
In chemistry and biology, fermentation is a biochemical process that obtains energy from carbohydrates without using oxygen. Many foods come from fermentation, plus the process has industrial applications. Here is the definition of fermentation, examples of fermented products, and a look at how fermentation works.
Key Takeaways: Fermentation
- Fermentation is an anaerobic (oxygen-free) energy-generating process.
- It converts carbohydrates into energy, alcohol, acids, or gases.
- Common fermentation products include ethanol, lactic acid, and carbon dioxide.
- It occurs in yeast, bacteria, and animal cells (e.g., muscle cells).
- Humans use fermentation to make bread, alcohol, yogurt, and biofuels.
Fermentation Definition
Fermentation is a metabolic process in organisms that converts carbohydrates into chemical energy, without requiring oxygen. In other words, it is an anaerobic process. In contrast, cellular respiration produces energy, but it is an aerobic process (requires oxygen). In addition to energy molecules (such as ATP), fermentation produces a variety of molecules, including ethanol, carbon dioxide, lactic acid, methanol, hydrogen, methane, butyric acid, acetone, and acetic acid. Examples of organisms that carry out fermentation include fungi (yeast), animals (humans, cattle), and bacteria (Clostridium).
The word fermentation comes from the Latin word fervere, which means “to boil.”
Fermentation Examples
While organisms use fermentation mainly for energy, people apply the process for making many products. You may know that beer, wine, and cheese come from fermentation, but some other examples may surprise you.
- Beer
- Wine
- Mead
- Liquor
- Cheese
- Yogurt
- Sour food containing lactic acid, such as kimchi, sauerkraut, pickles, and pepperoni
- Leavened bread
- Industrial alcohol, as for biofuels
- Sewage treatment involves fermentation.
- Human muscles initially use aerobic respiration, but switch to fermentation and produce lactic acid as an anaerobic energy supply.
- Bacteria in the human digestive tract perform fermentation, producing hydrogen gas and sometimes methane as flatus (farts). Herbivores, like cattle, release more methane.
Types of Fermentation
Fermentation takes several forms, depending on the organism and end products. While all fermentation is anaerobic and involves carbohydrate breakdown, each type has distinct biochemistry, products, and uses. Here are the main types of fermentation found in nature and industry:
Alcoholic Fermentation
Alcoholic fermentation converts sugars into ethanol and carbon dioxide. It is the most well-known type, especially in baking and alcohol production.
- Organisms: Saccharomyces cerevisiae (brewer’s and baker’s yeast), some bacteria
- Main products: Ethanol (C₂H₅OH) and CO₂
- Applications: Beer, wine, liquor, leavened bread, biofuels
Lactic Acid Fermentation
Lactic acid fermentation produces lactic acid instead of ethanol. It occurs in certain bacteria and in animal muscle cells under low oxygen.
- Organisms: Lactobacillus, Streptococcus, human muscle cells
- Main product: Lactic acid (C₃H₆O₃)
- Applications: Yogurt, sour cream, kimchi, sauerkraut, muscle fatigue response
There are two types of lactic acid fermentation:
- Homolactic (produces only lactic acid)
- Heterolactic (produces lactic acid, ethanol or acetic acid, and CO₂)
Acetic Acid Fermentation
Acetic acid fermentation oxidizes ethanol into acetic acid using aerobic bacteria.
- Organisms: Acetobacter, Gluconobacter
- Main product: Acetic acid (CH₃COOH)
- Applications: Vinegar production, kombucha brewing
Butyric Acid Fermentation
Butyric acid fermentation produces butyric acid and gases. It occurs in certain obligate anaerobic bacteria, especially in the gut and in spoiled food.
- Organisms: Clostridium butyricum, Clostridium acetobutylicum
- Main products: Butyric acid (C₄H₈O₂), CO₂, and hydrogen gas
- Applications: Cheese flavoring, industrial solvents (acetone, butanol)
Mixed Acid Fermentation
Mixed acid fermentation yields a complex mixture of acids and gases, typically seen in enteric bacteria.
- Organisms: Escherichia coli, other Enterobacteriaceae
- Main products: Lactic acid, acetic acid, ethanol, CO₂, H₂, and succinic acid
- Applications: Anaerobic waste digestion, microbial testing
Propionic Acid Fermentation
Propionic acid fermentation produces propionic acid and CO₂, giving Swiss cheese its distinctive holes and flavor.
- Organisms: Propionibacterium freudenreichii
- Main products: Propionic acid (C₃H₆O₂), acetic acid, CO₂
- Applications: Swiss-type cheeses, preservatives
| Fermentation Type | Main Product(s) | Example Organism | Example Use |
|---|---|---|---|
| Alcoholic | Ethanol, CO₂ | Saccharomyces cerevisiae | Beer, bread |
| Lactic acid | Lactic acid | Lactobacillus | Yogurt, kimchi |
| Acetic acid | Acetic acid | Acetobacter | Vinegar |
| Butyric acid | Butyric acid, gases | Clostridium butyricum | Intestinal flora |
| Mixed acid | Multiple acids, gases | E. coli | Anaerobic digestion |
| Propionic acid | Propionic acid | Propionibacterium | Swiss cheese |
Biochemistry of Yeast Fermentation – A Closer Look
The classic fermentation example is yeast fermentation of sucrose (a sugar) into ethanol and carbon dioxide. Each sucrose molecule consists of a glucose subunit and a fructose subunit. For every mole of glucose, fermentation produces two moles of ethanol, two moles of carbon dioxide, and two moles of adenosine triphosphate or ATP. The overall chemical reaction is as follows:
C6H12O6 → 2 C2H5OH + 2 CO2
But, fermentation is a process and not a single chemical reaction. It occurs in multiple steps.
(1) In the first step, the enzyme invertase breaks the glycosidic linkage between the glucose and fructose residues of sucrose.
C12H22O11 + H2O + invertase → 2 C6H12O6
(2) Next, glycolysis occurs. This is where each glucose molecule breaks into two pyruvate molecules. Glycolysis takes several steps, but here is the overall chemical equation:
glucose + 2 ADP + 2 inorganic phosphate → 2 pyruvate + 2 ATP + 2 NAD + 2 water + 2 protons
C6H12O6 + 2 ADP + 2 Pi + 2 NAD+ → 2 CH3COCOO− + 2 ATP + 2 NADH + 2 H2O + 2 H+
(3) Finally, pyruvate reacts and forms ethanol and carbon dioxide. This occurs in two steps and regenerates the oxidized NAD+ for glycolysis:
CH3COCOO− + H+ → CH3CHO + CO2 (catalyzed by pyruvate decarboxylase)
CH3CHO + NADH + H+ → C2H5OH + NAD+ (catalyzed by alcohol dehydrogenase)
These reactions convert two moles each of NAD+ and ADP into two moles each of NADH, ATP, and water.
Fermentation is not as efficient at energy production as cellular respiration, so organisms capable of both processes typically utilize respiration when oxygen is available. However, the presence of oxygen does not necessarily prevent fermentation from occurring. For example, yeast prefer fermentation over cellular respiration as long there is a sufficient sugar supply.
History
People have been using fermentation since at least the Neolithic (7000 to 6600 BCE), primarily for fermenting beverages and making cheese. However, it wasn’t until the nineteenth century that scientists started understanding the process. In 1837, Theodor Schwann observed yeast budding using a microscope and found that boiling grape juice prevented fermentation until new yeast was added. But, many chemists still believed fermentation was a simple chemical reaction that could occur without a living organism. In the 1850s and 1860s, Louis Pasteur repeated Schwann’s experiments and demonstrated fermentation came from living cells. However, he could not extract the enzyme responsible for the process. In 1897, German chemist Eduard Buechner ground up yeast, extracted fluid, and discovered this fluid fermented a sugar solution. His experiment earned him the 1907 Nobel Prize in Chemistry.
Related Terms
Studying fermentation for practical applications is called zymurgy. The name comes from a Greek word that literally means “the workings of fermentation.” The science of studying fermentation is zymology. A person who practices fermentation is a zymurgist, while a scientist specializing in fermentation is a zymologist.
Interesting Fermentation Facts
- Yeast fermentation produces carbon dioxide gas bubbles that expand during cooking and make baked goods rise. But, yeast also produces alcohol (ethanol). Less than 2% of this alcohol remains after baking.
- Overgrowth of yeast in the gut can cause auto-intoxication. This is where yeast produces ethanol that gets in the blood stream and causes intoxication even when a person has not been drinking.
- Yeast converts sugar into ethanol, which is safe for human consumption. But, if high levels of pectin are present, one fermentation product is toxic methanol.
Fermentation vs Cellular Respiration
Fermentation and cellular respiration are both metabolic pathways for converting glucose into energy, but they differ in oxygen use, energy yield, and byproducts. This section compares these processes side by side.
| Feature | Fermentation | Cellular Respiration |
|---|---|---|
| Oxygen requirement | Anaerobic (no oxygen) | Aerobic (requires oxygen) |
| Location in cell | Cytoplasm | Cytoplasm and mitochondria (in eukaryotes) |
| ATP yield per glucose | Low (2 ATP) | High (up to 36–38 ATP) |
| Final electron acceptor | Organic molecule (e.g., pyruvate or acetaldehyde) | Oxygen |
| Primary products | Lactic acid, ethanol, CO₂, etc. | CO₂ and water |
| Energy efficiency | Less efficient | More efficient |
| Organisms | Bacteria, yeast, muscle cells | Most plants, animals, fungi, protists |
| When it occurs | Anaerobic conditions or preference (e.g., yeast) | When oxygen is available |
| Role of mitochondria | Not involved | Crucial (site of Krebs cycle and ETC) |
Summary
- Fermentation is faster but less efficient than cellular respiration.
- Some organisms switch between both, depending on oxygen availability (facultative anaerobes).
- In fermentation, the goal is to regenerate NAD⁺ so glycolysis can continue producing ATP.
References
- Akhavan, Bobak; Luis Ostrosky-Zeichner; Thomas, Eric (2019). “Drunk Without Drinking: A Case of Auto-Brewery Syndrome.” ACG Case Reports Journal. 6(9): e00208. doi:10.14309/crj.0000000000000208
- Hui, Y. H. (2004). Handbook of Vegetable Preservation and Processing. New York: M. Dekker. ISBN 0-8247-4301-6.
- Klein, Donald W.; Lansing M.; Harley, John (2006). Microbiology (6th ed.). New York: McGraw-Hill. ISBN 978-0-07-255678-0.
- Purves, William K.; Sadava, David E.; Orians, Gordon H.; Heller, H. Craig (2003). Life, the Science of Biology (7th ed.). Sunderland, Mass.: Sinauer Associates. ISBN 978-0-7167-9856-9.
- Steinkraus, Keith (2018). Handbook of Indigenous Fermented Foods (2nd ed.). CRC Press. ISBN 9781351442510.
- Tortora, Gerard J.; Funke, Berdell R.; Case, Christine L. (2010). Microbiology: An Introduction (10th ed.). San Francisco, CA: Pearson Benjamin Cummings. ISBN 978-0-321-58202-7.
