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BIOL 443 Fermentation Biotechnology Nadine Kallweit (20288655)“Biology of brewing” November 16 th , 2007
Table of Contents
Introduction......................................................................................................................................2Yeast.................................................................................................................................................4Malting.............................................................................................................................................5Mashing............................................................................................................................................5Wort boiling......................................................................................................................................6Hops..................................................................................................................................................6Fermentation.....................................................................................................................................7Traditional Batch Systems................................................................................................................8
Top-fermentation skimming systems and their successors..........................................................8Yorkshire stone squares................................................................................................................8Burton Unions..............................................................................................................................8Lager fermentation.......................................................................................................................8
Continuous Fermentations................................................................................................................9A Specific Example: Cylindrical Fermenting Tower with Baffles to Inhibit Backflow..............9
Beer Treatment...............................................................................................................................11Haze Prevention..........................................................................................................................11Yeast Removal............................................................................................................................11Preservation against microbiological spoilage...........................................................................12Color and Clarity........................................................................................................................12Foam...........................................................................................................................................13Flavor..........................................................................................................................................13
Conclusion and Perspectives..........................................................................................................14References......................................................................................................................................15
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BIOL 443 Fermentation Biotechnology Nadine Kallweit (20288655)“Biology of brewing” November 16 th , 2007
Introduction
Beer was produced without any knowledge of micro-organisms or enzymes thousand of years
ago. Therefore, brewing is often mentioned as a typical example of traditional or old
biotechnology. Whereas, with today’s new developments, there are more possibilities to improve
the beer brewing process. Examples for this are, transgenic barleys, proper starter cultures in
malting, or continuous bioreactors with immobilized yeast cells. [1]
What is beer? First, in the Pocket Oxford Dictionary beer was defined as “an alcoholic beverage
made from fermented malt, etc. flavored with hops, etc.” or it is generously described as “barley
or other grains prepared by steeping and germination or otherwise, for brewing and distilling”.
Finally, beer is any liquid which is a fermented hop-flavored cereal product that contains four
important ingredients, such as, water, malt, hops and yeast. Especially, the German purity law of
1516 allows only water, barley malt and hops to avoid adding cheaper and poorer ingredients.
The newer Beer Tax Law permits also yeast, and for top-fermenting beers other cereals malts. [2],
[3]
In Figure 1 the conventional brewing process is illustrated. The whole process consists of four
steps: malting, word production, fermentation and down-stream processing. [1]
Fig. 1: Simplified Scheme for brewing [1]
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BIOL 443 Fermentation Biotechnology Nadine Kallweit (20288655)“Biology of brewing” November 16 th , 2007
Beer Brewing has a very long history. Before 6000 BC, beer was made from barley in Sumeria
and Babylonia. 2400 BC in Egyptain, barley was crushed, mixed with water, and dried into
cakes. The basis techniques of brewing came to Europe from the Middle East. Hops were in use
in Germany in the 11th century. During the 15th century, they were introduced into Britain from
Holland. In 1420, beer was firstly made in Germany by a bottom-fermentation process. [5]
In 1837, T. Schwann recognized the fungal nature of yeast and he called the organism
´Zuckerpilz´, which is translated into Saccharomyces. [7]
In the 1860s, Louis Pasteur established many of the microbiological practices. Additional, Emile
Hansen, a Danish botanist, devised methods for growing yeasts in culture. He discovered the pure
culture technology, which was a meaningful step to this time. And now, in the 21st century,
brewing is a large-scale industry, with stainless-steal equipment, computer-controlled and
automated operations. [5]
The main aim of brewers and researches is to understand the chemistry, biochemistry and
microbiology of the raw materials as well as the processes of malting, mashing and fermentation. [4]
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BIOL 443 Fermentation Biotechnology Nadine Kallweit (20288655)“Biology of brewing” November 16 th , 2007
Yeast
There is no satisfactory definition for yeasts. They are generally accepted as fungi. Three of the
four groups include yeasts: Ascomycetes, Basidiomycetes and Deuteromycetes.
Saccharomyces cerevisiae is mainly used in brewing processes; therefore, the vegetative growth
cycle and sporulation are very good investigated. Wild sporulation strains are normally diploid.
Different species are named according to their fermentation of sugars, industrial properties or cell
morphology. [8]
Yeast grows in simple media which contain fermentable carbohydrates, such as, glucose,
fructose, sucrose, maltose and maltotriose, ample minerals and accessory nutrients like
potassium, iron, magnesium, manganese, calcium, copper and zinc. A nitrogen source is met by
amino acids. Notably, for a starch-based alcoholic fermentation is the addition of exogenous
enzymes such as α- and β-amylases, β-glucanases and peptidases required. [9], [10]
The main pathway for the fermentation of glucose is the Embden-Meyerhof-Parnas (EMP) route.
It starts with α-D-glucose, which is phosphorylated to hexose monosaccharide glucose. The
major step for fermentation is decarboxylation of pyruvic acid to acetaldehyde and carbon
dioxide using the enzyme pyruvate decarboxylase. With the help of alcohol dehydrogenase and
NADH2 as cofactor the produced acetaldehyde is reduced to ethanol. [9]
Saccharomyces cerevisiae is a glucose-sensitive yeast type. That means, respiration is repressed
in the presence of small concentration of free glucose in the medium. Brewers´ wort typically
contains approximately 1% glucose. [9]
Many strains of Saccharomyces cerevisiae can attain ethanol concentrations of 12-14%. Thus, the
development of alcohol tolerant yeast strains, for increasing the plant productivity and decreasing
distillation costs, is becoming increasingly necessary. The knowledge about temperature- and pH-
optimum to achieve high cell growth and fermentation activity is not neglectable as well. [10]
A ´good´ brewing yeast should have: rapid fermentation rate, an efficient utilization of maltose
and maltotriose, an ability to withstand the stresses, an ideal flocculation character and good
´handling´ characteristics. [12]
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BIOL 443 Fermentation Biotechnology Nadine Kallweit (20288655)“Biology of brewing” November 16 th , 2007
Malting
The malting process consists of four steps. First, the freshly harvested barley goes through a
period of rest and ´sweating´ from about 30 to 60 days. Within the second step the barley kernels
are immersed in a tank of water at a temperature from about 50°F to 65°F. During the steeping
process, respiration of the kernels becomes noticeable. In the third step, the germination, the
respiration and growing process is accelerated. The barley is removed from the steeping tank to
appropriate conditions of temperature, moisture and oxygen. During this step the barley kernel
begins to ´chit´ at the base of the kernel by showing a white tip. The third step has three
important functions. With the aid of various hydrolytic enzymes including α- and β-amylases, β-
glucanases and peptidases, starches are converted to dextrins and sugars, insoluble proteins to
soluble proteins and a source of desirable taste and aroma is provided. In the fourth step, the
batch of green malt is conveyed to the drying or kiln compartments. Finally, the dried malt is
cleaned and stored. [6], [10], [11], [15]
The malting process is very important because the starch of the endosperm becomes mellow,
enzymes are developed, and the flavor and color ingredients are formed. [6]
Mashing
During the mashing process, the raw materials are extracted and carbohydrates and other
nutrients are solubilized and enzymatically hydrolyzed. The mash is started at a relatively low
temperature, e.g. 122°F, and held to allow sufficient time for the proteinase and β-glucanase to
act. After that, the temperature is raised up to 149°F level which is optimum for amylolysis.
β-Glucanase is an endo-enzyme, which acts on the cell walls of the endosperm and it keeps the
viscosity down. The proteinase is needed to release a range of nitrogenous compounds.
Aminoacids are needed for the yeast growth. Large polypeptides are the most important surface-
active compounds in beer foam. At higher temperatures α-amylase solubilises both the amylose
and the amylopectin compounds of starch. β-Amylase renders the straight-chain carbohydrates to
maltose and the branched-chain compounds to mixtures of maltose and small dextrins. It is
essential that all the added enzymes are inactivated during boiling. It is common, that the grist
contains a small proportion of unmalted grain, say 5-20%. In this case, some brewer may use
industrial enzymes as aids in the mashing process. [10], [11]
There are three systems of mashing: first, infusion mashing, evolved in Britain, where beer wort
is sterilized by boiling before the yeast is added, secondly, decoction mashing, the typical lager
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mashing system of continental Europe, which works at several temperatures to enable enzymatic
or bacterial action and thirdly, American development of combination or cooker mash, using
steam and the boiling adjunct to accomplish these temperatures. In particular, the higher the
temperature of conversion the faster the reaction, but the lower the quantity of sugar formed. [2], [6],
[10]
In the sparging process, the sweet wort is drawn into a vessel called an underback, and washed
with water at a temperature of about 167°F. The effect is to dissolute all residual soluble
materials and to inactivate any remaining enzymes. Silicates and tannins are concentrated in the
poorly buffered final sparge and phospholipids can be removed. The average gravity value for
final spargings from British mash tuns is 1.0001, for continental Europe 1.004 and for North
America 1.008. The specific gravity shows how many times heavier than water the liquid of
interest is. For instance, 1.050 SG wort is 1.05 times heavier than equal volume of water. [2]
Wort boiling
The medium which is fermented to produce beer or ale is called wort. [6]
The objectives of wort boiling are: inactivation of any enzymes, sterilization of the wort,
completion of ionic interactions, concentration of the wort, denaturation and precipitation of
proteins, dissolution of any additional sugars, isomerization of hop α-acids and volatilization and
removal of unwanted flavor components. [2], [10]
A small portion of malt is first prepared by boiling in a deep closed vessel (cocker). While the
main portion is mixed with water in a large vessel and heated. The cooker mash is then added, the
temperature increases and the conversion of starch to carbohydrates of lower molecular weight
takes place. In the usual European practice, the cooker mash may be omitted. [15], [6]
Hops
Important compounds present in the hope cone are the resins, the tannins and the essential oils.
There are although some other ingredients like pectins, hop waxes and water. The essential oils
are responsible for the beer flavor, known as hop character. The tannins combine with protein
during normal kettle boiling. Not precipitated tannins contribute to haze formation in the finished
beer. There are two classes of hop resins, the hard and the more important soft resins. The other
compounds like pectins, waxes and water have an improving effect on beer stability, the choice
of extraction and isomerization conditions and the foam retention of the beverage. [6], [12], [15]
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Fermentation
After adjustment to the desired specific gravity, the clarified wort is cooled to about 50°F.
Then it is often aerated with sterile air and injected with yeast for the fermentation stage of the
process. This process is called ‘pitching’. For a period of one or two weeks, the yeast ferments or
breaks down the fermentable sugars of the wort into alcohol and carbon dioxide. The
fermentation process is performed either batch or continuous. [15]
After the fermentation, when more than 95% of the fermentable sugars in a wort are converted,
the yeast cells tend to flocculate and precipitate to form a sediment layer along the bottom of the
tank (Saccharomyces karlsbergensis) or rise to the surface (Saccharomyces cerevisiae). This
process is usually enhanced by cooling the liquid in the fermenting vessels down to about 40°F.
The fermentation of the sugars in the wort is finished after five or nine days. Three or six days are
additionally required for cooling and further yeast sedimentation. The freshly fermented beer is
then removed from the fermentation vessel. After filtration, pasteurization and other treatments,
the beer is ready for dispatch. [9], [15]
According to the system of fermentation is employed, beers may be classified in three different
groups. First, there are bottom-fermented beer types, as the German Lager Beers (Pilsner,
Wiener, Muenchener), the American Lager Beers and the American Steam Beers. On the other
hand, the British beer types like Ale, Porter and Stout and the German Weiss Beer belong to the
top-fermented beer types. The last group comprises the spontanously-fermented beer types, for
instance, the Belgian beers, Lambic and Faro. Basically, the fermentation process consists of two
main parts. During the ‘main fermentation’, maltose is splitted at relative high temperatures. In
the further step, called the ‘after fermentation’, the residual malto-dextrin is digested at lower
temperatures. [14]
There are seven main factors, which influence the fermentation performance and beer quality. 1.
choice of yeast strain, 2. condition of the yeast at the time of pitching, 3. amount of yeast added
to the wort, 4. distribution of the yeast in the fermentation wort throughout the fermentation, and
the size and geometry of the fermentation vessel, 5. Aeration, 6. Wort composition and pH and 7.
Fermentation temperature and pressure. [7]
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BIOL 443 Fermentation Biotechnology Nadine Kallweit (20288655)“Biology of brewing” November 16 th , 2007
Traditional Batch Systems
Top-fermentation skimming systems and their successors
This is the traditional ale fermentation in a modern enclosed vessel, usually constructed of
stainless steel, with wall attemperation, collection points for carbon dioxide and facilities for in-
place cleaning. The skimming process is difficult, very expansive but very fast. [2]
Yorkshire stone squares
There is a deck or platform slightly below the level of the wort which represent the skimming
system. The yeast collects on the deck and leaves the beer. Therefore is highly flocculent yeast
required. [2]
Burton Unions
Fermenting wort is transferred after about 36 hours to a row of casks each provide with a swan-
neck tube which opens into a common inclined through above the set of casks. [2]
Lager fermentation
Lager Fermentation and ale fermentation differ in the species of yeast used and in the
fermentation temperature, which is not allowed to exceed 50°F. [2]
The starting yeast count is 107 cells/ per ml, the initial carbohydrate concentration is 12°P, the
specific gravity is 1.050 and the initial temperature is about 50°F. During the lager fermentation
process the temperature is rising and cooling is required. The formation of many acids, such as,
acetic acid, drops the pH about 1 unit from an initial value of approximately 5.2. The whole
fermentation takes about 10 days. 70-80% of the wort carbohydrates are converted to ethanol.
The residual 20-30% consists mainly of higher dextrins or limit dextrins which are not
susceptible to attack by malt amylases. With the help of microbial amyloglucosidase, dextrins can
be hydrolyzed. [10]
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BIOL 443 Fermentation Biotechnology Nadine Kallweit (20288655)“Biology of brewing” November 16 th , 2007
Continuous Fermentations
A Specific Example: Cylindrical Fermenting Tower with Baffles to Inhibit Backflow
During the cylindrical fermenting tower process, is used by M.G. Royston (U.S. Patent, 1967),
the hopped wort moves continuously into and through a fermentation vessel. The vessel
construction inhibits the backflow of the wort and prevents the intermixing of the progressively
fermented wort with the new one. [15]
The high overall rate of fermentation leads to lower costs and low floor space requirements. For
instance, the vessel has a measured value of 1 foot diameter by 20 feet high and the whole
fermentation is completed after 3 to 5 hours. [15]
Fermentation vessel comprises a cylindrical tower (1). At the lower end of the tower (1), inlet
means (2) are provided for introducing the hopped wort, so that the inflow of wort to the tower is
distributed over the cross-selection of the tower. A perforated baffle (3) is may be provided.
For the introduction of air or oxygen at the base of the tower (1), a sparging ring (4), is may be
provided with connection to a source of air or oxygen. The spaced baffle (5) serves to inhibit
backflow of the ascending liquid and assists in maintaining a forward flow of the liquid. When
the apparatus is in full operation, the cell concentration of the yeast will be high and will form a
yeast plug above the level of the perforated wort-distributing baffle (3). The yeast growth
increases in direct proportion to the sugar conversion taking place as the process proceeds. [15]
Fig.2: Cylindrical Fermenting Tower [15]
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BIOL 443 Fermentation Biotechnology Nadine Kallweit (20288655)“Biology of brewing” November 16 th , 2007
1: cylindrical tower
2: wort inlet
3: perforated baffles
4: sparging ring
5: spaced baffles
6: separating zone
7: beer outlet
8+9: water jacket
The yeast culture is isolated on a wort agar plate and finally transferred in liquid wort in a 2-liter
Pasteur flask. There is a sterile connection between the Pasteur flask and a sample point on the
tower. The flow introduction is continued and aerated with sterile air. The fermentation
temperature is held constant by 60°F. [15]
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BIOL 443 Fermentation Biotechnology Nadine Kallweit (20288655)“Biology of brewing” November 16 th , 2007
Beer Treatment
After the primary fermentation beer is said to be ´green´. It must undergo a number of changes,
including the elimination of certain volatile fermentation products, super-saturation which carbon
dioxide, separation of yeast cells, and removal of some of the polyphenolic and other materials. [2], [7]
The maturation process is carried out in close containers. Beer treatment involves six processes:
1. Carbonation, 2. Flavor/aroma changes, 3. Additions of coloring and flavoring materials, 4.
Stabilization against non-biological haze and flavor change, 5. Clarification, 6. Biological
stabilization. [2]
Haze Prevention
Haze consists of polyphenolic compounds and polypeptides of high molecular weight, together
with trace quantities of metallic ions. There are two types of protein-tannin haze; chill haze, when
beer is cooled to about 37,4°F and it redissolves on warming, and the permanent haze, which is
formed at temperatures of und does not redissolve in beer. [2]
A widely used method of delaying the onset of non-biological haze is to add an endoprotease to
the beer in the final stages of preparation. The most used enzyme is papain which acts through
cleavage of the polypeptide chains present in the beer. A critical concentration is necessary
before the polymerising polyphenol/polypeptide system becomes visible as a haze. To achieve
the best results, the enzyme should be kept in a reducing environment free from metal ions and be
added to the beer as late as possible. [11]
Yeast Removal
Centrifugation and filtration are common procedures to remove residual yeast. Care must be
taken to avoid any rise in temperature lest the material deposited previously redissolve. Filtration
occurs through pulp, kieselguhr, composite filter sheets or more rarely, through cellulose ester
membranes. [2]
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BIOL 443 Fermentation Biotechnology Nadine Kallweit (20288655)“Biology of brewing” November 16 th , 2007
Preservation against microbiological spoilage
Wort has a natural resistance to many infectious microorganisms because of its low and acidic pH
between 5.0 and 5.4. [5] Nevertheless, the microbiological spoilage of beer is a big problem. [11]
Acetic-acid bacteria (Acetobacter spp. and Acetomonas spp.), the lactic acid-forming
(Lactobacillus spp. and Pediococcus spp.) and Zymomonas anaerobia are the commonest beer
spoilage organisms. [2]
Pasteurization is very expensive. More important is the preservation with synthetic organic
chemical compounds because there is a reduced amount of bottle breakage. The beer contains
both yeast and bacteria, thus necessitating the use of a fungistatic chemical and a bacterostatic
chemical. As for instance, Imidazoline Hydrobromide, p-Hydrocybenzoic Acid Esters, 2,3
Epoxypropanol or Diethanolamides are effective chemicals. [15]
In the pasteurizing process the beverage is heated to the required temperature and is then filled
into sterilized transport containers or small containers in which the liquid is kept under a certain
pressure. The flash pasteurization, in which the entering beer flows countercurrent, first to hot
pasteurized beer and then to steam, until it reaches a holding area where it remains for 20sec at
158°F, is the commonest procedure for bulk beer. Tunnel pasteurizers are used with bottled or
canned beer. The Containers travel through an enclosed area in which they are sprayed with hot
water, left for an appropriate time, and then treated with a cold-water spray. This procedure
reduces the incidence of bottle breakage. There are also some other methods of pasteurization
such as the plate pasteuriser. [2, [15], [16]
Color and Clarity
Melanoidins from kilned malt introduce color to beer. Roasted barley is also used as a source of
color, which makes a characteristic contribution to the flavor and head retention of stout. The
control of beer clarity is now not difficult. Metal hazes, oxalate hazes, Silica hazes can occur. [2]
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BIOL 443 Fermentation Biotechnology Nadine Kallweit (20288655)“Biology of brewing” November 16 th , 2007
Foam
The foamy head on a cool glass is an honored characteristic. In the formation of the foam
aromatic constituents and hop flavoring materials are adsorbed. That is why the foam has a great
importance to the consumer to prolong the taste enjoyment of the beverage. The amount of foam
is directly related to its carbon dioxide content. [15], [2]
The foam of different beers collapses at different rates. This process is not completely understood
but the size of the gas bubbles is an important factor. The goal is to produce a beer with a long-
lasting head of foam. Detergents have a harmful effect on beer foam. [15]
Beside the foam stability, the foam adhesion, or ´cling´, is fundamental. This is the material
deposited on the wall of the drinking glass as beer foam collapses. The iso-α-acids are implicated
in the ability of beer foam to cling to the glass. To enhance the ´cling´ chemical compounds like
Polysaccharide, Zinc plus Chelating Agents or Propylene are added to the beer. [2],[15]
The condition of the glass containing the beer is highly important in relation to the head stability
and also the presence of traces of grease or dishwashing agent is a crucial factor. [2]
Flavor
The beer flavor is examined either within detailed chemical analysis or within some individual
attribute of its impact on the human sense organs. A lot of different groups of flavor compounds
have been reported. [2]
Beer flavor is never constant because during the aging process subtle changes occur. During the
first stage a papery, or cardboard-like flavor, causes by the formation of volatile, long chain,
unsaturated carbonyls, develops and the product decreased in fruity/estery and floral character.
During the next step the flavor becomes bready, sweeter and toffee-like. The sweetness of a beer
is, naturally, related to its sugar content. In the last step the beer attains a winey, sherry-like
character and the bitterness decreases. Generally, ales become a sweet with a molasses-like,
cloying character. However, darker, heavier beers have a better flavor stability. Additional,
oxidation is the principal cause of flavor instability, therefore brewers try to minimize the
introduction of oxygen into the product after the fermentation and during packaging.[2], [17], [18]
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BIOL 443 Fermentation Biotechnology Nadine Kallweit (20288655)“Biology of brewing” November 16 th , 2007
Conclusion and Perspectives
About Biotechnology is often told, that it was ´as old as civilization itself´. But now malting and
brewing is evolved into a multi-million dollar industry. [19]
As aforementioned there are new methods developed, so that the brewing process has been
reformed during the last years. Malting barley should give higher yields on the field, high extract
yield in the brewhouse, suitable activities of many enzymes, free as possible of harmful micro-
organisms etc. Therefore, transgenic barleys become more and more important. The main interest
has been to insert a thermostable β-glucanase gene into the genome. The produced heat-stable β-
glucanase reduces the viscosity of the wort in mashing experiments. [1]
Otherwise, a novel method is using lactic acid bacteria or Geotrichium Candidum as starter
cultures in malting to reduce the fungal contamination and to improve the malt quality. They also
restrict the growth of harmful Gram-negative and –positve bacteria. [1]
A key compound of maturation is diacetyl, which is formed as an unpleasant by-product during
the main fermentation. With the help of the enzym α-acetolactate decarboxylase, diacetyl is
reduced to acetoin and further to 2,3-butanediol. The taste threshold of acetoin is much higher
than that of diacetyl. But unfortunately brewer’s yeast lacks the enzyme α-acetolactate
decarboxylase. To solve this problem there are either exogenous enzymes during the process
added or the encoding gene for α-acetolactate decarboxylase is integrated into the yeast genome. [1]
Finally, big breweries produce 500 million liters of beer in a year. They must be efficient to
survive but the quality of the beer must be good and uniform. As well, they are interested in all
technical novelties making the processes cheaper, easier and safer. But also small breweries are
very popular although they are producing with higher costs and limited quantities. [1]
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BIOL 443 Fermentation Biotechnology Nadine Kallweit (20288655)„Biology of brewing” November 16 th , 2007
References
[1] Matti Linko et al., 1998
Recent advances in the malting and brewing industry
Journal of Biotechnology 65, Finland: 85–98
[2] A. H. Rose, 1977
Economic Microbiology - Alcoholic Beverages
Volume 1, Academic Press, London/
M. Macleod
Beer
Department of Brewing and Biological Sciences, Edinburgh, Scotland: 43-137
[3] http://www.garshol.priv.no/blog/4.html
[4] Tor-Magnus Enari, 1999
From Beer to Molecular Biology - The Evolution of Industrial Biotechnology
Fachverl. Carl, Nürnberg: 11-31
[5] http://www.britannica.com/eb/article-9106004/beer
[6] L. A. Underkofler and R. J. Hickey, 1954,
Industrial Fermentations/
R. I. Tenney
The Brewing Industry
Volume 1, Chemical Publishing Co., INC., New York: 172-195
[7] J.S. Hough et al., 1982
Malting and Brewing Science - Volume 2 Hopped Wort and Beer
2nd Edition, University Press, Cambridge
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BIOL 443 Fermentation Biotechnology Nadine Kallweit (20288655)„Biology of brewing” November 16 th , 2007
[8] F.G. Priest and I. Campbell, 1996
Brewing Microbiology
2nd Edition, Chapman and Hall, London/
I. Campbell
Chapter 1: Systematics of yeasts: 1-11
[9] F.G. Priest and I. Campbell, 1996,
Brewing Microbiology
2nd Edition, Chapman and Hall, London/
T.W. Young
Chapter 2: The biochemistry and physiology of yeast growth: 12-42
[10] Ward, O.P., 1989
Fermentation Biotechnology: Principles, Processes and Products
Pp. i-xii, 1-227, Prentice-Hall
[11] G. G. Birch and M.G. Lindley, 1985
Alcoholic Beverages
Elsevier Applied Science Publishers LTD, London/
J.C. Slaughter
2. Enzymes in the Brewing Industry
Department of Brewing and Biological Sciences, Edinburgh, UK: 15 – 28
[12] G. G. Birch and M.G. Lindley, 1985
Alcoholic Beverages
Elsevier Applied Science Publishers LTD, London/
A.L. Whitear and R. Sharpe
3. Hop Constituents and Beer Character
Whitbread and Company plc, Luton, UK: 29 - 50
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[13] F.G. Priest and I. Campbell, 1996
Brewing Microbiology
2nd Edition, Chapman and Hall, London/
J.R.M Hammond
Chapter 3: Yeast genetics: 43-82
[14] R.Wahl and M. Henius, 1902
American Handbook of the Brewing, Malting and Auxiliary Trades
2nd Edition, Chicago
[15] M.Gutscho, 1969
Alcoholic Malt Beverages
Noyes Development Corporation, United States
[16] G. B. Härnulv et al., 1995,
Beer Pasteurization – Manual of Good Practice
EPC Technology and Engineering Forum, Getränke - Fachverlag Hans Carl, Netherlands
[17] B. H. Gump and D. J. Pruett, 1992
Beer and Wine Production – Analysis, Characterization, and Technology Advances ACS
Symposium Series
San Francisco/
N. J. Huige
Chapter 5 - Progress in Beer Oxidation Control
Miller Brewing Company, Milwaukee
[18] B. Vanderhaegen et al., 2007
Aging characteristics of different beer types
Food Chemistry 103, Centre for Malting and Brewing Science, Belgium: 404-412
[19] D. Mc Elroy and J. Jacobensen, 1995
What is Brewing in Barley Biotechnology?
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BIOL 443 Fermentation Biotechnology Nadine Kallweit (20288655)„Biology of brewing” November 16 th , 2007
Biotechnology Vol 13, Nature Publishing Group, Australia: 245-249
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