Harvesting and Cropping Yeast: Flocculation and Centrifugation

  • Graham G. Stewart
Chapter
Part of the The Yeast Handbook book series (YEASTHDB)

Abstract

Brewers employ a number of methods to crop their yeast which varies depending on whether one is dealing with traditional ale top-cropping, accepted lager bottom-cropping and cylindroconical fermentation systems (also called a Nathan fermenter) where the yeast (ale or lager) is recovered from the cone (sometimes repitched cone to cone), a non-flocculent culture where the yeast, still in suspension, is cropped with a centrifuge or a portion of the yeast, still in suspension, is blended into the fresh wort of a subsequent fermentation. With the traditional ale top-cropping fermentation system, although there are many variations of this process, a single, dual or multi-strain culture can be employed. Yeast quality is influenced by the manner that yeast is cropped and centrifuges play an important part in this regard. The cell wall structure of S. cerevisiae is the critical parameter involved in yeast flocculation. Details of the cell wall have already been discussed in Chap.  5. However, here it is discussed that the wall consists of an inner layer composed predominantly of β-glucose and chitin and a fibrillar outer layer consisting primarily of α-mannan (highly glycosylated) associated with mannoproteins. Yeast management and handling systems, including culture harvesting, are influential in determining the physiological status of yeast.

References

  1. Abramova N, Sertil O, Mehta S, Lowry CV (2001) Reciprocal regulation of anaerobic and aerobic cell wall mannoprotein gene expression in Saccharomyces cerevisiae. J Bacteriol 183:2881–2887PubMedPubMedCentralCrossRefGoogle Scholar
  2. Alwine JC, Kemp DJ, Stark GR (1977) Methods for detection of specific RNAs in agarose gels by transfer to diazobenzyloxymethyl-paper and hybridization with DNA probes. Proc Natl Acad Sci U S A 74:5350–5354PubMedPubMedCentralCrossRefGoogle Scholar
  3. Amory DE, Genet MJ, Rouxhet PG (1988a) Application of XPS to the surface analysis of yeast cells. Surf Interface Anal 11:478–486CrossRefGoogle Scholar
  4. Amory DE, Rouxhet PG, Dufour JP (1988b) Flocculence of brewery yeasts and their surface properties: chemical composition, electrostatic charge and hydrophobicity. J Inst Brew 94:79–84CrossRefGoogle Scholar
  5. Armstrong K, Bendiak D (2007) PYF malt: practical brewery observations of fermentability. MBAA Tech Q 44:40–46Google Scholar
  6. Ashbee R, Bignell EM (eds) (2010) Pathogenic yeasts. The yeast handbook. Springer, BerlinGoogle Scholar
  7. Axcell BC, van Nierop S, Vundla W (2000) Malt induced premature yeast flocculation. MBAA Tech Q 37:501–504Google Scholar
  8. Baker DA, Kirsop BH (1972) Flocculation in Saccharomyces cerevisiae as influenced by wort composition and by actidione. J Inst Brew 78:454–458CrossRefGoogle Scholar
  9. Bamforth C (2009) Beer: tap into the art and science of brewing. Oxford University Press, OxfordGoogle Scholar
  10. Bamforth C (2017) Freshness: practical guides to beer quality. American Society of Brewing Chemists, Minneapolis, MNGoogle Scholar
  11. Bayly JC, Douglas LM, Pretorius IS, Bauer FF, Dranginis AM (2005) Characteristics of Flo11-dependent flocculation in Saccharomyces cerevisiae. FEMS Microbiol Lett 5:1151–1156Google Scholar
  12. Beavan MJ, Belki D, Stewart GG, Rose AH (1979) Changes in electrophoretic mobility and lytic enzyme activity associated with developments of flocculating ability in Saccharomyces cerevisiae. Can J Microbiol 25:888–895PubMedCrossRefGoogle Scholar
  13. Bester MC, Pretorius IS, Bauer FF (2006) The regulation of Saccharomyces cerevisiae FLO gene expression and Ca 2+-dependent flocculation by Flo8p and Mss11p. Curr Genet 49:375–383PubMedCrossRefGoogle Scholar
  14. Bhattacharyya MK, Lustig AJ (2006) Telomere dynamics in genome stability. Trends Biochem Sci 31:114–122PubMedCrossRefGoogle Scholar
  15. Bing J, Han PJ, Liu WQ, Wang QM, Bai FY (2014) Evidence for a Far East Asian origin of lager beer yeast. Curr Biol 24:R380–R381PubMedCrossRefGoogle Scholar
  16. Bird A (2002) DNA methylation patterns and epigenetic memory. Genes Dev 16:6–21PubMedCrossRefGoogle Scholar
  17. Botstein D, Fink GR (2011) Yeast: an experimental organism for 21st century biology. Genetics 189:695–704PubMedPubMedCentralCrossRefGoogle Scholar
  18. Boulton C (2011) Yeast handling. Brew Dist Int 7:7–10Google Scholar
  19. Boulton C (2012) Advances in analytical methodology in brewing. J Inst Brew 118:255–263; MBAA Tech Q 50:53–61Google Scholar
  20. Boulton C, Quain E (eds) (2001) Brewing yeast. In: Brewing yeast and fermentation. Blackwell Science, OxfordGoogle Scholar
  21. Briggs DE, McGuinness G (1992) Microbes on barley grains. J Inst Brew 98:249–255Google Scholar
  22. Calleja GB (1984) Microbial aggregation. CRC Press, Boca Raton, FLGoogle Scholar
  23. Calleja GB (1987) Cell aggregation. In: Rose AH (ed) The yeasts, vol 2. Academic, London, pp 165–237Google Scholar
  24. Chen EH, Grote E, Mohler W, Vignery A (2007) Cell–cell fusion. FEBS Lett 581:2181–2193PubMedCrossRefGoogle Scholar
  25. Chlup PH, Stewart GG (2011) Centrifuges in brewing. MBAA Tech Q 48:48–50Google Scholar
  26. Chlup PH, Bernard D, Stewart GG (2007a) The disc stack centrifuge and its impact on yeast and beer quality. J Am Soc Brew Chem 65:29–37Google Scholar
  27. Chlup PH, Conery J, Stewart GG (2007b) Detection of mannan from Saccharomyces cerevisiae by flow cytometry. J Am Soc Brew Chem 65:151–155Google Scholar
  28. Chlup PH, Wang T, Lee EG, Stewart GG (2007c) Assessment of the physiological status of yeast during high-and low-gravity wort fermentations determined by flow cytometry. MBAA Tech Q 44:286–295Google Scholar
  29. Chlup PH, Bernard D, Stewart GG (2008) Disc stack centrifuge operating parameters and their impact on yeast physiology. J Inst Brew 114:45–61CrossRefGoogle Scholar
  30. Claro FB, Rijsbrack K, Soares EV (2007) Flocculation onset in Saccharomyces cerevisiae: effect of ethanol, heat and osmotic stress. J Appl Microbiol 102:693–700PubMedCrossRefGoogle Scholar
  31. Cooper DJ, Stewart GG, Bryce JH (2000) Yeast proteolytic activity during high and low gravity wort fermentations and its effect on head retention. J Inst Brew 106:197–201CrossRefGoogle Scholar
  32. Cormack B (2004) Can you adhere me now? Good. Cell 116:353–354PubMedCrossRefGoogle Scholar
  33. D’Amore T, Panchal CJ, Stewart GG (1988) Intracellular ethanol accumulation in Saccharomyces cerevisiae during fermentation. J Appl Environ Microbiol 54:1471–1510Google Scholar
  34. Damas-Buenrostro LC, Gracia-González G, Hernández-Luna CE, Galán-Wong LJ, Pereyra-Alférez B, Sierra-Benavides JA (2008) Detection of FLO genes in lager and wild yeast strains. J Am Soc Brew Chem 66:184–187Google Scholar
  35. Day AW, Poon NH, Stewart GG (1975) Fungal fimbriae. III. The effect of flocculation in Saccharomyces. Can J Microbiol 21:558–564PubMedCrossRefGoogle Scholar
  36. Dengis PB, Nélissen LR, Rouxhet PG (1995) Mechanisms of yeast flocculation: comparison of top- and bottom-fermenting strains. Appl Environ Microbiol 61:718–728PubMedPubMedCentralGoogle Scholar
  37. Dietvorst J, Brandt A (2008) Flocculation in Saccharomyces cerevisiae is repressed by the COMPASS methylation complex during high-gravity fermentation. Yeast 25:891–901PubMedCrossRefGoogle Scholar
  38. Dudbridge M (2011) Handbook of lean manufacturing in the food industry. Wiley, New York, NYCrossRefGoogle Scholar
  39. Dunn B, Sherlock G (2008) Reconstruction of the genome origins and evolution of the hybrid lager yeast Saccharomyces pastorianus. Genome Res 18:1610–1623PubMedPubMedCentralCrossRefGoogle Scholar
  40. Eddy AA (1958) Composite nature of the flocculation process of top and bottom strains of Saccharomyces. J Inst Brew 64:143–151CrossRefGoogle Scholar
  41. Eddy AA, Rudin AD (1958) Part of the yeast surface apparently involved in flocculation. J Inst Brew 64:19–21CrossRefGoogle Scholar
  42. Fidalgo M, Barrales RR, Jimenez J (2008) Coding repeat instability in the FLO11 gene of Saccharomyces yeasts. Yeast 25:879–889PubMedCrossRefGoogle Scholar
  43. Fink SL, Cookson BT (2005) Apoptosis, pyroptosis, and necrosis: mechanistic description of dead and dying eukaryotic cells. Infect Immun 73:1907–1916PubMedPubMedCentralCrossRefGoogle Scholar
  44. Gibson BR (2011) 125th anniversary review: improvement of higher gravity brewery fermentation via wort enrichment and supplementation. J Inst Brew 117:268–284CrossRefGoogle Scholar
  45. Gilliland RB (1951) The flocculation characteristics of brewing yeasts during fermentation. Proceedings of the European Brewery Convention Congress, Brighton, pp 35–58Google Scholar
  46. Gilliland RB (1959) Determination of yeast viability. J Inst Brew 65:424CrossRefGoogle Scholar
  47. Gimeno CJ, Ljungdahl PO, Styles CA, Fink GR (1992) Unipolar cell divisions in the yeast S. cerevisiae lead to filamentous growth: regulation by starvation and RAS. Cell 68:1077–1090PubMedCrossRefGoogle Scholar
  48. Goffeau A, Barrell BG, Bussey H, Oliver SG (1996) Life with 6000 genes. Science 274(5287):546, 563–546, 567CrossRefGoogle Scholar
  49. Goldstein IJ, Poretz RD (1986) Isolation, physicochemical characterization and carbohydrate-binding specificity of lectins. In: Liener IE, Sharon N, Goldstein IJ (eds) The lectins. Academic, Orlando, FL, p 52Google Scholar
  50. Gouveia C, Soares EV (2004) Pb2+ inhibits competitively flocculation of Saccharomyces cerevisiae. J Inst Brew 110:141–145CrossRefGoogle Scholar
  51. Govender P, Domingo JL, Bester MC, Pretorius IS, Bauer FF (2008) Controlled expression of the dominant flocculation genes FLO1, FLO5, and FLO11 in Saccharomyces cerevisiae. Appl Environ Microbiol 74:6041–6052PubMedPubMedCentralCrossRefGoogle Scholar
  52. Guinard J-X, Lewis MJ (1993) Study of the phenomenon by agglomeration in the yeast Saccharomyces cerevisiae. J Inst Brew 99:487–503CrossRefGoogle Scholar
  53. Guo B, Styles CA, Feng Q, Fink G (2000) A Saccharomyces gene family involved in invasive growth, cell–cell adhesion, and mating. Proc Natl Acad Sci U S A 97:12158–12163PubMedPubMedCentralCrossRefGoogle Scholar
  54. Halina S, Nathan L (2007) Lectins. Springer, NetherlandsGoogle Scholar
  55. Halme A, Bumgarner S, Styles C, Fink GR (2004) Genetic and epigenetic regulation of the FLO gene family generates cell–surface variation in yeast. Cell 116:405–415PubMedCrossRefGoogle Scholar
  56. Heine F, Stahl F, Sträuber H, Wiacek C, Benndorf D, Repenning C, Schmidt F, Scheper T, von Bergen M, Harms H, Müller S (2009) Prediction of flocculation ability of brewing yeast inoculates by flow cytometry, proteome analysis and mRNA profiling. Cytometry A 75:140–147PubMedCrossRefGoogle Scholar
  57. Helm E, Nohr B, Thome RSW (1953) Measurement of yeast flocculation and its significance in brewing. Wallerstein Laboratory Communications 16:315–326Google Scholar
  58. Hough JS (1959) Flocculation characteristics of strains present in some typical British pitching yeasts. J Inst Brew 65:479–482CrossRefGoogle Scholar
  59. Hoyer LL (2001) The ALS gene family of Candida albicans. Trends Microbiol 9:176–180PubMedCrossRefGoogle Scholar
  60. Hoyer LL, Green CB, Oh SH, Zhao X (2008) Discovering the secrets of the Candida albicans agglutinin-like sequence (ALS) gene family—a sticky pursuit. Med Mycol 46:1–15PubMedPubMedCentralCrossRefGoogle Scholar
  61. Hutter K-J, Miedl M, Kushmann B, Nitzsche F, Bryce JH, Stewart GG (2005) Detection of proteinases in Saccharomyces cerevisiae by flow cytometry. J Inst Brew 111:26–32CrossRefGoogle Scholar
  62. Inagaki H, Yamazumi K, Uehara H, Mochzuki K (1994) Determination of fermentation behaviour-malt evaluation system based on the original small scale fermentation test. Eur Brew Conv 23:111–136Google Scholar
  63. Ishimaru S, Kudo S, Hattan M, Yoshida T, Kataoka J (1967) Selection of small vessels for fermentation tests in the laboratory. Rep Res Lab Kirin Brew Co 10:61–65Google Scholar
  64. Jarvis P, Jefferson B, Parsons SA (2005) Measuring flocstructural characteristics. Environ Sci Biotechnol 4:1–18CrossRefGoogle Scholar
  65. Jibiki M, Ishibiki T, Yuuki T, Kagami N (2001) Application of polymerase chain reaction to determine the flocculation properties of brewer’s lager yeast. J Am Soc Brew 59:107–110Google Scholar
  66. Jibiki M, Sasaki K, Kagami N, Kawatsura K (2006) Application of a newly developed method for estimating the premature yeast flocculation potential of malt samples. J Am Soc Brew Chem 64:79–85Google Scholar
  67. Jin Y, Speers RA (2000) Effect of environmental conditions on the flocculation of Saccharomyces cerevisiae. J Am Soc Brew Chem 58:108–116Google Scholar
  68. Jin Y, Ritcey LL, Speers RA (2001) Effect of cell surface hydrophobicity, charge, and zymolectin density on the flocculation of Saccharomyces cerevisiae. J Am Soc Brew Chem 59:1–9Google Scholar
  69. Kaur R, Domergue R, Zupancic M, Cormack BP (2005) A yeast by any other name: Candida glabrata and its interaction with the host. Curr Opin Microbiol 8:378–384PubMedCrossRefGoogle Scholar
  70. Kida K, Yamadaki M, Asno S, Nakata T, Sonoda Y (1989) The effect of aeration on stability of continuous ethanol fermentation by a flocculating yeast. J Ferment Bioeng 68:107–111CrossRefGoogle Scholar
  71. Kihn JC, Masy CL, Mestdagh MM (1988a) Yeast flocculation: competition between nonspecific repulsion and specific bonding in cell adhesion. Can J Microbiol 34:773–778PubMedCrossRefGoogle Scholar
  72. Kihn JC, Masy CL, Mestdagh MM, Rouxhet PG (1988b) Yeast flocculation: factors affecting the measurement of flocculence. Can J Microbiol 34:779–781PubMedCrossRefGoogle Scholar
  73. Klis FM, Boorsma A, De Groot PWJ (2006) Cell wall construction in Saccharomyces cerevisiae. Yeast 23:185–202PubMedCrossRefGoogle Scholar
  74. Kobayashi O, Hayashi N, Kuroki R, Sone H (1998) Region of Flo1 proteins responsible for sugar recognition. J Bacteriol 180:6503–6510PubMedPubMedCentralGoogle Scholar
  75. Kock JLF, Venter P, Smith DP, Van Wyk PWJ, Botes PJ, Coetzee DJ, Pohl CH, Botha A, Ridel K-H, Nigam S (2000) A novel oxylipin-associated ‘ghosting’ phenomenon in yeast flocculation. Antonie Van Leeuwenhoek 77:401–406PubMedCrossRefGoogle Scholar
  76. Koizumi H, Ogawa T (2005) Rapid and sensitive method to measure premature yeast flocculation activity in malt. J Am Soc Brew Chem 63:147–150Google Scholar
  77. Krogerus A, Gibson BR (2013) 125th anniversary review: diacetyl and its control during brewery fermentation. J Inst Brew 119:86–97Google Scholar
  78. Kruger L, Ryder DS, Alcock C, Murray JP (1982) Malt quality: prediction of malt fermentability. Part I. Tech Q Master Brew Assoc Am 19:45–51Google Scholar
  79. Kukuruzinska MA, Bergh MLE, Jackson BJ (1987) Protein glycosylation in yeast. Annu Rev Biochem 56:915–944PubMedCrossRefGoogle Scholar
  80. Kuřec M, Baszczyňski M, Lehnert R, Brányik T (2009) Flow cytometry for age assessment of a yeast population and its application in beer fermentations. J Inst Brew 115:253–258CrossRefGoogle Scholar
  81. Lake JC, Speers RA (2008) A discussion of malt-induced premature yeast flocculation. MBAA Tech Q 4:253–262Google Scholar
  82. Lake JC, Speers RA, Porter AV, Gill TA (2008) Miniaturizing the fermentation assay: effect of fermentor size and fermentation kinetics on detection of premature yeast flocculation. J Am Soc Brew Chem 66:94–102Google Scholar
  83. Lange C, Nett JH, Trumpower BL, Hunte C (2001) Specific roles of protein-phospholipid interactions in the yeast cytochrome bc1 complex structure. EMBO J 20:6591–6600PubMedPubMedCentralCrossRefGoogle Scholar
  84. Leiper KA, Miedl M (2009) Colloidal stability of beer. In: Bamforth CW, Russell I, Stewart GG (eds) Beer: a quality perspective. Boston, MA, Elsevier, pp 111–161CrossRefGoogle Scholar
  85. Lewis MJ, Poerwantaro WM (1991) Release of haze material from the cell walls of agitated yeast. J Am Soc Brew Chem 49:43–46Google Scholar
  86. Lewis CW, Johnston JR, Martin PA (1976) The genetics of yeast flocculation. J Inst Brew 82:158–160CrossRefGoogle Scholar
  87. Libkind D, Hittinger CT, Valério E, Gonçalves C, Dover J, Johnston M, Gonçalves P, Sampaio JP (2011) Microbe domestication and the identification of the wild genetic stock of lager-brewing yeast. Proc Natl Acad Sci U S A 108:14539–14544PubMedPubMedCentralCrossRefGoogle Scholar
  88. Lin CH, MacGurn JA, Chu T, Stefan CJ, Emr SD (2008) Arrestin-related ubiquitin-ligase adaptors regulate endocytosis and protein turnover at the cell surface. Cell 135:714–725PubMedCrossRefGoogle Scholar
  89. Loney ER, Inglis PW, Sharp S, Pryde FE, Kent NA, Mellor J, Louis EJ (2009) Repressive and non-repressive chromatin at native telomeres in Saccharomyces cerevisiae. Epigenetics Chromatin 2:18PubMedPubMedCentralCrossRefGoogle Scholar
  90. Lorenz RT, Parks LW (1991) Involvement of heme components in sterol metabolism of Saccharomyces cerevisiae. Lipids 26:598–603PubMedCrossRefGoogle Scholar
  91. Lyons TP, Hough JS (1970a) Flocculation of brewer’s yeast. J Inst Brew 76:564–571CrossRefGoogle Scholar
  92. Lyons TP, Hough JS (1970b) The role of yeast cell walls in brewing. Brew Digest 45:52–60Google Scholar
  93. Machado MD, Santos MSF, Gouveia C, Soares HMVM, Soares EV (2008) Removal of heavy metals using a brewer’s yeast strain of Saccharomyces cerevisiae: the flocculation as a separation process. Bioresour Technol 99:2107–2115PubMedCrossRefGoogle Scholar
  94. Mamvura TA, Paterson AE, Fanucchi D (2017) The impact of pipe geometry variations on hygiene and success of orbital welding of brewing industry equipment. J Inst Brew 123:81–97CrossRefGoogle Scholar
  95. Masy CL, Henquinet A, Mestdagh MM (1992) Flocculation of Saccharomyces cerevisiae: inhibition by sugars. Can J Microbiol 38:1298–1306PubMedCrossRefGoogle Scholar
  96. Meaden PG (1996) DNA fingerprinting of brewer’s yeast. Ferment:9267–9272Google Scholar
  97. Miedl M, Stewart GG, Bryce JH, Kuchman B, Hutter K-H (2005) A novel procedure for the determining yeast pitching rates employing flow cytometry. In: Proceedings of the 29th European Brewery Convention Congress, Prague, CD Paper No 33Google Scholar
  98. Miki BLA, Poon NH, James AP, Seligy VL (1982a) Possible mechanism for flocculation interactions governed by gene FLO1 in Saccharomyces cerevisiae. J Bacteriol 150:878–889PubMedPubMedCentralGoogle Scholar
  99. Miki BLA, Poon NH, Seligy VL (1982b) Repression and induction of flocculation interactions in Saccharomyces cerevisiae. J Bacteriol 150:890–899PubMedPubMedCentralGoogle Scholar
  100. Miller T, Krogan NJ, Dover J (2001) COMPASS: A complex of proteins associated with a trithorax-related SET domain protein. Proc Natl Acad Sci U S A 98:12902–12907PubMedPubMedCentralCrossRefGoogle Scholar
  101. Mortier A, Soares EV (2007) Separation of yeasts by addition of flocculent cells of Saccharomyces cerevisiae. World J Microbiol Biotechnol 23:1401–1407CrossRefGoogle Scholar
  102. Mundy RD, Cormack B (2009) Expression of Candida glabrata adhesions following exposure to chemical preservatives. J Infect Dis 199:1891–1898PubMedPubMedCentralCrossRefGoogle Scholar
  103. Nakao Y, Kanamori T, Itoh T, Kodama Y, Rainieri S, Nakamura N, Shimonaga T, Hattori M, Ashikari T (2009) Genome sequence of the lager brewing yeast, an interspecies hybrid. DNA Res 16:115–129PubMedPubMedCentralCrossRefGoogle Scholar
  104. Nathan L (1930) Improvements in the fermentation and maturation of beers. J Inst Brew 36:538–544CrossRefGoogle Scholar
  105. Nayar A, Walker G, Wardrob F, Adya A (2017) Flocculation in industrial strains of Saccharomyces cerevisiae: role of cell wall polysaccharides and lectin-like receptors. J Inst Brew 123:211–218CrossRefGoogle Scholar
  106. Nishihara H, Toraya T, Fukui S (1982) Flocculation of cell-walls of brewers-yeast and effects of metal-ions, protein-denaturants and enzyme treatments. Arch Microbiol 131:112–115CrossRefGoogle Scholar
  107. Nishihara H, Kio K, Imamura M (2000) Possible mechanism of co-flocculation between non-flocculent yeasts. J Inst Brew 106:7–10CrossRefGoogle Scholar
  108. Ogata T, Izumikawa M, Kohno K, Shibata K (2008) Chromosomal location of Lg-FLO1 in bottom-fermenting yeast and the FLO5 locus of industrial yeast. J Appl Microbiol 105:1186–1198PubMedCrossRefGoogle Scholar
  109. Ogur M, St. John R (1956) A differential and diagnostic plating method for population studies of respiration deficiency in yeast. J Bacteriol 72:500–504PubMedPubMedCentralGoogle Scholar
  110. Ogur M, St. John R, Nagai S (1957) Tetrazolium overlay technique for population studies of respiration deficiency in yeast. Science 125:928–929PubMedCrossRefGoogle Scholar
  111. Osumi M (2012) Visualization of yeast cells by electron microscopy. J Electron Microsc 61:343–365Google Scholar
  112. Panchal CJ, Whitney GK, Stewart GG (1984a) Susceptibility of Saccharomyces spp. and Schwanniomyces spp. to the aminoglycoside antibiotic G418. Appl Environ Microbiol 47:1164–1166PubMedPubMedCentralGoogle Scholar
  113. Panchal CJ, Russell I, Sills AM, Stewart GG (1984b) Fermentation ethanol production – application of the new genetics to an ancient art. In: Proceedings of the 11th energy technology conference, Washington, DC, pp 1270–1273Google Scholar
  114. Panchal CJ, Russell I, Sills AM, Stewart GG (1984c) Genetic manipulation of brewing and related yeast strains. Food Technol 111:99–106Google Scholar
  115. Paula L, Birrer F (2006) Including public perspectives in industrial biotechnology and the biobased economy. J Agric Environ Ethics 19:253–267PubMedCrossRefGoogle Scholar
  116. Peng X, Sun J, Iserentant D, Michiels C, Verachtert H (2001) Flocculation and coflocculation of bacteria by yeasts. Appl Microbiol Biotechnol 55:777–781PubMedCrossRefGoogle Scholar
  117. Pomper S, Burkholder PR (1949) Studies on the biochemical genetics of yeast. Proc Natl Acad Sci U S A 35:456–464PubMedPubMedCentralCrossRefGoogle Scholar
  118. Potter G, Budge SM, Speers RA (2015) Flocculation, cell surface hydrophobicity and 3-OH oxylipins in the SMA strain of Saccharomyces pastorianus. J Inst Brew 121:31–37CrossRefGoogle Scholar
  119. Powell CD, Diacetis AN (2007) Long term serial repitching and the genetic and phenotypic stability of brewer’s yeast. J Inst Brew 113:67–74CrossRefGoogle Scholar
  120. Powell CD, Quain DE, Smart KA (2003) The impact of brewing yeast cell age on fermentation performance, attenuation and flocculation. FEMS Yeast Res 3:149–157PubMedCrossRefGoogle Scholar
  121. Raspor P, Russell I, Stewart GG (1990) An update of zinc ion as an effector of flocculation in brewer’s yeast strains. J Inst Brew 96:303–305CrossRefGoogle Scholar
  122. Rees EMR, Stewart GG (1997a) The effects of divalent ions magnesium and calcium on yeast fermentation performance in conventional (12°P) and high (20°P) gravity worts in both static and shaking fermentations. In: Proceedings of the 26th congress – European Brewery Convention, Maestricht, The Netherlands, pp 461–468Google Scholar
  123. Rees EMR, Stewart GG (1997b) The effects of increased magnesium and calcium concentrations on yeast fermentation performance in high gravity worts. J Inst Brew 103:287–291CrossRefGoogle Scholar
  124. Rees EMR, Stewart GG (1998) Strain specific response of brewer’s yeast strains to zinc concentrations in conventional and high gravity worts. J Inst Brew 104:255–264CrossRefGoogle Scholar
  125. Rhymes MR, Smart KA (2001) Effect of storage conditions on the flocculation and cell wall characteristics of an ale brewing yeast strain. J Am Soc Brew Chem 59:32–38Google Scholar
  126. Richards M (1967) The use of giant-colony morphology for the differentiation of brewing yeasts. J Inst Brew 73:162–166CrossRefGoogle Scholar
  127. Rose AH (1980) Saccharomyces cerevisiae as a model eukaryote. In: Stewart GG, Russell I (eds) Current developments in yeast research. Pergamon, Toronto, pp 645–652Google Scholar
  128. Rose AH (1984) Physiology of cell aggregation: flocculation by Saccharomyces cerevisiae as a model system. In: Marshall CK (ed) Physiology of cell aggregation. Springer, New York, pp 323–335Google Scholar
  129. Rose AH, Harrison JF (1987) The yeasts, vol 1–5. Academic, LondonGoogle Scholar
  130. Rossouw D, Bagheri B, Setati ME, Bauer FF (2015) Co-flocculation of yeast species, a new mechanism to govern population dynamics in microbial ecosystems. PLoS One 10(8):e0136249PubMedPubMedCentralCrossRefGoogle Scholar
  131. Russell I, Stewart GG (1980) Revised nomenclature of genes that control yeast flocculation. J Inst Brew 86:120–121CrossRefGoogle Scholar
  132. Russell I, Stewart GG (eds) (2014) Whisky: technology, production and marketing, 2nd edn. Academic (Elsevier), Boston, MAGoogle Scholar
  133. Russell I, Dowhanick T, Raspor P, Stewart GG (1989) Yeast flocculation – the influence of divalent ions. In: Proceedings of the 22nd congress – European Brewery Convention, Zurich. IRL Press, Oxford, pp 529–536Google Scholar
  134. Sato M, Watari J, Shinotsuka K (2001) Genetic instability in flocculation of bottom-fermenting yeast. J Am Soc Brew Chem 59:130–134Google Scholar
  135. Schlee C, Miedl M, Leiper KA, Stewart GG (2006) The potential of confocal imaging for measuring physiological changes in brewer’s yeast. J Inst Brew 112:134–147CrossRefGoogle Scholar
  136. Sherman F, Fink GR, Hicks JB (1986) Methods in yeast genetics. Cold Spring Harbor Laboratory Press, New YorkGoogle Scholar
  137. Siebert KJ, Stenroos LE, Reid DS, Grabowski D (1987) Filtration difficulties resulting from damage to yeast during centrifugation. MBAA Tech Q 24:1–8Google Scholar
  138. Siero C, Reboredo NM, Villa TH (1994) Flocculation of industrial and laboratory strains of Saccharomyces cerevisiae. J Ind Microbiol 14:461–466CrossRefGoogle Scholar
  139. Smart KA, Whisker S (1996) Effect of serial repitching on the fermentation properties and condition of brewing yeast. J Am Soc Brew Chem 54:41–44Google Scholar
  140. Smit G, Straver MH, Lugtenberg BJJ, Kijne JW (1992) Flocculence of Saccharomyces cerevisiae cells is induced by nutrient limitation, with cell surface hydrophobicity as a major determinant. Appl Environ Microbiol 58:3709–3714PubMedPubMedCentralGoogle Scholar
  141. Soares EV (2010) Flocculation in Saccharomyces cerevisiae: a review. Appl Microbiol 110:1–18CrossRefGoogle Scholar
  142. Soares EV, Duarte AA (2002) Addition of nutrients induce a fast loss of flocculation in starved cells of Saccharomyces cerevisiae. Biotechnol Lett 24:1957–1960CrossRefGoogle Scholar
  143. Soares EV, Mota M (1996) Flocculation onset, growth phase, and genealogical age in Saccharomyces cerevisiae. Can J Microbiol 42:539–547PubMedCrossRefGoogle Scholar
  144. Soares EV, Seynaeve J (2000a) The use of succinic acid, as a pH buffer, expands the potentialities of utilisation of a chemically defined medium in Saccharomyces cerevisiae flocculation studies. Biotechnol Lett 22:859–863CrossRefGoogle Scholar
  145. Soares EV, Seynaeve J (2000b) Induction of flocculation of brewer’s yeast strains of Saccharomyces cerevisiae by changing the calcium concentration and pH of culture medium. Biotechnol Lett 22:1827–1832CrossRefGoogle Scholar
  146. Soares E, Teixeira JA, Mota M (1991) Influence of aeration and glucose concentration in the flocculation of Saccharomyces cerevisiae. Biotechnol Lett 13:207–212CrossRefGoogle Scholar
  147. Soares EV, Teixeira JA, Mota M (1994) Effect of cultural and nutritional conditions on the control of flocculation expression in Saccharomyces cerevisiae. Can J Microbiol 40:851–857PubMedCrossRefGoogle Scholar
  148. Song Q, Kumar A (2012) An overview of autophagy and yeast pseudohyphal growth: integration of signaling pathways during nitrogen stress. Cells 1:263–283PubMedPubMedCentralCrossRefGoogle Scholar
  149. Sousa MJ, Teixeira JA, Mota M (1992) Differences in the flocculation mechanism of Kluyveromyces marxianus and Saccharomyces cerevisiae. Biotechnol Lett 14:213–218CrossRefGoogle Scholar
  150. Southern EM (1975) Detection of specific sequences among DNA fragments separated by gel electrophoresis. J Mol Biol 98:503–517PubMedCrossRefGoogle Scholar
  151. Speers RA (2012) A review of yeast flocculation. In: Speers RA (ed) Proceedings of the 2nd international brewers symposium: yeast flocculation, vitality and viability. MBAA, St Paul, MNGoogle Scholar
  152. Speers RA (2016) Brewing fundamentals, Part 3: Yeast settling and flocculation. MBAA Tech Q 53:17–22Google Scholar
  153. Speers RA, Stokes S (2009) Effects of vessel geometry, fermenting volume and yeast repitching on fermenting beer. J Inst Brew 115:148–150CrossRefGoogle Scholar
  154. Speers RA, Durance TD, Odense P, Owen S, Tung MA (1993) Physical properties of commercial brewing yeast suspensions. J Inst Brew 99:159–164CrossRefGoogle Scholar
  155. Speers RA, Wan Y-Q, Jin Y, Stewart RJ (2006) Effects of fermentation parameters and cell wall properties on yeast flocculation. J Inst Brew 112:246–254CrossRefGoogle Scholar
  156. Sprague GF Jr, Thorner JW (1992) Pheromone response and signal transduction during the mating process of Saccharomyces cerevisiae. In: Jones EW, Pringle JR, Broach JR (eds) The molecular and cellular biology of the yeast saccharomyces: gene expression. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, NY, pp 657–744Google Scholar
  157. Stewart GG (1972) Co-flocculation of brewer’s yeast. MBAA Tech Q 9:25Google Scholar
  158. Stewart GG (1973) Recent developments in the characterization of brewery yeast strains. MBAA Tech Q 9:183–191Google Scholar
  159. Stewart GG (1974) Some thoughts on the microbiological aspects of brewing and other industries utilizing yeast. Adv Appl Microbiol 17:233–262CrossRefGoogle Scholar
  160. Stewart GG (1988) Twenty-five years of yeast research. Dev Ind Microbiol 29:1–21. SIM Charles Thom Award LectureGoogle Scholar
  161. Stewart GG (1996) Yeast performance and management. The Brewer 82:211–215Google Scholar
  162. Stewart GG (2006) Studies on the uptake and metabolism of wort sugars during brewing fermentations. MBAA Tech Q 43:265–269Google Scholar
  163. Stewart GG (2009) The IBD horace brown medal lecture – forty years of brewing research. J Inst Brew 115:3–29CrossRefGoogle Scholar
  164. Stewart GG (2010a) MBAA award of merit lecture. A love affair with yeast. MBAA Tech Q 47:4–11Google Scholar
  165. Stewart GG (2010b) The ASBC award of distinction lecture – high gravity brewing and distilling – past experiences and future prospects. J Am Soc Brew Chem 68:1–9Google Scholar
  166. Stewart GG (2014a) Brewing intensification. American Society for Brewing Chemists, St. Paul, MNGoogle Scholar
  167. Stewart GG (2014b) The concept of nature – nurture applied to brewer’s yeast and wort fermentations. MBAA Tech Q 51:69–80Google Scholar
  168. Stewart GG (2014c) Yeast mitochondria – their influence on brewer’s yeast fermentation and medical research. MBAA Tech Q 51:3–11Google Scholar
  169. Stewart GG (2014d) Saccharomyces. In: Catt C, Tortorello ML (eds) Encyclopedia of food microbiology, vol 3, 2nd edn. Elsevier, Oxford, pp 297–315CrossRefGoogle Scholar
  170. Stewart GG (2015a) Seduced by yeast. J Am Soc Brew Chem 73:1–21Google Scholar
  171. Stewart GG (2015b) Yeast quality assessment, management and culture maintenance, Chap. 2. In: Hill AE (ed) Brewing microbiology: managing microbes, ensuring quality and valorising waste. Elsevier, Oxford, pp 11–29CrossRefGoogle Scholar
  172. Stewart GG, Garrison IF (1972) Some observations on co-flocculation in Saccharomyces cerevisiae. Am Soc Brew Chem Proc:118–131Google Scholar
  173. Stewart GG, Goring TE (1976) Effect of some monovalent and divalent metal ions on the flocculation of brewer’s yeast strains. J Inst Brew 82:341–342CrossRefGoogle Scholar
  174. Stewart GG, Murray JP (2011) Using brewing science to make good beer. MBAA Tech Q 48:13–19Google Scholar
  175. Stewart GG, Murray JP (2012) Brewing intensification – successes and failures. MBAA Tech Q 49:111–120Google Scholar
  176. Stewart GG, Russell I (1977) The identification, characterization, and mapping of a gene for flocculation in Saccharomyces sp. Can J Microbiol 23:441–447PubMedCrossRefGoogle Scholar
  177. Stewart GG, Russell I (1981) Yeast flocculation. In: Pollock JAR (ed) Brewing science, food science and technology. Academic, New York, pp 61–92Google Scholar
  178. Stewart GG, Russell I (1983) Aspects of the biochemistry and genetics of sugar and carbohydrate uptake by yeasts. In: Spencer JFT, Spencer DM, Smith ARW (eds) Yeast genetics: fundamental and applied aspects. Springer, New York, pp 461–484CrossRefGoogle Scholar
  179. Stewart GG, Russell I (1986) The relevance of the flocculation properties of yeast in today’s brewing industry. In: European Brewing Convention – Symposium on ‘Brewers’ yeast, Vuoranta, Helsinki, Finland, pp 24–25, 53–68Google Scholar
  180. Stewart GG, Russell I (2009) An introduction to brewing science and technology, Series lll, Brewer’s yeast, 2nd edn. The Institute of Brewing and Distilling, LondonGoogle Scholar
  181. Stewart GG, Russell I, Garrison IF (1973) Further studies on flocculation and co-flocculation in brewer’s yeast strains. Am Soc Brew Chem Proc 31:100–106Google Scholar
  182. Stewart GG, Russell I, Garrison IF (1974) Factors influencing the flocculation of brewers’ yeast strains. MBAA Tech Q II:xiiiGoogle Scholar
  183. Stewart GG, Russell I, Garrison IF (1975a) Some considerations of the flocculation characteristics of ale and lager yeast strains. J Inst Brew 81:248–257CrossRefGoogle Scholar
  184. Stewart GG, Russell I, Goring IF (1975b) Nature-nurture anomalies – further studies in yeast flocculation. Am Soc Brew Chem Proc 33:137–147Google Scholar
  185. Stewart GG, Goring TE, Russell I (1983a) (issued October 11, 1983) Yeast strain for fermenting high plato value wort. US Patent 4,409,246Google Scholar
  186. Stewart GG, Panchal CJ, Russell I (1983b) Current developments in the genetic manipulation of brewing yeast strains – a review. J Inst Brew 89:170–188CrossRefGoogle Scholar
  187. Stewart GG, Murray CR, Panchal CJ, Russell I, Sills AM (1984a) The selection and modification of brewer’s yeast strains. Food Microbiol 1:289–302CrossRefGoogle Scholar
  188. Stewart GG, Panchal CJ, Russell I, Sills AM (1984b) Biology of ethanol producing microorganisms. CRC Crit Rev Biotechnol 1:161–188CrossRefGoogle Scholar
  189. Stewart GG, Russell I, Panchal CJ (1984c) Genetically stable allopolyploid somatic fusion product useful in the production of fuel alcohols. Australian Patent: 570,260 (issued August 15, 1984)Google Scholar
  190. Stewart GG, Hill A, Russell I (2013) 125th anniversary review - developments in brewing and distilling yeast strains. J Inst Brew 119:202–220CrossRefGoogle Scholar
  191. Stoupis T, Stewart GG, Stafford RA (2002) Mechanical agitation and rheological considerations of ale yeast slurry. J Am Soc Brew Chem 60:58–62Google Scholar
  192. Stratford M (1989) Yeast flocculation: calcium specificity. Yeast 5:487–496CrossRefGoogle Scholar
  193. Stratford M (1992a) Lectin-mediated aggregation of yeasts- yeast flocculation. Biotechnol Genet Eng Rev 10:283–341PubMedCrossRefGoogle Scholar
  194. Stratford M (1992b) Yeast flocculation – a new perspective. Adv Microb Physiol 33:1–71CrossRefGoogle Scholar
  195. Stratford M (1992c) Yeast flocculation – receptor definition by mnn mutants and concanavalin-A. Yeast 8:635–645PubMedCrossRefGoogle Scholar
  196. Stratford M (1992d) Yeast flocculation: calcium specificity. Yeast 5:487–496CrossRefGoogle Scholar
  197. Stratford M (1996) Induction of flocculation in brewing yeasts by change in pH value. FEMS Microbiol Lett 136:13–18PubMedCrossRefGoogle Scholar
  198. Stratford M, Assinder S (1991) Yeast flocculation: Flo1 and new Flo phenotypes and receptor structure. Yeast 7:559–574PubMedCrossRefGoogle Scholar
  199. Stratford M, Keenan MMJ (1988) Yeast flocculation: Quantification. Yeast 4:107–115PubMedCrossRefGoogle Scholar
  200. Strauss CJ, Kock JLF, van Wyk PWJ, Lodolo EJ, Pohl CH, Botes PJ (2005) Bioactive oxylipins in Saccharomyces cerevisiae. J Inst Brew 111:304–308CrossRefGoogle Scholar
  201. Strauss CJ, van Wyk PWJ, Lodolo EJ, Botes PJ, Pohl CH, Nigam S, Kock JLF (2007) Mitochondrial associated yeast flocculation – the effect of acetylsalicylic acid. J Inst Brew 113:42–47CrossRefGoogle Scholar
  202. Straver MH, Aar PCVD, Smit G, Kijne JW (1993) Determinants of flocculence of brewer’s yeast during fermentation in wort. Yeast 9:527–532PubMedCrossRefGoogle Scholar
  203. Taylor NW, Orton WI (1978) Aromatic compounds and sugars in flocculation of Saccharomyces cerevisiae. J Inst Brew 84:113–114CrossRefGoogle Scholar
  204. Teixeira JM, Teixeira JA, Mota M, Manuela M, Guerra B, Machado Cruz JM, S’Almeida AM (1991) The influence of cell wall composition of a brewer’s flocculant lager yeast on sedimentation during successive industrial fermentations. In: Proceedings of the European Brewery Convention Congress, Lisbon, pp 241–248Google Scholar
  205. Teunissen AWRH, Steensma HY (1995) Review: the dominant flocculation genes of Saccharomyces cerevisiae constitute a new subtelomeric gene family. Yeast 11:1001–1013PubMedCrossRefGoogle Scholar
  206. Teunissen AWRH, Holub E, Van Der Hucht J, Van Den Berg JA, Steensma HY (1993) Sequence of the open reading frame of the FLO1 gene from Saccharomyces cerevisiae. Yeast 9:423–427PubMedCrossRefGoogle Scholar
  207. Teunissen AWRH, Van Den Berg JA, Teunissen SHY (1995) Transcriptional regulation of flocculation genes in Saccharomyces cerevisiae. Yeast 11:435–446PubMedCrossRefGoogle Scholar
  208. Thorne RSW (1951) Some aspects of yeast flocculence. In: Proceedings of the European Brewery Convention Congress, Brighton, pp 21–34Google Scholar
  209. van Hamersveld EH, van der Lans RG, Luyben KC (1997) Quantification of brewers’ yeast flocculation in a stirred tank: effect of physical parameters on flocculation. Biotechnol Bioeng 56:190–200PubMedCrossRefGoogle Scholar
  210. van Holle A, Machado MD, Soares EV (2011) Flocculation in ale brewing strains of Saccharomyces cerevisiae: re-evaluation of the role of cell surface charge and hydrophobicity. Appl Microbiol Biotechnol 93:1221–1229PubMedCrossRefGoogle Scholar
  211. Van Lersel MFM, Meersman E, Arntz M, Rombouts FM, Abee T (1998) Effect of environmental conditions on flocculation and immobilisation of brewer’s yeast during production of alcohol-free beer. J Inst Brew 104:131–136CrossRefGoogle Scholar
  212. Van Mulders SE, Christianen E, Saerens SM, Daenen L, Verbelen PJ, Willaert R, Verstrepen KJ, Delvaux FR (2009) Phenotypic diversity of Flo protein family-mediated adhesion in Saccharomyces cerevisiae. FEMS Yeast Res 9:178–190PubMedCrossRefGoogle Scholar
  213. Van Mulders SE, Ghequire M, Daenen L, Verbelen PJ, Verstrepen KJ, Delvaux FR (2010) Flocculation gene variability in industrial brewer’s yeast strains. Appl Microbiol Biotechnol 88:1321–1331PubMedCrossRefGoogle Scholar
  214. Van Nierop SNE, Cameron-Clarke A, Axcell BC (2004) Enzymatic generation of factors malt responsible for premature yeast flocculation. J Am Soc Brew Chem 62:108–116Google Scholar
  215. Van Nierop SNE, Rautenbach M, Axcell BC, Cantrell IC (2006) The impact of microorganisms on barley and malt quality – a review. J Am Soc Brew Chem 62:69–79Google Scholar
  216. Verstrepen KJ, Fink GR (2009) Genetic and epigenetic mechanisms underlying cell-surface variability in protozoa and fungi. Annu Rev Genet 43:1–24PubMedCrossRefGoogle Scholar
  217. Verstrepen KJ, Klis FM (2006) Flocculation, adhesion and biofilm formation in yeasts. Mol Microbiol 60:5–15PubMedCrossRefGoogle Scholar
  218. Verstrepen KJ, Bauer FF, Winderickx J, Derdelinckx G, Dufour JP, Thevelein JM, Pretorius IS, Delvaux FR (2001a) Genetic modification of Saccharomyces cerevisiae: fitting the modern brewer’s needs. Cerevisia 26:89–97Google Scholar
  219. Verstrepen KJ, Derdelinckx G, Delvaux FR, Winderickx J, Thevelein JM, Bauer FF, Pretorius IS (2001b) Late fermentation expression of FLO1 in Saccharomyces cerevisiae. J Am Soc Brew Chem 59:69–76Google Scholar
  220. Verstrepen KJ, Van Laere SD, Vanderhaegen BM, Derdelinckx G, Dufour JP, Pretorius IS, Winderickx J, Thevelein JM, Delvaux FR (2003) Expression levels of the yeast alcohol acetyltransferase genes ATF1, Lg-ATF1, and ATF2 control the formation of a broad range of volatile esters. Appl Environ Microbiol 69:5228–5237PubMedPubMedCentralCrossRefGoogle Scholar
  221. Verstrepen KJ, Reynolds TB, Fink GR (2004) Origins of variation in the fungal cell surface. Nat Rev Microbiol 2:533–540PubMedCrossRefGoogle Scholar
  222. Verstrepen KJ, Jansen A, Lewitter F, Fink GR (2005) Intragenic tandem repeats generate functional variability. Nat Genet 37:986–990PubMedPubMedCentralCrossRefGoogle Scholar
  223. Vidgren V, Londesborough J (2011) 125th anniversary review: yeast flocculation and sedimentation in Brewing. J Inst Brew 117:475–487CrossRefGoogle Scholar
  224. Wang A, Raniga PP, Lane S, Lu Y, Liu H (2009) Hyphal chain formation in Candida albicans: Cdc28-Hgc1 phosphorylation of Efg1 represses cell separation genes. Mol Cell Biol 29:4406–4416PubMedPubMedCentralCrossRefGoogle Scholar
  225. Watari J, Takata Y, Ogawa M, Sahara H, Koshino S, Onnela M, Airaksinen U, Jaatinen R (1994) Molecular cloning and analysis of the yeast flocculation gene FlO1. Yeast 10:211–225PubMedCrossRefGoogle Scholar
  226. Watari J, Sato M, Ogawa M, Shinotsuka K (1999) Genetic and physiological instability of brewing yeast. Eur Brew Conv Monogr 28:148–160Google Scholar
  227. White FH, Kidney E (1979) The influence of yeast strain on beer spoilage bacteria. Proceedings of the 17th European Brewery Convention Congress, Berlin, DSW, Dordrecht, The Netherlands, pp 801–815Google Scholar
  228. Wickerham LJ (1951) Taxonomy of yeasts. Tech Bull 27.8. Dep Agric no 1029Google Scholar
  229. Wightman P, Quain DE, Meaden PG (1996) Analysis of production brewing strains of yeast by DNA fingerprinting. Lett Appl Microbiol 22:90–94PubMedCrossRefGoogle Scholar
  230. Wilcocks KL, Smart KA (1995) The importance of surface charge and hydrophobicity for the flocculation of chain-forming brewing yeast strains and resistance of these parameters to acid washing. FEMS Microbiol Lett 15:293–297CrossRefGoogle Scholar
  231. Williams LJ, Barnett GR, Ristow JL, Pitkin J, Perriere M, Davis RH (1992) Ornithine decarboxylase gene of Neurospora crassa: isolation, sequence and polyamine-mediated regulation of its mRNA. Mol Cell Biol 12:347–359PubMedPubMedCentralCrossRefGoogle Scholar
  232. Zarattini RA, Williams JW, Ernandes JR, Stewart GG (1993) Bacterial-induced flocculation in selected brewing strains of Saccharomyces. Cerevisia Biotechnol 18:65–70Google Scholar
  233. Zhao XQ, Bai FW (2009) Yeast flocculation: new story in fuel ethanol production. Biotechnol Adv 27:849–856PubMedCrossRefGoogle Scholar
  234. Zhuang S, Smart K, Powell C (2017) Impact of extracellular osmolarity on Saccharomyces yeast population during brewing fermentation. J Am Soc Brew Chem 2017:244–254Google Scholar

Copyright information

© Springer International Publishing AG 2017

Authors and Affiliations

  • Graham G. Stewart
    • 1
    • 2
  1. 1.International Centre for Brewing and DistillingHeriot Watt UniversityEdinburghUK
  2. 2.GGStewart AssociatesCardiff, WalesUK

Personalised recommendations