In biochemistry, chemosynthesis is the biological conversion of one or more carbon-containing molecules (usually carbon dioxide or methane) and nutrients into organic matter using the oxidation of inorganic compounds (e.g., hydrogen gas, hydrogen sulfide) or ferrous ions as a source of energy, rather than sunlight, as in photosynthesis. Chemoautotrophs, organisms that obtain carbon from carbon dioxide through chemosynthesis, are phylogenetically diverse. Groups that include conspicuous or biogeochemically important taxa include the sulfur-oxidizing Gammaproteobacteria, the Campylobacterota, the Aquificota, the methanogenic archaea, and the neutrophilic iron-oxidizing bacteria.
Many microorganisms in dark regions of the oceans use chemosynthesis to produce biomass from single-carbon molecules. Two categories can be distinguished. In the rare sites where hydrogen molecules (H2) are available, the energy available from the reaction between CO2 and H2 (leading to production of methane, CH4) can be large enough to drive the production of biomass. Alternatively, in most oceanic environments, energy for chemosynthesis derives from reactions in which substances such as hydrogen sulfide or ammonia are oxidized. This may occur with or without the presence of oxygen.
Many chemosynthetic microorganisms are consumed by other organisms in the ocean, and symbiotic associations between chemosynthesizers and respiring heterotrophs are quite common. Large populations of animals can be supported by chemosynthetic secondary production at hydrothermal vents, methane clathrates, cold seeps, whale falls, and isolated cave water.
It has been hypothesized that anaerobic chemosynthesis may support life below the surface of Mars, Jupiter's moon Europa, and other planets. [1] Chemosynthesis may have also been the first type of metabolism that evolved on Earth, leading the way for cellular respiration and photosynthesis to develop later.
Giant tube worms use bacteria in their trophosome to fix carbon dioxide (using hydrogen sulfide as their energy source) and produce sugars and amino acids. [2] Some reactions produce sulfur:
Instead of releasing oxygen gas while fixing carbon dioxide as in photosynthesis, hydrogen sulfide chemosynthesis produces solid globules of sulfur in the process. In bacteria capable of chemoautotrophy (a form a chemosynthesis), such as purple sulfur bacteria, [4] yellow globules of sulfur are present and visible in the cytoplasm.
In 1890, Sergei Winogradsky proposed a novel type of life process called "anorgoxydant". His discovery suggested that some microbes could live solely on inorganic matter and emerged during his physiological research in the 1880s in Strasbourg and Zürich on sulfur, iron, and nitrogen bacteria.
In 1897, Wilhelm Pfeffer coined the term "chemosynthesis" for the energy production by oxidation of inorganic substances, in association with autotrophic carbon dioxide assimilation—what would be named today as chemolithoautotrophy. Later, the term would be expanded to include also chemoorganoautotrophs, which are organisms that use organic energy substrates in order to assimilate carbon dioxide. [5] Thus, chemosynthesis can be seen as a synonym of chemoautotrophy.
The term "chemotrophy", less restrictive, would be introduced in the 1940s by André Lwoff for the production of energy by the oxidation of electron donors, organic or not, associated with auto- or heterotrophy. [6] [7]
The suggestion of Winogradsky was confirmed nearly 90 years later, when hydrothermal ocean vents were predicted to exist in the 1970s. The hot springs and strange creatures were discovered by Alvin , the world's first deep-sea submersible, in 1977 at the Galapagos Rift. At about the same time, then-graduate student Colleen Cavanaugh proposed chemosynthetic bacteria that oxidize sulfides or elemental sulfur as a mechanism by which tube worms could survive near hydrothermal vents. Cavanaugh later managed to confirm that this was indeed the method by which the worms could thrive, and is generally credited with the discovery of chemosynthesis. [8]
A 2004 television series hosted by Bill Nye named chemosynthesis as one of the 100 greatest scientific discoveries of all time. [9] [10]
In 2013, researchers reported their discovery of bacteria living in the rock of the oceanic crust below the thick layers of sediment, and apart from the hydrothermal vents that form along the edges of the tectonic plates. Preliminary findings are that these bacteria subsist on the hydrogen produced by chemical reduction of olivine by seawater circulating in the small veins that permeate the basalt that comprises oceanic crust. The bacteria synthesize methane by combining hydrogen and carbon dioxide. [11]
Despite the fact that the process of chemosynthesis has been known for more than a hundred years, its significance and importance are still relevant today in the transformation of chemical elements in biogeochemical cycles. Today, the vital processes of nitrifying bacteria, which lead to the oxidation of ammonia to nitric acid, require scientific substantiation and additional research. The ability of bacteria to convert inorganic substances into organic ones suggests that chemosynthetics can accumulate valuable resources for human needs.
Chemosynthetic communities in different environments are important biological systems in terms of their ecology, evolution and biogeography, as well as their potential as indicators of the availability of permanent hydrocarbon- based energy sources. In the process of chemosynthesis, bacteria produce organic matter where photosynthesis is impossible. Isolation of thermophilic sulfate-reducing bacteria Thermodesulfovibrio yellowstonii and other types of chemosynthetics provides prospects for further research. Thus, the importance of chemosynthesis remains relevant for use in innovative technologies, conservation of ecosystems, human life in general. Sergey Winogradsky helped discover the phenomenon of chemosynthesis. [12]
Primary nutritional groups are groups of organisms, divided in relation to the nutrition mode according to the sources of energy and carbon, needed for living, growth and reproduction. The sources of energy can be light or chemical compounds; the sources of carbon can be of organic or inorganic origin.
A heterotroph is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but not producers. Living organisms that are heterotrophic include all animals and fungi, some bacteria and protists, and many parasitic plants. The term heterotroph arose in microbiology in 1946 as part of a classification of microorganisms based on their type of nutrition. The term is now used in many fields, such as ecology, in describing the food chain.
The green sulfur bacteria are a phylum, Chlorobiota, of obligately anaerobic photoautotrophic bacteria that metabolize sulfur.
Hydrothermal vents are fissures on the seabed from which geothermally heated water discharges. They are commonly found near volcanically active places, areas where tectonic plates are moving apart at mid-ocean ridges, ocean basins, and hotspots. The dispersal of hydrothermal fluids throughout the global ocean at active vent sites creates hydrothermal plumes. Hydrothermal deposits are rocks and mineral ore deposits formed by the action of hydrothermal vents.
Riftia pachyptila, commonly known as the giant tube worm and less commonly known as the giant beardworm, is a marine invertebrate in the phylum Annelida related to tube worms commonly found in the intertidal and pelagic zones. R. pachyptila lives on the floor of the Pacific Ocean near hydrothermal vents. The vents provide a natural ambient temperature in their environment ranging from 2 to 30 °C, and this organism can tolerate extremely high hydrogen sulfide levels. These worms can reach a length of 3 m, and their tubular bodies have a diameter of 4 cm (1.6 in).
Sulfate-reducing microorganisms (SRM) or sulfate-reducing prokaryotes (SRP) are a group composed of sulfate-reducing bacteria (SRB) and sulfate-reducing archaea (SRA), both of which can perform anaerobic respiration utilizing sulfate (SO2−
4) as terminal electron acceptor, reducing it to hydrogen sulfide (H2S). Therefore, these sulfidogenic microorganisms "breathe" sulfate rather than molecular oxygen (O2), which is the terminal electron acceptor reduced to water (H2O) in aerobic respiration.
A chemotroph Greek words “chemo” and “troph” is an organism that obtains energy by the oxidation of electron donors in their environments. These molecules can be organic (chemoorganotrophs) or inorganic (chemolithotrophs). The chemotroph designation is in contrast to phototrophs, which use photons. Chemotrophs can be either autotrophic or heterotrophic. Chemotrophs can be found in areas where electron donors are present in high concentration, for instance around hydrothermal vents.
Sergei Nikolaevich Winogradsky (Russian: Сергей Николаевич Виноградский; Ukrainian: Сергій Миколайович Виноградський; 13 September [O.S. 1 September] 1856, Kyiv – 24 February 1953, Brie-Comte-Robert), also published under the name Sergius Winogradsky, was a Ukrainian and Russian microbiologist, ecologist and soil scientist who pioneered the cycle-of-life concept. Winogradsky discovered the first known form of lithotrophy during his research with Beggiatoa in 1887. He reported that Beggiatoa oxidized hydrogen sulfide (H2S) as an energy source and formed intracellular sulfur droplets. This research provided the first example of lithotrophy, but not autotrophy. Born in the capital of present-day Ukraine, his legacy is also celebrated by this nation.
The important sulfur cycle is a biogeochemical cycle in which the sulfur moves between rocks, waterways and living systems. It is important in geology as it affects many minerals and in life because sulfur is an essential element (CHNOPS), being a constituent of many proteins and cofactors, and sulfur compounds can be used as oxidants or reductants in microbial respiration. The global sulfur cycle involves the transformations of sulfur species through different oxidation states, which play an important role in both geological and biological processes. Steps of the sulfur cycle are:
Lithotrophs are a diverse group of organisms using an inorganic substrate to obtain reducing equivalents for use in biosynthesis or energy conservation via aerobic or anaerobic respiration. While lithotrophs in the broader sense include photolithotrophs like plants, chemolithotrophs are exclusively microorganisms; no known macrofauna possesses the ability to use inorganic compounds as electron sources. Macrofauna and lithotrophs can form symbiotic relationships, in which case the lithotrophs are called "prokaryotic symbionts". An example of this is chemolithotrophic bacteria in giant tube worms or plastids, which are organelles within plant cells that may have evolved from photolithotrophic cyanobacteria-like organisms. Chemolithotrophs belong to the domains Bacteria and Archaea. The term "lithotroph" was created from the Greek terms 'lithos' (rock) and 'troph' (consumer), meaning "eaters of rock". Many but not all lithoautotrophs are extremophiles.
Microbial metabolism is the means by which a microbe obtains the energy and nutrients it needs to live and reproduce. Microbes use many different types of metabolic strategies and species can often be differentiated from each other based on metabolic characteristics. The specific metabolic properties of a microbe are the major factors in determining that microbe's ecological niche, and often allow for that microbe to be useful in industrial processes or responsible for biogeochemical cycles.
Hydrogen-oxidizing bacteria are a group of facultative autotrophs that can use hydrogen as an electron donor. They can be divided into aerobes and anaerobes. The former use hydrogen as an electron donor and oxygen as an acceptor while the latter use sulphate or nitrogen dioxide as electron acceptors. Species of both types have been isolated from a variety of environments, including fresh waters, sediments, soils, activated sludge, hot springs, hydrothermal vents and percolating water.
Sulfur is metabolized by all organisms, from bacteria and archaea to plants and animals. Sulfur can have an oxidation state from -2 to +6 and is reduced or oxidized by a diverse range of organisms. The element is present in proteins, sulfate esters of polysaccharides, steroids, phenols, and sulfur-containing coenzymes.
An autotroph is an organism that produces complex organic compounds using carbon from simple substances such as carbon dioxide, generally using energy from light (photosynthesis) or inorganic chemical reactions (chemosynthesis). They convert an abiotic source of energy into energy stored in organic compounds, which can be used by other organisms. Autotrophs do not need a living source of carbon or energy and are the producers in a food chain, such as plants on land or algae in water. Autotrophs can reduce carbon dioxide to make organic compounds for biosynthesis and as stored chemical fuel. Most autotrophs use water as the reducing agent, but some can use other hydrogen compounds such as hydrogen sulfide.
A trophosome is a highly vascularised organ found in some animals that houses symbiotic bacteria that provide food for their host. Trophosomes are contained by the coelom of tube worms and in the body of symbiotic flatworms of the genus Paracatenula.
Microbial oxidation of sulfur is the oxidation of sulfur by microorganisms to build their structural components. The oxidation of inorganic compounds is the strategy primarily used by chemolithotrophic microorganisms to obtain energy to survive, grow and reproduce. Some inorganic forms of reduced sulfur, mainly sulfide (H2S/HS−) and elemental sulfur (S0), can be oxidized by chemolithotrophic sulfur-oxidizing prokaryotes, usually coupled to the reduction of oxygen (O2) or nitrate (NO3−). Anaerobic sulfur oxidizers include photolithoautotrophs that obtain their energy from sunlight, hydrogen from sulfide, and carbon from carbon dioxide (CO2).
The sulfate-methane transition zone (SMTZ) is a zone in oceans, lakes, and rivers typically found below the sediment surface in which sulfate and methane coexist. The formation of a SMTZ is driven by the diffusion of sulfate down the sediment column and the diffusion of methane up the sediments. At the SMTZ, their diffusion profiles meet and sulfate and methane react with one another, which allows the SMTZ to harbor a unique microbial community whose main form of metabolism is anaerobic oxidation of methane (AOM). The presence of AOM marks the transition from dissimilatory sulfate reduction to methanogenesis as the main metabolism utilized by organisms.
The hydrothermal vent microbial community includes all unicellular organisms that live and reproduce in a chemically distinct area around hydrothermal vents. These include organisms in the microbial mat, free floating cells, or bacteria in an endosymbiotic relationship with animals. Chemolithoautotrophic bacteria derive nutrients and energy from the geological activity at Hydrothermal vents to fix carbon into organic forms. Viruses are also a part of the hydrothermal vent microbial community and their influence on the microbial ecology in these ecosystems is a burgeoning field of research.
The deep biosphere is the part of the biosphere that resides below the first few meters of the surface. It extends down at least 5 kilometers below the continental surface and 10.5 kilometers below the sea surface, at temperatures that may reach beyond 120 °C (248 °F) which is comparable to the maximum temperature where a metabolically active organism has been found. It includes all three domains of life and the genetic diversity rivals that on the surface.
Hydrogen sulfide chemosynthesis is a form of chemosynthesis which uses hydrogen sulfide. It is common in hydrothermal vent microbial communities Due to the lack of light in these environments this is predominant over photosynthesis
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