑制作用，而对互花米草湿地土壤的有机碳的累积矿化量则多表现为促进作用。水分条件是影响闽江河口潮汐湿地土壤有机碳矿化的重要因素，淹水条件下两种湿地土壤的有机碳矿化速率均显著低于好气条件。土壤累积矿化量（52d）、C0值、k值、C0k与土壤DOC含量和pH均呈极显著相关关系，外源氮输入引起的土壤DOC含量和pH的变化是导致土壤有机碳矿化差异的重要原因。
（5）不同处理下互花米草湿地土壤的CH_4产生潜力均显著高于短叶茳芏湿地。氮输入对两种湿地土壤的CH_4产生潜力具有显著影响，外源氮输入引起的土壤DOC和矿质氮含量以及EC的变化是导致CH_4产生潜力变化的重要原因。
（6）氮输入对两种湿地土壤CH_4氧化潜力的影响与外源氮的浓度和类型有关，且盐分和氮输入对湿地土壤CH_4氧化速率存在一定的交互影响，高盐输入对短叶茳芏和互花米草湿地土壤的CH_4氧化均具有抑制作用，高盐和高铵态氮输入对其抑制作用增强，而高盐和高硝态氮共同作用却对其产生了促进。
（7）硝化细菌反硝化作用是短叶茳芏和互花米草湿地土壤N_2O产生的关键过程，对湿地N_2O排放的贡献不容忽视。硝态氮输入主要通过促进反硝化作用和其他过程从而促进了两种湿地土壤的N_2O总生产。低浓度的铵态氮输入则通过促进反硝化作用、硝化细菌反硝化作用和其他过程从而增加了短叶茳芏湿地土壤N_2O的总产生量，而中氮和高氮输入则通过抑制硝化作用、硝化细菌反硝化作用以及其他过程从而降低了湿地土壤N_2O的总产生量。而对互花米草湿地土壤来说，三种铵态氮均通过促进硝化细菌反硝化作用和其他过程从而促进了其N_2O产生，但显著低于硝态氮的促进作用。
（8）不同形态的外源氮输入对短叶茳芏和互花米草各器官生物量、植物株高和基茎粗均具有一定程度的促进作用，且铵态氮的促进作用大于硝态氮。氮输入对两种植物不同器官的碳含量均没有显著影响，对氮含量则均具有促进作用。两种氮输入影响下短叶茳芏各器官的C/N均有所降低，且铵态氮输入影响下氮含量的增加和C/N的降低均大于硝态氮输入，但两种形态的氮对互花米草各器官C/N的影响却并不一致。
（9）氮输入对短叶茳芏、互花米草叶和叶鞘的相对分解速率均具有一定程度的促进，但对互花米草茎的分解速率影响不大。残体相对分解速率还受分解环境变化、残体基质质量、温度条件、土壤盐分和pH等的影响。四种残体在不同分解条件下均一直表现为碳的净释放，氮输入和环境条件变化对残体碳释放的影响呈波动变化，不同处理下残体碳绝对量的变化受物质残留量的影响程度明显。在各分解小区植物残体的氮在分解过程中多表现为不同程度的净释放，氮输入在分解初期抑制了短叶茳芏残体的氮净释放，而在分解后期则促进了其氮的净释放，但氮输入对互花米草植物残体多表现为抑制其氮净释放。
Coastal wetland, located in the interaction areas between land and sea, is a veryimportant ecosystem, which is sensitive to the global change and human activities.With the population growth and rapid development of industrial and agriculturalproduction in coastal zones, the estuarine wetlands often receive more and morenitrogen (N) nutrition originated from atmospheric deposition, river input, surface andunderground runoff and pollutant discharge, which may affect the carbon (C) and Nbiogeochemical process in the wetlands. In this paper, the native Cyperusmalaccensis and invaded Spartina alterniflora wetlands in the Min River estuarywere selected as study objects in order to understand the response of C and N cycle toexogenous N addition. The observations in situ, microcosm experiments andlaboratory control experiment were performed to study the effects of N addition ongreenhouse gas emission, transformation characteristics of C and N in wetland soils,the plant growth and C and N accumulation, the litter decomposition rules and C andN dynamics in decomposition process. The main results were drawn as follows:
(1) The C. malaccensis and S. alternifora wetlands acted as sinks of CO_2,sources of CH_4and weak sources or sinks of N_2O in the growing season. Theinvasion of S. alternifora into the Min River estuary stimulated CO_2uptake andCH_4emission, while no significant effects were found on N_2O emission. Exogenous N(N1,21g N m-2a-1; N_2,42g N m-2a-1) promoted CO_2, CH_4and N_2O emissions bothin native and in invaded tidal marsh. Compared to N0treatment, the mean gas fluxesof N1and N_2treatments increased by25.80%and29.72%for CO_2,31.05%and123.50%for CH_4, and1604%and2706%for N_2O in the C. malaccensis marsh.However, in the S. alternifora marsh, the mean gas fluxes of N1and N_2treatmentsincreased by30.38%and28.35%for CO_2,63.88%and7.55%for CH_4, and1384%and1443%for N_2O.
(2) Significant temporal variability of CO_2, CH_4and N_2O fuxes were observedafter the N was gradually added to the native and invaded marshes. Within3hours ofN addition, N_2O fuxes were signifcantly higher in plots receiving N additionsrelative to controls. After8days, when N concentration decreased, no significant differences were found between treatments. We conclude that the overall increase ofCO_2, CH_4and N_2O emissions affected by N addition in coastal regions may besignificantly underestimated due to the short-term temporal variations of gas fuxes. Inorder to better assess the global climatic role of salt marshes as affected by N addition,the short-term temporal variability of greenhouse gas emission should receive muchmore attention.
(3) EC, pH and Eh in different soil depths were not significantly affected by Naddition during the entire experiment. There were some significant correlationbetween CO_2(CH_4) fluxes and temperature (Ec, pH and Eh), while the correlationdecreased when N was added to the wetlands. The N_2O fluxes were not significantlycorrelated with temperature, Ec, pH and Eh.
(4) Under60%WHC and submerged conditions, the added N of differentconcentrations and forms restrained the mineralization of soil organic carbon (SOC)in the C. malaccensis marsh and promoted the SOC mineralization in the S.alternifora marsh. Water condition was an important factor affecting the SOCmineralization in the Min River estuarine wetlands, the SOC mineralization ratesunder submerged condition were significantly lower than those under60%WHCcondition. There were significant correlations between the amounts of cumulative C,C0, k or C0k and the DOC contents or pH, suggesting that the changes of DOC and pHaffected by N input might be two of the important reasons that led to the differencesof SOC mineralization.
(5) The CH_4production potential of different treatments in the S. alterniforamarsh were higher than those in the C. malaccensis marsh. N addition significantlyaffected the the CH_4production potential of the two marsh soils. The changes ofDOC and mineral N contents and EC in soils affected by N input might be theimportant reasons that led to the differences of CH_4production potential.
(6) The CH_4oxidation potential were affected by the concentrations and forms ofadded N. There were interaction effects of N and salinity addition. Salinity additiondecreased the CH_4oxidation rate both in C. malaccensis and in S. alterniforawetland soils. The interaction of salinity and NH_4+-N addition inhibited the CH_4oxidation, while the interaction of salinity and NO3--N addition increased the CH_4oxidation.
(7) The nitrifier denitrification is an important process of N_2O production inthe C. malaccensis and S. alternifora wetland soils, which can not be ignored when we study the N_2O production in wetland ecosystems. Nitrate N inputincreased the total N_2O production by promoting denitrification and other process.Low ammonium N addition increased the total N_2O production by promotingdenitrification, nitrifier denitrification and other process, however, the middle andhigh ammonium N addition decreased the total N_2O production by inhibitingnitrification, nitrifier denitrification and other proces in the C. malaccensiswetland soils. Three concentrations of ammonium N input increased the totalN_2O production by promoting nitrifier denitrification and other process in the S.alternifora wetland soils, but the increase were significantly lower than those ofnitrate N addition treatments.
(8) Different forms of exogenous N input promoted the biomass of different organs,plant height and stem diameter of C. malaccensis and S. alternifora, and thepromotions of ammonium N were greater than those of nitrate N. There were nosignificant difference between treatments in TC contents of different organs of plants,while the N addition increased the N content in different organs. The C/N of differentorgans of C. malaccensis decreased as affected by the N addition, and the decreaseas effected by the ammonium N input were greater than those of nitrate N. The effectsof two forms of added N on C/N of S. alternifora were not consistent.
(9) Nitrogen addition promoted the relative decomposition rate of C. malaccensislitters, leaf and leaf sheath litters of S. alternifora, while little effects were observedon the decomposition of stem litter of S. alternifora. In addition, relativedecomposition rate of litters were also affected by environment change, substratequality, temperature conditions, salinity and pH of soils. The C in litters in differentdecomposition sub-zones showed release at all times, the effects of N addition andchange of decompositon environment on C release in litters showed fluctuated change.The mass remaining rates significantly affected the C absolute contents. The N inlitters in different decomposition sub-zones showed release, the effects of N additionpromoted N release of C. malaccensis litter in the period of0~43d, while inhibited Nrelease in the periods of72~345d. However, N addition inhibited N release in S.alternifora litters.
引文
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