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1 微生物学通报 OCT 20, 2009, 36(10): 1604~1608 Microbiology 专论与综述 2009 by Institute of Microbiology, CAS 酿酒酵母乙醇耐受性机理研究进展 * 张穗生黄日波周兴黎贞崇黄志民 ( ) 摘要 : 酿酒酵母 (Sacchromyces cerevisiae) 一直是主要的生物乙醇和酿酒业发酵菌株, 具有发酵速度快乙醇产量高特性然而, 产物乙醇积累造成的毒性效应是限制乙醇产量的主要因素之一, 研究酿酒酵母乙醇耐受性为解决这一工业难题奠定了理论基础本文从乙醇对酵母细胞生理细胞结构和组分的影响, 以及酿酒酵母乙醇耐受性的遗传基础方面综述了酿酒酵母乙醇耐受性机理的研究进展关键词 : 酿酒酵母, 乙醇耐受性, 机理 Advances in Research on the Mechanisms of Saccharomyces cerevisiae Ethanol Tolerance ZHANG Sui-Sheng * HUANG Ri-Bo ZHOU Xing LI Zhen-Chong HUANG Zhi-Min (Guangxi Academy of Sciences, Nanning, Guangxi , China) Abstract: The yeast Sacchromyces cerevisiae is most widely used for producing bioethanol in alcoholic industry due to its higher ethanol yield and fermentation rate. However, the toxic effect of accumulated ethanol is one of the main factors, which limit high ethanol production. Thus, investigating the mechanisms of yeast ethanol tolerance will provide the basis for solving the industrial problem. This article reviewed the mechanisms of Sacchromyces cerevisiae ethanol tolerance focusing on its cell physiological behaviors, structure and biochemical composition, as well as its genetic basis. Keywords: Saccharomyces cerevisiae, Ethanol tolerance, Mechanism,,, [1],,,, 基金项目 : (No ); (No. 08YJ16SW05) * 通讯作者 :Tel: ; : zss99@vip.163.com 收稿日期 : ; 接受日期 :

2 : 1605,,, [2], [3], 1 乙醇对酿酒酵母细胞生理的影响,,, :,,, [4] : ;,,, 2%~10%(V/V),, 5%~ 10%(V/V),,,,, ; ;, [5,6] 2 乙醇耐受性与酿酒酵母的细胞结构和组分相关,,, 2.1 线粒体结构完整性与酿酒酵母乙醇耐受性相关, [7], 2.2 细胞质膜组分变化与酿酒酵母乙醇耐受性相关,, ( ) ( ), ( ), : 1), ;, ( ), ( ) [8], [9] ; 2), [10] ; 3),, ; 4), / [7] H + -ATPase, H + -ATPase, ATP,,,, H + -ATPase, H + -ATPase, [8] 2.3 细胞内保护性物质变化与酿酒酵母乙醇耐受性相关 (Heat shock protein, HSP), HSP,,,,,

3 1606 微生物学通报 2009, Vol.36, No.10 HSP [11] ;,,1,1-,,,, [12,13] 3 酿酒酵母乙醇耐受性的遗传基础 DNA,, :,,,,, (HSP ), : 3.1 与酿酒酵母乙醇耐受性相关的信号转导有关基因,,, (Mitogen- activated protein kinase, MAPK), slt2p gpd1 hor2 gre3 dak1 hor7 [14] ; :, ume6 [6] msn2p ars1p [14] 3.2 与酿酒酵母乙醇耐受性相关的能量利用有关基因,, glk1 hxk1 ydr516c tdh1 ald4 cit2 mcr1 pyc1 [15], F- ATP atp14 atp18 atp19 ATP [16] ptk2 yor137c [17] ATP, 3- gpd1 3- hor2 dak1 [17] 3.3 与酿酒酵母乙醇耐受性相关的物质代谢与运输有关基因,,, rrl9 arpl32 kar4 yef3, nop1 nop5 [15] ;, ole1 opi3 faa1 eht1 [15], -1- ino1 [9] ;, mep2 dip5 hxt6 hxt7 [15] ccc2 ctr2 [17] clc1 vps34 [6] btn2 [18] rat8p [19] 3.4 与酿酒酵母乙醇耐受性相关的细胞形态建成有关基因, tdp3 [6] swe1 [20] bem2 pat1 rom2 ada2 [21], swe1 ; bem2 pat1 rom2 ada2, calcofluor white zymolyase [21], 3.5 与酿酒酵母乙醇耐受性相关的多种逆境应激有关基因,,, pro1 put1 car2 put4 mrna [13] ; tps1 tps2 [15] ; Hsp hsp26 hsp42 hsp30 hsp104 hsp12 hsp78 hsp70 ssa1 ssa2 ssa3 ssa4 sse1 hsp30 yro2 [15] ;,, sod1 [1] sod2 ctt1 ahp1 grx1 grx4

4 : 1607 ttr1 [15] mpr1 [22] 3.6 酿酒酵母乙醇耐受性相关基因表达的复杂性,,,,,,,, [15] ;, ATPase ptk2 yor137c, ATPase hsp30, [16], 4 研究展望 : [17], [23], [24] ;,,,, [25 27],, Alper (Global transcription machine engineering, gtme), spt15, [28],,, 参考文献 [1] Gibson BR, Lawrence SJ, Leclaire JP, et al. Yeast responses to stresses associated with industrial brewery handling. FEMS Microbiol Rev, 2007, 31(5): [2] Maiorella B, Wilke CR, Blanch HW. Alcohol production and recovery. Adv Biochem Eng, 1981, 20: [3] Taylor M, Tuffin M, Burton S, et al. Microbial responses to solvent and alcohol stress. Biotechnol J, 2008, 3(11): [4] Dinh TN, Nagahisa K, Hirasawa T, et al. Adaptation of Saccharomyces cerevisiae cells to high ethanol concentration and changes in fatty acid composition of membrane and cell size. PLoS ONE, 2008, 3(7): e2623. [5] Alexandre H, Charpentier C. Biochemical aspects of stuck and sluggish fermentation in grape must. J Ind Microbiol Biotech, 1998, 20: [6] Kumar GR, Goyashiki R, Ramakrishnan V, et al. Genes required for ethanol tolerance and utilization in saccharomyces cerevisiae. Am J Enol Vitic, 2008, 59(4): [7] Chi Z, Arneborg N. Relationship between lipid composition, frequency of ethanol-induced respiratory deficient mutants, and ethanol tolerance in Saccharomyces cerevisiae. J Appl Microbiol, 1999, 86(6): [8] Aguilera F, Peinado RA, Millán C, et al. Relationship between ethanol tolerance, H+-ATPase activity and the lipid composition of the plasma membrane in different wine yeast strains. Int J Food Microbiol, 2006, 110(1): [9] Furukawa K, Kitano H, Mizoguchi H, et al. Effect of cellular inositol content on ethanol tolerance of Saccharomyces cerevisiae in sake brewing. J Biosci Bioeng, 2004, 98(2): [10] Malhotra R, Singh B. Ethanol-induced changes in glycolipids of Saccharomyces cerevisiae. Appl Biochem Biotechnol, 2006, 128(3): [11] Sanchez Y, Taulien J, Borkovich KA, et al. Hspl04 is required for tolerance to many forms of stress. EMBO J, 1992, 11(6): [12] Singer MA, Lindquist S. Thermotolerance in Saccharomyces cerevisiae: the Yin and Yang of trehalose. Trends Biotechnol, 1998, 16(11): [13] Kaino T, Takagi H. Gene expression profiles and intracellular contents of stress protectants in Saccharomyces cerevisiae under ethanol and sorbitol stresses. Appl Microbiol Biotechnol, 2008, 79(2): [14] Van Voorst F, Houghton-Larsen J, JΦnson L, et al. Genome-wide identification of genes required for growth of Saccharomyces cerevisiae under ethanol stress. Yeast, 2006, 23(5): [15] Chandler M, Stanley GA, Rogers P, et al. A genomic approach to defining the ethanol stress response in the yeast Saccharomyces cerevisiae. Annals of Microbiology, 2004, 54(4): [16] Dinh TN, Nagahisa K, Yoshikawa K, et al. Analysis of adaptation to high ethanol concentration in Saccharomyces cerevisiae using DNA microarray. Bioprocess Biosyst Eng, 2009, DOI /s [17] Alexandre H, Ansanay-Galeote V, Dequin S, et al. Global gene expression during short-term ethanol stress in Saccharomyces cerevisiae. FEBS Lett, 2001, 498(1): [18] Espinazo-Romeu M, Cantoral JM, Matallana E, et al.

5 1608 微生物学通报 2009, Vol.36, No.10 Btn2p is involved in ethanol tolerance and biofilm formation in flor yeast. FEMS Yeast Res, 2008, 8(7): [19] Takemura R, Inoue Y, Izawa S. Stress response in yeast mrna export factor: reversible changes in Rat8p localization are caused by ethanol stress but not heat shock. J Cell Sci, 2004, 117(pt18): [20] Kubota S, Takeo I, Kume K, et al. Effect of ethanol on cell growth of budding yeast: genes that are important for cell growth in the presence of ethanol. Biosci Biotechnol Biochem, 2004, 68(4): [21] Takahashi T, Shimoi H, Ito K. Identification of genes required for growth under ethanol stress using transposon mutagenesis in Saccharomyces cerevisiae. Mol Genet Genomics, 2001, 265(6): [22] Du X, Takagi H. N-Acetyltransferase Mpr1 confers ethanol tolerance on Saccharomyces cerevisiae by reducing reactive oxygen species. Appl Microbiol Biotechnol, 2007, 75(6): [23] Hu XH, Wang MH, Tan T, et al. Genetic dissection of ethanol tolerance in the budding yeast Saccharomyces cerevisiae. Genetics, 2007, 175(3): [24] Lei J, Zhao X, Ge X, et al. Ethanol tolerance and the variation of plasma membrane composition of yeast floc populations with different size distribution. J Biotechnol, 2007, 131(3): [25] Takagi H, Takaoka M, Kawaguchi, et al. Effect of L-proline on sake brewing and ethanol stress in Saccharomyces cerevisiae. Appl Environ Microbiol, 2005, 71(12): [26] Hirasawa T, Yoshikawa K, Nakakura Y, et al. Identification of target genes conferring ethanol stress tolerance to Saccharomyces cerevisiae based on DNA microarray data analysis. J Biotechnol, 2007, 131(1): [27] Zhao XQ, Xue C, Ge XM, et al. Impact of zinc supplementation on the improvement of ethanol tolerance and yield. J Biotechnol, 2009, 139(1): [28] Alper H, Moxley J, Nevoigt E, et al. Engineering yeast transcription machinery for improved ethanol tolerance and production. Science, 2006, 314(5805): 编辑部公告关于微生物学通报专题刊申请的通知,,,,, 2008,,,,,,, 1. ; 2. () ;, tongbao@im.ac.cn, ; 3.,, tongbao@im.ac.cn

Microsoft Word - 793-797 tb20150504赵宏宇s-高校教改纵横.doc

Microsoft Word - 793-797 tb20150504赵宏宇s-高校教改纵横.doc 微生物学通报 Microbiology China tongbao@im.ac.cn Apr. 20, 2016, 43(4): 793 797 http://journals.im.ac.cn/wswxtbcn DOI: 10.13344/j.microbiol.china.150504 高校教改纵横生物工程专业发酵课程群建设探索 * 赵宏宇