Review: Beer Production

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Aroh, Kenechukwu. Uchechukwu
September, 2018.
Kenearoh12@gmail.com

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A. INTRODUCTION
Beer is the most popular and most consumed alcoholic drink around the world, and it is also one
of the oldest. Beer drinking and brewing have been part of human activities since the dawn of civilization.
The first beer was basically made from grain, water and spontaneous fermentation due to wild yeast present
in the air, just before bread was invented (Campbell, 2017). It has been reported that the Egyptians were the

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first to document the brewing process around 5000 B.C, it is also believed that the first brewers were part
of primitive cultures of Mesopotamia. The birth of modern beer was during the early Middle Ages, when
German monks introduced hop as a bittering and flavouring agent. Beer brewing used to be on a domestic
scale, but with the arrival of the Industrial revolution, its production moved to mass manufacture, allowing

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beer to be produced on a much larger scale (Sánchez, 2017).

Beer is produced commercially by the controlled fermentation of wort, a liquid-rich in sugars,

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nitrogenous compounds, sulphur compounds and trace elements extracted from malted barley. Fermentation is
the process by which glucose is converted to ethanol and carbon dioxide and is expressed chemically as:

C6H12O6 + 2PO43- + 2ADP → 2C2H5OH + 2CO2 + 2ATP

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Behind this simplified chemical reaction is a series of complex biochemical reactions. These
reactions (known as the ‘Glycolytic pathway’ or ‘Embden-Myerhof-Parnas pathway’) involve a number of
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enzymes and the reactions take place anaerobically inside the cells of brewing yeast (Campbell, 2017).
Breweries carry out this fermentation by a continuous process in which the beer moves through a series of
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stirred vessels for a period of 40 to 120 hours. After the ethanol has formed the beer is transferred to
maturation vessels and the flavour is naturally refined. Following this the product is developed into a variety
of different brands (Campbell, 2017). Brewing beer involves microbial activity at every stage, from raw
material production and malting to stability in the package. Most of these activities are desirable, but others
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represent threats to the quality of the final product and must be controlled actively through careful
management (Bokulichn and Bamforth, 2013).

B. BEER
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Beer is an alcoholic beverage made by brewing and fermentation from cereals, usually malted barley, as
well as maize and flavored with hops and the like for a slightly bitter taste. Barley beer is said to have many health
benefits if it is consumed in moderate amounts. Some of these interesting health benefits include anti-cancer
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properties, reduced risk of cardiovascular diseases, increased bone density, diabetes, prevention of anemia,
hypertension, anti-aging properties, gallstones, prevention of dementia and coronary disease, aids digestive
system, kidney stones and osteoporosis, stress buster and diuretic (Ore, Mironov and Shootov, 2018).

This preprint research paper has not been peer reviewed. Electronic copy available at: https://ssrn.com/abstract=3458983
 There are several types of strains of microorganisms used for the production of beer, such as lactic
bacteria and yeast, but the yeast strain Saccharomyces cerevisiae is used as it be found easily and cheaply. The

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manufacturing processes of beer essentially involves treatment of grains, malting or germination, mashing or
extraction with water, filtration and fermentation. Germination process is halted at desired malt quality, green
brown malt is converted to stable, storable product, colour and flavor are also developed, enzymes are stabilized
and preserved, and unwanted flavours are removed. The objectives of mashing are solubilization and dissolution

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of grain components, breakdown of grain cell wall structure extraction and hydrolysis of starch, sugars, proteins
and non-starch polysaccharides and fermentable sugar profile is established. During the fermentation, alcohol
level is established, flavor profile of beer is established and carbonation level is established. At the end of
fermentation, yeast flocculates and can be easily separated. Cold maturation temperatures will influence beer
clarity. Using a systematic procedure to solve material balance problems, mass balances in all six steps in this

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production process were solved (Ore, Mironov and Shootov, 2018).

Brewing of beer has been used in ancient history and has continued to be consumed in the present day

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with relatively few changes to the basic recipe. Malted barley is the main ingredient, which, when milled and
heated in water to extract its nutrients, provides a nourishing sugar and protein-rich solution named wort
(pronounced as wert). It is an ideal medium in which yeast may grow and ferment. In recent times hops is added
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to the boiling wort as it was discovered that hops had anti -bacterial properties which preserved the wort and
fermented beer, giving the beer a refreshing bitter taste (Campbell, 2017).

For many years the only known method of fermenting beer was a slow batch fermentation process
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carried out in a single fermentation vessel. This method had disadvantages in economic and quality aspects. The
slow fermentation times meant that large numbers of tanks were required to house all the fermenting batches of
beer (resulting in high costs of vessels and the associated costs for holding these vessels at the required
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temperatures and testing the quality of each batch). In addition, there was no guarantee that the beer would have
a consistent flavor. Recently, continuous fermentation is used which involves recycling part of the fermented
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beer back to the wort at the start of the fermentation process, the result is a continuous flow of beer out the other
end of the process. The wort brewing stage in a continuous system may be carried out at a time appropriate for
the brewery. Continuous fermentation employs a system of cold wort storage; the boiled wort is chilled to 0˚C
(the wort does not freeze at this temperature because of its high sugar content) and held in storage tanks where
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protein material (which would otherwise make the beer appear cloudy or ‘hazy’) precipitates out. The wort
remains in the storage vessel until it is required to be steadily transferred to the fermentation. One wort storage
tank will continuously feed into the fermentation for several days (Campbell, 2017).
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The flavour and aroma of any beer is, in large part is determined by the yeast strain employed together
with the wort composition. In addition, yeast properties such as flocculation, fermentation ability (including
the uptake of wort sugars, amino acids, small peptides, and ammonium ions), osmotic pressure, ethanol
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tolerance, and oxygen requirements have a critical impact on fermentation performance. Proprietary strains,
belonging to individual breweries, are usually jealously guarded and conserved (Stewart, 2016). With the
genetic manipulation of yeasts, numerous varietal strains have been bred, this, along with modifications in the
brewing process have led to different types of beers. Those most often seen beers include:

This preprint research paper has not been peer reviewed. Electronic copy available at: https://ssrn.com/abstract=3458983
  Lager: Beers made with yeast that settle on the bottom (Saccharomyces carlsbergensis) of the container used.
Thus, all the yeast and other material settles on the bottom which results in a clear beer.

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 Pilsner: A colorless lager beer originally brewed in the city of Pilsen. Water used for this style of beer tend to be
harder, with a higher calcium and magnesium content than water used for lager. The color of pilsner is also lighter
than that of lager beer.
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Ale: Beers made with yeast that floats (Saccharomyces cerevisiae) to the top of the brewing vats, resulting in a
cloudier beer. They tend to have a higher alcohol content than lagers.
 Porter: A very dark ale. The darker color and special flavor comes from toasting the malt before brewing. This
usually results in a stronger taste and higher alcohol content.
 Stout: A very dark, almost black ale. The dark color and roasted flavor is derived from the roasted barley, and/or

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roasted malt (Wong, 2003)

Beer like any fermented food, is a microbial product. Microbial activity is involved in every step
of its production, defining the many characteristics that contribute to final quality. While fermentation of

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cereal extracts by Saccharomyces is the most important microbial process involved in brewing, a vast array
of other microbes affects the complete process (Bokulichn and Bamforth,2013). The diagram below shows
an overview of bacterial and fungal species previously reported at all major stages of beer production.
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FIG 1: Microbiota of malting and brewing (Bokulich, Bamforth and Mills, 2012).

C. PROCESS OF BEER PRODUCTION: BREWING

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Brewing is a huge-scale complex process that transforms water, grains and hop to produce what we call
beer, and this is achieved mostly with the help of yeast. Basically the large variety of beer is due to the different
conditions (temperature, kind of grain, etc.) established during the stages of production (Sánchez, 2017). The body
of the beer is provided by barley, more specifically barley malt and in general, a few hundreds of grams are used

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 for one litre of beer. The malt may be partly substituted by starch-rich adjuncts such as rice, corn or wheat. When
a slurry of barley malt and brewing water (called ‘mash’) is heated at a temperature around 60˚C, the malt

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enzymes, mainly amylases but also proteases, degrade starch and proteins, leading to a mixture of sugars and
peptides or amino acids. For that purpose, barley must be subjected to a controlled germination, during which
these enzymes are formed in the barley grain prior to mashing. Such germinated barley is known as barley malt.
(Keukeleire,2000).

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The starch-to-sugar conversion is stopped by heating. Depending on the conditions (time, temperature),
pale or amber-coloured or even dark malts are obtained, the colour being due to caramelization of sugars. It is
important to notice that the colour of beer is derived from the colour of the malt(s) used. Furthermore, it is obvious
that coloured malts exhibited a distinct taste, which often is characteristic of particular dark beers. After filtration,

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the sugar solution in brewers’ jargon called ‘wort’ is transferred to the brewing kettle, where it is boiled during at
least one hour with the addition of hops (Humulus lupulus L.). The amount of hops needed is only a fraction of
the substantial quantities of malt used in the brewery. Besides the formation of insoluble complexes with proteins

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and polypeptides (contributing to the stability of beer) hops sterilize the wort solution. The most important asset
of hops is the bitter taste conferred to, particularly blond beers. Furthermore, hops are necessary for the
stabilization of beer foam (Keukeleire, 2000).
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After cooling and removal of spent hops, the liquid known as ‘hopped wort’ is pumped to the
fermentation vessels and yeast is added under aeration for growth. During the anaerobic phase yeast cells convert
sugars to ethanol and carbon dioxide. Depending on the temperature during fermentation and the nature of yeast
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collection at the end of the fermentation period, beers are distinguished as being produced by ‘bottom
fermentation’ or ‘top fermentation’. Yeast strains, appropriate for bottom-fermented beers (Saccharomyces
carlsbergensis), are active below 5˚ C and they settle to the bottom of the fermenter after production of about 5%
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ethanol. Conversely, yeasts typical for the production of top-fermented beers (Saccharomyces cerevisiae), operate
at ambient temperature and resist higher concentrations of ethanol up to 12%. When the activity stops, the yeast
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cells collect to the top as a dense foam (Keukeleire, 2000)

A typical fermentation takes about one week thereby delivering a so-called ‘green beer’ or ‘young beer’
which is not drinkable, as a number of offending (bad taste and smell) compounds are formed during fermentation.
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Consequently, beers need a maturation or lagering period of several weeks at about 0˚C, during which the
unwanted components are slowly decomposed. Only after the content has decreased below critical values can beer
be packaged. For prolonged conservation beers may be pasteurized. Special beers often require a slow (several
months) second fermentation, usually in oak kegs, to generate sour flavours. (Keukeleire, 2000). Below, a
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simplified diagram of beer production illustrates the whole process and the explanation of each stage is given
below:
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This preprint research paper has not been peer reviewed. Electronic copy available at: https://ssrn.com/abstract=3458983
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FIG 2: Simplified diagram of Beer production (Sánchez, 2017) rr
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In the production of beer there are 8 stages of brewing, these are represented schematically below:
1. STAGE ONE: MALTING & MILLING
The barley has been modified to malt by the maltster and its milled immediately prior to use.
Each beer has its own formulation with regards to the blend of different malts.
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2. STAGE TWO: MASHING & LAUTERING

 The tank has a mixing paddle to ensure that the mix of water and malt is constantly agitated during mashing. The
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malt is mixed with hot water to allow the starch to be converted into sugar by enzymes.
 The temperature of the mixture is crucial, as the type of sugar converted is temperature dependent: some sugars
are fermentable, while others are non-fermentable, giving richness feel to the finished beer.
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 There are steam jackets on the outside of the tanks to heat the mash in stages. This mashing process takes about
two hours.
 The sweet liquid, now called wort, can pass through the screens of the false bottom while the grain stays behind.
The wort is then pumped into the kettle. This process takes about 1.5 hours
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 Afterwards, the spent grain is collected for cattle feed.

3. STAGE THREE: BOILING & WHIRLPOOL
· Once all the wort is in the kettle, the liquid is boiled for 60-90 minutes. This ensures that the wort is sterile.
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Boiling also evaporates some water, concentrating the wort and intensifying the colour somewhat.
· Hops are added to the kettle at the start of boil for bitterness and at the end of boil for aroma and flavour
· The wort is then recirculated through a whirlpool effect which ensures the residual hop product and proteins and
enzymes are coagulated, and settle out of the liquid as a sludge called trub. This trub is partially removed from
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the bottom of the kettle which ensures the bitter wort is nice and clear when transferred through to the next stage.
4. STAGE FOUR: COOLING THROUGH HEAT EXCHANGER
 Once boiling is complete, the wort is cooled to around 20 ˚C through a heat exchanger on its way to the fermenter.
This process takes about 1 hour.

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  By heat exchanging, we recover the energy used to boil the wort, i.e. cold water becomes hot water, and returned
to the Hot Liquor Tank which is then used to brew more beer or for cleaning.

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5. STAGE FIVE: FERMENTATION & MATURATION
 Once all the wort is in the tank, the yeast is added. The yeast will ferment the wort and turn it into beer.
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Primary fermentation will take about 3 – 4 days to complete. Fermentation temperatures will vary depending on
beer styles, a lager is fermented below 16 ˚C, ales are fermented above 20˚C.
 At the end of fermentation, the finished beer is chilled to 10˚ C and then 4˚ C and kept in the tank for maturation,
usually around three weeks.
 Yeast is harvested from the cone section at the bottom of the fermenter, to be used to ferment another batch of

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beer.
6. STAGE SIX: FILTERING INTO A BRIGHT BEER TANK
 When the beer is required for packaging, either in kegs or bottles, the beer is earth filtered into a Bright beer tank
(BBT).
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Filtering removes yeast, leaving the beer crystal clear. Some beers are not filtered at all and still contain yeast.
The CO2 (carbon dioxide) is adjusted in the BBT and it is then ready to be packaged.
Beer is kept freezing cold.
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FIG 2: The 8 stages of brewing (Eight degrees brewing, 2018)

7. STAGE SEVEN: PACKAGING
 Beer is packaged into either kegs or bottled in 330ml glass bottles.
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 Each keg holds 30 litres or 50 pints or 50 litres, about 85 pints.

 If bottled, the beer is counter pressure filled (double pre-evacuation) to reduce oxidation and capped on foam to
ensure it is free of any nasty microbes and will remain stable in the bottle.
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8. STAGE EIGHT: DISTRIBUTION

 Kegs are delivered to the bars and pubs and also used for festivals and promotions.
 Bottles are delivered to off-licences, restaurants, pubs, sporting clubs etc.
 Beer is consumed and enjoyed (Eight degrees brewing, 2018).

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 D. PROCEDURE FOR A KNOWN BEER PRODUCTION

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Maize beer production procedures: The main source maize beer is starch and its production includes the following
steps:
1. TREATMENT OF MAIZE GRAINS
The aim of this stage is to have safety grains. Therefore, detoxification of grain before malting may not be practical

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unless further growth of the mold is also prevented. Physical, biological, chemical and biological methods are
used for inhibiting mold growth in grain.
2. GERMINATION OR MALTING
Grain is soaked in water which is absorbed by the grain. There are hydration of grain embryo and stimulation of
grain for start of germination. Activities of embryo are awakened, and enzymes will be distributed evenly

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throughout the kernel. The grain is placed in shallow vessels and water is removed. Aleurone layer is stimulated
to produce enzymes which act to degrade cell wall structures. Enzymes are stimulated from the aleurone layer
and there is hydrolysis of proteins, carbohydrates, hemicellulose and lipids in maize grain. The temperature of the

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malt is raised from 15 to 85°C in a controlled manner (temperature, airflow, time). Germination process is halted
at desired malt quality, green brown malt is converted to stable, storable product, colour and flavor are developed,
enzymes are stabilized and preserved and unwanted flavours are removed.
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3. EXTRACTION WITH WATER OR MASHING
Milled grain is mixed with water at various temperatures, rest times and agitation. Temperatures are optimized for
the malt enzymes: 50°C for proteolysis, 62°C for gelatinization/liquefaction, 72°C for saccharification and 78°C
for mashing-off and malt enzyme inactivation. There are solubilization and dissolution of grain components, break
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down of grain cell wall structure, extraction and hydrolysis of starch, sugars, proteins and non-starch
polysaccharides and fermentable sugar profile is also established. Wort (aqueous solution) is separated from the
spent grains (78°C). Grain bed is spared with hot water to extract as much extract as possible. Degree of clarity
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of wort is established and extract level of wort is established. Wort is boiled at 100°C and hops are added during
the boil. We have evaporation of water, coagulation of proteins, isomerization of hop bitter components,
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inactivation of enzymes, wort sterilization, evaporation of undesirable volatiles, formation of flavour compounds
and colour development.
4. FIRST FILTRATION
Insoluble material is separated from the wort solution. Coagulated proteins and insoluble hops material are
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removed.
5. FERMENTATION
Wort solution is then cooled to yeast pitching temperature (21-25°C). Temperature is reduced to allow for yeast
addition, yeast is added to sterile cooled wort (cold temperatures encourage precipitation of proteins and beer
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clarification). Wort components of sugars and proteins are utilized by yeast to form alcohol, CO 2 and flavor
components according to the following chemical reaction:
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C6 H 12 O6 → 2C 2 H 5OH + 2CO2

At the end; alcohol level is established, flavor profile of beer is established, carbonation level is established and
yeast flocculates and can be easily separated (Cold maturation temperatures will influence beer clarity).

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 6. SECOND FILTRATION
After the completion of fermentation, the beer is filtered in order to remove the yeast and all the other solid

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derivatives. After filtration, the clear product is driven to bottling (Ore, Mironov and Shootov, 2018).

E. THE CONTINUOUS FERMENTATION PROCESS

Fermentation is the process whereby “sugars” are converted by yeast to alcohol, carbon dioxide, and

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heat. In the brewing of most traditional beer, the sugars are derived mainly from malted barley, although other
cereal sources and other plant sugars can also be used. These materials also contribute proteinaceous substances
which in combination with the sugars and added flavoring agents, notably hops, generate the alcohol, flavors, and
aromas that we know and love as beer. (Wong, 2003)

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The fermentation system itself is made up of a cascading series of three stirred vessels and a fourth
unstirred vessel where the beer is separated from the yeast. The system uses a flocculent yeast strain which settles
quickly at the end of fermentation. From the fourth vessel the clarified beer flows to a warm Maturation Vessel

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where the flavour is refined by yeast action (from the small amount of residual yeast in the beer). The total
residence time in these four vessels can be anything from 40 to 120 hours depending on production requirements.
All the different types of beer are produced on the same continuous process line, and their respective differences
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are produced after the maturation stage. (Campbell, 2017). The various vessel actions include:
1. STEP 1 - THE HOLD-UP VESSEL (HUV)
The incoming wort is oxygenated to stimulate yeast growth and a steady flow of yeast and beer from
later in the fermentation process is mixed with the wort as it flows into this first small vessel. The introduction of
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yeast into wort can be somewhat stressful for the yeast because of the high nutrient levels; by mixing the wort
with partially fermented beer the concentration of nutrients is reduced and this allows for a more rapid
commencement of fermentation. The yeast recycled back to the hold-up vessel is still in an active fermentation
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state, so again there is no significant lag phase before the fermentation begins. The recycled partially fermented
beer reduces the pH in this vessel (from pH 5.0 to pH 4.3) and increases the concentration of ethanol. This creates
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an unfavourable environment for any competing micro-organisms such as bacteria or wild yeast (any yeast which
is not the ‘culture’ yeast of a given brewery) and thus minimizes the potential for microbiological spoilage. The
beer/wort mixture has a residency time in this first vessel of approximately 3 to 4 hours.
2. STEP 2 - CONTINUOUS FERMENTER 1 (CF1)
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Continuous Fermenter 1 is the main fermentation vessel, from which the partially fermented beer is
recycled back into the hold-up vessel. The residence time in this vessel is normally 30 hours or more, depending
on production demands.
3. STEP 3 - CONTINUOUS FERMENTER 2 (CF2)
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Continuous Fermenter 2 is an important vessel for the fine-tuning of the finished fermented beer (that is,
ensuring that the desired amount of ethanol is produced). Residence time in this vessel is 12 hours or more.
4. STEP 4 - YEAST SEPARATOR
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This is an unstirred vessel with a cone-shaped base. As the beer flows into the vessel most of the yeast
settles to the bottom of the cone and is piped back to the beginning of the fermentation where it mixes with the
incoming wort. Normally, more yeast is produced during fermentation than is required by the brewery. The
surplus yeast is washed to recover as much beer as possible (this wash-water is used for dilution later in the beer-

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 making process) and the yeast may then be sold.
5. STEP 5 - MATURATION VESSEL

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The purpose of this vessel is largely to remove a fermentation by-product with a disagreeable
toffee/butterscotch-type flavour. The flavour belongs to a diketone compound, 2,3-butanedione. The precursor to
this flavour compound, "- acetolactate”’, forms during the fermentation. The warm temperature and low pH of
the maturation vessel hasten the conversion of α- acetolactate to 2,3-butanedione. Once the flavour is formed it is

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rapidly absorbed by the low level of remaining yeast cells to produce a compound with no noticeable flavour, 2,3-
butanediol. After two day’s maturation the beer is ready for cold storage to aid sedimentation and removal of haze
particles (Haze is due to complexes of polyphenols and proteins, if not removed they reduce the clarity of the
beer). Brand diversification also occurs during these post-fermentation steps. Finally, the beer is filtered and
packaged (Campbell, 2017).

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At first the processes seem pretty simple. Yeast are added to your cooled wort and they begin consuming
oxygen and nutrients and multiply. They will continue to multiply until all the oxygen is consumed and then they

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attack the sugars in the beer. Beer fermentation can be so vigorous and so much CO2 can be produced that a foamy
head can rise a foot or more above the top of the beer (Winning-Homebrew, 2018). Alcohol, CO2, and other
byproducts such as fusel alcohols (or fusel oils) are being produced by the yeast fermentation. Fusel oils are a
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mixture of organic acids, higher alcohols (propyl, butyl, and amyl), aldehydes, and esters, known collectively as
congeners. It is these Congeners that are purported to cause hangovers. This process continues until all the simple
sugars are consumed in the beer. The yeast begins to settle down and some are going dormant and falling to the
bottom, while others continue to eat the more complex sugars and other byproducts in the beer. At some point, the
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yeast would have eaten everything there is to eat and have nothing to do now but settle to the bottom of the
fermenter and wait for more food. One decides if they need to transfer your beer to a secondary fermenter at this
point. (Winning-Homebrew, 2018).
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This beer fermentation process usually takes about a week for most beers. At higher fermentation
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temperatures, more of the byproducts (from fruity esters to solvent-like fusel alcohols) are formed giving the beer
a fruity or phenolic flavor and aroma. At cooler temperatures, the yeast act much slower and the byproducts are
minimal. This is one of the main difference between lagers (fermented cold) and ales (fermented warm).
6. Lagers are much cleaner tasting than ales with the flavors favoring malt and hops. Ales on the other hand are
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much more complex in their aromas and flavors favoring the fruity, spicy characters. There is a somewhat linear
relationship with temperature though. Ales fermented at colder temperatures will have less "ale character" and
more "lager character". Conversely, lagers fermented at warmer temperatures will show more of the "ale
character" with fruitier/spicy ester production.
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7. The other difference and what usually separates the two types of yeast, is that lager yeasts ferment on or near the
bottom of the beer while ale yeasts ferment at the top, then fall out. Since yeasts are everywhere, it makes sense
that some would be adapted to the colder climates (Winning-Homebrew, 2018).
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F. FERMENTATION CONTROL
Management of the fermentation process is dependent on a number of factors, including the composition
and oxygen content of the wort, the quality and quantity of the yeast used for pitching, temperature control in the

This preprint research paper has not been peer reviewed. Electronic copy available at: https://ssrn.com/abstract=3458983
 fermenter, time, and the fermenter design. Healthy yeast is the heart of sound fermentation. The specific strain of
yeast is critical to the outcome of the fermentation, not only in the ability of the yeast to metabolize the wort

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contents to produce alcohol and distinctive flavor characteristics but also in the capacity of the yeast to tolerate its
own products of metabolism, most notably alcohol and the particular attribute of aggregation (flocculation) or
otherwise that the yeast strain may normally exhibit (Wong, 2003). Consistent fermentations demands control of
the key variables of yeast quantity and health, oxygen input, wort nutritional status, temperature, and yeast-wort

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contact (mixing). Fermentations may be monitored in various ways, including measuring the decrease in specific
gravity of the wort (including in-process measurements), CO2 evolution, the pH decrease, and ethanol formation,
as well as cam-era-based observation of events in the fermenter (Bokulichn and Bamforth, 2013).

Brewing strains can utilize a wide variety of carbohydrate sources, although individual strains will vary

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in their particular appetites. Ale strains of S. cerevisiae are able to ferment glucose, sucrose, fructose, galactose,
raffinose, maltotriose, and occasionally trehalose. Lager strains of S. cerevisiae (sometimes also called
Saccharomyces carlsbergensis) are also able to ferment the disaccharide melibiose, whereas S. cerevisiae var

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diastaticus is also able to utilize some of the higher glucose polymers called limit dextrins, which are out of reach
of the other strains. Ale strains are generically described as “top fermenters” on account of their tendency to form
a head or crust on the top of traditional open fermenters at the end of fermentation. Lager strains on the other
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hand, tend to separate out at the bottom of the fermentation vessel and hence enjoy the description of “bottom
fermenters.” With modern fermentation systems employing fermenters for both ales and lagers, this differentiation
is less clear-cut. Primary function of fermentation is to convert sugars into alcohol, for the yeast to fulfill this
requirement it must be present in sufficient quantity to effect the transformation (Wong, 2003).
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The yeast used to ferment is usually one or several generations old and, as a consequence of storage prior
to pitching and the physiological condition at the end of the previous fermentation, it is depleted of nutrients for
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growth and fermentation. It is said to be in a stationary phase of growth and requires the stimulus of fresh wort
nutrients, particularly oxygen, to rebuild its nutrient store and recommence growth and multiplication. Pitched
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brewing yeast will normally take several hours to adapt to its new environment before growth begins. This period
is known as the lag phase and precedes a period of very active growth and metabolism, known as the exponential
or logarithmic phase. The yeast will multiply four- or fivefold by a process of budding and build up its nutrient
store while at the same time commencing the conversion of sugars in the wort. The oxygen present at the start of
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pitching is rapidly used up by the yeast and is not involved in the fermentative process. The sugars present in the
wort are taken in to the yeast cell and broken down into smaller units, ultimately producing alcohol, carbon
dioxide, heat, and a vast range of other compounds, many of which contribute distinctive flavors and aromas to
beer. At the same time, the nitrogenous compounds in the wort are also assimilated by the yeast and as well as
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being used in the growth of yeast, they are metabolized and contribute to the rich flavor spectrum. When all
available sugars have been utilized, the yeast will begin to use its own carbohydrate reserves (glycogen and
trehalose) and effectively shut down its metabolism. This is known as the stationary phase of growth (Wong,
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2003).

The fermentation temperature is critical in controlling the outcome of fermentation and has a significant
impact on the development of flavor. Ales are generally fermented in the temperature range of 16°C to 22°C

This preprint research paper has not been peer reviewed. Electronic copy available at: https://ssrn.com/abstract=3458983
 (61°F–72°F) using top-fermenting strains, whereas lagers are fermented much cooler, 9°C–14°C (48°F–57°F)
using bottom-fermenting strains. Some beers, may be fermented very warm with temperatures reaching almost

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32°C (90°F). The combination of specific yeast strains and temperature generates very distinctive flavor profiles
in the beers, with the ales and particularly stouts generally producing fruity/estery characters, whereas the lagers
feature much lower ester levels, enabling more of the delicate pale malt characters and hop aromas to manifest on
the nose. Fermentation is an exothermic process in that heat is produced and control of the heat generated is

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essential in fermentation control. Brewing vessels are equipped with cooling equipment of varying levels of
sophistication, designed to effect cooling at the appropriate times in fermentation. Cooling is important in
moderating yeast flocculation in that it tends to encourage yeast to flocculate. This is necessary at the end of
fermentation to facilitate yeast separation but if applied too early, it can cause incomplete fermentation (Wong,
2003).

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The progress of the fermentation is usually monitored by following the specific gravity drop and/or
increase in alcohol content. Yeast growth and alcohol production deplete the sugar concentration and the pH falls

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as nitrogenous materials are used up and the yeast secretes organic acids. Flavor compounds are generated during
yeast growth, although some volatile components are lost with the exhaust carbon dioxide whereas other
compounds (notably diacetyl) are absorbed and metabolized by the yeast. There are many different types of
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fermentation vessels used in the brewing of beer. This reflects the beer being brewed, the volume required,
tradition, the relative age of the equipment, and the type of yeast being used, particularly in relation to the use of
top- or bottom-fermenting yeasts. At the end of fermentation, the yeast was collected from the top of the fermenter
and used to pitch subsequent brews (Wong, 2003).
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Beer at the end of the primary fermentation process is sometimes referred to as “green beer.” Before it
can be packaged and dispensed, the beer is usually subject to a further processing, which can include secondary
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fermentation, conditioning, and maturation. In secondary fermentation. At the end of the primary fermentation,
most of the yeast is removed but a small concentration, usually 0.5 to 2 million cells/ml, is left. A small amount
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of additional sugar, known as “priming sugar” or “primings,” either in the form of sucrose or glucose, is added,
which stimulates a secondary fermentation. Very little alcohol is produced (about 0.1%) but the main reason for
priming is the generation of additional carbon dioxide, which gives the beer extra carbonation, or “condition”. A
similar process occurs in the production of “bottle-conditioned” beers, where secondary fermentation takes place
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in the bottle. In addition to giving the beer condition, the residual yeast also scavenges any small amounts of
dissolved oxygen picked up at the time of filling the bottle. This antioxidant effect can help to extend the shelf
life of the beer. (Wong, 2003). At the completion of fermentation, yeast is recovered either for disposal (commonly
to animal feed or production of yeast extracts or for repitching (Ferreira, Pinho, Vieira and Tavarela, 2010).
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Harvested yeast may either be pumped to the next fermenter filling with fresh wort (cone-to-cone pitching) or
stored in either a pressed or slurry form (Bokulichn and Bamforth, 2013). It may receive acid washing to kill any
bacteria that may have developed in the slurry. Its collection from fermenters is often through the use of
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centrifuges, creating damage that has implications for subsequent performance (Chlup, Bernard and Stewart,
2008).

This preprint research paper has not been peer reviewed. Electronic copy available at: https://ssrn.com/abstract=3458983
 ed
G. PRODUCTS OF YEAST METABOLISM IN BREWERY FERMENTATIONS
Brewer’s yeast strains are facultative anaerobes they are able to grow in the presence or absence of
oxygen and this ability supports their property as an important industrial microorganism (Stewart, 2016). During
fermentation, yeast excretes a range of molecules, in addition to ethanol and CO2, that can affect flavor. All

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brewing strains produce glycerol, vicinal dik-etones (VDKs), alcohols, esters, short-chain fatty acids, organic
acids and diverse sulfur-containing substances. The levels of each category that are found in beer are dependent
in part upon the yeast strain but also important are the precise fermentation conditions that exist, including pitching
rate (Verbelen, Dekoninck, Saerens, Van Mulders, Thev-elein and Delvaux, 2009), temperature, extent of oxygen
addition, duration of fermentation and maturation. VDKs, diacetyl and pentanedi-one afford a buttery or honey-

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like character that is undesirable for most beers (Bokulichn and Bamforth, 2013). They are produced during
fermentation by the non-enzymatic degradation of acetolactate and acetohydroxybutyrate, which are metabolic
intermediates in pathways of amino acid synthesis that leak out into fermenting wort.

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A range of esters are produced by brewing yeast, with perhaps the most important being isoamyl acetate,
owing to its very low flavor threshold. Such esters are produced by the action of the enzyme alcohol
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acetyltransferase (AAT) on higher alcohols and acetyl-coenzyme A (acetyl-CoA) (Procopio, Qian and Becker,
2011). A major factor affecting the extent of lipid production and by extension, ester formation is the amount of
oxygen and unsaturated fatty acids in wort. AAT is also responsible for the production of thioesters (Bokulichn
and Bamforth, 2013). Esters are also formed by the combination of reactive acids and alcohols in the beer. Since
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the alcohol present in by far the largest quantity is ethanol, most of the esters produced are ethyl esters. Two
examples are ethyl acetate (solventy, slightly gluey aroma) and ethyl hexanoate (red apples and aniseed aroma).
The flavour balance of the esters produced is dependent on the yeast strain used. For instance, some strains of
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Bretannomyces yeast (wild yeast) produce enormous quantities of ethyl acetate, causing a UHU glue-type off-
flavour in beer (Campbell, 2017).
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There is some interest in selecting yeast strains with elevated -glycosidase (-G) activity for enhancing
the aroma of specialty beers. -Glycosidases in Saccharomyces cleave non-volatile glycosides derived from hops,
fruit, and other plants used in brewing, cleaving a sugar moiety from the aglycon. The free aglycon may exhibit
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aromatic activity in this state and represents a largely untapped source of aroma in beer (Daenen, Saison, Sterckx,
Delvaux, Verachtert and Derdelinckx, 2008)

Production of SO2 by yeast is significant for its direct contribution to aroma and also on account of its
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role in protecting against flavor deterioration, notably by scavenging the carbonyl substances that afford staling.
Dimethyl sulfide (DMS) is often a significant contributor to the character of lager beers, the bulk of the DMS
originates from thermal degradation of a malt-derived precursor, some yeast strains are capable of reducing
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dimethyl sulfoxide that also originates in malt (Bokulichn and Bamforth, 2013).

The production of alcohols other than ethanol is linked with nitrogen uptake by yeast. The yeast requires
nitrogen (in the form of amino acids extracted from the malt) in order to make protein and other nitrogenous cell

This preprint research paper has not been peer reviewed. Electronic copy available at: https://ssrn.com/abstract=3458983
 components. Examples of higher alcohols formed as by - products of nitrogen metabolism are propanol, isobutanol
and isoamyl alcohol. Shortages of critical amino acids can lead to the development of off-flavours such as diacetyl

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(2,3-butanedione), a buttery flavour which tends to be produced if valine levels are low in the wort. This flavour
can be avoided by allowing the beer adequate maturation time after the fermentation is complete (Campbell,
2017).

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1. SUGAR METABOLISM - THE GLYCOLYTIC PATHWAY
There are five sugars which may be present in wort which are readily utilised by standard brewer’s yeast
in fermentation; glucose, fructose, sucrose, maltose and maltotriose. These sugars are the main source of carbon
compounds for all the structural materials of yeast cells. The sugars are always taken up by the yeast in the same
sequence; first glucose, fructose and sucrose then maltose and lastly maltotriose. Sucrose is hydrolysed by the

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invertase enzyme in the yeast’s cell wall and splits into one glucose molecule and one fructose molecule both of
which may be assimilated into the glycolytic pathway. The enzymes responsible for the transport of maltose and
maltotriose through the yeast cell membrane (permeases) are ‘blocked’ by the presence of the simpler

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monosaccharides and so their uptake is delayed. Once within the yeast cell, both maltose and maltotriose are
converted to glucose by the enzyme ‘maltase’ (Campbell, S. L., 2017). Below is a simplified schematic summary
of the main metabolic pathways linked to beer flavor modulation by Saccharomyces:
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FIG 5: Overview of Saccharomyces metabolic activities influencing beer quality (Bokulichn and Bamforth,
2013)
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H. IMPLICATIONS AND BENEFITS OF BEER

In the environment, carbon dioxide is a major fermentation product which has the potential to pose a risk
as a ‘greenhouse gas’. Not all the CO2 produced remains dissolved in the beer since the fermentation operates at

This preprint research paper has not been peer reviewed. Electronic copy available at: https://ssrn.com/abstract=3458983
 only a low head pressure (high CO2 concentrations in the fermenting beer can have adverse effects on yeast
performance and production of flavour components such as esters). The excess CO 2 is collected off the top of the

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fermenting vessels, rather than allowing it to vent to atmosphere. The CO2 is ‘scrubbed’ to remove impurities and
the surplus may be sold for a profit to gas suppliers or in some cases, injected back into the finished product to
boost the carbonation level (Campbell, 2017). For economic implications, the continuous fermenter or CF avoids
losses (reduced wastage) caused by stop-start operations. Surplus yeast from the CF is collected and washed

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countercurrent with de-aerated water to extract any beer residues. The water (containing beer extracts) is then
used to dilute the fermented beer and the yeast is sold as a food material. Excess CO2 is collected and may be
purified and re-used. CF provides a continuous supply of CO2 which can be used in other parts of the process.
Energy, production labour and testing costs are reduced since only five vessels are involved in the fermentation
process (Campbell, 2017).

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I. HEALTH BENEFITS OF BEER
Beer has many health benefits if it is consumed in moderate amounts. Some of these interesting health benefits

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would include the following (Ore, Mironov and Shootov, 2018):
1. Anti-cancer properties
Hops used in beer, through its flavonoid compound called Xanthohumol plays a major role in the
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chemoprevention of cancer, including prostate cancer.
2. Reduced risk of cardiovascular diseases
Beer contains vitamin B6 which protects against heart diseases by preventing the build-up of a compound
called homocysteine. Moderate beer consumption increases bone density, thereby preventing the risk of fractures
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and osteoporosis.
3. Diabetes
Moderate beer consumption would have a lower prevalence of type 2diabetes.
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4. Prevention of anemia
Beer is a good source of vitamin B12 and folic acid which prevent anemia. Vitamin B12 is also essential for
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maintaining normal growth, good memory and concentration.

5. Hypertension
According to Biomedicine, regular beer drinkers would have lower blood pressure, compared to people that
consume similar amounts of wine or other spirits.
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6. Anti-aging properties
Beer increases the potency and impact of vitamin E, which is a major antioxidant in the body. It is an important
part of the maintenance of healthy skin, while also slowing down the aging process.
7. Gallstones
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Regular consumption of moderate amounts of beer would affect the cholesterol levels and decreases bile
concentration, leading to a reduced risk of developing gallstones.
8. Prevention of dementia and coronary disease
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Beer consumption also boosts the level of “good cholesterol” by 10-20%, thus reducing the risk of dementia
and cardiovascular diseases.
9. Aids digestive system
Beer possesses a number of digestive properties, which include the stimulation of gastrin, gastric acid,

This preprint research paper has not been peer reviewed. Electronic copy available at: https://ssrn.com/abstract=3458983
 cholecystokinin and pancreatic enzymes.
10. Kidney stones and osteoporosis

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Potassium, sodium and magnesium present in beer is important in reducing risk of kidney stones. The silicon
also present in beer is readily absorbed by the body, further explaining the protective effect of beer against
osteoporosis.
11. Stress buster

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Beer reduce stress and facilitates sleep like other alcohols.
12. Diuretic
Beer acts as a diuretic and significantly increase urination. This facilitates the increased removal of toxins and
waste materials from the body.

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REFERENCES
[1] Bokulich, N. A., Bamforth, C. W., and Mills, D. A., 2012. A review of molecular methods for microbial
community profiling of beer and wine. Journal of American Society Brewing Chemists, 70:150 –162.

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[2] Bokulichn, N. A., and Bamforth, C. W., 2013. The Microbiology of Malting and Brewing. Microbiology
and Molecular Biology Reviews, 2 (77):157–172.
[3] Campbell, S. L., 2017. The continuous brewing of beer. VI-Food-A-Beer:1-8
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[4] Chlup, P. H., Bernard, D., and Stewart, G. G., 2008. Disc stack centrifuge operating parameters and their
impact on yeast physiology. Journal of Institute of Brewing, 114:45– 61.
[5] Daenen, L., Saison, D., Sterckx, F., Delvaux, F. R., Verachtert, H., and Derdelinckx, G., 2008.
Screening and evaluation of the glucoside hydrolase activity in Saccharomyces and Brettanomyces brewing
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yeasts. Journal of Applied Microbiology, 104:478 – 488.

[6] Eight degrees brewing, 2018. Brewing Process. Available @ https://www.eightdegrees.ie/brewing-
process-2/. Accessed on 18/09/2018.
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[7] Ferreira, I., Pinho, O., Vieira, E., and Tavarela, J. G., 2010. Brewer’s Saccharomyces yeast biomass:
characteristics and potential applications. Trends in Food Science and Technology, 21:77– 84.
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[8] George Wong, 2003. Role of Yeast in Production of Alcoholic Beverage. Botany135, 1: 30-45.
[9] Keukeleire, D., 2000. Fundamentals of beer and hop chemistry. QUÍMICA NOVA, 23(1):108-112.
[10] Ore, G., Mironov, M., and Shootov, A., 2018. Design and production of maize beer. MOJ Food
Processing and Technology, 6(1) :78‒87.
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[11] Procopio, S., Qian, F., and Becker, T., 2011. Function and regulation of yeast genes involved in higher
alcohol and ester metabolism during beverage fermentation. European Food Research and Technology,
233:721–729.
[12] Sánchez, H. C., 2017. The mathematics of brewing. Available @ http://chalkdustmagazine.com/blog/the-
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mathematics-of-brewing/. Accessed on 18/ 09/ 2018.

[13] Stewart, G. G., 2016. Saccharomyces species in the Production of Beer. Beverages, (2) 34: 1-18.
[14] Verbelen, P. J., Dekoninck, T. M., Saerens, S.M., Van Mulders, S. E., Thev-elein, J. M., and Delvaux, F.
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R., 2009. Impact of pitching rate on yeast fermentation performance and beer flavour. Applied
Microbiology and Biotechnology, 82:155– 167.
[15] Winning-Homebrew, 2018. Controlling Beer Fermentation and Fermentability. Available @
https://www.winning-homebrew.com/beer-fermentation.html. Accessed on 17/09/2018.

This preprint research paper has not been peer reviewed. Electronic copy available at: https://ssrn.com/abstract=3458983