Purple sulfur bacteria

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Purple sulfur bacteria
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Bacteria
Phylum: Pseudomonadota
Class: Gammaproteobacteria
Order: Chromatiales
Imhoff 2005 [1]
Families

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. [5] [6] 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. [5] The sulfur is oxidized to produce granules of elemental sulfur. This, in turn, may be oxidized to form sulfuric acid.

The purple sulfur bacteria are largely divided into two families, the Chromatiaceae and the Ectothiorhodospiraceae, which produce internal and external sulfur granules respectively, and show differences in the structure of their internal membranes. [5] They make up part of the order Chromatiales, included in the Gammaproteobacteria. The genus Halothiobacillus is also included in the Chromatiales, in its own family, but it is not photosynthetic.

Characteristics of purple sulfur bacteria

Major photosynthetic pigments: Bacteriochlorophylls a or b

Location of photosynthetic pigments: Plasma membrane and chromatophore (lamellar membrane complexes that are continuous with the plasma membrane)

Photosynthetic electron donors: H2, H2S, S

Sulfur deposition: Inside the cell

Metabolic type: Photolithoautotroph [7]

Ecology

Habitat

Purple sulfur bacteria are generally found in illuminated anoxic zones of lakes and other aquatic habitats where hydrogen sulfide accumulates and also in "sulfur springs" where geochemically or biologically produced hydrogen sulfide can trigger the formation of blooms of purple sulfur bacteria. Anoxic conditions are required for photosynthesis; these bacteria cannot thrive in oxygenated environments. [8]

The most favorable lakes for the development of purple sulfur bacteria are meromictic (permanently stratified) lakes. [9] Meromictic lakes stratify because they have denser (usually saline) water in the bottom and less dense (usually fresh water) nearer the surface. Growth of purple sulfur bacteria is also supported by the layering in holomictic lakes. [9] These lakes are thermally stratified; in the spring and summer time, water at the surface is warmed making it less dense than underlying colder water which provides sufficiently stable stratification for purple sulfur bacteria growth. If sufficient sulfate is present to support sulfate reduction, the sulfide, produced in the sediments, diffuses upward into the anoxic bottom waters, where purple sulfur bacteria can form dense cell masses, called blooms, usually in association with green phototrophic bacteria.

Purple sulfur bacteria can also be found and are a prominent component in intertidal microbial mats. Mats, such as the Sippewissett Microbial Mat, have dynamic environments due to the flow of tides and incoming fresh water leading to similarly stratified environments as meromictic lakes. Purple sulfur bacteria growth is enabled as sulfur is supplied from the death and decomposition of microorganisms located above them within these intertidal pools. [5] The stratification and sulfur source allows the PSB to grow in these intertidal pools where the mats occur. The PSB can help stabilize these microbial mat environment sediments through the secretion of extracellular polymeric substances that can bind the sediments in the pools. [10] [11]

Ecological significance

Purple sulfur bacteria are able to affect their environment by contributing to nutrient cycling, and by using their metabolism to alter their surroundings. They are able to play a significant role in primary production suggesting that these organisms affect the carbon cycle through carbon fixation. [12] Purple sulfur bacteria also contribute to the phosphorus cycle in their habitat, [13] and the iron cycle. [14] Through upwelling of these organisms, phosphorus, a limiting nutrient in the oxic layer of lakes, is recycled and provided to heterotrophic bacteria for use. [13] This indicates that although purple sulfur bacteria are found in the anoxic layer of their habitat, they are able to promote the growth of many heterotrophic organisms by supplying inorganic nutrients to the above oxic layer. Another form of recycling of inorganic nutrients and dissolved organic matter by purple sulfur bacteria is through the food chain; they act as a source of food to other organisms. [13]

Some purple sulfur bacteria have evolved to optimize their environmental conditions for their own growth. For example, in the South Andros Black Hole in the Bahamas, purple sulfur bacteria adopted a new characteristic in which they are able to use their metabolism to radiate heat energy into their surroundings. [15] Due to the inefficiency of their carotenoids, or light-harvesting centres, the organisms are able to release excess light energy as heat energy. [15] This adaptation allows them to compete more effectively within their environment. By raising the temperature of the surrounding water, they create an ecological niche which supports their own growth, while also allowing them to outcompete other non-thermotolerant organisms.

Growth in meromictic lakes

Meromictic lakes are permanently stratified lakes produced by a gradient of saline concentrations. The highly salinated bottom layer is separated from the top layer of fresh water by the chemocline, where the salinity changes drastically. Due to the large difference in density, the upper and lower layers do not mix, resulting in an anoxic environment below the chemocline. [16] This anoxic environment with light and sufficient sulfide availability is ideal for purple sulfur bacteria. [17] [16]

A study done at the Mahoney Lake suggested that purple sulfur bacteria contributes to the recycling of the inorganic nutrient, phosphorus. [16] The upwelling of purple sulfur bacteria into the top layer of water creates a source of bound phosphorus, and phosphatase activity releases this phosphorus into the water. The soluble phosphorus is then incorporated into heterotrophic bacteria for use in developmental processes. In this way, purple sulfur bacteria participates in the phosphorus cycle and minimizes nutrient loss. [16]

Biomarkers

Purple sulfur bacteria make conjugated pigments called carotenoids that function in the light harvesting complex. When these organisms die and sink, some pigment molecules are preserved in modified form in the sediments. One carotenoid molecule produced, okenone, is diagenetically altered to the biomarker okenane. The discovery of okenane in marine sediments implies the presence of purple sulfur bacteria during the time of burial. Okenane has been identified in one sedimentary outcrop from Northern Australia dating to 1640 million years ago. [18] The authors of the study concluded that, based on the presence of purple sulfur bacteria's biomarker, the Paleoproterozoic ocean must have been anoxic and sulfidic at depth. This finding provides evidence for the Canfield Ocean hypothesis.

Bioremediation

Purple sulfur bacteria can contribute to a reduction of environmentally harmful organic compounds and odour emission in manure wastewater lagoons where they are known to grow. Harmful compounds such as methane, a greenhouse gas, and hydrogen sulfide, a pungent, toxic compound, can be found in wastewater lagoons. PSB can help lower the concentration of both, and others. [19]

Harmful organic compounds can be removed through photoassimilation, the uptake of carbon by organisms through photosynthesis. [20] When PSB in the lagoons perform photosynthesis they can utilize the carbon from harmful compounds, such as methane, [21] as their carbon source. This removes methane, a greenhouse gas, from the lagoon and reduces the lagoons' atmospheric pollution affect.

H2S can act as a sulfur source for PSB during these same photosynthetic processes that remove the organic compounds. The use of H2S as a reducing agent by PSB removes it from the lagoon and leads to a reduction of odour and toxicity in the lagoons. [22] [23] [24]

See also

Related Research Articles

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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">Green Lake (New York)</span> Lake in New York

Green Lake is the larger of the two lakes in Green Lakes State Park, which lies about 9 miles (14 km) east of downtown Syracuse in Onondaga County, New York. Round Lake is the smaller lake located west of Green Lake. Both lakes are meromictic, which means no seasonal mixing of surface and bottom waters occurs. Meromictic lakes are fairly rare; they have been extensively studied, in part because their sediments can preserve a historical record extending back thousands of years, and because of the euxinic conditions which can form in the deep water.

<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.

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.

<i>Beggiatoa</i> Genus of bacteria

Beggiatoa is a genus of Gammaproteobacteria belonging to the order Thiotrichales, in the Pseudomonadota phylum. These bacteria form colorless filaments composed of cells that can be up to 200 µm in diameter, and are one of the largest prokaryotes on Earth. Beggiatoa are chemolithotrophic sulfur-oxidizers, using reduced sulfur species as an energy source. They live in sulfur-rich environments such as soil, both marine and freshwater, in the deep sea hydrothermal vents, and in polluted marine environments. In association with other sulfur bacteria, e.g. Thiothrix, they can form biofilms that are visible to the naked eye as mats of long white filaments; the white color is due to sulfur globules stored inside the cells.

Anoxic waters are areas of sea water, fresh water, or groundwater that are depleted of dissolved oxygen. The US Geological Survey defines anoxic groundwater as those with dissolved oxygen concentration of less than 0.5 milligrams per litre. Anoxic waters can be contrasted with hypoxic waters, which are low in dissolved oxygen. This condition is generally found in areas that have restricted water exchange.

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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">Soda lake</span> Lake that is strongly alkaline

A soda lake or alkaline lake is a lake on the strongly alkaline side of neutrality, typically with a pH value between 9 and 12. They are characterized by high concentrations of carbonate salts, typically sodium carbonate, giving rise to their alkalinity. In addition, many soda lakes also contain high concentrations of sodium chloride and other dissolved salts, making them saline or hypersaline lakes as well. High pH and salinity often coincide, because of how soda lakes develop. The resulting hypersaline and highly alkalic soda lakes are considered some of the most extreme aquatic environments on Earth.

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

Isorenieratene /ˌaɪsoʊrəˈnɪərətiːn/ is a carotenoid light harvesting pigment produced exclusively by the genus Chlorobium. Chlorobium are the brown-colored strains of the family of green sulfur bacteria (Chlorobiaceae). Green sulfur bacteria are anaerobic photoautotrophic organisms meaning they perform photosynthesis in the absence of oxygen using hydrogen sulfide in the following reaction:

The Sippewissett microbial mat is a microbial mat in the Sippewissett Salt Marsh located along the lower eastern Buzzards Bay shoreline of Cape Cod, about 5 miles north of Woods Hole and 1 mile southwest of West Falmouth, Massachusetts, in the United States. The marsh has two regions, the Great Sippewisset Marsh to the north and Little Sippewisset Marsh to the south, separated from each other by a narrow tongue of land. The marsh extends into an estuary in which the intertidal zone provides a dynamic environment that supports a diverse ecology, including threatened and endangered species such as the roseate tern. The ecology of the salt marsh is based in and supported by the microbial mats which cover the ground of the marsh.

Arsenate-reducing bacteria are bacteria which reduce arsenates. Arsenate-reducing bacteria are ubiquitous in arsenic-contaminated groundwater (aqueous environment). Arsenates are salts or esters of arsenic acid (H3AsO4), consisting of the ion AsO43−. They are moderate oxidizers that can be reduced to arsenites and to arsine. Arsenate can serve as a respiratory electron acceptor for oxidation of organic substrates and H2S or H2. Arsenates occur naturally in minerals such as adamite, alarsite, legrandite, and erythrite, and as hydrated or anhydrous arsenates. Arsenates are similar to phosphates since arsenic (As) and phosphorus (P) occur in group 15 (or VA) of the periodic table. Unlike phosphates, arsenates are not readily lost from minerals due to weathering. They are the predominant form of inorganic arsenic in aqueous aerobic environments. On the other hand, arsenite is more common in anaerobic environments, more mobile, and more toxic than arsenate. Arsenite is 25–60 times more toxic and more mobile than arsenate under most environmental conditions. Arsenate can lead to poisoning, since it can replace inorganic phosphate in the glyceraldehyde-3-phosphate --> 1,3-biphosphoglycerate step of glycolysis, producing 1-arseno-3-phosphoglycerate instead. Although glycolysis continues, 1 ATP molecule is lost. Thus, arsenate is toxic due to its ability to uncouple glycolysis. Arsenate can also inhibit pyruvate conversion into acetyl-CoA, thereby blocking the TCA cycle, resulting in additional loss of ATP.

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.

Okenane, the diagenetic end product of okenone, is a biomarker for Chromatiaceae, the purple sulfur bacteria. These anoxygenic phototrophs use light for energy and sulfide as their electron donor and sulfur source. Discovery of okenane in marine sediments implies a past euxinic environment, where water columns were anoxic and sulfidic. This is potentially tremendously important for reconstructing past oceanic conditions, but so far okenane has only been identified in one Paleoproterozoic rock sample from Northern Australia.

Euxinia or euxinic conditions occur when water is both anoxic and sulfidic. This means that there is no oxygen (O2) and a raised level of free hydrogen sulfide (H2S). Euxinic bodies of water are frequently strongly stratified; have an oxic, highly productive, thin surface layer; and have anoxic, sulfidic bottom water. The word "euxinia" is derived from the Greek name for the Black Sea (Εὔξεινος Πόντος (Euxeinos Pontos)) which translates to "hospitable sea". Euxinic deep water is a key component of the Canfield ocean, a model of oceans during part of the Proterozoic eon (a part specifically known as the Boring Billion) proposed by Donald Canfield, an American geologist, in 1998. There is still debate within the scientific community on both the duration and frequency of euxinic conditions in the ancient oceans. Euxinia is relatively rare in modern bodies of water, but does still happen in places like the Black Sea and certain fjords.

<span class="mw-page-title-main">Microbial oxidation of sulfur</span>

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).

Halorhodospira neutriphila is a bacterium from the genus of Halorhodospira which has been isolated from a microbial mat from a marine saltern from Rhone Delta in France. The microbial mat forms at the sediment surface and is between 10 and 20 mm thick, below a fine layer (2–3 cm) of gypsum crust. The mat is composed of a red layer of purple bacteria strains below a green layer of cyanobacteria, interspersed with sulfur globules, and occasionally covered by halite deposits. These mat forming microbes live in anoxic muds and sediments and form a benthic mat in a hypersaline lagoon environment where the salinity of the water ranges from 240-320‰ of total salinity. H. neutriphila was isolated from the red layer of the microbial layer and found to be extremely halophilic and well adapted to withstand the extreme saline conditions of their modified marine habitat. The type strain was identified as strain SG 3301T.

Chlorobactane is the diagenetic product of an aromatic carotenoid produced uniquely by green-pigmented green sulfur bacteria (GSB) in the order Chlorobiales. Observed in organic matter as far back as the Paleoproterozoic, its identity as a diagnostic biomarker has been used to interpret ancient environments.

Thiodictyon is a genus of gram-negative bacterium classified within purple sulfur bacteria (PSB).

References

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