Below are some definitions of fermentation.

They range from informal, general usages to more scientific

definitions.[4]

1. Preservation methods for food via microorganisms (general use).

2. Any large-scale microbial process occurring with or without air (common definition used in
industry).

3. Any process that produces alcoholic beverages or acidic dairy products (general use).

4. Any energy-releasing metabolic process that takes place only under anaerobic conditions
(somewhat scientific).

5. Any metabolic process that releases energy from a sugar or other organic molecule, does not
require oxygen or an electron transport system, and uses an organic molecule as the final
electron acceptor (most scientific).

Biological role[edit]

Along with aerobic respiration, fermentation is a method to extract energy from molecules. This method
is the only one common to all bacteria and eukaryotes. It is therefore considered the oldest metabolic
pathway, suitable for primeval environments – before plant life on Earth, that is, before oxygen in the
atmosphere.[5]: 389 

Yeast, a form of fungus, occurs in almost any environment capable of supporting microbes, from the
skins of fruits to the guts of insects and mammals to the deep ocean. Yeasts convert (break down) sugar-
rich molecules to produce ethanol and carbon dioxide. [6][7]

Basic mechanisms for fermentation remain present in all cells of higher

organisms. Mammalian muscle carries out fermentation during periods of intense exercise where
oxygen supply becomes limited, resulting in the creation of lactic acid.[8]: 63  In invertebrates, fermentation
also produces succinate and alanine.[9]: 141 

Fermentative bacteria play an essential role in the production of methane in habitats ranging from
the rumens of cattle to sewage digesters and freshwater sediments. They produce hydrogen, carbon
dioxide, formate and acetate and carboxylic acids. Then consortia of microbes convert the carbon
dioxide and acetate to methane. Acetogenic bacteria oxidize the acids, obtaining more acetate and
either hydrogen or formate. Finally, methanogens (in the domain Archea) convert acetate to methane.
[10]

Biochemical overview[edit]
Comparison of a aerobic respiration and most known fermentation types in eukaryotic cell.[11] Numbers
in circles indicate counts of carbon atoms in molecules, C6 is glucose C6H12O6, C1 carbon
dioxide CO2. Mitochondrial outer membrane is omitted.

Fermentation reacts NADH with an endogenous, organic electron acceptor.[2] Usually this

is pyruvate formed from sugar through glycolysis. The reaction produces NAD+ and an organic product,
typical examples being ethanol, lactic acid, and hydrogen gas (H2), and often also carbon dioxide.
However, more exotic compounds can be produced by fermentation, such as butyric acid and acetone.
Fermentation products are considered waste products, since they cannot be metabolized further
without the use of oxygen.[12]

Fermentation normally occurs in an anaerobic environment. In the presence of O2, NADH, and pyruvate
are used to generate ATP in respiration. This is called oxidative phosphorylation. This generates much
more ATP than glycolysis alone. It releases the chemical energy of O 2.[12] For this reason, fermentation is
rarely used when oxygen is available. However, even in the presence of abundant oxygen, some strains
of yeast such as Saccharomyces cerevisiae prefer fermentation to aerobic respiration as long as there is
an adequate supply of sugars (a phenomenon known as the Crabtree effect).[13] Some fermentation
processes involve obligate anaerobes, which cannot tolerate oxygen.[citation needed]

Although yeast carries out the fermentation in the production of ethanol in beers, wines, and other

alcoholic drinks, this is not the only possible agent: bacteria carry out the fermentation in the production
of xanthan gum.[citation needed]

Products of fermentation[edit]

Ethanol[edit]

Main article:  Ethanol fermentation

In ethanol fermentation, one glucose molecule is converted into two ethanol molecules and two carbon
dioxide molecules.[14][15] It is used to make bread dough rise: the carbon dioxide forms bubbles,
expanding the dough into a foam. [16][17] The ethanol is the intoxicating agent in alcoholic beverages such
as wine, beer and liquor.[18] Fermentation of feedstocks, including sugarcane, corn, and sugar beets,
produces ethanol that is added to gasoline.[19] In some species of fish, including goldfish and carp, it
provides energy when oxygen is scarce (along with lactic acid fermentation). [20]

The figure illustrates the process. Before fermentation, a glucose molecule breaks down into two
pyruvate molecules (Glycolysis). The energy from this exothermic reaction is used to bind
inorganic phosphates to ADP, which converts it to ATP, and convert NAD + to NADH. The pyruvates break
down into two acetaldehyde molecules and give off two carbon dioxide molecules as waste products.
The acetaldehyde is reduced into ethanol using the energy and hydrogen from NADH, and the NADH is
oxidized into NAD+ so that the cycle may repeat. The reaction is catalyzed by the enzymes pyruvate
decarboxylase and alcohol dehydrogenase.[14]

Lactic acid[edit]

Main article:  Lactic acid fermentation

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Homolactic fermentation (producing only lactic acid) is the simplest type of fermentation. Pyruvate from
glycolysis[21] undergoes a simple redox reaction, forming lactic acid.[22][23] Overall, one molecule of glucose
(or any six-carbon sugar) is converted to two molecules of lactic acid:

C6H12O6 → 2 CH3CHOHCOOH

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 the type of bacteria that
convert lactose into lactic acid in yogurt, giving it its sour taste. These lactic acid bacteria can carry out
either homolactic fermentation, where the end-product is mostly lactic acid, or heterolactic
fermentation, where some lactate is further metabolized to ethanol and carbon dioxide [22] (via
the phosphoketolase pathway), acetate, or other metabolic products, e.g.:

C6H12O6 → CH3CHOHCOOH + C2H5OH + CO2

If lactose is fermented (as in yogurts and cheeses), it is first converted into glucose and galactose (both
six-carbon sugars with the same atomic formula):

C12H22O11 + H2O → 2 C6H12O6

Heterolactic fermentation is in a sense intermediate between lactic acid fermentation and other types,

e.g. alcoholic fermentation. Reasons to go further and convert lactic acid into something else include:

 The acidity of lactic acid impedes biological processes. This can be beneficial to the fermenting
organism as it drives out competitors that are unadapted to the acidity. As a result, the food will
have a longer shelf life (one reason foods are purposely fermented in the first place); however,
beyond a certain point, the acidity starts affecting the organism that produces it.
  The high concentration of lactic acid (the final product of fermentation) drives the equilibrium
backwards (Le Chatelier's principle), decreasing the rate at which fermentation can occur and
slowing down growth.

 Ethanol, into which lactic acid can be easily converted, is volatile and will readily escape,
allowing the reaction to proceed easily. CO2 is also produced, but it is only weakly acidic and even
more volatile than ethanol.

 Acetic acid (another conversion product) is acidic and not as volatile as ethanol; however, in the
presence of limited oxygen, its creation from lactic acid releases additional energy. It is a lighter
molecule than lactic acid, forming fewer hydrogen bonds with its surroundings (due to having
fewer groups that can form such bonds), thus is more volatile and will also allow the reaction to
proceed more quickly.

 If propionic acid, butyric acid, and longer monocarboxylic acids are produced (see mixed acid
fermentation), the amount of acidity produced per glucose consumed will decrease, as with
ethanol, allowing faster growth.

Hydrogen gas[edit]

Main article:  Fermentative hydrogen production

Hydrogen gas is produced in many types of fermentation as a way to regenerate NAD + from

NADH. Electrons are transferred to ferredoxin, whi