Lager啤酒酵母起源的历史足迹及基因组学研究
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摘要
Lager酵母(Saccharomyces pastorianus)是由酿酒酵母(S. cerevisiae)与真贝氏酵母(S. eubayanus)通过自然杂交获得的异源多倍体菌株。Lager酵母保留了来自于两个亲本的染色体,同时也产生了新的重组片段,这就赋予Lager酵母适合低温发酵、麦芽三糖利用率高等重要特性。根据菌株的染色体含量,可以将Lager酵母分为两个类型:GroupⅠ型(Saaz型)与GroupⅡ型(Frohberg型)。近年来,多株Lager酵母菌株陆续完成了全基因组测序,为探索不同类型Lager酵母起源提供了基因组学基础。该文综述了Lager酵母的起源及基因组信息,并讨论了不同类型Lager酵母进化的模型。
Lager yeasts,which belong to the species of Saccharomyces pastorianus,are an interspecific hybrid of Saccharomyces cerevisiae and Saccharomyces eubayanus. Lager yeasts retain the chromosomes from both parental species and also include sets of novel hybrid chromosomes,resulting in their strong fermentative ability with cold tolerance and good maltotriose utilizing ability. Lager yeasts can be divided into two groups: Group I( Saaz) and and Group II( Frohberg),based on their S. cerevisiae: S. eubayanus gene content and the types and numbers of hybrid chromosomes. As whole genome sequences for several Lager yeasts have become available recently,this paper summarized the initial findings on this hybrid nature and their available genome data,and discussed the likely origins of the parental species that gave rise to S. pastorianus and considered several evolutionary models. This review lays a genomic foundation for exploring the origins of different types of Lager yeast.
Lager yeasts,which belong to the species of Saccharomyces pastorianus,are an interspecific hybrid of Saccharomyces cerevisiae and Saccharomyces eubayanus. Lager yeasts retain the chromosomes from both parental species and also include sets of novel hybrid chromosomes,resulting in their strong fermentative ability with cold tolerance and good maltotriose utilizing ability. Lager yeasts can be divided into two groups: Group I( Saaz) and and Group II( Frohberg),based on their S. cerevisiae: S. eubayanus gene content and the types and numbers of hybrid chromosomes. As whole genome sequences for several Lager yeasts have become available recently,this paper summarized the initial findings on this hybrid nature and their available genome data,and discussed the likely origins of the parental species that gave rise to S. pastorianus and considered several evolutionary models. This review lays a genomic foundation for exploring the origins of different types of Lager yeast.
引文
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[3] WANG J,LIU L,BALL T,et al. Revealing a 5,000-y-old beer recipe in China[J]. Proceedings of the National Academy of Sciences,2016,113:6 444-6 448.
[4] HORNSEY I S. A History of Beer and Brewing[M]. Royal Society of Chemistry,2003.
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[6] HORNSEY I S. Alcohol and its role in the evolution of human society[M]. Royal Society of Chemistry,2012.
[7] RUSCHE L N,NGUYEN H V,LEGRAS J L,et al. Deciphering the hybridisation history leading to the lager lineage based on the mosaic genomes of Saccharomyces bayanus strains NBRC1948 and CBS380T[J]. PLo S ONE,2011,6(10):e25821.
[8] DUNN B,SHERLOCK G. Reconstruction of the genome origins and evolution of the hybrid lager yeast Saccharomyces pastorianus[J]. Genome Res,2008,18:1 610-1 623.
[9] LEGRAS J L,Merdinoglu D,CORNUET J,et al. Bread,beer and wine:Saccharomyces cerevisiae diversity reflects human history[J]. Molecular Ecology,2007,16(10):2 091-2 102.
[10] MONERAWELAC,BOND U. Brewing up a storm:The genomes of lager yeasts and how they evolved[J]. Biotechnology Advances,2017,35(4):512-519.
[11] GALLONE B,STEENSELS J,PRAHL T,et al. Domestication and divergence of Saccharomyces cerevisiae beer yeasts[J]. Cell,2016,166(6):1 397-1 410.
[12] GONCALVES M,PONTES A,ALMEIDA P,et al. Distinct domestication trajectories in top-fermenting beer yeasts and wine yeasts[J]. Current Biology,2016,26(20):2 750-2 761.
[13] NILSSN T,PETERSEN J,HOLMBERG S,et al. Transfer of chromosome III during mediated cytoduction in yeast[J]. Carlsberg Res Commun,1980,45:113-117.
[14] TAMAI Y,MOMMA T,YOSHIMOTO H,et al. Co-existence of two types of chromosome in the bottom fermenting yeast,Saccharomyces pastorianus[J]. Yeast,1998,14(10):923-933.
[15] BOND U,NEAL C,DONNELLY D,et al. Aneuploidy and copy number breakpoints in the genome of lager yeasts mapped by microarray hybridisation[J]. Current Genetics,2004,45:360-370.
[16] NAKAO Y,KANAMORI T,ITOH T,et al. Genome sequence of the lager brewing yeast,an interspecies hybrid[J]. DNA Research,2009,16:115-129.
[17] RAINIERI S,KODAMA Y,KANEKO Y,et al. Pure and mixed genetic lines of Saccharomyces bayanus and Saccharomyces pastorianus and their contribution to the lager brewing strain genome[J]. Applied Environmental Microbiology,2006,72:3 968-3 974.
[18] LIBKIND D,HITTINGER C T,VALERIO E,et al. Microbe domestication and the identification of the wild genetic stock of lager-brewing yeast[J]. Proceedings of the National Academy of Sciences of the United States of America,2011,108(35):14 539-14 544.
[19] PERIS D,SYLVESTER K,LIBKIND D,et al. Population structure and reticulate evolution of Saccharomyces eubayanus and its lager-brewing hybrids[J]. Molecular Ecology,2014,23(8):2 031-2 045.
[20] PERIS D,LANGDON Q K,MORIARTY R V,et al.Complex ancestries of lager-brewing hybrids were shaped by standing variation in the wild yeast Saccharomyces eubayanus[J]. PLo S Genetics,2016,12(7):e1006155.
[21] RODRIGUEZ M E,PEREZ L,SANGORRIN M P,et al.Saccharomyces eubayanus and Saccharomyces uvarum associated with the fermentation of Araucaria araucana seeds in Patagonia[J]. FEMS Yeast Research,2014,14(6):948-965.
[22] BING J,HAN P J,LIU W Q,et al. Evidence for a Far East Asian origin of lager beer yeast[J]. Current Biology,2014,24(10):380-381.
[23] GAYEVSKIY V,GODDARD M R. Saccharomyces eubayanus and Saccharomyces arboricola reside in North Island native New Zealand forests[J]. Environmental Microbiology,2015,18(4):1 137-1 147
[24] GIBSON B R,STORGARDS E,KROGERUS K,et al.Comparative physiology and fermentation performance of Saaz and Frohberg lager yeast strains and the parental species Saccharomyces eubayanus[J]. Yeast,2013,30:255-266.
[25] WENDLAND J. Lager yeast comes of age[J]. Eukaryotic cell,2014,13(10):1 256-1 265.
[26] GALLONE B,MERTENS S,GORDON J L,et al. Origins,evolution,domestication and diversity of Saccharomyces beer yeasts[J]. Current Opinion in Biotechnology,2018,49:148-155.
[27] BAKER E C,WANG B,BELLORA N,et al. The genome sequence of Saccharomyces eubayanus and the domestication of lager-brewing yeasts[J]. Molecular Biology and Evolution,2015,32(11):2 818-2 831.
[28] MONERAWELA C,JAMES T C,WOLFE K H,et al.Loss of lager specific genes and subtelomeric regions define two different Saccharomyces cerevisiae lineages for Saccharomyces pastorianus GroupⅠandⅡstrains[J].FEMS Yeast Research,2015,15(2):fou008.
[29] LITI G,PERUFFO A,JAMES SA,et al. Inferences of evolutionary relationships from a population survey of LTR-retrotransposons and telomeric-associated sequences in the Saccharomyces sensu stricto complex[J]. Yeast,2005,22:177-192.
[30] WALTHER A,HESSELBART A,WENDLAND J. Genome sequence of Saccharomyces carlsbergensis, the world’s first pure culture lager yeast[J]. G3(Bethesda),2014,4:783-793.
[31] DE LEON-MEDINA P M,ELIZONDO R,DAMAS L C,et al. Genome annotation of a Saccharomyces sp. lager brewer's yeast[J]. Genome Data,2016,9:25-29.
[32] HEWITT S K,DONALDSON I,LOVELL S C,et al. Sequencing and characterisation of rearrangements in three S. pastorianus strains reveals the presence of chimeric genes and gives evidence of breakpoint reuse[J]. PLo S ONE,2013,9:e92203.
[33] VANDEN M,BOLAT I,NIJKAMP J F,et al. Chromosomal copy number variation in Saccharomyces pastorianus is evidence for extensive genome dynamics in industrial lager brewing strains[J]. Applied Environmental Microbiology,2015,81:6 253-6 267.
[34] OKUNO M,KAJITANI R,RYUSUI R,et al. Next-generation sequencing analysis of lager brewing yeast strains reveals the evolutionary history of interspecies hybridization[J]. DNA Research,2016,23:67-80.
[35] YAMAGISHI H,OHNUKI S,NOGAMI S,et al. Role of bottom-fermenting brewer’s yeast KEX2 in high temperature resistance and poor proliferation at low temperatures[J]. Journal of General and Applied Microbiology,2010,56:297-312.
[36] OGATA T,YAMAGISHI H,UKIBE K,et al. Construction of bottom-fermenting yeasts that over express the ubiquitin ligase RSP5 gene and their effects on high-gravity fermentation[J]. J Brew Dist,2012,3:1-5.
[37] YANG H,JIANJUN D,HUA Y,et al. Monitoring of the production of flavor compounds by analysis of the gene transcription involved in higher alcohol and ester formation by the brewer’s yeast Saccharomyces pastorianus using a multiplex RT-qPCR assay[J]. Journal of the Institute of Brewing,2014,120:119-126.
[38] COUSSEAU F E,ALVES S L,TRICHEZ D,et al..Characterization of maltotriose transporters from the Saccharomyces eubayanus subgenome of the hybrid Saccharomyces pastorianus lager brewing yeast strain Weihenstephan 34/70[J]. Letters of Applied Microbiology,2013,56:21-29.
[39] VIDGREN V,MULTANEN J P,RUOHONEN L,et al.The temperature dependence of maltose transport in ale and lager strains of brewer’s yeast[J]. FEMS Yeast Research,2010,10:402-411.
[40] VIDGREN V,LONDESBOROUGH J. Characterization of the Saccharomyces bayanus-type Agt1 transporter of lager yeast[J]. Journal of the Institute of Brewing,2012,118:148-151.
[41] BRIAN G,GIANNI L. Saccharomyces pastorianus:genomic insights inspiring innovation for industry[J].Yeast,2015,32(1):17-27.
[43] BLIEK L,TOYE G,DUMORTIER F,et al. Isolation and characterization of brewer’s yeast variants with improved fermentation performance under high gravity conditions[J]. Applied Environmental Microbiology,2007,73:815-824.
[44] HUUSKONEN A,MARKULA T,VIDGREN V,et al.Selection from industrial lager yeast strains of variants with improved fermentation performance in very-high-gravity worts[J]. Applied Environmental Microbiology,2010,76:1 563-1 573.
[45] YU Z,ZHAO H,LI H,et al. Selection of Saccharomyces pastorianus variants with improved fermentation performance under very high-gravity wort conditions[J]. Biotechnology Letters,2012,34:365-370.
[46] JAMES T C,USHER J,CAMPBELL S,et al. Lager yeasts possess dynamic genomes that undergo rearrangements and gene amplification in response to stress[J].Current Genetics,2008,53:139-152.
[47] EKBERG J,RAUTIO J,MATTINEN L,et al. Adaptive evolution of the lager brewing yeast Saccharomyces pastorianus for improved growth under hyperosmotic conditions and its influence on fermentation performance[J]. FEMS Yeast Research,2013,13(3):335-349.
[48] STREJC J,SIRISTOVA L,KARABIN M,et al. Production of alcoholfree beer with elevated amounts of flavouring compounds using lager yeast mutants[J]. Journal of the Institute of Brewing,2013,119:149-155.
[3] WANG J,LIU L,BALL T,et al. Revealing a 5,000-y-old beer recipe in China[J]. Proceedings of the National Academy of Sciences,2016,113:6 444-6 448.
[4] HORNSEY I S. A History of Beer and Brewing[M]. Royal Society of Chemistry,2003.
[5] BARNETT J A. A history of research on yeasts 1:Work by chemists and biologists 1789-1850[J]. Yeast,1998,14(16):1 439.
[6] HORNSEY I S. Alcohol and its role in the evolution of human society[M]. Royal Society of Chemistry,2012.
[7] RUSCHE L N,NGUYEN H V,LEGRAS J L,et al. Deciphering the hybridisation history leading to the lager lineage based on the mosaic genomes of Saccharomyces bayanus strains NBRC1948 and CBS380T[J]. PLo S ONE,2011,6(10):e25821.
[8] DUNN B,SHERLOCK G. Reconstruction of the genome origins and evolution of the hybrid lager yeast Saccharomyces pastorianus[J]. Genome Res,2008,18:1 610-1 623.
[9] LEGRAS J L,Merdinoglu D,CORNUET J,et al. Bread,beer and wine:Saccharomyces cerevisiae diversity reflects human history[J]. Molecular Ecology,2007,16(10):2 091-2 102.
[10] MONERAWELAC,BOND U. Brewing up a storm:The genomes of lager yeasts and how they evolved[J]. Biotechnology Advances,2017,35(4):512-519.
[11] GALLONE B,STEENSELS J,PRAHL T,et al. Domestication and divergence of Saccharomyces cerevisiae beer yeasts[J]. Cell,2016,166(6):1 397-1 410.
[12] GONCALVES M,PONTES A,ALMEIDA P,et al. Distinct domestication trajectories in top-fermenting beer yeasts and wine yeasts[J]. Current Biology,2016,26(20):2 750-2 761.
[13] NILSSN T,PETERSEN J,HOLMBERG S,et al. Transfer of chromosome III during mediated cytoduction in yeast[J]. Carlsberg Res Commun,1980,45:113-117.
[14] TAMAI Y,MOMMA T,YOSHIMOTO H,et al. Co-existence of two types of chromosome in the bottom fermenting yeast,Saccharomyces pastorianus[J]. Yeast,1998,14(10):923-933.
[15] BOND U,NEAL C,DONNELLY D,et al. Aneuploidy and copy number breakpoints in the genome of lager yeasts mapped by microarray hybridisation[J]. Current Genetics,2004,45:360-370.
[16] NAKAO Y,KANAMORI T,ITOH T,et al. Genome sequence of the lager brewing yeast,an interspecies hybrid[J]. DNA Research,2009,16:115-129.
[17] RAINIERI S,KODAMA Y,KANEKO Y,et al. Pure and mixed genetic lines of Saccharomyces bayanus and Saccharomyces pastorianus and their contribution to the lager brewing strain genome[J]. Applied Environmental Microbiology,2006,72:3 968-3 974.
[18] LIBKIND D,HITTINGER C T,VALERIO E,et al. Microbe domestication and the identification of the wild genetic stock of lager-brewing yeast[J]. Proceedings of the National Academy of Sciences of the United States of America,2011,108(35):14 539-14 544.
[19] PERIS D,SYLVESTER K,LIBKIND D,et al. Population structure and reticulate evolution of Saccharomyces eubayanus and its lager-brewing hybrids[J]. Molecular Ecology,2014,23(8):2 031-2 045.
[20] PERIS D,LANGDON Q K,MORIARTY R V,et al.Complex ancestries of lager-brewing hybrids were shaped by standing variation in the wild yeast Saccharomyces eubayanus[J]. PLo S Genetics,2016,12(7):e1006155.
[21] RODRIGUEZ M E,PEREZ L,SANGORRIN M P,et al.Saccharomyces eubayanus and Saccharomyces uvarum associated with the fermentation of Araucaria araucana seeds in Patagonia[J]. FEMS Yeast Research,2014,14(6):948-965.
[22] BING J,HAN P J,LIU W Q,et al. Evidence for a Far East Asian origin of lager beer yeast[J]. Current Biology,2014,24(10):380-381.
[23] GAYEVSKIY V,GODDARD M R. Saccharomyces eubayanus and Saccharomyces arboricola reside in North Island native New Zealand forests[J]. Environmental Microbiology,2015,18(4):1 137-1 147
[24] GIBSON B R,STORGARDS E,KROGERUS K,et al.Comparative physiology and fermentation performance of Saaz and Frohberg lager yeast strains and the parental species Saccharomyces eubayanus[J]. Yeast,2013,30:255-266.
[25] WENDLAND J. Lager yeast comes of age[J]. Eukaryotic cell,2014,13(10):1 256-1 265.
[26] GALLONE B,MERTENS S,GORDON J L,et al. Origins,evolution,domestication and diversity of Saccharomyces beer yeasts[J]. Current Opinion in Biotechnology,2018,49:148-155.
[27] BAKER E C,WANG B,BELLORA N,et al. The genome sequence of Saccharomyces eubayanus and the domestication of lager-brewing yeasts[J]. Molecular Biology and Evolution,2015,32(11):2 818-2 831.
[28] MONERAWELA C,JAMES T C,WOLFE K H,et al.Loss of lager specific genes and subtelomeric regions define two different Saccharomyces cerevisiae lineages for Saccharomyces pastorianus GroupⅠandⅡstrains[J].FEMS Yeast Research,2015,15(2):fou008.
[29] LITI G,PERUFFO A,JAMES SA,et al. Inferences of evolutionary relationships from a population survey of LTR-retrotransposons and telomeric-associated sequences in the Saccharomyces sensu stricto complex[J]. Yeast,2005,22:177-192.
[30] WALTHER A,HESSELBART A,WENDLAND J. Genome sequence of Saccharomyces carlsbergensis, the world’s first pure culture lager yeast[J]. G3(Bethesda),2014,4:783-793.
[31] DE LEON-MEDINA P M,ELIZONDO R,DAMAS L C,et al. Genome annotation of a Saccharomyces sp. lager brewer's yeast[J]. Genome Data,2016,9:25-29.
[32] HEWITT S K,DONALDSON I,LOVELL S C,et al. Sequencing and characterisation of rearrangements in three S. pastorianus strains reveals the presence of chimeric genes and gives evidence of breakpoint reuse[J]. PLo S ONE,2013,9:e92203.
[33] VANDEN M,BOLAT I,NIJKAMP J F,et al. Chromosomal copy number variation in Saccharomyces pastorianus is evidence for extensive genome dynamics in industrial lager brewing strains[J]. Applied Environmental Microbiology,2015,81:6 253-6 267.
[34] OKUNO M,KAJITANI R,RYUSUI R,et al. Next-generation sequencing analysis of lager brewing yeast strains reveals the evolutionary history of interspecies hybridization[J]. DNA Research,2016,23:67-80.
[35] YAMAGISHI H,OHNUKI S,NOGAMI S,et al. Role of bottom-fermenting brewer’s yeast KEX2 in high temperature resistance and poor proliferation at low temperatures[J]. Journal of General and Applied Microbiology,2010,56:297-312.
[36] OGATA T,YAMAGISHI H,UKIBE K,et al. Construction of bottom-fermenting yeasts that over express the ubiquitin ligase RSP5 gene and their effects on high-gravity fermentation[J]. J Brew Dist,2012,3:1-5.
[37] YANG H,JIANJUN D,HUA Y,et al. Monitoring of the production of flavor compounds by analysis of the gene transcription involved in higher alcohol and ester formation by the brewer’s yeast Saccharomyces pastorianus using a multiplex RT-qPCR assay[J]. Journal of the Institute of Brewing,2014,120:119-126.
[38] COUSSEAU F E,ALVES S L,TRICHEZ D,et al..Characterization of maltotriose transporters from the Saccharomyces eubayanus subgenome of the hybrid Saccharomyces pastorianus lager brewing yeast strain Weihenstephan 34/70[J]. Letters of Applied Microbiology,2013,56:21-29.
[39] VIDGREN V,MULTANEN J P,RUOHONEN L,et al.The temperature dependence of maltose transport in ale and lager strains of brewer’s yeast[J]. FEMS Yeast Research,2010,10:402-411.
[40] VIDGREN V,LONDESBOROUGH J. Characterization of the Saccharomyces bayanus-type Agt1 transporter of lager yeast[J]. Journal of the Institute of Brewing,2012,118:148-151.
[41] BRIAN G,GIANNI L. Saccharomyces pastorianus:genomic insights inspiring innovation for industry[J].Yeast,2015,32(1):17-27.
[43] BLIEK L,TOYE G,DUMORTIER F,et al. Isolation and characterization of brewer’s yeast variants with improved fermentation performance under high gravity conditions[J]. Applied Environmental Microbiology,2007,73:815-824.
[44] HUUSKONEN A,MARKULA T,VIDGREN V,et al.Selection from industrial lager yeast strains of variants with improved fermentation performance in very-high-gravity worts[J]. Applied Environmental Microbiology,2010,76:1 563-1 573.
[45] YU Z,ZHAO H,LI H,et al. Selection of Saccharomyces pastorianus variants with improved fermentation performance under very high-gravity wort conditions[J]. Biotechnology Letters,2012,34:365-370.
[46] JAMES T C,USHER J,CAMPBELL S,et al. Lager yeasts possess dynamic genomes that undergo rearrangements and gene amplification in response to stress[J].Current Genetics,2008,53:139-152.
[47] EKBERG J,RAUTIO J,MATTINEN L,et al. Adaptive evolution of the lager brewing yeast Saccharomyces pastorianus for improved growth under hyperosmotic conditions and its influence on fermentation performance[J]. FEMS Yeast Research,2013,13(3):335-349.
[48] STREJC J,SIRISTOVA L,KARABIN M,et al. Production of alcoholfree beer with elevated amounts of flavouring compounds using lager yeast mutants[J]. Journal of the Institute of Brewing,2013,119:149-155.