Fig. 1 - Glycolysis |
Fermentation, according to Louis Pasteur, is
essentially respiration in the absence of oxygen. This, however, is an
oversimplification of the process.
More
technically, fermentation involves the breakdown of carbohydrate molecules to
form an intermediate product, pyruvate and a small amount of energy. This stage
of the process is known as glycolysis, and it occurs inside all living cells
(see figure 1).
The
pyruvate molecule may then be changed into products such as ethanol, lactic
acid, carbon dioxide, hydrogen, citric or acetic acid, depending on the type of
organism and the conditions. Further metabolism of these products may occur if
oxygen becomes available – one example of this is the oxidation of ethanol to
acetic acid when wine is spoiled.
The Production of Alcohol
Beers,
wines and spirits are all produced by fermenting various carbohydrates. Yeasts
do this naturally to sugars; a property that has been utilised by humans for
thousands of years. Wine was probably first produced in the Middle East as long
ago as 5000 BC, and evidence of a process involving crushed grapes dating from
6500 years ago has been found in Macedonia.
Fig, 2 - Fermentation Vats |
Modern
wine makers combine additional yeast varieties to those present naturally in
the harvested grapes, and the process of fermentation from sugar to ethanol and
carbon dioxide occurs in large steel or plastic vats or oak barrels (see figure
2). The carbon dioxide produced is either allowed to escape, or trapped to form
sparkling wines.
Yeasts
are also instrumental in the production of beer, manufactured since around 6000
BC. Unlike wine making, the source of carbohydrates is partly germinated malt
barley rather than grape sugar (see figure 3). The reason malt grain needs to
be germinated is that this procedure produces enzymes that begin the conversion
of starches into sugars. Hops are added to produce a bitter flavour during the
brewing process, and also to act as a preservative.
Spirits
such as whiskey, brandy and Cognac are produced using distillation to further
concentrate the ethanol to around 37-43% (as opposed to around 12% in wines).
Additional sugar may be added to produce liqueurs.
Ethanol
is also produced industrially on a large scale for use as a biofuel. This has
traditionally involved a two step fermentation procedure using aerated tanks
containing the yeast Saccharomyces cerevisciae and substrate carbohydrates.
Recent studies have shown that the bacterium Zymomonas mobilis produces a
larger quantity of ethanol and, unlike yeast, does not become inhibited by high
ethanol concentrations. Interestingly, Z. mobilis can also be genetically
modified to convert a larger range of sugars into ethanol.
The Production of Citric Acid
Fig. 3 - Germinated Malt Barley |
Citric
acid is a useful product in both the food and pharmaceutical industries; it is
used in food as a preservative and to produce an acidic, sour taste in soft
drinks and other beverages. In the pharmaceutical industry it can be used as a
buffering agent and to clean equipment.
Citric
acid is formed by the fermentation of a molasses substrate by the fungus Aspergillus
niger. The biochemical pathway involved includes the production of pyruvate
in glycolysis, followed by its conversion to citric acid via the condensation
of acetyly co-enzyme A and oxaloacetate. The presence of specific amounts of
trace elements such as iron have proved to be important in this process.
Acetic Acid Production
In
the presence of the Acetobacter bacterium and oxygen, fermented
carbohydrates, ciders or wines can be converted to vinegar (acetic acid). The
result is usually a 5% solution of acetic acid.
Acetic
acid is used in diluted form in the food industry as a condiment and pickling
agent. It is also employed in industry as a solvent and an important reagent in
many organic synthesis reactions.
Lactic Acid Production
Again,
this product is formed after pyruvate has been produced in the glycolysis
pathway. The presence of lactic acid bacteria (Lactobacillus bulgaricus,
Streptococcus thermophilus or similar species) usually results in the
conversion of two pyruvate molecules to lactic acid or lactic acid, ethanol and
carbon dioxide.
In
the food industry, lactic acid fermentation is used in the production of
yoghurt, sauerkraut, pickles and cheeses. Other uses for lactic acid include
tanning leather, manufacturing lacquers and inks, as a moisturising agent in
cosmetics and in the production of polylactic acid. Polylactic acid, which can
be used to make biodegradable plastics, is finding many uses in the biomedical
industry, as well as in areas such as clothing and food packaging.
A Versatile Reaction
Fermentation
certainly produces a diverse range of chemicals and is obviously a key reaction
in many industries. The one thing all these processes have in common is an
initial culture containing carbohydrates and a particular species of
microorganism. Although in all cases pyruvate is produced as an intermediate
step, the microorganisms - whether bacteria, yeasts or fungi - will each
determine the specific end fermentation products.
References
Gunasekaran,
P., and Chandra Raj.,K., 'Ethanol Fermentation Technology', Madurai Kamaraj
University, ias.ac.in, accessed 14/4/2010
Mudgeee
Grape Grower's Association, 2010, 'How is Wine Made?', mudgeewine.com, accessed
13/4/2010
Temph.info,
2010, 'Acetic Acid Fermentation', tempeh.info, accessed 12/4/2010
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