Heterotrophic metabolism

As stated above, heterotrophic (or organotrophic) bacteria require organic molecules to provide their carbon and energy. The energy-yielding catabolic reactions can be of many different types, although they all involve electron-transfer reactions in which the movement of an electron from one molecule to another is coupled with an energy-trapping reaction that yields ATP. Some heterotrophic bacteria can metabolize sugars or complex carbohydrates to produce energy. These bacteria must produce a number of specific proteins, including enzymes that degrade the polysaccharides into their constituent sugar units, a transport system to accumulate the sugar inside the cell, and enzymes to convert the sugar into one of the central intermediates of metabolism, such as glucose-6-phosphate. There are several central pathways for carbohydrate utilization, including the Embden-Meyerhof pathway of glycolysis and the pentose phosphate pathway, both of which are also present in eukaryotic cells. Some bacteria possess the Entner-Doudoroff pathway, which converts glucose primarily to pyruvate, as well as other pathways that accomplish the conversion of glucose into smaller compounds with fewer enzyme-catalyzed steps.

Sugar metabolism produces energy for the cell via two different processes, fermentation and respiration. Fermentation is an anaerobic process that takes place in the absence of any external electron acceptor. The organic compound, such as a sugar or amino acid, is broken down into smaller organic molecules, which accept the electrons that had been released during the breakdown of the energy source. These catabolic reactions include a few steps that result in the direct formation of ATP. When glucose is broken down to lactic acid, as occurs in some Lactococcus and Lactobacillus species, as well as in muscle cells in higher eukaryotes, each molecule of glucose yields only two molecules of ATP, and considerable quantities of glucose must be degraded to provide sufficient energy for bacterial growth. Because organic molecules are only partially oxidized during fermentation, the growth of fermentative bacteria results in the production of large quantities of organic end products and a relatively small output of energy per glucose molecule consumed. Few bacteria produce only lactic acid, which is fairly toxic for bacteria and limits the growth of a colony. A variety of additional fermentation pathways are used by specific bacteria to break down glucose; the characteristic end products of these pathways assist in the identification of the bacteria. These end products are often less toxic than lactic acid or are formed with the harnessing of additional metabolic energy. For example, the products of mixed-acid fermentation in E. coli include lactic acid, succinic acid, acetic acid, formic acid, ethyl alcohol, carbon dioxide, and hydrogen gas. Enterobacter aerogenes produces most of the same set of fermentation products, as well as large amounts of 2,3-butylene glycol, which is nonacidic and permits more bacterial growth.

Considerably more energy is available to the cell from respiration, a process in which the electrons from molecules of sugar are transferred not to another organic molecule but to an inorganic molecule. The most familiar respiratory process (aerobic respiration) uses oxygen as the final electron acceptor. The sugar is completely broken down to carbon dioxide and water, yielding a maximum of 38 molecules of ATP per molecule of glucose. Electrons are transferred to oxygen using the electron transport chain, a system of enzymes and cofactors located in the cell membrane and arranged so that the passage of electrons down the chain is coupled with the movement of protons (hydrogen ions) across the membrane and out of the cell. Electron transport induces the movement of positively charged hydrogen ions to the outside of the cell and negatively charged ions to its interior. This ion gradient results in the acidification of the external medium and an energized plasma membrane with an electrical charge of 150 to 200 millivolts. The generation of ion gradients, including this protonmotive force (gradient of protons), is a common aspect of energy generation and storage in all living organisms. The gradient of protons is used directly by the cell for many processes, including the active transport of nutrients and the rotation of flagella. The protons also can move from the exterior of the cell into the cytoplasm by passing through a membrane enzyme called the F₁F₀-proton-translocating ATPase, which couples this proton movement to ATP synthesis in a process identical to that which occurs in the mitochondria of eukaryotic cells (see metabolism: The combustion of food materials).

Bacteria that are able to use respiration produce far more energy per sugar molecule than do fermentative cells, because the complete oxidation (breakdown) of the energy source allows complete extraction of all of the energy available as shown by the substantially greater yield of ATP for respiring organisms than for fermenting bacteria. Respiring organisms achieve a greater yield of cell material using a given amount of nutrient; they also generate fewer toxic end products. The solubility of oxygen in water is limited, however, and the growth and survival of populations of aerobic bacteria are directly proportional to the available supply of oxygen. Continuous supplies of oxygen are available only to bacteria that come into contact with air, as occurs when bacteria are able to float on a surface that exposes them to air or when the medium in which the bacteria live is stirred vigorously.

Respiration can also occur under anaerobic conditions by processes called anaerobic respiration, in which the final electron acceptor is an inorganic molecule, such as nitrate (NO₃⁻), nitrite (NO₂⁻), sulfate (SO₄²⁻), or carbon dioxide (CO₂). The energy yields available to the cell using these acceptors are lower than in respiration with oxygen—much lower in the case of sulfate and carbon dioxide—but they are still substantially higher than the energy yields available from fermentation. The ability of some bacteria to use inorganic molecules in anaerobic respiration can have environmental significance. E. coli can use oxygen, nitrate, or nitrite as an electron acceptor, and Pseudomonas stutzeri is of major global importance for its activity in denitrification, the conversion of nitrate to nitrite and dinitrogen gas (N₂). Desulfovibrio and Desulfuromonas reduce sulfate and elemental sulfur (S), respectively, yielding sulfide (S²⁻), and the bacterium Acetobacterium woodii and methanogenic archaea, such as Methanothermobacter thermautotrophicus, reduce carbon dioxide to acetate and methane, respectively. The Archaea typically use hydrogen as an electron donor with carbon dioxide as an electron acceptor to yield methane or with sulfate as an electron acceptor to yield sulfide.