A REPORT FROM

THE AMERI CAN ACADEMY

OF MI CROBI OLOGY
IF THE
YEAST AINT
HAPPY,
AINT
NOBODY
HAPPY
The Microbiology of Beer
ABOUT ASM FAQs
The American Academy of Microbiology
is the honoric branch of the American
Society for Microbiology, a non-prot
scientic society with almost 40,000
members. Fellows of the AAM have been
elected by their peers in recognition of
their outstanding contributions to the eld
of microbiology. Through its colloquium
program, the AAM draws on the expertise
of these fellows to address critical issues
in microbiology.
FAQ reports are based on the
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who gather for a day to develop science-
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might have about topics in microbiology.
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every effort is made to ensure that the
information is accurate and complete.
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further so long as the authorship of the
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Copyright 2012
American Academy
of Microbiology
1752 N Street, NW
Washington, DC 20036
www.asm.org
STEERING
COMMITTEE:
Charles Bamforth,
Ph.D., D.Sc.
University of
California Davis
Department of
Food Science and
Technology
Morten Kielland
Brandt, Ph.D., Dr.H.C.
DTU Systems Biology
Chris White, Ph.D.
White Labs
Ann Reid
Director
American Academy
of Microbiology
Michael Ingerson-
Mahar, Ph.D.
Colloquia Fellow
American Academy
of Microbiology
PARTICIPANTS:
Nicholas Bokulich
University of
California Davis
Department of
Viticulture and
Enology
Chris Boulton, Ph.D.
University of
Nottingham
Department of
Brewing Science
Kyria Boundy
Mills, Ph.D.
University of
California Davis
Department of
Food Science and
Technology
Sue James, Ph.D.
Heineken
Supply Chain
William Maca
MillerCoors LLC.
John Sheppard, Ph.D.
North Carolina
State University
Dept. of Food,
Bioprocessing and
Nutrition Sciences
Bill Simpson, Ph.D.
Cara Technology
Limited
Katherine
Smart, Ph.D.
SABMiller
Jan Steensels
CMPG lab for
Genetics and
Genomics, KULeuven
VIB Laboratory for
Systems Biology
Graham Stewart,
Ph.D., D.Sc.
GGStewart Associates
Erna Storgards, Ph.D.
VTT Technical
Research Centre
of Finland
Graeme Walker,
B.Sc., Ph.D.,
D.Sc., FIBD
University of
Abertay Dundee
Food Division
Barry Ziola, Ph.D.
University of
Saskatchewan
Written by Ann Reid and Michael Ingerson-Mahar
In a word yeast. Without yeast, there would be no beer. Its that simple. And yeast is a
microbe, that is, a living organism that is too small to see with the human eye. Every time
someone brews a batch of beer, in a very real sense he or she is doing a microbiology
experiment. If you brew beer at home, youre a microbiologist! (In fact, you might even
want to join the American Society for Microbiology!) So lets talk about yeast.
1. What does microbiology have to do with beer?
IF THE YEAST AINT HAPPY, AINT NOBODY HAPPY: THE MICROBIOLOGY OF BEER 1
YEAST YOUR SOPHISTICATED BEER-MAKING
PARTNER, AND NOT SO DISTANT COUSIN:
You may remember from high school a description
of biodiversity that divided the living world into ve
kingdoms: animals, plants, fungi, monera and protists.
This classication, proposed in 1969 and now seriously
outdated, is heavily inuenced by what humans can
see plants, animals and fungi are all visible and their
diversity is obvious. Monera and protists are single-
celled microscopic organisms that are invisible to the
naked eye and cannot all be reliably distinguished
visually, even under the microscope. New molecular
techniques that look at an organisms genes to see
how closely they are related have shed new light on
lifes diversity. Now instead of ve kingdoms, scientists
recognize three domains of life: bacteria, archaea, and
eukaryotes. Animals, plants, fungi and protists actually
all belong to just one large domain, the Eukaryotes. As
you can see in the tree of life in Figure 1, the group that
used to be called monera in reality represents a huge
proportion of overall biodiversity.
But back to yeast. Yeasts are members of the fungal
kingdom, so they, like humans, are eukaryotes. In
addition to approximately 1500 species of yeast, the
fungal kingdom also includes mushrooms and molds.
Yeasts, while unicellular, are much more closely
related to humans than they are to bacteria. Each yeast
cell has about 6,000 genes (compared to 30,000 in a
human cell) and most of its basic life processes (DNA
replication, cell cycle control, gene regulation) are so
similar to humans that for decades yeast has been
used as the quintessential model organism in labs
studying how cells work. In the wild, the yeast family is
diverse. Many kinds of yeast seem to have evolved to
ll ecological niches based on feeding on simple sugars
produced by animals and plants. Yeasts are found in
every environment: on grape skins and human skin, in
the stomachs of bees and in tree sap, from the bottom
of the ocean to the soil under our feet.
Once you realize that the yeast youre pitching into
your beer is a highly sophisticated, complex organism,
and a member of a diverse and evolutionarily ancient
family, you will begin to appreciate how each step of
the brewing process is essentially an elaborate effort to
ensure that a very particular, highly specialized yeast
strain is given precisely the conditions that will help it
to produce the kind of beer you want.
Figure 1: The Three Domain Tree of Life
A REPORT FROM THE AMERICAN ACADEMY OF MICROBIOLOGY 2
There are two main yeasts used to brew beer,
each with several closely related varieties.
Commonly called brewers yeasts, they are
very different from any yeasts found in the
wild, just as house cats are very different
from lions and corn is very different from its
ancestor, teosinthe. Brewers yeast is different
for exactly the same reason like other
domesticated animals and plants, humans
have selected and used the strains that best
provide the characteristics that make it much
better for brewing beer than wild yeast. No one
knows exactly when the rst beer was brewed
or by whom. The discovery might have been
accidental, but it probably happened almost
as soon as humans began growing grain. The
rst historical evidence of beer brewing comes
from ancient Sumaria. Most grains in the right
conditions can be fermented by wild yeast that
live in virtually all environments and are easily
carried in air currents. Grain has long been
allowed to sprout because this makes it easier
to digest (this is still a common practice with
chicken feed, for example). If a store of sprouted
grain was left exposed to the elements, it could
have been ooded and seeded with wild yeasts
carried with the wind. A few weeks later, the
mixture would have fermented, producing the
rst beer. Brewing beer caught on very quickly.
It made good use of grain especially of grain
that had begun to sprout; it was safer to drink
than water (more on that later), and, of course,
it contained alcohol, which many animals,
including humans, nd appealing. Recognition
that brewing beer required expertise came early
as well; at least in Europe, communities often
had a designated brewer, a tradition that later
passed to monks in monasteries.
Brewers yeast is a domesticated organism. For centuries, humans have been
selecting yeast for particular characteristics including the ability to grow
quickly on a diet of malted grain, to produce satisfactory levels of alcohol, and
of course, to make a nal product that keeps well and tastes good!
2. Whats so special
about brewers yeast?
IF THE YEAST AINT HAPPY, AINT NOBODY HAPPY: THE MICROBIOLOGY OF BEER 3
Our ancestors did not know that an invisible living
organism was crucial to their beer-making success, but
they learned by experience that material skimmed from
the top of the fermenting beer, called barm, would
kick-start the next batch. An indication of brewers
appreciation for barm was that it was also called
Godesgood, meaning God is good. Home brewers
used a designated beer stick to stir their brews
unwittingly transferring yeast from one batch to the
next. It was not until the early- and mid-1800s when
rst Charles Cagniard-Latour and Friedrich T. Ktzing
and later Louis Pasteur studied yeast, that it became
clear that beer making was not just a chemical process
but rather that living organisms were responsible for
fermentation (please see the sidebar: Fermentation: a
transformative experience (and a lifesaver)). In the late
1800s, Emil Christian Hansen, a scientist and brewer at
the Carlsberg laboratories made a startling discovery.
The yeast that brewers had been using to produce beer
was actually a mixture of several different strains of
yeast. He was the rst brewer to isolate and propagate
one specic strain of yeast. Hansens accomplishment
helped brewers standardize the brewing process,
making beer more consistent from batch to batch.
What makes brewers yeast so special? The most
important answer is that unlike wild yeast, brewers
yeast grows very effectively on a particular sugar,
maltose, which just happens to be the predominant
sugar in the mash that brewers make from sprouted
barley. Brewers yeast also exhibits a variation on the
two distinct lifestyles that are typical of all yeast. In
the presence of oxygen and nutrients, yeast will use
available nutrients to reproduce that is, make more
yeast. When the oxygen becomes scarce, yeast will
switch over to another lifestyle based on fermentation
in which they consume sugars to produce alcohol
and carbon dioxide instead of just making more yeast.
Many yeasts do not turn to fermentation until oxygen
is entirely absent but brewers yeast make this switch
when there is still some oxygen around. Brewers
yeasts ability to ferment when there is still some
oxygen around is essential to brewing beer. Early beers
were probably weakly alcoholic because most common
wild yeasts stop fermenting at quite a low alcohol
concentration, around 3%. Brewers yeast will continue
to ferment even when the alcohol content rises to 7-9%.
In fact, if provided with supplemental nutrients, some
strains of brewers yeast will continue to ferment to an
alcohol concentration of over 16%!
Fermentation: a transformative experience (and a lifesaver)
Very broadly, fermentation is a biological reaction
that harvests energy from the chemical bonds in
sugars. When yeast carries out fermentation, it
breaks down simple sugars, and produces carbon
dioxide and alcohol as waste products. It is likely
that yeast evolved this life strategy to maintain a
competitive advantage over other microbes that
also feed on simple sugars. Most of those microbes
cannot tolerate alcohol or the high acidity caused
by the carbon dioxide and organic acids produced
by the yeast. Most beers have a pH in the range
of 4.0-4.5 intermediate between pure water
(pH = 7.0) and lemon juice (pH = 2.0) too acidic
for most bacteria. So by switching to fermentation,
the yeast ensures that it keeps the sugar source to
itself. For humans, this turned out to be a public
health bonanza. Before the development of water
sanitation systems in the late 1800s, water sources
were frequently contaminated with disease-causing
bacteria and viruses. All of these organisms are
killed by the brewing process (especially when
the grain extract is boiled) and cannot grow in
the nished product. The disease-causing bacteria
that can contaminate water are inhibited by beers
alcohol content, high acidity, low oxygen content
and by antibacterial compounds that come from
hops. Thus for much of human history, beer was
often safer to drink than the local water supply.
There are other microbes that can carry out
fermentation, but they produce different waste
products; for example, when Lactobacillus bacteria
ferment the sugars in milk to make yogurt, they
produce lactic acid, not alcohol. Again, the high
acidity inhibits bacterial growth. In both cases,
humans have taken advantage of microbial
fermentation to convert food into a form that is safer
to consume and can be stored longer.
A REPORT FROM THE AMERICAN ACADEMY OF MICROBIOLOGY 4
Yes and no. Brewers yeasts are all closely
related, but there are hundreds of different
strains with subtle differences that affect
what conditions they prefer and what kind
of beer they produce. Just as you wouldnt
select a Great Dane to be a lap dog, you
wouldnt select a lager yeast to produce ale.
The story of how brewers yeast came to be is complex
and still unfolding. In general, there are two major kinds
of brewers yeast ale yeasts and lager yeasts. Ale yeasts
are members of the species Saccharomyces cerevisiae
the same species used to make wine and bread.
However, after millennia of domestication and selection for
optimal performance in a particular growth environment,
the strains of S. cerevisiae now known as bakers yeast
(used for bread) and brewers yeast are genetically quite
different. Yeast has a lot of ways to generate genetic
diversity; it can reproduce both asexually (a single cell
buds off a nearly identical copy) and sexually (two yeast
strains make new combinations of their genetic material,
resulting in offspring with characteristics of both parents).
The yeast genome is carried on 16 separate chromosomes,
but yeast can tolerate quite a high degree of chromosomal
variation, including carrying more than 2 copies of some or
all of its chromosomes and multiple copies of large chunks
of chromosomes. Yeast can also maintain additional genetic
material on pieces of DNA that reproduce separately from the
chromosomes. These pieces of DNA, called plasmids, can be
shared between different yeast cells. Whats important about
all this genetic variety is that genes carry the instructions for
various life processes. When a yeast strain picks up or loses
genes, it gains and loses various capabilities. Not surprisingly,
brewers yeast is genetically enriched for many of the genes that
are important for fermentation, and especially for the ability to
ferment maltose. It also has extra copies of the genes that allow
it to tolerate high levels of alcohol.
3. Is all
brewers yeast
the same?
IF THE YEAST AINT HAPPY, AINT NOBODY HAPPY: THE MICROBIOLOGY OF BEER 5
Ale yeasts prefer a warm temperature (18-25
o
C,
65-77
o
F) for fermentation and rise to the top of the
fermentation tank when they are nished fermenting.
The origin of the cold-fermenting (7-15
o
C, 45-60
o
F), or
lager yeast, strain is more genetically complex. Lager
yeasts scientic name is Saccharomyces pastorianus
(to honor the great microbiologist Louis Pasteur), or
Saccharomyces carlsbergensis (to honor Emil Christian
Hansen, who rst isolated the strain while working for
the Carlsberg brewery in the late 1800s). It has long
been known that S. pastorianus was a hybrid between
traditional ale yeast and another yeast strain, but it was
only in 2011 that the other member of the hybrid was
isolated from beech forests on the southern tip of South
America. Researchers recently showed that the genome
of the newly identied Saccharomyces eubayanus is
virtually identical to the parts of the S. pastorianus
that differ from ale yeast. Why is this important?
S. eubayanus is what scientists call cryotolerant, that
is, it grows well at low temperatures. For brewers in
Northern Europe, this quality was important. When and
where S. eubayanus and S. cerevisiae mated to form
S. pastorianus is unknown, but cold-fermented beer
began to be produced in Bavaria in the 15th century.
The new style of beer spread quickly as Northern
European brewers recognized the value of a strain that
could ferment at cooler temperatures. The resultant
clear, crisp product that came to be known as lager
proved very popular. Today it accounts for the lions
share of beer production worldwide.
Ale and lager yeasts are responsible for virtually all
beer production, but there are a few styles of beer that
rely on other kinds of yeast for fermentation. Of these,
the lambics of Belgium, are the best-known. To make
a lambic beer, the boiled wort (see below) is exposed
to the open air for about a day before going into the
fermentation tank. Fermentation is carried out not only
by wild yeast but also by bacteria that settled on the
wort while it was exposed to the air. Clearly, this leaves a
great deal to chance, and true lambics can only be made
around Brussels, and only during certain times of the
year. In the summer, the air contains too many bacteria
that make the beer taste bad. Even in its home territory,
batches of lambic are usually blended to produce a more
consistent nal product. Using the lambic technique
will make quite a different product from place to place,
depending on what yeast and bacteria are found in
the local atmosphere. Fortunately for those who love
lambics, but dont live in Brussels, some of the microbes
specic to the area of Belgium where lambics originated
have been characterized, so brewers wanting to re-create
the avor can purchase pure strains of Brettanomyces
(the yeast genus found in Belgian lambics) and its two
predominate bacterial strains: Lactobacillus delbrkii and
Pediococcus damnosus.
A REPORT FROM THE AMERICAN ACADEMY OF MICROBIOLOGY 6
STEP 1: MALTING
The main ingredient in beer is
water, followed by grain usually
barley, sometimes wheat, and
occasionally rice or corn. But in
order for barley to make good
beer, it has to be sprouted, which
is called malting. Why? The part
of the barley plant that is used in
beer is the seed which is like a
miniature storage container for the
baby barley plant. In addition to the
plant embryo, the seed contains
starch that the baby plant will feed
on when it sprouts. When the seed
is moistened (think spring rain),
enzymes in the seed are activated
that start to break down cell
walls and proteins to expose the
seeds stored starch. Brewers take
advantage of this process because
yeast cannot feed directly on barley
seeds. During the malting process,
the barley seed is moistened and
allowed to begin sprouting. Before
the sprout breaks through the
seeds hull, the barley is roasted,
or kilned (to stop the sprouting
process and develop avor). The
kilning must be conducted in such
a way as to maintain the seeds
enzyme activity.
STEP 2: MASHING
During the mashing process, the
ground malt is mixed with water
and heated. This provides an
ideal environment for the various
enzymes that were produced in
the barley seed during sprouting
to break down the barley starch
into simpler, fermentable sugars.
The enzymes are quite specic
each has a particular job to do in
breaking down starch and each has
a temperature at which it is most
efcient. As a result, beer recipes
contain detailed instructions for
how long to hold the mash at
particular temperatures.
STEP 3: PREPARING THE WORT
Now that the sugars have been
released from the barley, the
yellow-brown sweet liquid called
the wort can be drained away from
the remains of the barley seeds
(spent grains). If you taste the wort
now, you will nd it very sweet.
The sugary liquid designed to
be yeast food also provides a
highly attractive environment for
the growth of any bacteria or wild
yeast that might have survived the
mashing process. Therefore the
wort is now boiled. At the same
time, hops are added. Hops are the
dried owers of the hops plant; they
release a number of compounds
some of which are toxic to bacteria.
The same compounds also add
a bitter avor to the wort that
balances out the sweetness of the
barley sugars. Hops also provide
oils that contribute to the charac-
teristic aroma of many beers. Bitter
herbs have been added to beer for
centuries because early brewers
probably recognized that the beer
with added herbs was less likely to
spoil. The hops are boiled with the
wort for about an hour, and then
the wort is ltered and cooled.
Barley, hops, water, and yeast. Thats all it takes to make beer. But every step in the process is
important, and most of them have a direct impact on how happy and productive the yeast will be.
Because if the yeast aint happy, aint nobody happy.
4. How is beer made?
IF THE YEAST AINT HAPPY, AINT NOBODY HAPPY: THE MICROBIOLOGY OF BEER 7
STEP 4: FINALLY! ADDING THE YEAST
At this point the yeast nally enters the brewing process,
but the preceding steps have produced a nourishing
broth that is ideal for yeast growth. One nal step is
necessary for the wort to be an ideal environment for the
yeast. Yeast needs oxygen to grow and reproduce, and
boiling drives most of the oxygen out of the wort, so it is
important to re-aerate the wort before adding the yeast.
This is usually accomplished by bubbling air or oxygen
through the wort (as you would with an aquarium),
although oxygenation can also be accomplished by
vigorously shaking the wort. Yeast is now added (the
brewing term is pitched) into the oxygenated wort, and
the mixture goes into a fermentation tank. What happens
next is a two-step process. In the presence of plentiful
nutrients and oxygen, yeast reproduces rapidly. In the
case of brewers yeast, this means making lots of new
yeast cells through a process called budding.
When conditions are just right, a yeast cell can bud
every 80 minutes. As the oxygen is used up, growth
slows and the yeast cells switch to another kind of
lifestyle fermentation. Successful brewing requires
both steps the growth phase to increase the number
of yeast cells, and the fermentation phase to convert
much of the remaining sugar in the wort to alcohol and
carbon dioxide. As the fermentation proceeds, available
sugar declines and the alcohol concentration rises.
Eventually most of the yeast stop trying to extract any
more energy from the wort and enter a hibernation-
like stage. Entering this stage triggers physiological
changes that cause the yeast to clump together a
process called occulation. Some strains of yeast drop
to the bottom of the fermenting vessel when they
occulate, and others rise to the surface. If they rise,
they can be skimmed off the top of the beer. If they
drop, the beer can be carefully poured off the layer of
yeast cells and other debris (called trub) that settles at
the bottom of the tank.
STEP 5: SECONDARY FERMENTATION
Even after the main fermentation has ceased and most
of the yeast has occulated out of suspension, a small
amount of yeast remains in the beer and continues to
break down sugars and produce other compounds. This
stage is called secondary fermentation, although very
little actual fermentation is occurring. It is important,
however, because it is during this time that the yeast
metabolizes (or digests) diacetyl and acetaldehyde that
have been produced during primary fermentation. The
tastes imparted by these compounds butterscotch
and green apples, respectively are considered
undesirable in most beers. Depending on the type of
beer, this secondary fermentation may be prolonged for
weeks or even months at quite a cool temperature
about 7
o
C (45
o
F). If the secondary fermentation is going
to be very lengthy, it is often recommended to decant
the beer into a different vessel to get it away from the
spent yeast. Yeast cells can remain dormant for a long
time, but eventually they will begin to die off and the
cells will burst, releasing their not-very-tasty cellular
contents into the beer. After secondary fermentation,
beer is often held at an even lower temperature which
promotes settling of any remaining solid residues or
yeast and makes the beer less cloudy.
STEP 6: BOTTLING
One last way in which yeast contributes to the quality of
nished beer; it adds natural carbonation. After the beer
is bottled, any remaining yeast will continue to ferment
any remaining sugar, and add carbon dioxide to the
beer in the bottle. Sometimes, a little more sugar and
some fresh yeast is added just before bottling to ensure
an appropriate amount of carbonation (a process called
conditioning). Restraint is called for, though; too much
additional carbon dioxide can cause bottles to explode.
More commonly these days, brewers add carbon dioxide
just before bottling to bring the level of carbonation up
to desired levels.
A REPORT FROM THE AMERICAN ACADEMY OF MICROBIOLOGY 8
The yeast you pitch into your wort is made up of
billions of individual yeast cells. Ideally each cell is
alive and ready to begin fermenting as soon as you
add it to your cooled and oxygenated wort. The
individual cells are far too small to see and count
without a microscope, but it is important to get the
number right. Imagine you were planting a eld of
corn. You want to plant exactly the right amount of
seeds to optimize your yield: if you plant too few, you
will end up with less corn and more open space for
weeds to occupy. If you plant too many, the plants
will crowd each other and may exhaust the nutrients
in the soil, reducing the yield. Exactly the same is
true for yeast: add too little and fermentation will
take a long time to get started, increasing the risk
that bacteria will outcompete the yeast. Add too
much, and the yeast will turn quickly to fermentation
without having made many new yeast cells. Since
all those growing yeast cells release complex
compounds like esters, the nished beer will have a
less complex avor and aroma.
Brewers use formulas to gure out how many yeast
cells to add. They are slightly different depending
on the type of beer being brewed, but all have the
following factors: number of yeast cells per milliliter
of wort per degrees Plato of the wort. Degrees
Plato is a measurement of the specic gravity of
the wort, which in turn is simply a measurement
of how much sugar and other compounds are
dissolved in the wort. For comparisons sake, the
specic gravity of water is (by denition) equal to
one. The specic gravity of, say, molasses, is around
1.4. Most worts have a specic gravity of about 1.04
to 1.05 in other words, about 4-5% higher than
pure water. Pitch rates are calculated to add enough
yeast to reproduce for a few days with that amount
of nutrients before turning to fermentation. The
fermentation, therefore, will take place with many
more yeast cells than were originally added, and
those new yeast cells will be young and healthy,
leading to a vigorous fermentation.
Companies that sell yeast to home brewers grow it
from pure cultures under standardized conditions
and provide it in packets that contain the correct
number of yeast cells to inoculate 5 gallons of
standard wort (thats about 100 billion yeast
cells!). Buying new yeast for each batch of beer
ensures that you know exactly how many cells
youre adding, and that the cells are at the right
life stage and healthy. However, it is also possible
to re-use or re-pitch yeast from batch to
batch. After all, not only are most of the original
yeast cells still alive, but they will have budded off
several generations of new yeast cells. The risk,
however, is that the yeast that is left at the end of
fermentation is not exactly the same as the yeast
you started with; along with young, newly budded
cells, there will be dead yeast cells, cells that have
already budded off so many offspring that they can
no longer reproduce, and cells that have undergone
various mutations. Furthermore, the yeast that was
originally pitched was not genetically identical,
but contained billions of individuals that each
differed very slightly from each other. The genetic
variants that were most successful in the particular
conditions of that fermentation will be slightly
more frequent in the next generation; thats
evolution at work. Over time, the character of the
yeast can change substantially. Judging how much
of the leftover yeast to re-pitch, and how often, is
something of an art. In general, it is recommended
to re-pitch no more than 5 to 10 times.
How much yeast does it take to make beer?
IF THE YEAST AINT HAPPY, AINT NOBODY HAPPY: THE MICROBIOLOGY OF BEER 9
Remember, yeast is a sophisticated, living
organism. In addition to fermentation, it
carries out thousands of additional reactions
as part of its general internal housekeeping
and its interaction with the environment.
During the fermentation process, yeast
produces hundreds of chemical compounds
that can affect the nal product in many ways
including avor, appearance and stability.
Even the foam is affected by chemical
compounds and enzymes made by the yeast.
FLAVOR:
Flavor is a complex characteristic that includes
taste, aroma, and mouthfeel, and it is this quality
that beer drinkers are most concerned with beer.
Of the thousands of other compounds produced
by yeast, about twenty ve to thirty are critical in
giving a distinctive avor to a given beer style. These
avor compounds include acids, aldehydes, sulfur
compounds, ketones, and esters. The exact mix of
these components is different in every type of beer and
even from batch to batch if conditions are not carefully
controlled. Yeast can also change beers avor when
they do not consume certain components, for example
complex sugars, contributed by the wort or hops.
Acids: The acids in beer are not the kind that you have
to worry about burning your skin at least not in the
levels present in beer, but they do contribute to the
taste of the beer. The most commonly found acids in
beer are citric, acetic, malic, pyruvic, and succinic acids.
They generally confer sour avors to the beer. These
acids also affect the beers pH.
Aldehydes: The most common aldehyde in beer is
acetaldehyde. Acetaldehyde confers a green apple taste
to the beer and, as mentioned above, is considered
undesirable at high levels. Levels of acetaldehyde are
highest just after the start of fermentation because
yeast must convert sugars into acetaldehyde before
they can be converted into ethanol. As the fermentation
5. How else does the yeast affect the beer?
A REPORT FROM THE AMERICAN ACADEMY OF MICROBIOLOGY 10
progresses, yeast will generally complete the process
of converting acetaldehyde (and other aldehydes
and ketones that might make the beer taste stale) to
ethanol. Many essential yeast life processes require
zinc, including the conversion of aldehydes, so brewers
will add trace amounts of the metal if the wort has a
low zinc level in part to ensure that acetaldehyde levels
do not become too high. Brewers have another reason
to make sure acetaldehyde is at acceptable levels in
the beer acetaldehyde has been implicated as one
component of beer that causes hangovers!
Sulfur compounds: There are several sulfur compounds
that can be introduced into beer by the yeast. Like the
acids and aldehydes small amounts of many sulfur
compounds are acceptable or even desirable in beer,
but at higher levels, can be off-putting. Hydrogen
sulde, for example can improve the taste of pale ale
at low levels, but in greater quantities can confer the
avor of rotten eggs. Dimethyl sulde is a similar story,
as low levels are an expected part of the taste of Pilsner-
style beers, but higher levels can leave beer tasting like
cabbage or corn.
Ketones: Ketones can show up in both beers aroma
and taste. Diacetyl, one common ketone, can confer a
butterscotch avor to beer, however too much diacetyl
can result in an off-avor. Another ketone, with the
chemical name 2,3-pentanedione, can also give beer a
buttery or stale milk avor.
Esters: Humans are thought to be able to distinguish
about 10,000 different chemical compounds by their
aroma, so many of the compounds produced by, or not
digested by, yeast could contribute to a beers aroma.
However, one particular family of chemical compounds,
the esters, has a disproportionate effect on the overall
aroma of beer.
Many fruits produce high levels of distinctive esters.
The easily identied smells of oranges, pears, lavender
and cinnamon all come from esters. Small amounts of
such esters may be desirable in a beer. Some others,
like the esters that give glue and nail polish remover
their distinctive smells would not be so welcome.
Yeast produces as many as 90 different esters, and
while they are not identical to those produced by
plants or herbs, they can have a powerful effect on
the beers aroma. For example, yeast can produce the
esters isoamyl acetate, which smells like banana, and
phenylethyl acetate, which smells like roses.
MOUTHFEEL:
Mouthfeel, the sensation of the beer coming in contact
with the mouth, is the nal component of avor.
Carbon dioxide is a major component of the mouthfeel,
giving the tingly feeling associated with beer and
other carbonated beverages like soft drinks. Both the
size and numbers of carbon dioxide bubbles affect
the mouthfeel (imagine the difference in carbonation
between a dark stout beer and champagne, for
example). Sugars can also affect the mouthfeel of
the beer. S. cerevisiae is very good at metabolizing
maltose, but cannot consume certain other types of
larger sugars. These sugars remain in the beer. Different
amounts and types of residual sugars can give the beer
a fuller or thinner body and are responsible for much of
the caloric content of the beer.
FOAM:
One nal characteristic of beer that is affected by yeast
is the foam, or head, that forms on the top of a glass of
beer. The head on beer is deemed important by most
drinkers and can make or break its acceptability.
Yeast can have positive and negative effects on beer
foam. Clearly the carbon dioxide produced by the
yeast is what is inside the bubbles, but the bubbles
themselves are stabilized by proteins from the grains
and the bitter substances from the hops. Yeast can
seriously damage the foam in two ways. If the initial
fermentation is too vigorous, too many of these
foam-stabilizing proteins may be lost in the head that
forms as the carbon dioxide rises. Second, if yeast
deteriorates by becoming stressed (for example,
because the alcohol level gets too high, or the yeast
gets too old) then it can start to secrete enzymes called
proteases that chew up the foam proteins.
IF THE YEAST AINT HAPPY, AINT NOBODY HAPPY: THE MICROBIOLOGY OF BEER 11
While yeast is essential to brewing and
affects the nal product in many ways, many
other factors affect the avor, color and
stability of beer. What kind of grain is used,
the mashing characteristics, the variety and
amount of hops, other avoring components,
the length and temperature of fermentation
and many other factors also have important
effects. Hopefully, you now see that part of
the reason these factors make a difference
is because they affect yeast health and
activity but they also have impacts that are
not related to their interaction with yeast.
Also, yeast is not the only microbe that affects beer, for
good or ill. Some of these other microbial effects are
quite interesting. For example, barley can be infected
by a fungus called Fusarium that produces proteins that
survive the entire malting, mashing, and fermentation
process. Their presence is revealed only when the beer
bottle is opened; the beer will foam over, or gush,
upon opening when these proteins are present.
As discussed above, beers high alcohol and low
oxygen content, plus the antibacterial compounds
produced by hops, tend to inhibit the growth of
most bacteria. Bacteria, however, are extraordinarily
adaptable, and given time and opportunity, many of
them are able to evolve in ways that allow them to
grow under once-hostile conditions. For example, just
as many human pathogens have developed resistance
to antibiotics, there are bacteria that have developed
resistance to antimicrobial compounds from hops.
These bacteria are not human pathogens beer that
contains them will not make you sick but they do
produce compounds that make the beer taste bad.
Proper sanitation procedures are crucial in commercial
breweries and for home brewers; once a beer spoiler
organism nds its way into a brewery, it can be
practically impossible to eliminate.
6. Is it really only about the yeast?
A REPORT FROM THE AMERICAN ACADEMY OF MICROBIOLOGY 12
IF THE YEAST AINT HAPPY, AINT NOBODY HAPPY: THE MICROBIOLOGY OF BEER 13
THIS REPORT WAS DESIGNED BY
PENSAR DESIGN GROUP
WWW.PENSAREDESIGN.COM