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Biochemistry of Beer Fermentation
Ronnie Willaert
Introduction
The Beer Brewing Process
Carbohydrate Metabolism—Ethanol Production
Wort Carbohydrates Uptake and Metabolism
Maltose and Maltotriose Metabolism
Glycogen and Trehalose Metabolism
Wort Fermentation
Metabolism of Bioflavoring by-Products
Biosynthesis of Higher Alcohols
Biosynthesis of Esters
Biosynthesis of Organic Acids
Biosynthesis of Vicinal Diketones
Secondary Fermentation
Vicinal Diketones
Hydrogen Sulfide
Acetaldehyde
Development of Flavor Fullness
Beer Fermentation Using Immobilized Cell Technology
Carrier Materials
Applications of ICT in the Brewing Industry
Flavor Maturation of Green Beer
Production of Alcohol-Free or Low-Alcohol Beer
Production of Acidified Wort Using Immobilized
Lactic Acid Bacteria
Continuous Main Fermentation
Acknowledgments
References
Abstract:The carbohydrate metabolism and flavor formation dur-
ing yeast primary and secondary fermentation (maturation) is re-
viewed. Carbohydrate metabolism and ethanol production during
the primary fermentation is discussed firstly. Next, the metabolism
of the bioflavoring by-product formation, that is, higher alcohols,
esters, organic acids, and vicinal diketones, is elaborated. The next
step of the fermentation process is the maturation process where
the vicinal diketones, acetaldehyde, and hydrogen sulfide concen-
tration needs to be reduced to acceptable levels. The chapter is
concluded with a discussion about the use of immobilized cell tech-
nology to intensify the fermentation process and its impact on flavor
production.
INTRODUCTION
The production of alcoholic beverages is as old as history.
Wine may have an archeological record going back more than
7500 years, with the early suspected wine residues dating from
early to mid-fifth millenniumbc(McGovern et al. 1996). Clear
evidence of intentional winemaking first appears in the repre-
sentations of wine presses that date back to the reign of Udimu
in Egypt, some 5000 years ago. The direct fermentation of
fruit juices, such as that of grape, had doubtlessly taken place
for many thousands of years before early thinking man devel-
oped beer brewing and, probably coincidentally, bread baking
(Hardwick 1995). The oldest historical evidence of formal brew-
ing dates back to about 6000bcin ancient Babylonia is a piece
of pottery found there, which shows workers either stirring or
skimming a brewing vat.
Nowadays, alcoholic beverage production represents a sig-
nificant contribution to the economies of many countries. The
most important beverages today are beer, wine, distilled spir-
its, cider, sake, and liqueurs (Lea and Piggott 1995). In Bel-
gium (“the beer paradise”), beer is the most important alcoholic
beverage, although the beer consumption declined in the last
40 years: from 11,096,717 hL in 1965 to 9,703,000 hL in 2004
(NN 2005). In this time frame, wine consumption doubled from
1,059,964 to 2,215,579 hL. Another trend is the spectacular in-
crease in waters and soft drinks consumption (from 5,215,056 to
26,395,000 hL).
In this chapter, the biochemistry and fermentation of beer is
reviewed. First, the carbohydrate metabolism in brewer’s yeast
is discussed. The maltose metabolism is of major importance in
beer brewing, since this sugar is in a high concentration present in
wort. For the production of a high-quality beer, a well-controlled
Food Biochemistry and Food Processing, Second Edition. Edited by Benjamin K. Simpson, Leo M.L. Nollet, Fidel Toldr ́a, Soottawat Benjakul, Gopinadhan Paliyath and Y.H. Hui.
©C2012 John Wiley & Sons, Inc. Published 2012 by John Wiley & Sons, Inc.
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