Overview of Fermentative Metabolism
Fermentation enables bacteria to generate energy in the absence of oxygen. In this process, hydrogen removed from an organic substrate is transferred to other organic compounds rather than inorganic compounds, which occurs during respiration.
The end products of this metabolism include various alcohols, organic acids, and gases. Fermentation substrates can include carbohydrates, polyols, purines, pyrimidines, and most amino acids. However, aromatic compounds, fatty acids, and steroids remain stable under anaerobic conditions and are not fermented.
Biochemistry of Glucose Fermentation
A key feature of fermentation is the partial oxidation of pyruvate. The tricarboxylic acid (TCA) cycle does not serve an energetic function here, but rather a strictly biosynthetic one, supplying precursors for anabolic metabolism.
- ATP production: 2 ATP molecules are synthesized via substrate-level phosphorylation (during the oxidation of triose phosphate to pyruvate).
- Fate of hydrogen: Hydrogen from reduced NADH is transferred onto pyruvate or its derivatives, converting them into ethanol, acids, or gases.
Major Types of Fermentation
There are several classic pathways of carbohydrate fermentation, each distinguished by its enzyme systems and end products:
- Alcoholic fermentation. Carried out by yeasts under anaerobic conditions via the glycolytic pathway. End products are ethanol and carbon dioxide. In the presence of oxygen, this process shifts to the more energetically favorable respiration (Pasteur effect).
- Lactic acid fermentation. Divided into homofermentative (producing only lactic acid via glycolysis) and heterofermentative (breaking down via the pentose phosphate pathway to produce lactic acid, acetic acid, and ethanol). Typical for lactobacilli, streptococci, and bifidobacteria.
- Mixed acid fermentation. Characteristic of Enterobacteriaceae and vibrios. Divided into acid pathway (producing mixed acids with possible gas evolution) and butanediol pathway (synthesizing acetoin and 2,3-butanediol).
- Butyric acid fermentation. Characteristic of strict anaerobes (saccharolytic clostridia). Products include butyric acid, other volatile fatty acids, and solvents (butanol, acetone).
Protein Fermentation (Peptolytic Metabolism)
Some microorganisms do not utilize carbohydrates, obtaining energy through an alternative pathway via amino acid fermentation and protein hydrolysis. These bacteria are called peptolytic. They include certain members of the genus Clostridium (e.g., Clostridium botulinum and Clostridium histolyticum).