Hydrogenosome

hydrogenosome, membrane-bound organelle found in the cytoplasm of eukaryotic cells (cells with clearly defined nuclei) that is so named because it releases molecular hydrogen (H₂) as a by-product of energy generation under anaerobic (oxygen-deficient) conditions.

The term hydrogenosome was introduced in 1973 to describe a unique structure found in Tritrichomonas foetus, a parasite that lives in the gastrointestinal tracts of cats and in the reproductive tracts of cattle. Since then a variety of organisms have also been found to possess hydrogenosomes, including multiple species of flagellated trichomonads, many of which are parasitic in animals; several free-living anaerobic ciliates, such as Trimyema, Plagiopyla, and Metopus; and some anaerobic chytridiomycete fungi, including Neocallimastix, which lives in the rumen of herbivores. Hydrogenosome-like organelles have been found in several small multicellular marine organisms known as loriciferans—namely, members of the genera Pliciloricus, Spinoloricus, and Rugiloricus.

In hydrogenosome-containing organisms, the hydrogenosomes take the place of the energy-producing mitochondria, and, similar to organisms with mitochondria, organisms with hydrogenosomes make use of the by-products of metabolic reactions that occur in the cell cytoplasm. For example, in anaerobically respiring trichomonads, pyruvate (pyruvic acid) that is generated from glycolysis (the breakdown of glucose) in the cytoplasm enters the hydrogenosome, where it is acted on by enzymes and ultimately produces energy in the form of adenosine triphosphate (ATP). The breakdown of pyruvate in the hydrogenosome begins with the enzyme pyruvate:ferredoxin oxidoreductase, which converts pyruvate into acetyl coenzyme A (acetyl CoA) and carbon dioxide. During this process, electrons are transferred from pyruvate to the ferredoxin portion of the enzyme, and hence the enzyme assumes a reduced state. The enzyme then reacts with protons (H⁺) and the enzyme hydrogenase, resulting in the removal of electrons from ferredoxin and the generation of molecular hydrogen. The enzymes acetate:succinate CoA transferase and succinate thiokinase (succinyl-CoA synthetase) catalyze the subsequent metabolism of acetyl CoA into acetate (acetic acid) and ATP.

In the presence of oxygen, trichomonads and other hydrogenosome-containing organisms switch to aerobic (oxygen-dependent) metabolism. Scientists suspect that in this instance, rather than ferredoxin reacting with protons to form molecular hydrogen, the molecule reacts with molecular oxygen (O₂) to form water.

The similarity of hydrogenosomes to other energy-producing organelles, including mitochondria and chloroplasts, has led to various hypotheses concerning the evolutionary origin of hydrogenosomes. Among the leading hypotheses is that hydrogenosomes and mitochondria share a common evolutionary ancestor. This idea has gained some support from studies that have identified proteins of similar structure and function within hydrogenosome and mitochondrial membranes. However, hydrogenosomes differ significantly from mitochondria and chloroplasts because they do not possess their own DNA. A unique respiratory organelle containing a genome in the anaerobic microorganism Nyctotherus ovalis, which lives in the hindgut of termites and cockroaches, is thought to be representative of an organelle intermediate between mitochondria and hydrogenosomes, lending support to the notion that these organelles might share a common ancestor.