土地利用方式对中国东南部红壤微生物特性及氮转化作用的影响
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摘要
我国东南部的热带和亚热带红壤总面积113.3万km2,占全国总面积的11.8%,供给着全国22.5%的人口。红壤具有低pH、低养分含量、氮相对富集、硝化与反硝化活性低、N20排放量却较大的特点,红壤的分布具有明显的地域特征,而土地利用方式的变化可能引起对该地区的土壤肥力以及土壤功能变化,甚至衰退。因此,研究该地区农业、林地等不同利用方式对红壤微生物特性及氮转化作用的影响对维持我国红壤的可持续发展具有重要的科学意义。
本文以我国东南部热带和亚热带地区典型利用方式的红壤为研究对象,选取了海南琼中香蕉林(BAN)、桉树林(EUC)、橡胶林(RUB)和天然次生林(NSF)4种利用方式土壤,福建建瓯万木林自然保护区内观光木(TSO)、细柄阿丁枫(ALG)、椤浮栲(CAF)、浙江桂(CIC)4种天然林和杉木(CUL)、桔园(ORG)2种人工种植林土壤,以及江西鹰潭市周边地区花岗岩(G)、第三纪红砂岩(R)和第四纪红粘土(Q)3种母质和阔叶林(B)、针叶林(C)、灌丛(S)、农用地(U/F)4种利用方式红壤。本文测定了土壤微生物生物量碳、氮,土壤酶活性；硝化潜势(NP)、硝化回复势(RNP)；用淹水厌氧密闭培养-乙炔抑制法测定了反硝化势(DNP)及其气态产物；利用实时荧光定量PCR (Real-time PCR, qPCR)检测了氨氧化细菌(AOB)和氨氧化古菌(AOA)的amoA基因拷贝数,反硝化功能基因即硝酸盐还原酶基因(narG)、亚硝酸还原酶基因(nirK)、一氧化氮还原酶基因(norB)和氧化亚氮还原酶基因(nosZ)的拷贝数,以及细菌16SrRNA基因拷贝数；采用PCR-DGGE技术研究了土壤中细菌的群落结构差异；采用磷脂脂肪酸(PLFA)方法分析土壤微生物群落多样性。
土壤微生物生物量以及土壤酶活性在不同利用方式土壤中有显著差异,海南、福建和江西三个地区农用地中微生物生物量碳和氮、土壤脲酶和脱氢酶活性均低于天然林等其他利用方式土壤,相关性分析显示不同利用方式对土壤营养因子(土壤有机碳、全氮、水解氮、速效钾等)的改变是引起土壤微生物生物量和土壤酶活性的关键因素。对福建不同土地利用方式对微生物群落多样性的影响研究发现,天然林(CIC,CAF, ALG, TSO)的年落叶量、土壤养分含量、微生物群落多样性指标显著高于两种种植园土壤(CUL, Orchard),不同土壤利用方式对土壤微生物群落多样性有显著影响；天然林改造成为农用经济林地后减少了土壤微生物多样性；年落叶量、土壤有机碳和总氮等是引起天然林和种植园土壤微生物多样性差异的关键因素。
对于硝化微生物和硝化作用的研究发现,江西天然林的硝化势(NP)显著低于福建与海南地区土壤的NP,所有调查地区农用利用方式土壤的NP较天然林大,说明人为管理方式提高了土壤硝化活性；AOA和AOB对于红壤硝化作用的相对贡献率在不同地区不同和并受不同利用方式的影响：qPCR结果与NP的相关性分析显示福建和海南地区NP可能主要由AOB承担,而江西鹰潭地区NP可能主要由AOA承担；土壤RNPs对于抗生素的敏感性实验显示,海南地区BAN、EUC、RUB,福建地区CIC、CUL、ORG六种土壤的NP可能主要由AOB承担,而海南的NSF、福建的ALG、江西的RB、RC、RS和RU土壤NP主要由AOA承担。
对于反硝化微生物、反硝化作用及其气态产物的研究表明,不同母质和不同利用方式土壤反硝化势和气态产物中氮氧化物的比例差异较大：不同土壤的反硝化势顺序为：GF(0.120mg·kg-1·h-1)>GS (0.056mg·kg-1·h-1)>GC (0.052mg·kg-1·h-1)>QF (0.018mg·kg-1·h-1)>QC (0.010mg·kg-1·h-1)>QS (0.009mg·kg-1·h-1);红粘土的NO占反硝化气体比例较花岗岩大,而N20是针叶林与灌丛土壤反硝化的主要产物；而农用地土壤反硝化的主要产物是N2；农业利用方式的反硝化作用高于针叶林与灌丛,pH是影响亚热带红壤的主要影响因子；在四种编码反硝化酶的基因narG、nirK、norB和nosZ中,norB基因是引起不同土地利用方式土壤之间反硝化势和反硝化气态产物组成差异的关键因子。
Tropical and subtropical red soil in southeast China cover about1.13million km2or11.8%of the country's land surface, and support22.5%of the nation's population. Red soil is characterized with low pH, poor nutrition, higher nitrogen, low nitrification, low denitrification and abundant N2O emission. The distribution of red soil has obvious regional characteristic. The land use change, such as the conversion of the natural forests to agricultural purpose, might affect soil fertility and soil function. Therefore, it will be instructive to progress of the effect of land-use types on microbial properties and nitrogen transformation for sustainable development of red soil in China.
The representative land-use types of tropical and subtropical red soils in southeastern China were investigated in this paper:the first series including a natural secondary forest (NSF), a banana plantation (BAN), a eucalyptus forest (EUC), and a rubber plantation (RUB) in Qiongzhong, Hainan province; the sencond series including four natural forests, Altingia gralilipes (ALG), Cinnamomum chekiangense (CIC), Castanopsis fargesii (CAF), Tsoongiodendron odorum (TSO), and two plantations, Cunninghamia lanceolata (CUL) and Citrus reticulata orchard (ORG) in Jian'ou, Fujian province; the third series including three soil parent materials:granite (G), tertiary red sandstone (R), and quaternary red earth (Q) and four land use patterns:broad-leaf forest (B), conifer forest (C), shrub land (S) and upland/farmland (U/F) in Yingtan, Jiangxi province. In this study, soil nitrification potential (NP) and recovery of nitrification potential (RNP) were measured; soil denitrification and its gas products (N2O, NO, N2) were studied by acetylene inhibition method applied to laboratory incubation under flooded condition; real-time quantitative polymerase chain reaction (qPCR) was conducted to investigate amoA gene copies of soil ammonia-oxidizing bacteria (AOB) and ammonia-oxidizing archaea (AOA), narG, nirK, norB, nosZ gene copies of denitrifying bacteria and soil16S rRNA bacterial gene copies; denaturing gradient gel electrophoresis (DGGE) was conducted to investigate soil16S rRNA bacterial diversity; phospholipid fatty acid analysis (PLFA) was conducted to investigate soil microbial diversity.
Soil microbial biomass and soil enzyme activities were significantly differed in different land-use soils. The agricultural land-use had lower soil microbial biomass carbon and nitrogen and soil urease and dehydrogenase activities than other land-use soils. These significantle difference might be driven by changes of soil nutrition factors (soil organic carbon, total N. available N, available K et al.) according to correlation analysis. Land use types had great effects on soil microbial community structure in the mid-subtropic region of China. Soil microbial diversity were highest under four natural forests (CIC, CAF, ALG, TSO), followed by CUL platation, whilst least in orchard soil. The conversion from native forests to agricultural purpose could decrease soil microbial biomass, activity and diversity in mid-subtropical region of China. Litterfall, soil oganic C and soil total N distinguished the microbial community of forests from that of economic plantation in the mid-subtropical China.
Our result showed that soil nitrification potential (NP) of natural forest was significantly lower in Jiangxi than Fujian and Hainan; NP of framland use soils was higher than natural forest soils in all soil series. It indicated that deliberate management might increase soil denitrification activity. Contribution rate to nitrification of AOB and AOA was differed from different land-use types. Relative analysis between NP and amoA gene copies of AOB and AOA from qPCR analysis indicated that soil nitrification potential could be mediated by AOB in Fujian and Hainan soil samples:nitrification potential might be mediated by AOA in Jiangxi. Further study of sensitivity of RNPs to protein synthesis inhibitors suggested that soil NP of BAN, EUC, RUB in Hainan and CIC, CUL,ORG in Fujian could be attributed exclusively to AOB; in contrast, those nitrification potential NSF in Hainan, ALG in Fujian, RB, RC, RS, RU in Jiangxi were mediated by AOA.
Soil denitrification potential and composition of denitrification products varied greatly among the investigated soils with a following order of denitrification potential:GF (0.120 mg·kg-1·h-1)> GS (0.056mg·kg-1·h-1)> GC (0.052mg·kg-1·h-1)> QF (0.018mg·kg-1·h-1)> QC (0.010mg·kg-1·h-1)> QS (0.009mg·kg-1·h-1). NO/(N2O+NO+N2) was higher in quaternary red earth than granite soil. The dominant denitrification gas product was N2O in conifer forest (C) and shrub land (S) soils and N2in farmland (F) soil. Soil denitrification potential was higher in farmland soil than in conifer forest and shrub land soils.Soil pH was the important influence factor of the denitrification. Among the four genes:narG, nirK, norB and nosZ, the norB gene was the key factor of the difference of denitrification in different land-use soils.
本文以我国东南部热带和亚热带地区典型利用方式的红壤为研究对象,选取了海南琼中香蕉林(BAN)、桉树林(EUC)、橡胶林(RUB)和天然次生林(NSF)4种利用方式土壤,福建建瓯万木林自然保护区内观光木(TSO)、细柄阿丁枫(ALG)、椤浮栲(CAF)、浙江桂(CIC)4种天然林和杉木(CUL)、桔园(ORG)2种人工种植林土壤,以及江西鹰潭市周边地区花岗岩(G)、第三纪红砂岩(R)和第四纪红粘土(Q)3种母质和阔叶林(B)、针叶林(C)、灌丛(S)、农用地(U/F)4种利用方式红壤。本文测定了土壤微生物生物量碳、氮,土壤酶活性；硝化潜势(NP)、硝化回复势(RNP)；用淹水厌氧密闭培养-乙炔抑制法测定了反硝化势(DNP)及其气态产物；利用实时荧光定量PCR (Real-time PCR, qPCR)检测了氨氧化细菌(AOB)和氨氧化古菌(AOA)的amoA基因拷贝数,反硝化功能基因即硝酸盐还原酶基因(narG)、亚硝酸还原酶基因(nirK)、一氧化氮还原酶基因(norB)和氧化亚氮还原酶基因(nosZ)的拷贝数,以及细菌16SrRNA基因拷贝数；采用PCR-DGGE技术研究了土壤中细菌的群落结构差异；采用磷脂脂肪酸(PLFA)方法分析土壤微生物群落多样性。
土壤微生物生物量以及土壤酶活性在不同利用方式土壤中有显著差异,海南、福建和江西三个地区农用地中微生物生物量碳和氮、土壤脲酶和脱氢酶活性均低于天然林等其他利用方式土壤,相关性分析显示不同利用方式对土壤营养因子(土壤有机碳、全氮、水解氮、速效钾等)的改变是引起土壤微生物生物量和土壤酶活性的关键因素。对福建不同土地利用方式对微生物群落多样性的影响研究发现,天然林(CIC,CAF, ALG, TSO)的年落叶量、土壤养分含量、微生物群落多样性指标显著高于两种种植园土壤(CUL, Orchard),不同土壤利用方式对土壤微生物群落多样性有显著影响；天然林改造成为农用经济林地后减少了土壤微生物多样性；年落叶量、土壤有机碳和总氮等是引起天然林和种植园土壤微生物多样性差异的关键因素。
对于硝化微生物和硝化作用的研究发现,江西天然林的硝化势(NP)显著低于福建与海南地区土壤的NP,所有调查地区农用利用方式土壤的NP较天然林大,说明人为管理方式提高了土壤硝化活性；AOA和AOB对于红壤硝化作用的相对贡献率在不同地区不同和并受不同利用方式的影响：qPCR结果与NP的相关性分析显示福建和海南地区NP可能主要由AOB承担,而江西鹰潭地区NP可能主要由AOA承担；土壤RNPs对于抗生素的敏感性实验显示,海南地区BAN、EUC、RUB,福建地区CIC、CUL、ORG六种土壤的NP可能主要由AOB承担,而海南的NSF、福建的ALG、江西的RB、RC、RS和RU土壤NP主要由AOA承担。
对于反硝化微生物、反硝化作用及其气态产物的研究表明,不同母质和不同利用方式土壤反硝化势和气态产物中氮氧化物的比例差异较大：不同土壤的反硝化势顺序为：GF(0.120mg·kg-1·h-1)>GS (0.056mg·kg-1·h-1)>GC (0.052mg·kg-1·h-1)>QF (0.018mg·kg-1·h-1)>QC (0.010mg·kg-1·h-1)>QS (0.009mg·kg-1·h-1);红粘土的NO占反硝化气体比例较花岗岩大,而N20是针叶林与灌丛土壤反硝化的主要产物；而农用地土壤反硝化的主要产物是N2；农业利用方式的反硝化作用高于针叶林与灌丛,pH是影响亚热带红壤的主要影响因子；在四种编码反硝化酶的基因narG、nirK、norB和nosZ中,norB基因是引起不同土地利用方式土壤之间反硝化势和反硝化气态产物组成差异的关键因子。
Tropical and subtropical red soil in southeast China cover about1.13million km2or11.8%of the country's land surface, and support22.5%of the nation's population. Red soil is characterized with low pH, poor nutrition, higher nitrogen, low nitrification, low denitrification and abundant N2O emission. The distribution of red soil has obvious regional characteristic. The land use change, such as the conversion of the natural forests to agricultural purpose, might affect soil fertility and soil function. Therefore, it will be instructive to progress of the effect of land-use types on microbial properties and nitrogen transformation for sustainable development of red soil in China.
The representative land-use types of tropical and subtropical red soils in southeastern China were investigated in this paper:the first series including a natural secondary forest (NSF), a banana plantation (BAN), a eucalyptus forest (EUC), and a rubber plantation (RUB) in Qiongzhong, Hainan province; the sencond series including four natural forests, Altingia gralilipes (ALG), Cinnamomum chekiangense (CIC), Castanopsis fargesii (CAF), Tsoongiodendron odorum (TSO), and two plantations, Cunninghamia lanceolata (CUL) and Citrus reticulata orchard (ORG) in Jian'ou, Fujian province; the third series including three soil parent materials:granite (G), tertiary red sandstone (R), and quaternary red earth (Q) and four land use patterns:broad-leaf forest (B), conifer forest (C), shrub land (S) and upland/farmland (U/F) in Yingtan, Jiangxi province. In this study, soil nitrification potential (NP) and recovery of nitrification potential (RNP) were measured; soil denitrification and its gas products (N2O, NO, N2) were studied by acetylene inhibition method applied to laboratory incubation under flooded condition; real-time quantitative polymerase chain reaction (qPCR) was conducted to investigate amoA gene copies of soil ammonia-oxidizing bacteria (AOB) and ammonia-oxidizing archaea (AOA), narG, nirK, norB, nosZ gene copies of denitrifying bacteria and soil16S rRNA bacterial gene copies; denaturing gradient gel electrophoresis (DGGE) was conducted to investigate soil16S rRNA bacterial diversity; phospholipid fatty acid analysis (PLFA) was conducted to investigate soil microbial diversity.
Soil microbial biomass and soil enzyme activities were significantly differed in different land-use soils. The agricultural land-use had lower soil microbial biomass carbon and nitrogen and soil urease and dehydrogenase activities than other land-use soils. These significantle difference might be driven by changes of soil nutrition factors (soil organic carbon, total N. available N, available K et al.) according to correlation analysis. Land use types had great effects on soil microbial community structure in the mid-subtropic region of China. Soil microbial diversity were highest under four natural forests (CIC, CAF, ALG, TSO), followed by CUL platation, whilst least in orchard soil. The conversion from native forests to agricultural purpose could decrease soil microbial biomass, activity and diversity in mid-subtropical region of China. Litterfall, soil oganic C and soil total N distinguished the microbial community of forests from that of economic plantation in the mid-subtropical China.
Our result showed that soil nitrification potential (NP) of natural forest was significantly lower in Jiangxi than Fujian and Hainan; NP of framland use soils was higher than natural forest soils in all soil series. It indicated that deliberate management might increase soil denitrification activity. Contribution rate to nitrification of AOB and AOA was differed from different land-use types. Relative analysis between NP and amoA gene copies of AOB and AOA from qPCR analysis indicated that soil nitrification potential could be mediated by AOB in Fujian and Hainan soil samples:nitrification potential might be mediated by AOA in Jiangxi. Further study of sensitivity of RNPs to protein synthesis inhibitors suggested that soil NP of BAN, EUC, RUB in Hainan and CIC, CUL,ORG in Fujian could be attributed exclusively to AOB; in contrast, those nitrification potential NSF in Hainan, ALG in Fujian, RB, RC, RS, RU in Jiangxi were mediated by AOA.
Soil denitrification potential and composition of denitrification products varied greatly among the investigated soils with a following order of denitrification potential:GF (0.120 mg·kg-1·h-1)> GS (0.056mg·kg-1·h-1)> GC (0.052mg·kg-1·h-1)> QF (0.018mg·kg-1·h-1)> QC (0.010mg·kg-1·h-1)> QS (0.009mg·kg-1·h-1). NO/(N2O+NO+N2) was higher in quaternary red earth than granite soil. The dominant denitrification gas product was N2O in conifer forest (C) and shrub land (S) soils and N2in farmland (F) soil. Soil denitrification potential was higher in farmland soil than in conifer forest and shrub land soils.Soil pH was the important influence factor of the denitrification. Among the four genes:narG, nirK, norB and nosZ, the norB gene was the key factor of the difference of denitrification in different land-use soils.
引文
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