Everything about Fermentation Biochemistry totally explained
Fermentation is the process of deriving energy from the oxidation of organic compounds, such as carbohydrates, using an endogenous electron acceptor, which is usually an organic compound . This is in contrast to
cellular respiration, where electrons are donated to an exogenous electron acceptor, such as oxygen, via an electron transport chain .
Sugars are the 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.
History
French chemist
Louis Pasteur was the first
zymologist, when, in
1857, he connected yeast to fermentation. Pasteur originally defined fermentation as
respiration without air.
Pasteur performed careful research and concluded,
"I am of the opinion that alcoholic fermentation never occurs without simultaneous organization, development and multiplication of cells.... If asked, in what consists the chemical act whereby the sugar is decomposed ... I'm completely ignorant of it."
German chemist and
zymologist,
Eduard Buchner, winner of the
1907 Nobel Prize in chemistry, later determined that fermentation is actually caused by a yeast secretion that he termed
zymase.
Reaction
The reaction of fermentation differs according to the sugar being used and the product produced. In the reaction shown below, the sugar will be
glucose (C
6H
12O
6), the simplest sugar, and the product will be
ethanol (2C
2H
5OH). This is one of the fermentation reactions carried out by yeast, and used in food production.
Chemical equation » ::C
6H
12O
6 → 2 CH
3CH
2OH + 2 CO
2 + 2 ATP (energy released:118
kJ/
mol)
Word equation » ::Sugar (glucose or fructose) → alcohol (ethanol) + carbon dioxide + energy (ATP)
The actual
biochemical pathway the reaction takes varies depending on the sugars involved, but the most common involves part of the
glycolysis pathway, which is shared with the early stages of
aerobic respiration in most organisms. The later stages of the pathway vary considerably depending on the final product.
Energy source in anaerobic conditions
Fermentation is thought to have been the primary means of energy production in earlier organisms before oxygen was at high concentration in the atmosphere, and thus would represent a more ancient form of energy production in cells.
Fermentation products contain chemical energy (they are not fully
oxidized) but are considered waste products, since they can't 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. Fermentation produces 4 ATP molecules per molecule of glucose, compared to 38 by
aerobic respiration: 8 are produced from FADH
2, and 30 are produced from NADH, for a total of 38. Since 2 ATP molecules are used in glycolysis, the net yield for fermentation is 2 ATP versus 36 ATP from aerobic respiration.
Aerobic glycolysis is a method employed by muscle cells for the production of lower-intensity energy over a longer period of time when oxygen is plentiful. Under low-oxygen conditions, however, vertebrates use the less-efficient but faster
anaerobic glycolysis to produce ATP. The speed at which ATP is produced is about 100 times that of
oxidative phosphorylation. While fermentation is helpful during short, intense periods of exertion, it isn't sustained over extended periods in complex aerobic organisms. In humans, for example, lactic acid fermentation provides energy for a period ranging from 30 seconds to 2 minutes.
The final step of fermentation, the conversion of pyruvate to fermentation end-products, doesn't produce energy. However, it's critical for an anaerobic cell, since it regenerates nicotinamide adenine dinucleotide (
NAD+), which is required for glycolysis. This is important for normal cellular function, as glycolysis is a primary source of ATP production in anaerobic conditions.
Products
Products produced by fermentation are actually waste products produced during the reduction of pyruvate to regenerate NAD+ in the absence of oxygen. In general, bacteria produce acids: Vinegar (
acetic acid) is the direct result of bacterial metabolism (Bacteria need oxygen to convert the alcohol to acetic acid). In milk, the acid coagulates the
casein, producing curds. In pickling, the acid preserves the food from pathogenic and putrefactive bacteria.
When yeast ferments, it breaks down the
glucose (C
6H
12O
6) into exactly two molecules of
ethanol (CH
3CH
2OH) and two molecules of
carbon dioxide (CO
2).
- 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 produced.
- Lactic acid fermentation breaks down the pyruvate into lactic acid. It occurs in the muscles of animals when they need energy faster than the blood can supply oxygen. It also occurs in some bacteria and some fungi. It is this type of bacteria that converts lactose into lactic acid in yogurt, giving it its sour taste.
In vertebrates, during intense exercise,
cellular respiration will deplete oxygen in the muscles faster than it can be replenished. An associated burning sensation in muscles has been attributed to
lactic acid's causing a decrease in the pH during a shift to
anaerobic glycolysis. While this does partially explain
acute muscle soreness, lactic acid may also help delay muscle fatigue, although, eventually the lower pH will inhibit enzymes involved in glycolysis. Contrary to currently popular belief, the lactic acid isn't the primary cause for the drop in pH, but rather ATP-derived hydrogen ions.
Delayed onset muscle soreness can't be attributed to the lactic acid and other waste products as they're quickly removed after exercise. It is actually due to
microtrauma of the
muscle fibers. Eventually the
liver metabolizes the lactic acid back to pyruvate.
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
+ and FAD from NADH and FADH
2. Electrons are transferred to ferredoxin, which in turn is oxidized by hydrogenase, producing H
2. 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.
Some anaerobic eukaryotic microorganisms also produce hydrogen gas, in their
hydrogenosomes. The concentration of hydrogen gas is kept low by symbionts such as methanogens that reside in the cytosol of the eukaryote.
Enzymology and Zymology
Zymology is the scientific term for the study of the chemical process of fermentation, and includes fermentation by
yeast,
bacteria, and other lower species. Zymology includes the study of enzymes that facilitate the chemical process of fermentation, as well as engineering issues of
yeast selection, yeast
physiology, and the practical issues of
brewing.
Enzymology, the study of enzymes, shouldn't be confused with zymology, as the modern study of the
enzyme encompasses enzymes from a multitude of sources, and studies enzymes in many terms, as well as studying enzymes from biochemical pathways including fermentation and beyond. Historically, zymologists were the first enzymologists, and since enzymology has expanded beyond the study of yeast fermentation, zymology is infrequently regarded as an archaic word for enzymology.
Further Information
Get more info on 'Fermentation Biochemistry'.
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