29
Biochemistry and Fermentation of
Beer*
R. Willaert
659
Introduction
The Beer Brewing Process
Carbohydrate Metabolism—Ethanol Production
Carbohydrate Uptake
Maltose and Maltotriose 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 Sulphide
Acetaldehyde
Development of Flavor Fullness
Beer Fermentation Using Immobilized Cell Technology
Carrier Materials
Applications of Immobilized Cell Technology (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
References
INTRODUCTION
The production of alcoholic beverages is as old has
history. Wine may have an archeological record
going back more than 7500 years, with the early sus-
pected wine residues dating from early to mid-fifth
millennium BC (McGovern et al. 1996). Clear evi-
dence of intentional wine making first appears in the
representations 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 thou-
sands of years before early thinking man developed
beer brewing and, probably coincidentally, bread
baking (Hardwick 1995). The oldest historical evi-
dence of formal brewing dates back to about 6000
BC in ancient Babylonia: a piece of pottery found
there shows workers either stirring or skimming a
brewing vat.
Nowadays, alcoholic beverage production repre-
sents a significant contribution to the economies of
many countries. The most important beverages
today are beer, wine, distilled spirits, cider, sake,
and liqueurs (Lea and Piggott 1995). In Belgium
(“the beer paradise”), beer is the most important al-
coholic beverage, although the beer consumption
has declined in the last 40 years: from 11,096,717
hL in 1965 to 10,059,513 hL in 2001 (N.N. 2002).
In this time frame, wine consumption doubled from
1,059,964 to 2,215,579 hL. Another trend is the
spectacular increase in water and soft drink con-
sumption (from 5,215,056 hL to 24,628,781 hL).
In this chapter, the biochemistry and fermentation
of beer is reviewed. Firstly, the carbohydrate metab-
olism in brewer’s yeast is discussed. The maltose
metabolism is of major importance in beer brewing
since this sugar is present in a high concentration in
*The information in this chapter has been derived from
“Beer,” in Alcoholic Beverages Manual,©2004 by
Ronnie Willaert. Used with permission.
Food Biochemistry and Food Processing
Edited by Y. H. Hui
Copyright © 2006 by Blackwell Publishing