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+ 2 ATP + 2 NADH
+ 2 H
2
O + 2H
+
The chemical formula of pyruvate is CH
3
COCOO
. P
i
stands for the inorganic
phosphate. As shown by the reaction equation, glycolysis causes the reduction of
two molecules of NAD
+
to NADH.
[7]
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.
[7]
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
2
.
[4]
Hydrogen
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.
[9]
History
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.
[10]
Louis Pasteur (18221895), during the 1850s and 1860s, showed that fermentation
was initiated by living organisms in a series of investigations.
[6]
In 1857 Pasteur
showed that lactic acid fermentation is caused by living organisms.
[11]
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.
[12]
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.
[13]
He defined fermentation (incorrectly) as "Life without
air,"
[14]
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.
[14]
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
yeasts.
[15]
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
microorganisms.
[16]
In 1907 Buechner won the Nobel Prize in chemistry for his
work.
[17]
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.
[18]
Strain selection and hybridization developed as well,
affecting most modern food fermentations.
Etymology
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.
[19]
Uses
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
acids.
Food fermentation has been said to serve five main purposes:
[11]
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.
BIBLIOGRAPHY
Websites:
www.google.com
www.wikipedia-biology.com
www.encyclopedia.org