微生物对低温极端环境适应性的研究进展
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国家自然科学基金(41701085);重要生物资源保护与利用研究安徽省重点实验室开放课题基金(swzy202008)


Progress in research on the adaptability of microorganisms to extremely cold environments
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    摘要:

    嗜冷微生物是地球寒冷环境中最主要的生物类群,并且是驱动全球生物地球化学循环的关键环节。嗜冷微生物在适应策略上显示出应对多种极端环境因素的巨大潜力,研究其适应和进化机制有助于更好地理解微生物与环境之间相互作用过程,并有效利用极端环境微生物资源。近年来,随着分子生物学和基因组学技术的高速发展,对微生物适应寒冷环境的机制及嗜冷微生物在指示气候变化和工农业应用方面均有一系列的突破。在此,本文将从基因组的GC含量、蛋白质稳定性、转录翻译调控、细胞膜流动性、渗透压调节、抗氧化损失和基因组适应性进化等方面总结当前在微生物适应低温环境机制上所取得的进展,并展望低温环境微生物在指示气候变化和工农业应用中的前景。

    Abstract:

    Psychrophilic microorganisms are a major life form in cold environments of the Earth and drive critical global biogeochemical cycles. The survival strategies of these microorganisms have demonstrated great potential in overcoming the extreme environmental factors. Deciphering their adaptation and evolutionary mechanisms will improve our understanding of the interaction between microorganisms and the environment and facilitate the effective use of microbial resources from extreme environments. With the rapid development of molecular biology and genome sequencing in recent years, researchers have made great breakthroughs in research on the adaptation mechanisms of microorganisms to cold environments and application of these psychrophiles in climate change prediction, industry, and agriculture. In this review, we summarize the progress of research on microbial adaptation to cold environment in terms of genomic GC content, protein stability, transcriptional and translational regulation, cell membrane fluidity, osmotic pressure regulation, oxidation resistance, and genome adaptive evolution.

    参考文献
    [1] Priscu JC, Johnson L, Christner BC. Earth's icy biosphere. Washington, D.C.:ASM Press, 2004.
    [2] Xiang SR, Shang TC, Chen Y, Jing ZF, Yao TD. Changes in diversity and biomass of bacteria along a shallow snow pit from Kuytun 51 Glacier, Tianshan Mountains, China. Journal of Geophysical Research Atmospheres, 2009, 114(G4):G04008.
    [3] Jungblut AD, Lovejoy C, Vincent WF. Global distribution of cyanobacterial ecotypes in the cold biosphere. The ISME Journal, 2010, 4(2):191-202.
    [4] An L, Chen Y, Xiang SR, Shang TC, Tian LD. Differences in community composition of bacteria in four deep ice sheets in Western China. Biogeosciences Discussions, 2010, 7(1):1167.
    [5] Boetius A, Anesio AM, Deming JW, Mikucki JA, Rapp JZ. Microbial ecology of the cryosphere:sea ice and glacial habitats. Nature Reviews Microbiology, 2015, 13(11):677-690.
    [6] Delgado-Baquerizo M, Oliverio AM, Brewer TE, Benavent-González A, Eldridge DJ, Bardgett RD, Maestre FT, Singh BK, Fierer N. A global atlas of the dominant bacteria found in soil. Science, 2018, 359(6373):320-325.
    [7] Panikov NS, Flanagan PW, Oechel WC, Mastepanov MA, Christensen TR. Microbial activity in soils frozen to below-39℃. Soil Biology and Biochemistry, 2006, 38(4):785-794.
    [8] Shen L, Yao TD, Liu YQ, Jiao NZ, Kang SC, Xu BQ, Zhang SH, Liu XB. Downward-shifting temperature range for the growth of snow-bacteria on glaciers of the Tibetan Plateau. Geomicrobiology Journal, 2014, 31(9):779-787.
    [9] Scholze C, Jørgensen BB, Røy H. Psychrophilic properties of sulfate-reducing bacteria in Arctic marine sediments. Limnology and Oceanography, 2021, 66(S1):S293-S302.
    [10] Shen L, Liu YQ, Allen MA, Xu BQ, Wang NL, Williams TJ, Wang F, Zhou YG, Liu Q, Cavicchioli R. Linking genomic and physiological characteristics of psychrophilic Arthrobacter to metagenomic data to explain global environmental distribution. Microbiome, 2021, 9(1):136.
    [11] Cary SC, McDonald IR, Barrett JE, Cowan DA. On the rocks:the microbiology of Antarctic dry valley soils. Nature Reviews Microbiology, 2010, 8(2):129-138.
    [12] Cavicchioli R. Microbial ecology of Antarctic aquatic systems. Nature Reviews Microbiology, 2015, 13(11):691-706.
    [13] Jansson JK, Taş N. The microbial ecology of permafrost. Nature Reviews Microbiology, 2014, 12(6):414-425.
    [14] Smith HJ, Foster RA, McKnight DM, Lisle JT, Littmann S, Kuypers MMM, Foreman CM. Microbial formation of labile organic carbon in Antarctic glacial environments. Nature Geoscience, 2017, 10(5):356-359.
    [15] Cauvy-Fraunié S, Dangles O. A global synthesis of biodiversity responses to glacier retreat. Nature Ecology & Evolution, 2019, 3(12):1675-1685.
    [16] Yarzábal LA. Perspectives for using glacial and periglacial microorganisms for plant growth promotion at low temperatures. Applied Microbiology and Biotechnology, 2020, 104(8):3267-3278.
    [17] Kasana RC, Pandey CB. Exiguobacterium:an overview of a versatile genus with potential in industry and agriculture. Critical Reviews in Biotechnology, 2018, 38(1):141-156.
    [18] Bowman JP, McCammon SA, Brown MV, Nichols DS, McMeekin TA. Diversity and association of psychrophilic bacteria in Antarctic Sea ice. Applied and Environmental Microbiology, 1997, 63(8):3068-3078.
    [19] Morita RY. Psychrophilic bacteria. Bacteriological Reviews, 1975, 39(2):144-167.
    [20] Gounot AM. Effects of temperature on the growth of psychrophilic bacteria from glaciers. Canadian Journal of Microbiology, 1976, 22(6):839-846.
    [21] Cavicchioli R. On the concept of a psychrophile. The ISME Journal, 2016, 10(4):793-795.
    [22] Farrell J, Rose A. Temperature effects on microorganisms. Annual Review of Microbiology, 1967, 21:101-120.
    [23] D'Amico S, Collins T, Marx JC, Feller G, Gerday C. Psychrophilic microorganisms:challenges for life. EMBO Reports, 2006, 7(4):385-389.
    [24] Liu YQ, Priscu JC, Yao TD, Vick-Majors TJ, Michaud AB, Sheng L. Culturable bacteria isolated from seven high-altitude ice cores on the Tibetan Plateau. Journal of Glaciology, 2019, 65(249):29-38.
    [25] Shen L, Liu Y, Xu B, Wang N, Zhao H, Liu X, Liu F. Comparative genomic analysis reveals the environmental impacts on two Arcticibacter strains including sixteen Sphingobacteriaceae species. Scientific Reports, 2017, 7:2055.
    [26] Rivkina EM, Friedmann EI, McKay CP, Gilichinsky DA. Metabolic activity of permafrost bacteria below the freezing point. Applied and Environmental Microbiology, 2000, 66(8):3230-3233.
    [27] Panikov NS, Sizova MV. Growth kinetics of microorganisms isolated from Alaskan soil and permafrost in solid media frozen down to-35℃. FEMS Microbiology Ecology, 2007, 59(2):500-512.
    [28] Gadkari PS, McGuinness LR, Männistö MK, Kerkhof LJ, Häggblom MM. Arctic tundra soil bacterial communities active at subzero temperatures detected by stable isotope probing. FEMS Microbiology Ecology, 2019, 96(2):fiz192.
    [29] Rodrigues DF, Tiedje JM. Coping with our cold planet. Applied and Environmental Microbiology, 2008, 74(6):1677-1686.
    [30] Piette F, D'Amico S, Mazzucchelli G, Danchin A, Leprince P, Feller G. Life in the cold:a proteomic study of cold-repressed proteins in the Antarctic bacterium Pseudoalteromonas haloplanktis TAC125. Applied and Environmental Microbiology, 2011, 77(11):3881-3883.
    [31] Morgan-Kiss RM, Priscu JC, Pocock T, Gudynaite-Savitch L, Huner NPA. Adaptation and acclimation of photosynthetic microorganisms to permanently cold environments. Microbiology and Molecular Biology Reviews:MMBR, 2006, 70(1):222-252.
    [32] Tehei M, Zaccai G. Adaptation to extreme environments:macromolecular dynamics in complex systems. Biochimica et Biophysica Acta:BBA-General Subjects, 2005, 1724(3):404-410.
    [33] Hinnebusch J, Tilly K. Linear plasmids and chromosomes in bacteria. Molecular Microbiology, 1993, 10(5):917-922.
    [34] Galtier N, Lobry JR. Relationships between genomic G+C content, RNA secondary structures, and optimal growth temperature in prokaryotes. Journal of Molecular Evolution, 1997, 44(6):632-636.
    [35] Meyer MM. Revisiting the relationships between genomic G+C content, RNA secondary structures, and optimal growth temperature. Journal of Molecular Evolution, 2021, 89(3):165-171.
    [36] Liu Q, Song WZ, Zhou YG, Dong XZ, Xin YH. Phenotypic divergence of thermotolerance:molecular basis and cold adaptive evolution related to intrinsic DNA flexibility of glacier-inhabiting Cryobacterium strains. Environmental Microbiology, 2020, 22(4):1409-1420.
    [37] Wang HC, Susko E, Roger AJ. On the correlation between genomic G+C content and optimal growth temperature in prokaryotes:data quality and confounding factors. Biochemical and Biophysical Research Communications, 2006, 342(3):681-684.
    [38] Khachane AN, Timmis KN, Dos Santos VAPM. Uracil content of 16S rRNA of thermophilic and psychrophilic prokaryotes correlates inversely with their optimal growth temperatures. Nucleic Acids Research, 2005, 33(13):4016-4022.
    [39] Sato Y, Kimura H. Temperature-dependent expression of different guanine-plus-cytosine content 16S rRNA genes in Haloarcula strains of the class Halobacteria. Antonie Van Leeuwenhoek, 2019, 112(2):187-201.
    [40] Cavicchioli R. Cold-adapted archaea. Nature Reviews Microbiology, 2006, 4(5):331-343.
    [41] Saunders NFW, Thomas T, Curmi PMG, Mattick JS, Kuczek E, Slade R, Davis J, Franzmann PD, Boone D, Rusterholtz K, Feldman R, Gates C, Bench S, Sowers K, Kadner K, Aerts A, Dehal P, Detter C, Glavina T, Lucas S, Richardson P, Larimer F, Hauser L, Land M, Cavicchioli R. Mechanisms of thermal adaptation revealed from the genomes of the Antarctic archaea Methanogenium frigidum and Methanococcoides burtonii. Genome Research, 2003, 13(7):1580-1588.
    [42] Siddiqui KS, Williams TJ, Wilkins D, Yau S, Allen MA, Brown MV, Lauro FM, Cavicchioli R. Psychrophiles. Annual Review of Earth and Planetary Sciences, 2013, 41:87-115.
    [43] Siddiqui KS, Poljak A, Guilhaus M, De Francisci D, Curmi PMG, Feller G, D'Amico S, Gerday C, Uversky VN, Cavicchioli R. Role of lysine versus arginine in enzyme cold-adaptation:modifying lysine to Homo-arginine stabilizes the cold-adapted α-amylase from Pseudoalteramonas haloplanktis. Proteins:Structure, Function, and Bioinformatics, 2006, 64(2):486-501.
    [44] Huston AL, Haeggström JZ, Feller G. Cold adaptation of enzymes:structural, kinetic and microcalorimetric characterizations of an aminopeptidase from the Arctic psychrophile Colwellia psychrerythraea and of human leukotriene A4 hydrolase. Biochimica et Biophysica Acta:BBA-Proteins and Proteomics, 2008, 1784(11):1865-1872.
    [45] Michaux C, Massant J, Kerff F, Frère JM, Docquier JD, Vandenberghe I, Samyn B, Pierrard A, Feller G, Charlier P, Van Beeumen J, Wouters J. Crystal structure of a cold-adapted class C β-lactamase. The FEBS Journal, 2008, 275(8):1687-1697.
    [46] Bauvois C, Jacquamet L, Huston AL, Borel F, Feller G, Ferrer JL. Crystal structure of the cold-active aminopeptidase from Colwellia psychrerythraea, a close structural homologue of the human bifunctional leukotriene A4 hydrolase. Journal of Biological Chemistry, 2008, 283(34):23315-23325.
    [47] Sonan GK, Receveur-Brechot V, Duez C, Aghajari N, Czjzek M, Haser R, Gerday C. The linker region plays a key role in the adaptation to cold of the cellulase from an Antarctic bacterium. The Biochemical Journal, 2007, 407(2):293-302.
    [48] Aghajari N, Van Petegem F, Villeret V, Chessa JP, Gerday C, Haser R, Van Beeumen J. Crystal structures of a psychrophilic metalloprotease reveal new insights into catalysis by cold-adapted proteases. Proteins, 2003, 50(4):636-647.
    [49] Leiros HKS, Pey AL, Innselset M, Moe EL, Leiros I, Steen IH, Martinez A. Structure of phenylalanine hydroxylase from Colwellia psychrerythraea 34H, a monomeric cold active enzyme with local flexibility around the active site and high overall stability. Journal of Biological Chemistry, 2007, 282(30):21973-21986.
    [50] Paredes DI, Watters K, Pitman DJ, Bystroff C, Dordick JS. Comparative void-volume analysis of psychrophilic and mesophilic enzymes:structural bioinformatics of psychrophilic enzymes reveals sources of core flexibility. BMC Structural Biology, 2011, 11:42.
    [51] Jung SK, Jeong DG, Lee MS, Lee JK, Kim HK, Ryu SE, Park BC, Kim JH, Kim SJ. Structural basis for the cold adaptation of psychrophilic M37 lipase from Photobacterium lipolyticum. Proteins, 2008, 71(1):476-484.
    [52] Cramer P. AlphaFold2 and the future of structural biology. Nature Structural & Molecular Biology, 2021, 28(9):704-705.
    [53] Graumann P, Marahiel MA. Some like it cold:response of microorganisms to cold shock. Archives of Microbiology, 1996, 166(5):293-300.
    [54] Horn G, Hofweber R, Kremer W, Kalbitzer HR. Structure and function of bacterial cold shock proteins. Cellular and Molecular Life Sciences:CMLS, 2007, 64(12):1457-1470.
    [55] Chen ZJ, Yu HY, Li LY, Hu SN, Dong XZ. The genome and transcriptome of a newly described psychrophilic archaeon, Methanolobus psychrophilus R15, reveal its cold adaptive characteristics. Environmental Microbiology Reports, 2012, 4(6):633-641.
    [56] Gao HC, Wang Y, Liu XD, Yan TF, Wu LY, Alm E, Arkin A, Thompson DK, Zhou JZ. Global transcriptome analysis of the heat shock response of Shewanella oneidensis. Journal of Bacteriology, 2004, 186(22):7796-7803.
    [57] Frank S, Schmidt F, Klockgether J, Davenport CF, Gesell Salazar M, Völker U, Tümmler B. Functional genomics of the initial phase of cold adaptation of Pseudomonas putida KT2440. FEMS Microbiology Letters, 2011, 318(1):47-54.
    [58] Casanueva A, Tuffin M, Cary C, Cowan DA. Molecular adaptations to psychrophily:the impact of'omic'technologies. Trends in Microbiology, 2010, 18(8):374-381.
    [59] Dammel CS, Noller HF. Suppression of a cold-sensitive mutation in 16S rRNA by overexpression of a novel ribosome-binding factor,RbfA. Genes & Development, 1995, 9(5):626-637.
    [60] Brandi A, Piersimoni L, Feto NA, Spurio R, Alix JH, Schmidt F, Gualerzi CO. Translation initiation factor IF2 contributes to ribosome assembly and maturation during cold adaptation. Nucleic Acids Research, 2019, 47(9):4652-4662.
    [61] Deming JW. Psychrophiles and polar regions. Current Opinion in Microbiology, 2002, 5(3):301-309.
    [62] Chintalapati S, Kiran MD, Shivaji S. Role of membrane lipid fatty acids in cold adaptation. Cellular and Molecular Biology:Noisy Le Grand, France, 2004, 50(5):631-642.
    [63] Guan ZQ, Tian B, Perfumo A, Goldfine H. The polar lipids of Clostridium psychrophilum, an anaerobic psychrophile. Biochimica et Biophysica Acta:BBA-Molecular and Cell Biology of Lipids, 2013, 1831(6):1108-1112.
    [64] Nichols DS, Miller MR, Davies NW, Goodchild A, Raftery M, Cavicchioli R. Cold adaptation in the Antarctic archaeon Methanococcoides burtonii involves membrane lipid unsaturation. Journal of Bacteriology, 2004, 186(24):8508-8515.
    [65] Lauro FM, Tran K, Vezzi A, Vitulo N, Valle G, Bartlett DH. Large-scale transposon mutagenesis of Photobacterium profundum SS9 reveals new genetic loci important for growth at low temperature and high pressure. Journal of Bacteriology, 2008, 190(5):1699-1709.
    [66] Cacace G, Mazzeo MF, Sorrentino A, Spada V, Malorni A, Siciliano RA. Proteomics for the elucidation of cold adaptation mechanisms in Listeria monocytogenes. Journal of Proteomics, 2010, 73(10):2021-2030.
    [67] Bakermans C, Tollaksen SL, Giometti CS, Wilkerson C, Tiedje JM, Thomashow MF. Proteomic analysis of Psychrobacter cryohalolentis K5 during growth at subzero temperatures. Extremophiles:Life Under Extreme Conditions, 2007, 11(2):343-354.
    [68] Durack J, Ross T, Bowman JP. Characterisation of the transcriptomes of genetically diverse Listeria monocytogenes exposed to hyperosmotic and low temperature conditions reveal global stress-adaptation mechanisms. PLoS One, 2013, 8(9):e73603.
    [69] Dieser M, Greenwood M, Foreman CM. Carotenoid pigmentation in Antarctic heterotrophic bacteria as a strategy to withstand environmental stresses. Arctic, Antarctic, and Alpine Research, 2010, 42(4):396-405.
    [70] Chattopadhyay M?椀渀?瀀爀漀琀攀椀渀猀???漀甀爀渀愀氀?漀昀?倀爀漀琀攀漀洀椀挀猀??? ??????????????戀爀?嬀??崀??愀氀最愀渀搀?????漀甀??????攀????????瘀椀渀????圀攀椀欀?????愀搀攀爀渀????刀攀洀漀琀攀?漀砀椀搀愀琀椀瘀攀?洀漀搀椀昀椀挀愀琀椀漀渀猀?椀渀搀甀挀攀搀?戀礀?漀砀礀最攀渀?昀爀攀攀?爀愀搀椀挀愀氀猀?洀漀搀椀昀礀?吀?刀?愀氀氀漀猀琀攀爀椀挀?攀焀甀椀氀椀戀爀椀甀洀?漀昀?愀?栀礀瀀攀爀琀栀攀爀洀漀瀀栀椀氀椀挀?氀愀挀琀愀琀攀?搀攀栀礀搀爀漀最攀渀愀猀攀???漀甀爀渀愀氀?漀昀?匀琀爀甀挀琀甀爀愀氀??椀漀氀漀最礀??? ? ???? ????? ??????戀爀?嬀? 崀??戀愀爀愀?匀??匀栀椀最攀洀漀爀椀?夀???氀欀愀氀椀?琀漀氀攀爀愀渀琀?栀椀最栀?愀挀琀椀瘀椀琀礀?挀愀琀愀氀愀猀攀?昀爀漀洀?愀?琀栀攀爀洀漀瀀栀椀氀椀挀?戀愀挀琀攀爀椀甀洀?愀渀搀?椀琀猀?漀瘀攀爀攀砀瀀爀攀猀猀椀漀渀?椀渀??猀挀栀攀爀椀挀栀椀愀?挀漀氀椀??倀爀漀琀攀椀渀??砀瀀爀攀猀猀椀漀渀?愀渀搀?倀甀爀椀昀椀挀愀琀椀漀渀???  ??????????????? ??戀爀?嬀??崀??漀甀猀琀愀昀愀??????愀椀渀?一??匀爀椀爀愀渀最愀渀愀琀栀愀渀?一??嘀攀洀甀氀愀瀀愀氀氀椀?刀???搀攀渀琀椀昀椀挀愀琀椀漀渀?漀昀?愀?猀椀渀最氀攀?渀甀挀氀攀漀琀椀搀攀?椀渀猀攀爀琀椀漀渀?椀渀?琀栀攀?瀀爀漀洀漀琀攀爀?爀攀最椀漀渀?愀昀昀攀挀琀椀渀最?琀栀攀?猀漀搀??瀀爀漀洀漀琀攀爀?愀挀琀椀瘀椀琀礀?椀渀??爀甀挀攀氀氀愀?渀攀漀琀漀洀愀攀??倀?漀匀?伀渀攀??? ? ????????攀???????戀爀?嬀??崀??愀氀氀愀氀?????愀渀渀愀??????漀渀琀爀漀氀?漀昀?琀栀椀漀爀攀搀漀砀椀渀?爀攀搀甀挀琀愀猀攀?最攀渀攀??琀爀砀???琀爀愀渀猀挀爀椀瀀琀椀漀渀?戀礀?匀愀爀??椀渀?匀琀愀瀀栀礀氀漀挀漀挀挀甀猀?愀甀爀攀甀猀???漀甀爀渀愀氀?漀昀??愀挀琀攀爀椀漀氀漀最礀??? ? ??????????????????戀爀?嬀??崀????????搀椀最甀攀?????爀椀渀????倀愀猀挀愀氀?????愀爀戀攀?嘀???攀爀渀猀攀氀?????攀爀琀椀渀?倀一???栀攀甀渀最?????爀甀瘀攀椀氀氀攀爀?匀?????洀椀挀漀?匀???甀椀氀椀漀?????愀渀最?????攀氀氀攀爀?????漀?????愀渀最攀渀漀琀?匀???愀爀椀渀漀????一椀氀猀猀漀渀????倀愀爀爀椀氀氀椀????刀漀挀栀愀??倀???刀漀甀礀?娀??匀攀欀漀眀猀欀愀????吀甀琀椀渀漀?????嘀愀氀氀攀渀攀琀????嘀漀渀??攀椀樀渀攀?????愀渀挀栀椀渀?????漀瀀椀渀最?眀椀琀栀?挀漀氀搀?琀栀攀?最攀渀漀洀攀?漀昀?琀栀攀?瘀攀爀猀愀琀椀氀攀?洀愀爀椀渀攀??渀琀愀爀挀琀椀挀愀?戀愀挀琀攀爀椀甀洀?倀猀攀甀搀漀愀氀琀攀爀漀洀漀渀愀猀?栀愀氀漀瀀氀愀渀欀琀椀猀?吀????????攀渀漀洀攀?刀攀猀攀愀爀挀栀??? 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雷婷婷,陈良仲,陈绍兴,沈亮. 微生物对低温极端环境适应性的研究进展[J]. 微生物学报, 2022, 62(6): 2150-2164

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  • 收稿日期:2021-10-24
  • 最后修改日期:2022-01-15
  • 在线发布日期: 2022-06-13
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