海洋环境中铁的来源、微生物作用过程及生态效应
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摘要
铁在地壳中的含量位列第四,在海洋中常以微量元素的身份出现。铁拥有多变的价态和多样的功能,是调节海洋初级生产力和驱动海洋生物地球化学循环的重要力量。以往的研究表明,铁在维持初级生产力、耦合物质循环以及调节生源要素转化中具有重要作用。近年来微生物生态学的发展将铁的研究推向了新的高度,包括微生物驱动的铁氧化—还原行为、代谢过程以及与主要元素(C/N/P)的交互关系等。以近15年发表的文献为重点,尝试综述铁的最新进展。首先梳理了海洋中铁的来源和赋存状态(溶解态、胶体态、颗粒态和有机态);其次阐释了微生物介导的铁氧化还原类型和过程机制(如硝酸盐氧化、生物还原等);最后总结了铁与C/N/P循环的耦合关系以及在特定生态事件中的生态效应。此外,对海洋铁研究的"化学—生物—物理"理论框架也进行了展望,旨在为更好地认识铁循环及其在海洋微生态过程的作用提供资料借鉴。
Iron is the fourth most abundant element in the earth's crust,and it often appears as a trace element in the ocean. Iron has a variable valence and diverse functions,and is an important force to regulate marine primary productivity and drive the geochemical cycle in the ocean. Previous studies have shown that iron plays an important role in maintaining primary productivity,coupling matter cycles,and regulating the transformation of biogenic factors. In recent years,with the development of microbial ecology,iron research has enter more in-depth levels,including microbial-driven iron oxidation-reduction behavior,metabolic processes,and interactions with other major elements(C/N/P). This paper attempts to review the latest progress of iron,focusing on the published literatures of the past fifteen years. Firstly,we explained the sources and occurrence states of iron in the ocean(such as dissolved,colloidal,granular and organic state);Secondly,we reviewed the types and process mechanisms of microbial-mediated iron redoxbehavior(for example nitrate oxidation,and biological reduction,etc.);Finally,we summarized the coupling relationship between iron and C/N/P cycle.Additionally,the ecological roles of iron in specific ecological event(for instance algal bloom)has also been described. Furthermore,the "chemical-biological-physical" theoretical framework for marine iron research is also discussed. The purpose of this paper is to provide more information for marine microecological research and their effects on iron cycle under changing environment,such as global change.
Iron is the fourth most abundant element in the earth's crust,and it often appears as a trace element in the ocean. Iron has a variable valence and diverse functions,and is an important force to regulate marine primary productivity and drive the geochemical cycle in the ocean. Previous studies have shown that iron plays an important role in maintaining primary productivity,coupling matter cycles,and regulating the transformation of biogenic factors. In recent years,with the development of microbial ecology,iron research has enter more in-depth levels,including microbial-driven iron oxidation-reduction behavior,metabolic processes,and interactions with other major elements(C/N/P). This paper attempts to review the latest progress of iron,focusing on the published literatures of the past fifteen years. Firstly,we explained the sources and occurrence states of iron in the ocean(such as dissolved,colloidal,granular and organic state);Secondly,we reviewed the types and process mechanisms of microbial-mediated iron redoxbehavior(for example nitrate oxidation,and biological reduction,etc.);Finally,we summarized the coupling relationship between iron and C/N/P cycle.Additionally,the ecological roles of iron in specific ecological event(for instance algal bloom)has also been described. Furthermore,the "chemical-biological-physical" theoretical framework for marine iron research is also discussed. The purpose of this paper is to provide more information for marine microecological research and their effects on iron cycle under changing environment,such as global change.
引文
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[2]Tu Xiaoxia.The effect of iron to the primary production in the ocean[J].Journal of Agricultural Mechanization Research,2007,(3):18-20.[屠霄霞.铁对海洋初级生产力的影响[J].农机化研究,2007,(3):18-20.]
[3]Martin J H,Gordon M,Fitzwater S E.The case for iron[J].Limnology&Oceanography,1991,36(8):1 793-1 802.
[4]Hogle S L,Barbeau K A,Gledhill M.Heme in the marine environment:From cells to the iron cycle[J].Metallomics,2014,6(6):1 107-1 120.
[5]Emerson D.Biogenic iron dust:A novel approach to ocean iron fertilization as a means of large scale removal of carbon dioxide from the atmosphere[J].Frontiers in Marine Science,2019,6:22.
[6]Watson A J,Bakker D C E,Ridgwell A J,et al.Effect of iron supply on Southern Ocean CO2 uptake and implications for glacial atmospheric CO2[J].Nature,2000,407(6 805):730.
[7]Zhou L,Tan Y,Huang L,et al.Aluminum effects on marine phytoplankton:Implications for a revised iron hypothesis(IronAluminum Hypothesis)[J].Biogeochemistry,2018,139(2):123-137.
[8]Tagliabue A,Bowie A R,Boyd P W,et al.The integral role of iron in ocean biogeochemistry[J].Nature,2017,543(7 643):51-59.
[9]Norman L,Cabanesa D J,Blanco-Ameijeiras S,et al.Iron biogeochemistry in aquatic systems:From source to bioavailability[J].Chimia International Journal for Chemistry,2014,68(11):764-771.
[10]Havens S M,Hassler C S,North R L,et al.Iron plays a role in nitrate drawdown by phytoplankton in Lake Erie surface waters as observed in lake-wide assessments[J].Canadian Journal of Fisheries&Aquatic Sciences,2012,69(2):369-381.
[11]Qin Yanwen,Zhang Manping.Iron sources,existing forms and their limiting action on the primary productivity of phytoplankton in seawater[J].Advances in Marine Science,1998,16(3):67-75.[秦延文,张曼平.海洋中铁的来源、形态和对初级生产力的限制作用[J].海洋科学进展,1998,16(3):67-75.]
[12]Birchill A J,Hartner N T,Kunde K,et al.The eastern extent of seasonal iron limitation in the high latitude North Atlantic Ocean[J].Scientific Reports,2019,9(1):1 435.
[13]Naito K,Matsui M,Imai I.Ability of marine eukaryotic red tide microalgae to utilize insoluble iron[J].Harmful Algae,2005,4(6):1 021-1 032.
[14]McRose D L,Seyedsayamdost M R,Morel F M M.Multiple siderophores:Bug or feature?[J].Journal of Biological Inorganic Chemistry,2018,23(7):983-993.
[15]Kazamia E,Sutak R,Paz-Yepes J,et al.Endocytosis-mediated siderophore uptake as a strategy for Fe acquisition in diatoms[J].Science Advances,2018,4(5):eaar4 536.
[16]Blain S,Tagliabue A.Iron Cycle in Oceans[M].Hoboken,New Jersey:John Wiley&Sons,2016.
[17]Wells M L,Trick C G,Cochlan W P,et al.Domoic acid:The synergy of iron,copper,and the toxicity of diatoms[J].Limnology and Oceanography,2005,50(6):1 908-1 917.
[18]Decho A W,Gutierrez T.Microbial Extracellular Polymeric Substances(EPSs)in ocean systems[J].Frontiers in Microbiology,2017,8:922.
[19]Hassler C S,Schoemann V,Nichols C M,et al.Saccharides enhance iron bioavailability to Southern Ocean phytoplankton[J].Proceedings of the National Academy of Sciences,2011,108(3):1 076-1 081.
[20]Tortell P D,Maldonado M T,Price N M.The role of heterotrophic bacteria in iron-limited ocean ecosystems[J].Nature,1996,383(6 598):330-332.
[21]Bonnain C,Breitbart M,Buck K N.The ferrojan horse hypothesis:Iron-virus interactions in the ocean[J].Frontiers in Marine Science,2016,3:82.
[22]Mackey K R,Post A F,Mcilvin M R,et al.Divergent responses of Atlantic coastal and oceanic Synechococcus to iron limitation[J].Proceedings of the National Academy of Sciences of the United States of America,2015,112(32):9 944-9 949.
[23]Hogle S L,Bianca B,Barbeau K A.Direct heme uptake by phytoplankton-associated roseobacter bacteria[J].Msystems,2017,2(1):e00124-16.
[24]Maldonado M T,Price N M.Utilization of iron bound to strong organic ligands by plankton communities in the subarctic Pacific Ocean[J].Deep-Sea Research Part II:Topical Studies in Oceanography,1999,46(11/12):2 447-2 473.
[25]Johnson K S,Gordon R M,Coale K H.What controls dissolved iron concentrations in the world ocean?[J].Marine Chemistry,1997,57(3/4):137-161.
[26]Chen Lei,Zhang Hongxia,Li Ying,et al.The role of microorganisms in the geochemical iron cycle[J].Scientia Sinica Vitae,2016,46(9):1 069-1 078.[陈蕾,张洪霞,李莹,等.微生物在地球化学铁循环过程中的作用[J].中国科学:生命科学,2016,46(9):1 069-1 078.]
[27]Straub K L,Benz M,Schink B.Iron metabolism in anoxic environments at near neutral pH[J].Fems Microbiology Ecology,2000,34(3):181-186.
[28]Makita H.Iron-oxidizing bacteria in marine environments:Recent progresses and future directions[J].World Journal of Microbiology and Biotechnology,2018,34(8):110.
[29]Emerson D.The role of iron-oxidizing bacteria in biocorrosion:A review[J].Biofouling,2018,34(9):989-1 000.
[30]Nelson Y M,Lion LW,Ghiorse W C,et al.Production of biogenic Mn oxides by leptothrix discophora SS-1 in a chemically defined growth medium and evaluation of their Pb adsorption characteristics[J].Applied and Environmental Microbiology,1999,65(1):175-180.
[31]Makita H.Iron-oxidizing bacteria in marine environments:Recent progresses and future directions[J].World Journal of Microbiology and Biotechnology,2018,34(8):110.
[32]Bryce C,Blackwell N,Schmidt C,et al.Microbial anaerobic Fe(II)oxidation-ecology,mechanisms and environmental implications[J].Environmental Microbiology,2018,20(10):3 462-3 483.
[33]Kappler A,Schink B,Newman D K.Fe(III)mineral formation and cell encrustation by the nitrate-dependent Fe(II)-oxidizer strain BoFeN1[J].Geobiology,2010,3(4):235-245.
[34]Chakraborty A,Picardal F.Neutrophilic,nitrate-dependent,Fe(II)oxidation by a dechloromonas species[J].World Journal of Microbiology Biotechnology,2013,29(4):617-623.
[35]Fleming E J,Langdon A E,Martinez-Garcia M,et al.What's new is old:Resolving the identity of leptothrix ochracea using single cell genomics,pyrosequencing and fish[J].PLoS ONE,2011,6(3):e17769.
[36]Hu Min,Li Fangbai.Soil microbe mediated iron cycling and its environmental implication[J].Acta Pedologica Sinica,2014,5(14):683-698.[胡敏,李芳柏.土壤微生物铁循环及其环境意义[J].土壤学报,2014,5(14):683-698.]
[37]Luo Hailin,Tang Jia,Zhou Puxiong,et al.Influence of secondary iron-oxide mineralization induced by dissimilatory iron reduction bacteria on fraction transformation of heavy metals in soil[J].Chinese Journal of Ecology,2018,37(6):1 620-1 627.[罗海林,汤佳,周普雄,等.异化铁还原诱导次生铁矿对土壤重金属形态转化的影响[J].生态学杂志,2018,37(6):1 620-1 627.]
[38]Lovley D R.Dissimilatory Fe(III)and Mn(IV)reduction[J].Advances in Microbial Physiology,2004,49(2):259-287.
[39]Ebrahiminezhad A,Manafi Z,Berenjian A,et al.Iron-reducing bacteria and iron nanostructures[J].Journal of Advanced Medical Sciences and Applied Technologies,2017,3(1):9-16.
[40]Lovley D R.Microbial Fe(III)reduction in subsurface environments[J].Fems Microbiology Reviews,2010,20(3/4):305-313.
[41]Newsome L,Lopez Adams R,Downie H F,et al.NanoSIMSimaging of extracellular electron transport processes during microbial iron(III)reduction[J].FEMS Microbiology Ecology,2018,94(8):fiy104.
[42]Shi L,Richardson D J,Wang Z,et al.The roles of outer membrane cytochromes of Shewanella and Geobacter in extracellular electron transfer[J].Environmental Microbiology Reports,2009,1(4):220-227.
[43]Wang Wenyan,Quan Xiangchun,He Mengchang,et al.Review on the mechanism and development of ferric iron microbial reduction[J].Environmental Pollution&Control,2006,28(2):116-120.[王文燕,全向春,何孟常,等.Fe(Ⅲ)微生物还原机理及其研究进展[J].环境污染与防治,2006,28(2):116-120.]
[44]Conway T M,John S G.Quantification of dissolved iron sources to the North Atlantic Ocean[J].Nature,2014,511(7 508):212-215.
[45]Gruber N.Elusive marine nitrogen fixation[J].Proceedings of the National Academy of Sciences,2016,113(16):4 246-4 248.
[46]Weber T,Deutsch C.Local versus basin-scale limitation of marine nitrogen fixation[J].Proceedings of the National Academy of Sciences of the United States of America,2014,111(24):8 741-8 746.
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