Fermentation is the process of deriving energy from the oxidation of organic
compounds, such as carbohydrates, and using an endogenous electron acceptor,
which is usually an organic compound.
In contrast, respiration is where electrons
are donated to an exogenous electron acceptor, such as oxygen, via an electron
transport chain. Fermentation is important in anaerobic conditions when there is no
oxidative phosphorylation to maintain the production of ATP (Adenosine
triphosphate) by glycolysis. During fermentation, pyruvate is metabolised to
various different compounds. Homolactic fermentation is the production of lactic
acid from pyruvate; alcoholic fermentation is the conversion of pyruvate into
ethanol and carbon dioxide; and heterolactic fermentation is the production of
lactic acid as well as other acids and alcohols. Fermentation does not necessarily
have to be carried out in an anaerobic environment. For example, even in the
presence of abundant oxygen, yeast cells greatly prefer fermentation to oxidative
phosphorylation, as long as sugars are readily available for consumption.

Sugars are the most common substrate of fermentation, and typical examples of
fermentation products are ethanol, lactic acid, and hydrogen. However, more
exotic compounds can be produced by fermentation, such as butyric acid and
acetone. Yeast carries out fermentation in the production of ethanol in beers, wines
and other alcoholic drinks, along with the production of large quantities of carbon
dioxide. Fermentation occurs in mammalian muscle during periods of intense
exercise where oxygen supply becomes limited, resulting in the creation of lactic

 1 Chemistry
o 1.1 Ethanol fermentation
o 1.2 Lactic acid fermentation
o 1.3 Glycolysis
o 1.4 Aerobic respiration
 2 Hydrogen gas production in fermentation
 3 History
 4 Etymology

Fermentation products contain chemical energy (they are not fully oxidized), but
are considered waste products since they cannot be metabolized further without the
use of oxygen (or other more highly-oxidized electron acceptors). A consequence
is that the production of adenosine triphosphate (ATP) by fermentation is less
efficient than oxidative phosphorylation, whereby pyruvate is fully oxidized to
carbon dioxide.

Ethanol fermentation
Ethanol fermentation (performed by yeast and some types of bacteria) breaks the
pyruvate down into ethanol and carbon dioxide. It is important in bread-making,
brewing, and wine-making. Usually only one of the products is desired; in bread-
making, the alcohol is baked out, and, in alcohol production, the carbon dioxide is
released into the atmosphere or used for carbonating the beverage. When the
ferment has a high concentration of pectin, minute quantities of methanol can be
The chemical equation below summarizes the fermentation of glucose, whose
chemical formula is C
One glucose molecule is converted into two
ethanol molecules and two carbon dioxide molecules:
→ 2 C
OH + 2 CO

OH is the chemical formula for ethanol.
Before fermentation takes place, one glucose molecule is broken down into two
pyruvate molecules. This is known as glycolysis.

Lactic acid fermentation
Lactic acid fermentation is the simplest type of fermentation. Essentially, it is a
redox reaction. In anaerobic conditions, the cell’s primary mechanism of ATP
production is glycolysis. Glycolysis reduces – transfers electrons to – NAD
forming NADH. However, there is only a limited supply of NAD
available in a
cell. For glycolysis to continue, NADH must be oxidized – have electrons taken
away – to regenerate the NAD
. This is usually done through an electron transport
chain in a process called oxidative phosphorylation; however, this mechanism is
not available without oxygen.

Instead, the NADH donates its extra electrons to the pyruvate molecules formed
during glycolysis. Since the NADH has lost electrons, NAD
regenerates and is
again available for glycolysis. Lactic acid, for which this process is named, is
formed by the reduction of pyruvate.

In heterolactic acid fermentation, one molecule of pyruvate is converted to lactate;
the other is converted to ethanol and carbon dioxide. In homolactic acid
fermentation, both molecules of pyruvate are converted to lactate. Homolactic acid
fermentation is unique because it is one of the only respiration processes to not
produce a gas as a byproduct.
Homolactic fermentation breaks down the pyruvate into lactate. It occurs in the
muscles of animals when they need energy faster than the blood can supply
oxygen. It also occurs in some kinds of bacteria (such as lactobacilli) and some
fungi. It is this type of bacteria that converts lactose into lactic acid in yogurt,
giving it its sour taste. These lactic acid bacteria can be classed as
homofermentative, where the end product is mostly lactate, or heterofermentative,
where some lactate is further metabolized and results in carbon dioxide, acetate or
other metabolic products.
The process of lactic acid fermentation using glucose is summarized below.
homolactic fermentation, one molecule of glucose is converted to two molecules of
lactic acid:
→ 2 CH
In heterolactic fermentation, the reaction proceeds as follows, with one molecule of
glucose being converted to one molecule of lactic acid, one molecule of ethanol,
and one molecule of carbon dioxide:

→ CH

Before lactic acid fermentation can occur, the molecule of glucose must be split
into two molecules of pyruvate. This process is called glycolysis.

To extract chemical energy from glucose, the glucose molecule must be split into
two molecules of pyruvate.
This process also generates two molecules of
adenosine triphosphate as an immediate energy yield and two molecules of

+ 2 ADP + 2 P
+ 2 NAD
→ 2 CH

+ 2 ATP + 2 NADH
+ 2 H
O + 2H

The chemical formula of pyruvate is CH

. P
stands for the inorganic
phosphate. As shown by the reaction equation, glycolysis causes the reduction of
two molecules of NAD
to NADH.
Two ADP molecules are also converted to
two ATP and two water molecules via substrate-level phosphorylation.
Aerobic respiration
In aerobic respiration, the pyruvate produced by glycolysis is further oxidized
completely, generating additional ATP and NADH in the citric acid cycle and by
oxidative phosphorylation. However, this can only occur in the presence of
oxygen. Oxygen is toxic to organisms which are obligate anaerobes, and are not
required by facultative anaerobic organisms. In the absence of oxygen, one of the
fermentation pathways occurs in order to regenerate NAD
; lactic acid
fermentation is one of these pathways.

Hydrogen gas production in fermentation
Hydrogen gas is produced in many types of fermentation (mixed acid fermentation,
butyric acid fermentation, caproate fermentation, butanol fermentation, glyoxylate
fermentation), as a way to regenerate NAD
from NADH. Electrons are transferred
to ferredoxin, which in turn is oxidized by hydrogenase, producing H
gas is a substrate for methanogens and sulfate reducers, which keep the
concentration of hydrogen sufficiently low to allow the production of such an
energy-rich compound.

The first solid evidence of the living nature of yeast appeared between 1837 and
1838 when three publications appeared by C. Cagniard de la Tour, T. Swann, and
F. Kuetzing, each of whom independently concluded as a result of microscopic
investigations that yeast was a living organism that reproduced by budding. The
word "yeast," it should be noted, traces its origins back to the Sanskrit word
meaning “boiling.” It was perhaps because wine, beer, and bread were each basic
foods in Europe, that most of the early studies on fermentation were done on
yeasts, with which they were made. Soon bacteria were also discovered; the term
was first used in English in the late 1840s, but it did not come into general use until
the 1870s, and then largely in connection with the new germ theory of disease.

Louis Pasteur (1822–1895), during the 1850s and 1860s, showed that fermentation
was initiated by living organisms in a series of investigations.
In 1857 Pasteur
showed that lactic acid fermentation is caused by living organisms.
In 1860 he
demonstrated that bacteria cause souring in milk, a process formerly thought to be
merely a chemical change, and his work in identifying the role of microorganisms
in food spoilage led to the process of pasteurization.
In 1877, working to
improve the French brewing industry, Pasteur published his famous paper on
fermentation, Etudes sur la Biere , which was translated into English in 1879 as
Studies on Fermentation.
He defined fermentation (incorrectly) as "Life without
but correctly showed specific types of microorganisms cause specific types
of fermentations and specific end products.
Although showing that fermentation was generally the result of the action of living
microorganisms was a breakthrough, it did not explain the basic nature of the
fermentation process, or prove that it was caused by the microorganisms that were
apparently always present. Many scientists, including Pasteur, had attempted
unsuccessfully to extract the fermentation enzyme from yeast.
Success came in
1897 when the German chemist Eduard Buechner ground up yeast, extracted a
juice from them, then found to his amazement that this "dead" liquid would
ferment a sugar solution, forming carbon dioxide and alcohol much like living
The "unorganized ferments" behaved just like the organized ones. From
that time on the term enzyme came to be applied to all ferments. It was then
understood that fermentation is caused by enzymes which are produced by
In 1907 Buechner won the Nobel Prize in chemistry for his

Advances in microbiology and fermentation technology have continued steadily up
until the present. For example, in the late 1970s it was discovered that
microorganisms could be mutated with physical and chemical treatments to be
higher yielding, faster growing, tolerant of less oxygen, and able to use a more
concentrated medium.
Strain selection and hybridization developed as well,
affecting most modern food fermentations.
The word fermentation is derived from the Latin verb "fervere" which means "to
boil". It is thought to have been first used in the late fourteenth century in alchemy,
but only in a broad sense. It was not used in the modern scientific sense until
around 1600.

The primary benefit of fermentation is the conversion of sugars and other
carbohydrates, e.g., converting juice into wine, grains into beer, carbohydrates into
carbon dioxide to leaven bread, and sugars in vegetables into preservative organic
Food fermentation has been said to serve five main purposes:

 Enrichment of the diet through development of a diversity of flavors,
aromas, and textures in food substrates
 Preservation of substantial amounts of food through lactic acid, alcohol,
acetic acid and alkaline fermentations
 Biological enrichment of food substrates with protein, essential amino acids,
essential fatty acids, and vitamins
 Elimination of antinutrients
 A decrease in cooking times and fuel requirements
Some fermentation products (e.g., fusel alcohol) are deleterious.



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