67 Metabolism without Oxygen

In aerobic respiration, the final electron acceptor is an oxygen molecule, O2. If aerobic respiration occurs, then ATP will be produced using the energy of high-energy electrons carried by NADH or FADH2 to the electron transport chain. If aerobic respiration does not occur, NADH must be reoxidized to NAD+ for reuse as an electron carrier for the glycolytic pathway to continue. How is this done? Some living systems use an organic molecule as the final electron acceptor. Processes that use an organic molecule to regenerate NAD+ from NADH are collectively referred to as fermentation. In contrast, some living systems use an inorganic molecule as a final electron acceptor. Both methods are called anaerobic cellular respiration, in which organisms convert energy for their use in the absence of oxygen.

Anaerobic Cellular Respiration

Certain prokaryotes, including some species in the domains Bacteria and Archaea, use anaerobic respiration. For example, a group of archaeans called methanogens reduces carbon dioxide to methane to oxidize NADH. These microorganisms are found in soil and in the digestive tracts of ruminants, such as cows and sheep. Similarly, sulfate-reducing bacteria, most of which are anaerobic (Figure 7.14), reduce sulfate to hydrogen sulfide to regenerate NAD+ from NADH.

Lactic Acid Fermentation

The fermentation method used by animals and certain bacteria, such as those in yogurt, is lactic acid fermentation (Figure 7.15). This type of fermentation is used routinely in mammalian red blood cells, which do not have mitochondria, and in skeletal muscle that has an insufficient oxygen supply to allow aerobic respiration to continue (that is, in muscles used to the point of fatigue). In muscles, lactic acid accumulation must be removed by the blood circulation, and when the lactic acid loses a hydrogen, the resulting lactate is brought to the liver for further metabolism. The chemical reactions of lactic acid fermentation are the following:

Pyruvic acid+NADH↔lactic acid+NAD⁺

The enzyme used in this reaction is lactate dehydrogenase (LDH). The reaction can proceed in either direction, but the reaction from left to right is inhibited by acidic conditions. Such lactic acid accumulation was once believed to cause muscle stiffness, fatigue, and soreness, although more recent research disputes this hypothesis. Once the lactic acid has been removed from the muscle and circulated to the liver, it can be reconverted into pyruvic acid and further catabolized for energy.

Visual Connection

 

 

Tremetol, a metabolic poison found in the white snakeroot plant, prevents the metabolism of lactate. When cows eat this plant, tremetol is concentrated in the milk they produce. Humans who consume the milk can become seriously ill. Symptoms of this disease, which include vomiting, abdominal pain, and tremors, become worse after exercise. Why do you think this is the case?

Alcohol Fermentation

Another familiar fermentation process is alcohol fermentation (Figure 7.16), which produces ethanol. The first chemical reaction of alcohol fermentation is the following (CO2 does not participate in the second reaction):

pyruvic acid+H⁺→CO₂+acetaldehyde+NADH+H⁺→ethanol+NAD⁺

The first reaction is catalyzed by pyruvate decarboxylase, a cytoplasmic enzyme, with a coenzyme of thiamine pyrophosphate (TPP, derived from vitamin B₁ and also called thiamine). A carboxyl group is removed from pyruvic acid, releasing carbon dioxide as a gas. The loss of carbon dioxide reduces the size of the molecule by one carbon, producing acetaldehyde. The second reaction is catalyzed by alcohol dehydrogenase to oxidize NADH to NAD⁺ and reduce acetaldehyde to ethanol. The fermentation of pyruvic acid by yeast produces the ethanol found in alcoholic beverages. Ethanol tolerance of yeast is variable, ranging from about 5 percent to 21 percent, depending on the yeast strain and environmental conditions.