Anoxygenic photosynthesis

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Sulfide is used as a reducing agent during photosynthesis in green and sulfur bacteria.
Energy in the form of sunlight
The light dependent reactions take place when the light excites a reaction center, which donates an electron to another molecule and starts the electron transport chain to produce ATP and NADPH.
Once NADPH has been produced, the Calvin cycle proceeds as in oxygenic photosynthesis, turning CO2 into glucose. Anoxygenic Photosynthesis in Green Sulfur Bacteria.svg
Sulfide is used as a reducing agent during photosynthesis in green and sulfur bacteria.
  1. Energy in the form of sunlight
  2. The light dependent reactions take place when the light excites a reaction center, which donates an electron to another molecule and starts the electron transport chain to produce ATP and NADPH.
  3. Once NADPH has been produced, the Calvin cycle proceeds as in oxygenic photosynthesis, turning CO2 into glucose.

Anoxygenic photosynthesis is a special form of photosynthesis used by some bacteria and archaea, which differs from the better known oxygenic photosynthesis in plants in the reductant used (e.g. hydrogen sulfide instead of water) and the byproduct generated (e.g. elemental sulfur instead of molecular oxygen).

Contents

Bacteria and archaea

Several groups of bacteria can conduct anoxygenic photosynthesis: green sulfur bacteria (GSB), red and green filamentous phototrophs (FAPs e.g. Chloroflexia), purple bacteria, acidobacteriota, and heliobacteria. [2] [3]

Some archaea (e.g. Halobacterium ) capture light energy for metabolic function and are thus phototrophic but none are known to "fix" carbon (i.e. be photosynthetic). Instead of a chlorophyll-type receptor and electron transport chain, proteins such as halorhodopsin capture light energy with the aid of diterpenes to move ions against a gradient and produce ATP via chemiosmosis in the manner of mitochondria.

Pigments

The photopigments used to carry out anaerobic photosynthesis are similar to chlorophyll but differ in molecular detail and peak wavelength of light absorbed. Bacteriochlorophylls a through g absorb electromagnetic radiation maximally in the near-infrared within their natural membrane milieu. This differs from chlorophyll a, the predominant plant and cyanobacteria pigment, which has peak absorption wavelength approximately 100 nanometers shorter (in the red portion of the visible spectrum).

Reaction centers

There are two main types of anaerobic photosynthetic electron transport chains in bacteria. The type I reaction centers are found in GSB, Chloracidobacterium, and Heliobacteria, while the type II reaction centers are found in FAPs and purple bacteria.

Type I reaction centers

The electron transport chain of green sulfur bacteria — such as is present in the model organism Chlorobaculum tepidum — uses the reaction center bacteriochlorophyll pair, P840. When light is absorbed by the reaction center, P840 enters an excited state with a large negative reduction potential, and so readily donates the electron to bacteriochlorophyll 663, which passes it on down an electron transport chain. The electron is transferred through a series of electron carriers and complexes until it is used to reduce NAD+ to NADH. P840 regeneration is accomplished with the oxidation of a sulfide ion from hydrogen sulfide (or of hydrogen or ferrous iron) by cytochrome c555[ citation needed ].

Type II reaction centers

Although the type II reaction centers are structurally and sequentially analogous to Photosystem II (PSII) in plant chloroplasts and cyanobacteria, known organisms that exhibit anoxygenic photosynthesis do not have a region analogous to the oxygen-evolving complex of PSII.

The electron transport chain of purple non-sulfur bacteria begins when the reaction center bacteriochlorophyll pair, P870, becomes excited from the absorption of light. Excited P870 will then donate an electron to bacteriopheophytin, which then passes it on to a series of electron carriers down the electron chain. In the process, it will generate an electrochemical gradient which can then be used to synthesize ATP by chemiosmosis. P870 has to be regenerated (reduced) to be available again for a photon reaching the reaction-center to start the process anew. Molecular hydrogen in the bacterial environment is the usual electron donor.

Related Research Articles

<span class="mw-page-title-main">Photosynthesis</span> Biological process to convert light into chemical energy

Photosynthesis is a biological process used by many cellular organisms to convert light energy into chemical energy, which is stored in organic compounds that can later be metabolized through cellular respiration to fuel the organism's activities. The term mostly refer to oxygenic photosynthesis, where oxygen is produced as a byproduct, and some of the chemical energy produced is stored in carbohydrate molecules such as sugars, starch and cellulose, which are synthesized from endergonic reaction of carbon dioxide with water. Most plants, algae and cyanobacteria perform photosynthesis; such organisms are called photoautotrophs. Photosynthesis is largely responsible for producing and maintaining the oxygen content of the Earth's atmosphere, and supplies most of the biological energy necessary for complex life on Earth.

<span class="mw-page-title-main">Green sulfur bacteria</span> Family of bacteria

The green sulfur bacteria, Chlorobiota, are a phylum of obligately anaerobic photoautotrophic bacteria that metabolize sulfur.

<i>Chloroflexus aurantiacus</i> Species of bacterium

Chloroflexus aurantiacus is a photosynthetic bacterium isolated from hot springs, belonging to the green non-sulfur bacteria. This organism is thermophilic and can grow at temperatures from 35 °C to 70 °C. Chloroflexus aurantiacus can survive in the dark if oxygen is available. When grown in the dark, Chloroflexus aurantiacus has a dark orange color. When grown in sunlight it is dark green. The individual bacteria tend to form filamentous colonies enclosed in sheaths, which are known as trichomes.

The purple sulfur bacteria (PSB) are part of a group of Pseudomonadota capable of photosynthesis, collectively referred to as purple bacteria. They are anaerobic or microaerophilic, and are often found in stratified water environments including hot springs, stagnant water bodies, as well as microbial mats in intertidal zones. Unlike plants, algae, and cyanobacteria, purple sulfur bacteria do not use water as their reducing agent, and therefore do not produce oxygen. Instead, they can use sulfur in the form of sulfide, or thiosulfate (as well, some species can use H2, Fe2+, or NO2) as the electron donor in their photosynthetic pathways. The sulfur is oxidized to produce granules of elemental sulfur. This, in turn, may be oxidized to form sulfuric acid.

<span class="mw-page-title-main">Bacteriochlorophyll</span> Chemical compound

Bacteriochlorophylls (BChl) are photosynthetic pigments that occur in various phototrophic bacteria. They were discovered by C. B. van Niel in 1932. They are related to chlorophylls, which are the primary pigments in plants, algae, and cyanobacteria. Organisms that contain bacteriochlorophyll conduct photosynthesis to sustain their energy requirements, but the process is anoxygenic and does not produce oxygen as a byproduct. They use wavelengths of light not absorbed by plants or cyanobacteria. Replacement of Mg2+ with protons gives bacteriophaeophytin (BPh), the phaeophytin form.

<span class="mw-page-title-main">Photosystem</span> Structural units of protein involved in photosynthesis

Photosystems are functional and structural units of protein complexes involved in photosynthesis. Together they carry out the primary photochemistry of photosynthesis: the absorption of light and the transfer of energy and electrons. Photosystems are found in the thylakoid membranes of plants, algae, and cyanobacteria. These membranes are located inside the chloroplasts of plants and algae, and in the cytoplasmic membrane of photosynthetic bacteria. There are two kinds of photosystems: PSI and PSII.

Chlorophyll <i>a</i> Chemical compound

Chlorophyll a is a specific form of chlorophyll used in oxygenic photosynthesis. It absorbs most energy from wavelengths of violet-blue and orange-red light, and it is a poor absorber of green and near-green portions of the spectrum. Chlorophyll does not reflect light but chlorophyll-containing tissues appear green because green light is diffusively reflected by structures like cell walls. This photosynthetic pigment is essential for photosynthesis in eukaryotes, cyanobacteria and prochlorophytes because of its role as primary electron donor in the electron transport chain. Chlorophyll a also transfers resonance energy in the antenna complex, ending in the reaction center where specific chlorophylls P680 and P700 are located.

Heliobacteria are a unique subset of prokaryotic bacteria that process light for energy. Distinguishable from other phototrophic bacteria, they utilize a unique photosynthetic pigment, bacteriochlorophyll g and are the only known Gram-positive phototroph. They are a key player in symbiotic nitrogen fixation alongside plants, and use a type I reaction center like green-sulfur bacteria.

<span class="mw-page-title-main">Purple bacteria</span> Group of phototrophic bacteria

Purple bacteria or purple photosynthetic bacteria are Gram-negative proteobacteria that are phototrophic, capable of producing their own food via photosynthesis. They are pigmented with bacteriochlorophyll a or b, together with various carotenoids, which give them colours ranging between purple, red, brown, and orange. They may be divided into two groups – purple sulfur bacteria and purple non-sulfur bacteria. Purple bacteria are anoxygenic phototrophs widely spread in nature, but especially in aquatic environments, where there are anoxic conditions that favor the synthesis of their pigments.

<span class="mw-page-title-main">Chromatiaceae</span> Family of purple sulfur bacteria

The Chromatiaceae are one of the two families of purple sulfur bacteria, together with the Ectothiorhodospiraceae. They belong to the order Chromatiales of the class Gammaproteobacteria, which is composed by unicellular Gram-negative organisms. Most of the species are photolithoautotrophs and conduct an anoxygenic photosynthesis, but there are also representatives capable of growing under dark and/or microaerobic conditions as either chemolithoautotrophs or chemoorganoheterotrophs.

<span class="mw-page-title-main">Phototroph</span> Organism using energy from light in metabolic processes

Phototrophs are organisms that carry out photon capture to produce complex organic compounds and acquire energy. They use the energy from light to carry out various cellular metabolic processes. It is a common misconception that phototrophs are obligatorily photosynthetic. Many, but not all, phototrophs often photosynthesize: they anabolically convert carbon dioxide into organic material to be utilized structurally, functionally, or as a source for later catabolic processes. All phototrophs either use electron transport chains or direct proton pumping to establish an electrochemical gradient which is utilized by ATP synthase, to provide the molecular energy currency for the cell. Phototrophs can be either autotrophs or heterotrophs. If their electron and hydrogen donors are inorganic compounds they can be also called lithotrophs, and so, some photoautotrophs are also called photolithoautotrophs. Examples of phototroph organisms are Rhodobacter capsulatus, Chromatium, and Chlorobium.

Photoheterotrophs are heterotrophic phototrophs – that is, they are organisms that use light for energy, but cannot use carbon dioxide as their sole carbon source. Consequently, they use organic compounds from the environment to satisfy their carbon requirements; these compounds include carbohydrates, fatty acids, and alcohols. Examples of photoheterotrophic organisms include purple non-sulfur bacteria, green non-sulfur bacteria, and heliobacteria. These microorganisms are ubiquitous in aquatic habitats, occupy unique niche-spaces, and contribute to global biogeochemical cycling. Recent research has also indicated that the oriental hornet and some aphids may be able to use light to supplement their energy supply.

<span class="mw-page-title-main">Photosynthetic reaction centre</span> Molecular unit responsible for absorbing light in photosynthesis

A photosynthetic reaction center is a complex of several proteins, pigments and other co-factors that together execute the primary energy conversion reactions of photosynthesis. Molecular excitations, either originating directly from sunlight or transferred as excitation energy via light-harvesting antenna systems, give rise to electron transfer reactions along the path of a series of protein-bound co-factors. These co-factors are light-absorbing molecules (also named chromophores or pigments) such as chlorophyll and pheophytin, as well as quinones. The energy of the photon is used to excite an electron of a pigment. The free energy created is then used, via a chain of nearby electron acceptors, for a transfer of hydrogen atoms (as protons and electrons) from H2O or hydrogen sulfide towards carbon dioxide, eventually producing glucose. These electron transfer steps ultimately result in the conversion of the energy of photons to chemical energy.

<span class="mw-page-title-main">Chlorosome</span>

A chlorosome is a photosynthetic antenna complex found in green sulfur bacteria (GSB) and many green non-sulfur bacteria (GNsB), together known as green bacteria. They differ from other antenna complexes by their large size and lack of protein matrix supporting the photosynthetic pigments. Green sulfur bacteria are a group of organisms that generally live in extremely low-light environments, such as at depths of 100 metres in the Black Sea. The ability to capture light energy and rapidly deliver it to where it needs to go is essential to these bacteria, some of which see only a few photons of light per chlorophyll per day. To achieve this, the bacteria contain chlorosome structures, which contain up to 250,000 chlorophyll molecules. Chlorosomes are ellipsoidal bodies, in GSB their length varies from 100 to 200 nm, width of 50-100 nm and height of 15 – 30 nm, in GNsB the chlorosomes are somewhat smaller.

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.

<span class="mw-page-title-main">Pheophytin</span> Chlorophyll molecules lacking a central Mg2+ ion

Pheophytin or phaeophytin is a chemical compound that serves as the first electron carrier intermediate in the electron transfer pathway of Photosystem II in plants, and the type II photosynthetic reaction center found in purple bacteria. In both PS II and RC P870, light drives electrons from the reaction center through pheophytin, which then passes the electrons to a quinone (QA) in RC P870 and RC P680. The overall mechanisms, roles, and purposes of the pheophytin molecules in the two transport chains are analogous to each other.

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.

<span class="mw-page-title-main">Light-dependent reactions</span> Photosynthetic reactions

Light-dependent reactions is jargon for certain photochemical reactions that are involved in photosynthesis, the main process by which plants acquire energy. There are two light dependent reactions, the first occurs at photosystem II (PSII) and the second occurs at photosystem I (PSI),

Chlorobium chlorochromatii, originally known as Chlorobium aggregatum, is a symbiotic green sulfur bacteria that performs anoxygenic photosynthesis and functions as an obligate photoautotroph using reduced sulfur species as electron donors. Chlorobium chlorochromatii can be found in stratified freshwater lakes.

Photoautotrophs are organisms that use light energy and inorganic carbon to produce organic materials. Eukaryotic photoautotrophs absorb energy through the chlorophyll molecules in their chloroplasts while prokaryotic photoautotrophs use chlorophylls and bacteriochlorophylls present in free-floating thylakoids in their cytoplasm. All known photoautotrophs perform photosynthesis. Examples include plants, algae, and cyanobacteria.

References

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  2. Donald A. Bryant; Niels-Ulrik Frigaard (November 2006). "Prokaryotic photosynthesis and phototrophy illuminated". Trends in Microbiology. 14 (11): 488–496. doi:10.1016/j.tim.2006.09.001. PMID   16997562.
  3. Bryant DA, Costas AM, Maresca JA, Chew AG, Klatt CG, Bateson MM, Tallon LJ, Hostetler J, Nelson WC, Heidelberg JF, Ward DM (27 July 2007). "Candidatus Chloracidobacterium thermophilum: An Aerobic Phototrophic Acidobacterium". Science. 317 (5837): 523–6. Bibcode:2007Sci...317..523B. doi:10.1126/science.1143236. PMID   17656724. S2CID   20419870.