添加磷素对低磷稻田根际土壤固碳自养微生物数量的影响
|
摘要
为了探讨磷素添加对低磷稻田土壤固碳自养微生物数量的影响,以低磷水稻土为研究对象,设置添加磷素(P添加量为80 mg·kg~(-1))和不添加磷素两种处理土壤,种植水稻进行室内培养实验.利用实时荧光定量PCR(real-time PCR)技术分析了水稻分蘖期(移栽后14 d)和拔节期(移栽后22 d)添加磷素(P)和对照(CK)处理根际土壤固碳自养微生物cbb L、cbb M、acc A和acl B基因数量的差异,同时测定了土壤理化性质,并分析了不同处理方式下固碳功能基因丰度与土壤理化性质之间的关系.结果表明,分蘖期磷素添加降低了土壤MBC和NH+4-N含量,提高了土壤DOC、Olsen-P和p H;分蘖期P处理NO-3-N含量比CK处理低,而拔节期反而比CK处理高.分蘖期,磷素添加显著提高了cbb L、cbb M、acc A和acl B基因的数量,与CK处理相比,分别提高156%、99%、110%和193%.而磷素添加对拔节期cbb L、acc A和acl B基因数量的促进作用并不明显,对cbb M基因数量反而产生了抑制作用.冗余分析(RDA)显示,土壤Olsen-P含量是影响固碳自养微生物丰度最显著的环境因子.
A rice pot experiment was conducted to investigate the effect of phosphorus addition on the abundance of autotrophic CO_2-fixation microorganisms using phosphorus-limited paddy soil from the Changsha Observation and Research Station for the Agricultural Environment. Rice seedlings were transplanted in the paddy soil with or without phosphorus addition,corresponding to P-treated-pot(P) or control pot(CK),respectively. Rhizosphere soils were collected from the P and CK treatments during the tillering and shooting stages. The physical and chemical soil properties were measured and the abundance of autotrophic CO_2-fixation microorganisms was quantified with a real-time PCR technique based on four functional genes(cbb L,cbb M,acc A,and acl B) involved in three CO_2-fixation pathways(CBB cycle,r TCA cycle,and 3-hydroxypropionate/4-hydroxybutyrate cycle). The results show that phosphorus addition improves the concentrations of DOC and Olsen-P and the p H value,whereas negative effects on the MBC and NH+4-N concentrations are revealed during the tillering stage. The effect of phosphorus addition on the NO-3-N concentration in the tillering and shooting stages differs. Phosphorus addition significantly increases the abundances of the cbb L,cbb M,acc A,and acl B genes,which are 156%,99%,110%,and 193% higher than those of the CK treatment in the tillering stage. However,this positive effect is not notable for the cbb L,acc A,and acl B genes during the shooting stage. Redundancy analysis(RDA) shows that Olsen-P is the environmental factor that most significantly affects the abundance of autotrophic CO_2-fixation microorganisms.
A rice pot experiment was conducted to investigate the effect of phosphorus addition on the abundance of autotrophic CO_2-fixation microorganisms using phosphorus-limited paddy soil from the Changsha Observation and Research Station for the Agricultural Environment. Rice seedlings were transplanted in the paddy soil with or without phosphorus addition,corresponding to P-treated-pot(P) or control pot(CK),respectively. Rhizosphere soils were collected from the P and CK treatments during the tillering and shooting stages. The physical and chemical soil properties were measured and the abundance of autotrophic CO_2-fixation microorganisms was quantified with a real-time PCR technique based on four functional genes(cbb L,cbb M,acc A,and acl B) involved in three CO_2-fixation pathways(CBB cycle,r TCA cycle,and 3-hydroxypropionate/4-hydroxybutyrate cycle). The results show that phosphorus addition improves the concentrations of DOC and Olsen-P and the p H value,whereas negative effects on the MBC and NH+4-N concentrations are revealed during the tillering stage. The effect of phosphorus addition on the NO-3-N concentration in the tillering and shooting stages differs. Phosphorus addition significantly increases the abundances of the cbb L,cbb M,acc A,and acl B genes,which are 156%,99%,110%,and 193% higher than those of the CK treatment in the tillering stage. However,this positive effect is not notable for the cbb L,acc A,and acl B genes during the shooting stage. Redundancy analysis(RDA) shows that Olsen-P is the environmental factor that most significantly affects the abundance of autotrophic CO_2-fixation microorganisms.
引文
[1]郝文英,曹正邦.红壤中的微生物[A].见:中国红壤[M].北京:科学出版社,1983. 128-144.
[2] Vance C P,Uhde-Stone C,Allan D L. Phosphorus acquisition and use:critical adaptations by plants for securing a nonrenewable resource[J]. New Phytologist,2003,157(3):423-447.
[3]赵小蓉,周然,李贵桐,等.低磷石灰性土壤施磷和小麦秸秆后土壤微生物量磷的变化[J].应用生态学报,2009,20(2):325-330.Zhao X R,Zhou R,Li G T,et al. Effects of applying inorganic P and wheat straw on the microbial biomass P and microbial P concentration in a calcareous soil with low concentration available P[J]. Chinese Journal of Applied Ecology,2009,20(2):325-330.
[4]王超.不同供磷量对玉米根系与土壤微生物互作及吐丝后地上部碳磷分配的影响[D].北京:中国农业大学,2017.Wang C. Influences of phosphorus supply on interactions between roots and soil microbes and post-silking partitioning of shoot carbon/phosphorus in field-grown maize[D]. Beijing:China Agricultural University,2017.
[5]刘俊杰,王光华,金剑,等.磷浓度处理对大豆根际土壤微生物群落结构的影响[J].大豆科学,2008,27(5):801-805.Liu J J,Wang G H,Jin J,et al. Effect of different phosphorus concentrations on microbial communities in soybean rhizosphere[J]. Soybean Science,2008,27(5):801-805.
[6]施瑶,王忠强,张心昱,等.氮磷添加对内蒙古温带典型草原土壤微生物群落结构的影响[J].生态学报,2014,34(17):4943-4949.Shi Y,Wang Z Q,Zhang X Y,et al. Effects of nitrogen and phosphorus addition on soil microbial community composition in temperate typical grassland in Inner Mongolia[J]. Acta Ecologica Sinica,2014,34(17):4943-4949.
[7]张四海,黄健,骆争荣,等.添加秸秆和磷素对土壤微生物群落的影响[J].应用生态学报,2014,25(3):797-802.Zhang S H,Huang J,Luo Z R,et al. Effect of adding different amounts of wheat straw and phosphorus on soil microorganism community[J]. Chinese Journal of Applied Ecology,2014,25(3):797-802.
[8] Huang J S,Hu B,Qi K B,et al. Effects of phosphorus addition on soil microbial biomass and community composition in a subalpine spruce plantation[J]. European Journal of Soil Biology,2016,72:35-41.
[9]林先贵,冯有智,陈瑞蕊.农田潮土养分耦合循环的微生物学机理研究进展[J].植物营养与肥料学报,2017,23(6):1575-1589.Lin X G,Feng Y Z,Chen R R. Research progresses of soil microorganisms driven nutrient coupling cycles in fluvo-aquic soils of China[J]. Journal of Plant Nutrition and Fertilizer,2017,23(6):1575-1589.
[10] Yuan H Z,Ge T D,Chen C Y,et al. Significant role for microbial autotrophy in the sequestration of soil carbon[J].Applied and Environmental Microbiology,2012,78(7):2328-2336.
[11]钱明媚,肖永良,彭文涛,等.免耕水稻土固定CO2自养微生物多样性[J].中国环境科学,2015,35(12):3754-3761.Qian M M, Xiao Y L, Peng W T, et al. Diversities of autotrophic CO2-fixing microbes in no-tillage paddy soils[J].China Environmental Science,2015,35(12):3754-3761.
[12] Wu X H,Ge T D,Wang W,et al. Cropping systems modulate the rate and magnitude of soil microbial autotrophic CO2fixation in soil[J]. Frontiers in Microbiology,2015,6:379.
[13] Meyer O,Schlegel H G. Biology of aerobic carbon monoxideoxidizing bacteria[J]. Annual Review of Microbiology,1983,37:277-310.
[14]刘琼,魏晓梦,吴小红,等.稻田土壤固碳功能微生物群落结构和数量特征[J].环境科学,2017,38(2):760-768.Liu Q,Wei X M,Wu X H,et al. Characteristic of abundances and diversity of carbon dioxide fixation microbes in Paddy soils[J]. Environmental Science,2017,38(2):760-768.
[15]王群艳,吴小红,祝贞科,等.土壤质地对自养固碳微生物及其同化碳的影响[J].环境科学,2016,37(10):3987-3995.Wang Q Y,Wu X H,Zhu Z K,et al. Effects of soil texture on autotrophic CO2fixation bacterial communities and their CO2assimilation contents[J]. Environmental Science,2016,37(10):3987-3995.
[16] Selesi D,Schmid M,Hartmann A. Diversity of green-like and red-like ribulose-1,5-bisphosphate carboxylase/oxygenase largesubunit genes(cbb L)in differently managed agricultural soils[J]. Applied and Environmental Microbiology,2005,71(1):175-184.
[17] Yuan H Z,Ge T D,Wu X H,et al. Long-term field fertilization alters the diversity of autotrophic bacteria based on the ribulose-1,5-biphosphate carboxylase/oxygenase(RubisCO)largesubunit genes in paddy soil[J]. Applied Microbiology and Biotechnology,2012,95(4):1061-1071.
[18]方峰,王磊,席雪飞,等.南海水域不同深度非光合微生物的固碳潜能及其对不同电子供体的响应[J].环境科学,2015,36(5):1550-1556.Fang F,Wang L,Xi X F,et al. Potential carbon fixation capability of non-photosynthetic microbial community at different depth of the south china sea and its response to different electron donors[J]. Environmental Science,2015,36(5):1550-1556.
[19]巩伏雨,蔡真,李寅. CO2固定的合成生物学[J].中国科学:生命科学,2015,45(10):993-1002.Gong F Y,Cai Z,Li Y. Synthetic biology for CO2fixation[J].Science China Life Sciences,2016,59(11):1106-1114.
[20] Hügler M,Sievert S M. Beyond the Calvin cycle:autotrophic carbon fixation in the ocean[J]. Annual Review of Marine Science,2011,3:261-289.
[21] Ge T D,Yuan H Z,Zhu H H,et al. Biological carbon assimilation and dynamics in a flooded rice-soil system[J]. Soil Biology and Biochemistry,2012,48:39-46.
[22] Tolli J, King G M. Diversity and structure of bacterial chemolithotrophic communities in pine forest and agroecosystem soils[J]. Applied and Environmental Microbiology,2005,71(12):8411-8418.
[23] Alfreider A,Vogt C,Hoffmann D,et al. Diversity of ribulose-1,5-bisphosphate carboxylase/oxygenase large-subunit genes from groundwater and aquifer microorganisms[J]. Microbial Ecology,2003,45(4):317-328.
[24] Yakimov M M,La Cono V,Denaro R. A first insight into the occurrence and expression of functional amoA and accA genes of autotrophic and ammonia-oxidizing bathypelagic Crenarchaeota of Tyrrhenian Sea[J]. Deep Sea Research PartⅡ:Topical Studies in Oceanography,2009,56(11-12):748-754.
[25] Campbell B J, Stein J L, Cary S C. Evidence of chemolithoautotrophy in the bacterial community associated with Alvinella pompejana, a hydrothermal vent polychaete[J].Applied and Environmental Microbiology,2003,69(9):5070-5078.
[26]李景云,秦嗣军,葛鹏,等.不同生育期苹果园土壤氨氧化微生物丰度研究[J].植物营养与肥料学报,2016,22(4):1149-1156.Li J Y,Qin S J,Ge P,et al. Abundance of ammonia oxidizers in apple orchard soil at different growth stages[J]. Journal of Plant Nutrition and Fertilizer,2016,22(4):1149-1156.
[27] Jiang X J,Hou X Y,Zhou X,et al. p H regulates key players of nitrification in paddy soils[J]. Soil Biology and Biochemistry,2015,81:9-16.
[28] Beauregard M S, Hamel C,Atul-Nayyar,et al. Long-term phosphorus fertilization impacts soil fungal and bacterial diversity but not AM fungal community in Alfalfa[J]. Microbial Ecology,2010,59(2):379-389.
[29]王继红,刘景双,于君宝,等.氮磷肥对黑土玉米农田生态系统土壤微生物量碳、氮的影响[J].水土保持学报,2004,18(1):35-38.Wang J H,Liu J S,Yu J B,et al. Effect of fertilizing N and P on soil microbial biomass carbon and nitrogen of black soil corn agroecosystem[J]. Journal of Soil and Water Conservation,2004,18(1):35-38.
[30]侯云鹏,杨建,孔丽丽,等.施磷对苏打盐碱土区水稻养分吸收、转运及分配的影响[J].吉林农业大学学报,2017,39(1):60-66.Hou Y P,Yang J,Kong L L,et al. Effects of phosphorus fertilizer application on nutrient absorption, translocation and distribution of rice in saline-sodic soil region[J]. Journal of Jilin Agricultural University,2017,39(1):60-66.
[31]魏亮,汤珍珠,祝贞科,等.水稻不同生育期根际与非根际土壤胞外酶对施氮的响应[J].环境科学,2017,38(8):3489-3496.Wei L,Tang Z Z,Zhu Z K,et al. Responses of extracellular enzymes to nitrogen application in rice of various ages with rhizosphere and bulk soil[J]. Environmental Science,2017,38(8):3489-3496.
[32]王佩琦,周伟丽,何圣兵,等.磷对混养反硝化污泥活性和微生物群落结构的影响[J].环境科学,2018,39(3):1350-1356.Wang P Q,Zhou W L,He S B,et al. Effects of phosphorus on the activity and bacterial community in mixotrophic denitrification sludge[J]. Environmental Science,2018,39(3):1350-1356.
[33] Cavigelli M A,Robertson G P. The functional significance of denitrifier community composition in a terrestrial ecosystem[J].Ecology,2000,81(5):1402-1414.
[34]王海涛,郑天凌,杨小茹.土壤反硝化的分子生态学研究进展及其影响因素[J].农业环境科学学报,2013,32(10):1915-1924.Wang H T,Zheng T L,Yang X R. Molecular ecology research progress for soil denitrification and research status for its influencing factors[J]. Journal of Agro-Environment Science,2013,32(10):1915-1924.
[35] Wei X M,Hu Y J,Peng P Q,et al. Effect of P stoichiometry on the abundance of nitrogen-cycle genes in phosphorus-limited paddy soil[J]. Biology and Fertility of Soils,2017,53(7):767-776.
[36] Ge T D,Li B Z,Zhu Z K,et al. Rice rhizodeposition and its utilization by microbial groups depends on N fertilization[J].Biology and Fertility of Soils,2017,53(1):37-48.
[37] Su J Q,Ding L J,Xue K,et al. Long-term balanced fertilization increases the soil microbial functional diversity in a phosphoruslimited paddy soil[J]. Molecular Ecology,2015,24(1):136-150.
[38] Vejchasarn P,Lynch J P,Brown K M. Genetic variability in phosphorus responses of rice root phenotypes[J]. Rice,2016,9:29.
[39] Baltar F,Lundin D,Palovaara J,et al. Prokaryotic responses to ammonium and organic carbon reveal alternative CO2fixation pathways and importance of alkaline phosphatase in the mesopelagic North Atlantic[J]. Frontiers in Microbiology,2016,7:1670.
[40] Berg I A. Ecological aspects of the distribution of different autotrophic CO2fixation pathways[J]. Applied and Environmental Microbiology,2011,77(6):1925-1936.
[41]袁红朝,秦红灵,刘守龙,等.固碳微生物分子生态学研究[J].中国农业科学,2011,44(5):2951-2958.Yuan H Z,Qin H L,Liu S L,et al. Advances in research of molecular ecology of carbon fixation microorganism[J]. Scientia Agricultura Sinica,2011,44(5):2951-2958.
[42]陈晓娟,吴小红,简燕,等.农田土壤自养微生物碳同化潜力及其功能基因数量、关键酶活性分析[J].环境科学,2014,35(3):1144-1150.Chen X J,Wu X H,Jian Y,et al. Carbon dioxide assimilation potential, functional gene amount and RubisCO activity of autotrophic microorganisms in agricultural Soils[J].Environmental Science,2014,35(3):1144-1150.
[2] Vance C P,Uhde-Stone C,Allan D L. Phosphorus acquisition and use:critical adaptations by plants for securing a nonrenewable resource[J]. New Phytologist,2003,157(3):423-447.
[3]赵小蓉,周然,李贵桐,等.低磷石灰性土壤施磷和小麦秸秆后土壤微生物量磷的变化[J].应用生态学报,2009,20(2):325-330.Zhao X R,Zhou R,Li G T,et al. Effects of applying inorganic P and wheat straw on the microbial biomass P and microbial P concentration in a calcareous soil with low concentration available P[J]. Chinese Journal of Applied Ecology,2009,20(2):325-330.
[4]王超.不同供磷量对玉米根系与土壤微生物互作及吐丝后地上部碳磷分配的影响[D].北京:中国农业大学,2017.Wang C. Influences of phosphorus supply on interactions between roots and soil microbes and post-silking partitioning of shoot carbon/phosphorus in field-grown maize[D]. Beijing:China Agricultural University,2017.
[5]刘俊杰,王光华,金剑,等.磷浓度处理对大豆根际土壤微生物群落结构的影响[J].大豆科学,2008,27(5):801-805.Liu J J,Wang G H,Jin J,et al. Effect of different phosphorus concentrations on microbial communities in soybean rhizosphere[J]. Soybean Science,2008,27(5):801-805.
[6]施瑶,王忠强,张心昱,等.氮磷添加对内蒙古温带典型草原土壤微生物群落结构的影响[J].生态学报,2014,34(17):4943-4949.Shi Y,Wang Z Q,Zhang X Y,et al. Effects of nitrogen and phosphorus addition on soil microbial community composition in temperate typical grassland in Inner Mongolia[J]. Acta Ecologica Sinica,2014,34(17):4943-4949.
[7]张四海,黄健,骆争荣,等.添加秸秆和磷素对土壤微生物群落的影响[J].应用生态学报,2014,25(3):797-802.Zhang S H,Huang J,Luo Z R,et al. Effect of adding different amounts of wheat straw and phosphorus on soil microorganism community[J]. Chinese Journal of Applied Ecology,2014,25(3):797-802.
[8] Huang J S,Hu B,Qi K B,et al. Effects of phosphorus addition on soil microbial biomass and community composition in a subalpine spruce plantation[J]. European Journal of Soil Biology,2016,72:35-41.
[9]林先贵,冯有智,陈瑞蕊.农田潮土养分耦合循环的微生物学机理研究进展[J].植物营养与肥料学报,2017,23(6):1575-1589.Lin X G,Feng Y Z,Chen R R. Research progresses of soil microorganisms driven nutrient coupling cycles in fluvo-aquic soils of China[J]. Journal of Plant Nutrition and Fertilizer,2017,23(6):1575-1589.
[10] Yuan H Z,Ge T D,Chen C Y,et al. Significant role for microbial autotrophy in the sequestration of soil carbon[J].Applied and Environmental Microbiology,2012,78(7):2328-2336.
[11]钱明媚,肖永良,彭文涛,等.免耕水稻土固定CO2自养微生物多样性[J].中国环境科学,2015,35(12):3754-3761.Qian M M, Xiao Y L, Peng W T, et al. Diversities of autotrophic CO2-fixing microbes in no-tillage paddy soils[J].China Environmental Science,2015,35(12):3754-3761.
[12] Wu X H,Ge T D,Wang W,et al. Cropping systems modulate the rate and magnitude of soil microbial autotrophic CO2fixation in soil[J]. Frontiers in Microbiology,2015,6:379.
[13] Meyer O,Schlegel H G. Biology of aerobic carbon monoxideoxidizing bacteria[J]. Annual Review of Microbiology,1983,37:277-310.
[14]刘琼,魏晓梦,吴小红,等.稻田土壤固碳功能微生物群落结构和数量特征[J].环境科学,2017,38(2):760-768.Liu Q,Wei X M,Wu X H,et al. Characteristic of abundances and diversity of carbon dioxide fixation microbes in Paddy soils[J]. Environmental Science,2017,38(2):760-768.
[15]王群艳,吴小红,祝贞科,等.土壤质地对自养固碳微生物及其同化碳的影响[J].环境科学,2016,37(10):3987-3995.Wang Q Y,Wu X H,Zhu Z K,et al. Effects of soil texture on autotrophic CO2fixation bacterial communities and their CO2assimilation contents[J]. Environmental Science,2016,37(10):3987-3995.
[16] Selesi D,Schmid M,Hartmann A. Diversity of green-like and red-like ribulose-1,5-bisphosphate carboxylase/oxygenase largesubunit genes(cbb L)in differently managed agricultural soils[J]. Applied and Environmental Microbiology,2005,71(1):175-184.
[17] Yuan H Z,Ge T D,Wu X H,et al. Long-term field fertilization alters the diversity of autotrophic bacteria based on the ribulose-1,5-biphosphate carboxylase/oxygenase(RubisCO)largesubunit genes in paddy soil[J]. Applied Microbiology and Biotechnology,2012,95(4):1061-1071.
[18]方峰,王磊,席雪飞,等.南海水域不同深度非光合微生物的固碳潜能及其对不同电子供体的响应[J].环境科学,2015,36(5):1550-1556.Fang F,Wang L,Xi X F,et al. Potential carbon fixation capability of non-photosynthetic microbial community at different depth of the south china sea and its response to different electron donors[J]. Environmental Science,2015,36(5):1550-1556.
[19]巩伏雨,蔡真,李寅. CO2固定的合成生物学[J].中国科学:生命科学,2015,45(10):993-1002.Gong F Y,Cai Z,Li Y. Synthetic biology for CO2fixation[J].Science China Life Sciences,2016,59(11):1106-1114.
[20] Hügler M,Sievert S M. Beyond the Calvin cycle:autotrophic carbon fixation in the ocean[J]. Annual Review of Marine Science,2011,3:261-289.
[21] Ge T D,Yuan H Z,Zhu H H,et al. Biological carbon assimilation and dynamics in a flooded rice-soil system[J]. Soil Biology and Biochemistry,2012,48:39-46.
[22] Tolli J, King G M. Diversity and structure of bacterial chemolithotrophic communities in pine forest and agroecosystem soils[J]. Applied and Environmental Microbiology,2005,71(12):8411-8418.
[23] Alfreider A,Vogt C,Hoffmann D,et al. Diversity of ribulose-1,5-bisphosphate carboxylase/oxygenase large-subunit genes from groundwater and aquifer microorganisms[J]. Microbial Ecology,2003,45(4):317-328.
[24] Yakimov M M,La Cono V,Denaro R. A first insight into the occurrence and expression of functional amoA and accA genes of autotrophic and ammonia-oxidizing bathypelagic Crenarchaeota of Tyrrhenian Sea[J]. Deep Sea Research PartⅡ:Topical Studies in Oceanography,2009,56(11-12):748-754.
[25] Campbell B J, Stein J L, Cary S C. Evidence of chemolithoautotrophy in the bacterial community associated with Alvinella pompejana, a hydrothermal vent polychaete[J].Applied and Environmental Microbiology,2003,69(9):5070-5078.
[26]李景云,秦嗣军,葛鹏,等.不同生育期苹果园土壤氨氧化微生物丰度研究[J].植物营养与肥料学报,2016,22(4):1149-1156.Li J Y,Qin S J,Ge P,et al. Abundance of ammonia oxidizers in apple orchard soil at different growth stages[J]. Journal of Plant Nutrition and Fertilizer,2016,22(4):1149-1156.
[27] Jiang X J,Hou X Y,Zhou X,et al. p H regulates key players of nitrification in paddy soils[J]. Soil Biology and Biochemistry,2015,81:9-16.
[28] Beauregard M S, Hamel C,Atul-Nayyar,et al. Long-term phosphorus fertilization impacts soil fungal and bacterial diversity but not AM fungal community in Alfalfa[J]. Microbial Ecology,2010,59(2):379-389.
[29]王继红,刘景双,于君宝,等.氮磷肥对黑土玉米农田生态系统土壤微生物量碳、氮的影响[J].水土保持学报,2004,18(1):35-38.Wang J H,Liu J S,Yu J B,et al. Effect of fertilizing N and P on soil microbial biomass carbon and nitrogen of black soil corn agroecosystem[J]. Journal of Soil and Water Conservation,2004,18(1):35-38.
[30]侯云鹏,杨建,孔丽丽,等.施磷对苏打盐碱土区水稻养分吸收、转运及分配的影响[J].吉林农业大学学报,2017,39(1):60-66.Hou Y P,Yang J,Kong L L,et al. Effects of phosphorus fertilizer application on nutrient absorption, translocation and distribution of rice in saline-sodic soil region[J]. Journal of Jilin Agricultural University,2017,39(1):60-66.
[31]魏亮,汤珍珠,祝贞科,等.水稻不同生育期根际与非根际土壤胞外酶对施氮的响应[J].环境科学,2017,38(8):3489-3496.Wei L,Tang Z Z,Zhu Z K,et al. Responses of extracellular enzymes to nitrogen application in rice of various ages with rhizosphere and bulk soil[J]. Environmental Science,2017,38(8):3489-3496.
[32]王佩琦,周伟丽,何圣兵,等.磷对混养反硝化污泥活性和微生物群落结构的影响[J].环境科学,2018,39(3):1350-1356.Wang P Q,Zhou W L,He S B,et al. Effects of phosphorus on the activity and bacterial community in mixotrophic denitrification sludge[J]. Environmental Science,2018,39(3):1350-1356.
[33] Cavigelli M A,Robertson G P. The functional significance of denitrifier community composition in a terrestrial ecosystem[J].Ecology,2000,81(5):1402-1414.
[34]王海涛,郑天凌,杨小茹.土壤反硝化的分子生态学研究进展及其影响因素[J].农业环境科学学报,2013,32(10):1915-1924.Wang H T,Zheng T L,Yang X R. Molecular ecology research progress for soil denitrification and research status for its influencing factors[J]. Journal of Agro-Environment Science,2013,32(10):1915-1924.
[35] Wei X M,Hu Y J,Peng P Q,et al. Effect of P stoichiometry on the abundance of nitrogen-cycle genes in phosphorus-limited paddy soil[J]. Biology and Fertility of Soils,2017,53(7):767-776.
[36] Ge T D,Li B Z,Zhu Z K,et al. Rice rhizodeposition and its utilization by microbial groups depends on N fertilization[J].Biology and Fertility of Soils,2017,53(1):37-48.
[37] Su J Q,Ding L J,Xue K,et al. Long-term balanced fertilization increases the soil microbial functional diversity in a phosphoruslimited paddy soil[J]. Molecular Ecology,2015,24(1):136-150.
[38] Vejchasarn P,Lynch J P,Brown K M. Genetic variability in phosphorus responses of rice root phenotypes[J]. Rice,2016,9:29.
[39] Baltar F,Lundin D,Palovaara J,et al. Prokaryotic responses to ammonium and organic carbon reveal alternative CO2fixation pathways and importance of alkaline phosphatase in the mesopelagic North Atlantic[J]. Frontiers in Microbiology,2016,7:1670.
[40] Berg I A. Ecological aspects of the distribution of different autotrophic CO2fixation pathways[J]. Applied and Environmental Microbiology,2011,77(6):1925-1936.
[41]袁红朝,秦红灵,刘守龙,等.固碳微生物分子生态学研究[J].中国农业科学,2011,44(5):2951-2958.Yuan H Z,Qin H L,Liu S L,et al. Advances in research of molecular ecology of carbon fixation microorganism[J]. Scientia Agricultura Sinica,2011,44(5):2951-2958.
[42]陈晓娟,吴小红,简燕,等.农田土壤自养微生物碳同化潜力及其功能基因数量、关键酶活性分析[J].环境科学,2014,35(3):1144-1150.Chen X J,Wu X H,Jian Y,et al. Carbon dioxide assimilation potential, functional gene amount and RubisCO activity of autotrophic microorganisms in agricultural Soils[J].Environmental Science,2014,35(3):1144-1150.