Whisky Science pp 143-163 | Cite as


  • Gregory H. Miller


Fermentation is the breakdown of material by the action of microorganisms. In the present case, fermentation refers to the conversion of sugars to ethanol by the yeast Saccharomyces cerevisiea, a fungus. The main fermen sugars are maltose, glucose, fructose, sucrose, maltotriose, and maltotetrose. Dextrins, which are small molecular weight glucose polymers, may be fermentable to a degree by yeast strains that express amylase. The substrain S. diastaticus (Andrews and Gilliland, J Inst Brew 58:189–196, 1952; Gilliland, J Inst Brew 72:271–275, 1966), which produces extracellular glucoamylase (Erratt and Stewart, J Am Soc Brew Chem 36:151–161, 1978) is a notable example. Distiller’s wort will have some residual amylase activity from malting, whereas a brewer’s boiled wort will not. Nonfermentable sugars include the pentose sugars (e.g., arabinose and xylose), isomaltose (an α-(1-6) linked glucose dimer), and unconverted starches including β-glucans.


  1. 7.
    Adinoff B, Bone GHA, Linnoila M (1988) Acute ethanol poisoning and the ethanol withdrawal syndrome. Med Toxicol Adverse Drug Exp 3:172–196PubMedGoogle Scholar
  2. 22.
    Andrews J, Gilliland RB (1952) Super-attenuation of beer: a study of three organisms capable of causing abnormal attenuations. J Inst Brew 58:189–196CrossRefGoogle Scholar
  3. 23.
    Anness BJ (1980) The reduction of dimethyl sulphoxide to dimethyl sulfide during fermentation. J Inst Brew 86:134–137CrossRefGoogle Scholar
  4. 25.
    Anness BJ, Bamforth CW, Wainwright T (1979) The measurement of dimethyl sulphoxide in barley and malt and its reduction to dimethyl sulphide by yeast. J Inst Brew 85:346–349CrossRefGoogle Scholar
  5. 54.
    Aylward F, Coleman G, Haisman DR (1967) Catty odours in food: the reaction between mesityl oxide and sulfur compounds in foodstuffs. Chem Ind 37:1563–1564PubMedGoogle Scholar
  6. 55.
    Äyräpää T (1971) Biosynthetic formation of higher alcohols by yeast. Dependence on the nitrogenous nutrient level of the medium. J Inst Brew 77:266–276Google Scholar
  7. 83.
    Berry DR, Watson DC (1987) Production of organoleptic compounds. In: Berry DR, Russell I, Stewart GG (eds) Yeast biotechnology. Allen & Unwin, London, p 345–368CrossRefGoogle Scholar
  8. 125.
    Burgé G, Saulou-Bérion C, Moussa M, Pollet B, Flourat A, Allais F, Athès V, Spinnler HE (2015) Diversity of Lactobacillus reuteri strains in converting glycerol into 3-hydroxypropionic acid. Appl Biochem Biotechnol 177:923–939CrossRefGoogle Scholar
  9. 137.
    Cachat E, Priest FG (2005) Lactobacillus suntoryeus sp. nov., isolated from malt whisky distilleries. Int J Syst Evol Microbiol 55:31–34CrossRefGoogle Scholar
  10. 144.
    Capote T (1980) Music for chameleons. Random House, New YorkGoogle Scholar
  11. 154.
    Chen EC-H (1978) The relative contribution of Ehrlich and biosynthetic pathways to the formation of fusel alcohols. J Am Soc Brew Chem 36:39–43Google Scholar
  12. 161.
    Clarke S (1991) Protein isoprenylation and methylation at carboxyl-terminal cysteine residues. Annu Rev Biochem 61:355–386CrossRefGoogle Scholar
  13. 225.
    Deschenes RJ, Stimmel JB, Clarke S, Stock J, Broach JR (1989) RAS2 protein of Saccharomyces cerevisiae is methyl-esterified at its carboxyl terminus. J Biol Chem 264:11865–11873PubMedGoogle Scholar
  14. 232.
    Dittrich F, Zajonc D, Hühne K, Hoja U, Ekici A, Greiner E, Klein H, Hofmann J, Bessoule J-J, Sperling P, Schweizer E (1998) Fatty acid elongation in yeast. Biochemical characteristics of the enzyme system and isolation of elongation-defective mutants. Eur J Biochem 252:477–485CrossRefGoogle Scholar
  15. 246.
    Engan S (1981) Beer composition: volatile substances. In: Pollock JRA (ed) Brewing sciences, vol 2, Academic Press, London, p 98–165Google Scholar
  16. 247.
    Engel K-H, Flath RA, Buttery RG, Mon TR, Ramming DW, Teranishi R (1988) Investigation of volatile constituents in nectarines. 1. Analytical and sensory characterization of aroma components in some nectarine cultivars. J Agric Food Chem 36:549–553CrossRefGoogle Scholar
  17. 250.
    Erratt JA, Stewart GG (1978) Genetic and biochemical studies on yeast strains able to use dextrins. J Am Soc Brew Chem 36:151–161Google Scholar
  18. 286.
    Geddes PA (1986) The production of hydrogen sulfide by lactobacillus spp. in fermenting wort. In: Campbell I, Priest FG (eds) Proceedings of the 2nd Aviemore conference on malting, brewing, and distilling. Institute of Brewing, London, p 364–370Google Scholar
  19. 287.
    Geddes PA, Riffkin HL (1989) Influence of lactic acid bacteria on aldehyde, ester, and higher alcohol formation during Scotch whisky fermentations. In: Piggott JR, Paterson A (eds) Distilled beverage flavour, Ellis Horwood, Chichester, p 193–199Google Scholar
  20. 295.
    Gilliland RB (1966) Saccharomyces diastaticus—a starch-fermenting yeast. J Inst Brew 72:271–275CrossRefGoogle Scholar
  21. 317.
    Guichard E (1995) Chiral γ-lactones, key compounds to apricot flavour. In: Rouseff RL, Leahy MM (eds) Fruit flavors. American Chemical Society, Washington, DC, p 258–267CrossRefGoogle Scholar
  22. 324.
    Guymon JF, Ingraham JL, Crowell EA (1961) The formation of n-propyl alcohol by Saccharomyces cerevisiae. Arch Biochem Biophys 95:163–168CrossRefGoogle Scholar
  23. 331.
    Hammer SK, Avalos JL (2017) Uncovering the role of branched-chain amino acid transaminases in Saccharomyces cerevisiae isobutanol biosynthesis. Matab Eng 44:302–312CrossRefGoogle Scholar
  24. 332.
    Hansen J, Bruun SV, Bech LM, Gjermansen C (2002) The level of MXR1 gene expression in brewing yeast during beer fermentation is a major determinant for the concentration of dimethyl sulfide in beer. FEMS Yeast Res 2:137–149PubMedGoogle Scholar
  25. 342.
    Hazelwood LA, Daran J-M, van Maris AJA, Pronk JT, Dickinson JR (2008) The Ehrlich pathway for fusel alcohol production: a century of research on Saccharomyces cerevisiae metabolism. Appl Environ Microbiol 74:2259–2266, 3920CrossRefGoogle Scholar
  26. 359.
    Hou C-Y, Lin Y-S, Wang Y, Jiang C-M, Wu M-C (2008) Effect of storage conditions on methanol content offruit and vegetable juices. J Food Comp Anal 21:410–415CrossRefGoogle Scholar
  27. 402.
    Kächele M, Monakhova YB, Kuballa T, Lachenmeier DW (2014) NMR investigation of acrolein stability in hydroalcoholic solution as a foundation for the valid HS-SPME/GC-MS quantification of the unsaturated aldehyde in beverages. Anal Chim Acta 820:112–118CrossRefGoogle Scholar
  28. 406.
    Kahn JH, LaRoe EG, Conner HA (1968) Whiskey composition: identification of components by single-pass gas chromatography-mass spectrometry. J Food Sci 33:395–400CrossRefGoogle Scholar
  29. 421.
    Kinzurik MI, Herbst-Johnstone M, Gardner RC, Fedrizzi B (2016) Hydrogen sulfide production during yeast fermentation causes the accumulation of ethanethiol, S-ethyl thioacetate and diethyl disulfide. Food Chem 209:341–347CrossRefGoogle Scholar
  30. 439.
    Landaud S, Helinck S, Bonnarme P (2008) Formation of volatile sulfur compounds and metabolism of methionine and other sulfur compounds in fermented food. Appl Microbiol Biotechnol 77:1191–1205CrossRefGoogle Scholar
  31. 470.
    Lynen F (1967) The role of biotin-dependent carboxylations in biosynthetic reactions. Biochem J 102:381–399CrossRefGoogle Scholar
  32. 471.
    Maarse H, ten Noever de Brauw MC (1974) Another catty odour compound causing air pollution. Chem Ind 44:36–37Google Scholar
  33. 477.
    MacKenzie WM, Aylott RI (2004) Analytical strategies to confirm Scotch whisky authenticity. Analyst 129:607–612CrossRefGoogle Scholar
  34. 495.
    Makanjuola DB, Springham DG (1984) Identification of lactic acid bacteria isolated from different stages of malt whisky distillery fermentations. J Inst Brew 90:13–19CrossRefGoogle Scholar
  35. 537.
    Middlekauff JE (1995) Sanitation and pest control: Part B. Microbiological aspects. In: Hardwick WA (ed) Handbook of brewing. Marcel Dekker, New York, p 480–499Google Scholar
  36. 539.
    Mills DE, Baugh WD, Conner HA (1954) Studies on the formation of acrolein in distillery mashes. Appl Microbiol 2:9–13PubMedPubMedCentralGoogle Scholar
  37. 564.
    Narendranath NV, Power R (2004) Effect of yeast inoculation rate on the metabolism of contaminating lactobacilli during fermentation of corn mash. J Ind Microbiol Biotechnol 31:581–583CrossRefGoogle Scholar
  38. 569.
    Nemoto S (1975) Possibilities of utilization of butyric acid bacteria for rum making. Ann Technol Agric 24:397–410Google Scholar
  39. 571.
    Nettleton JA (1913) The manufacture of whisky and plain spirit. G. Cornwall & Sons, AberdeenGoogle Scholar
  40. 589.
    Nordström K (1963) Formation of esters from acids by brewer’s yeast II. Formation from lower fatty acids. J Inst Brew 70:42–55Google Scholar
  41. 590.
    Nordström K (1966) Yeast growth and glycerol formation. Acta Chem Scand 20:1016–1025CrossRefGoogle Scholar
  42. 594.
    Nykänen L, Nykänen I (1977) Production of esters by different yeast strains in sugar fermentations. J Inst Brew 83:30–31CrossRefGoogle Scholar
  43. 598.
    Nykänen L, Nykänen I, Suomalainen H (1977) Distribution of esters produced during sugar fermentation between the yeast cell and the medium. J Inst Brew 83:32–34CrossRefGoogle Scholar
  44. 617.
    Paine AJ, Dayan AD (2001) Defining a tolerable concentration of methanol in alcoholic drinks. Hum Exp Toxicol 20:563–568CrossRefGoogle Scholar
  45. 623.
    Pasteur ML (1857) Mémoire sur la fermentation alcoolique. C R Hebd Seances Acad Sci 45:1032–1036Google Scholar
  46. 628.
    Patterson RLS, Rhodes DN (1967) Catty odours in food: their production in meat stores from mesityl oxide in paint solvents. Chem Ind 37:2003–2004Google Scholar
  47. 629.
    Pearce TJP, Peacock JM, Aylward F, Haisman DR (1967) Catty odours in food: reactions between hydrogen sulfide and unsaturated ketones. Chem Ind 37:1562–1563PubMedGoogle Scholar
  48. 634.
    Perpéte P, Duthoid O, de Maeyer S, Imray L, Lawton AI, Stavropoulos KE, Gitonga VW, Hewlins MJE, Dickinson JR (2006) Methionine catabolism in Saccharomyces cerevisiae. FEMS Yeast Res 6:48–56CrossRefGoogle Scholar
  49. 678.
    Ramsay CM, Berry DR (1984) The effect of inoculum level on the formation of higher alcohols, fatty acids and esters in the malt whisky fermentation. Food Microbiol 1:111–115CrossRefGoogle Scholar
  50. 679.
    Ramsay CM, Berry DR (1984) Effect of temperature and pH on the formation of higher alcohols, fatty acids and esters in the malt whisky fermentation. Food Microbiol 1:117–121CrossRefGoogle Scholar
  51. 681.
    Rankine BC (1968) The importance of yeasts in determining the composition and quality of wines. Vitis 7:22–49Google Scholar
  52. 682.
    Rauhut D (2017) Usage and formation of sulfur compounds. In: König H, Unden G, Fröhlich J (eds) Biology of microorganisms on grapes, in must and in wine, 2nd edn, Springer, Cham, p 255–291CrossRefGoogle Scholar
  53. 687.
    Reazin GH, Scales H, Andreasen A (1973) Production of higher alcohols from theonine and isoleucine in alcoholic fermentations of different types of grain mash. J Agric Food Chem 21:50–54CrossRefGoogle Scholar
  54. 716.
    Ryan ED, Kohlhaw GB (1974) Subcellular localization of isoleucine-valine biosynthetic enzymes in yeast. J Bacteriol 120:631–637PubMedPubMedCentralGoogle Scholar
  55. 718.
    Saerens SMG, Delvaux F, Verstrepen KJ, Van Dijck P, Thevelein JM, Delvaux FR (2008) Parameters affecting ethyl ester production by Saccharomyces cerevisiae during fermentation. Appl Environ Microbiol 74:454–461CrossRefGoogle Scholar
  56. 746.
    Serjak WC, Day WH, van Lanen JM, Boruff CS (1954) Acrolein production by bacteria found in distillery grain mashes. Appl Microbiol 2:14–20PubMedPubMedCentralGoogle Scholar
  57. 757.
    Simpson KL, Pettersson B, Priest FG (2001) Characterization of lactobacilli from Scotch malt whisky distilleries and description of Lactobacillus ferintoshensis sp. nov., a new species isolated from malt whisky fermentations. Microbiology 147:1007–1016CrossRefGoogle Scholar
  58. 768.
    Sobolov M, Smiley KL (1960) Metabolism of glycerol by an acrolein-forming lactobacillus. J Bacteriol 79:261–266PubMedPubMedCentralGoogle Scholar
  59. 778.
    Spiropoulos A, Tanaka J, Flerianos I, Bisson LF (2000) Characterization of hydrogen sulfide formation in commercial and natural wine isolates of Saccharomyces. Am J Enol Vitic 51:233–248Google Scholar
  60. 781.
    Steinke RD, Paulson MC (1964) Phenols from grain. The production of steam-volatile phenols during the cooking and alcoholic fermentation of grain. J Agric Food Chem 12:381–387Google Scholar
  61. 788.
    Suomalainen H, Lehtonen M (1979) The production of aroma compounds by yeast. J Inst Brew 85:149–156CrossRefGoogle Scholar
  62. 790.
    Suomalainen H, Nykänen L (1970) Investigations on the aroma of alcoholic beverages. Naeringsmiddelindustrien 23:15–30Google Scholar
  63. 802.
    Taylor EH Jr (1882) Making whisky. Patent US262256, Aug 1882Google Scholar
  64. 803.
    Taylor GT, Thurston PA, Kirsop BH (1979) The influence of lipids derived from malt spent grains on yeast metabolism and fermentation. J Inst Brew 85:219–227CrossRefGoogle Scholar
  65. 834.
    Udo M (2006) The Scottish whisky distilleries. Black & White, EdinburghGoogle Scholar
  66. 865.
    Wang XD, Bohlscheid JC, Edwards CG (2003) Fermentative activity and production of volatile compounds by Saccharomyces grown in synthetic grape juice media deficient in assimable nitrogen and/or pantothenic acid. J Appl Microbiol 94:349–359CrossRefGoogle Scholar
  67. 866.
    Wanikawa A, Hosoi K, Kato T (2000) Conversion of unsaturated fatty acids to precursors of γ-lactones by lactic acid bacteria during the production of malt whisky. J Am Soc Brew Chem 58:51–56Google Scholar
  68. 867.
    Wanikawa A, Hosoi K, Takise I, Kato T (2000) Detection of γ-lactones in malt whisky. J Inst Brew 106:39–43CrossRefGoogle Scholar
  69. 868.
    Wanikawa A, Hosoi K, Shoji H, Nakagawa K-I (2001) Estimation of the distribution of enantiomers of γ-decalactone and γ-dodecalactone in malt whisky. J Inst Brew 107:253–259CrossRefGoogle Scholar
  70. 870.
    Wanikawa A, Shoji H, Hosoi K, Nakagawa K (2002) Stereospecificity of 10-hydroxystearic acid and formation of 10-ketostearic acid by lactic acid bacteria. J Am Soc Brew Chem 60:14–20Google Scholar
  71. 878.
    Watson DC (1981) The development of specialised yeast strains for use in Scotch malt whisky fermentations. In: Stewart GG, Russell I (eds) Current developments in yeast research. Pergamon, New York, p 57–62Google Scholar
  72. 895.
    Williams PJ, Strauss CR (1975) 3,3-diethoxybutan-2-one and 1,1,3-triethoxypropane—acetals in spirits distilled from vitis-vinifera grape wines. J Sci Food Agric 26:1127–1136CrossRefGoogle Scholar
  73. 897.
    Willkie HF, Prochaska JA (1943) Fundamentals of distillery practice. Joseph E. Seagram & Sons, LouisvilleGoogle Scholar
  74. 899.
    Wilson CW III, Shaw PE, Knight RJ Jr (1990) Importance of some lactones and 2,5-dimethyl-4-hydroxyl-3(2h)-furanone to mango (Mangifera indica L.) aroma. J Agric Food Chem 38:1556–1559CrossRefGoogle Scholar
  75. 918.
    Yuan J, Mishra P, Ching CB (2017) Engineering the leucine biosynthetic pathway for isoamyl alcohol overproduction in Saccharomyces cerevisiae. J Ind Microbiol Biotechnol 44:107–117CrossRefGoogle Scholar

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© Springer Nature Switzerland AG 2019

Authors and Affiliations

  • Gregory H. Miller
    • 1
  1. 1.University of California, Professor of Chemical EngineeringDavisUSA

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