Purple 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.

Taxonomy

All purple bacteria belong in the phylum of Pseudomonadota. This phylum was established as Proteobacteria by Carl Woese in 1987 calling it "purple bacteria and their relatives". Purple bacteria are distributed between 3 classes: Alphaproteobacteria, Betaproteobacteria, Gammaproteobacteria each characterized by a photosynthetic phenotype. All these classes also contain numerous non-photosynthetic members, such as the nitrogen-fixing Rhizobium and the human gut bacterium Escherichia coli.
Purple non-sulfur bacteria are found in Alphaproteobacteria and Betaproteobacteria. The families are:

Class Alphaproteobacteria
*Order Rhodospirillales
** Family Rhodospirillaceae, e.g. Rhodospirillum rubrum
** Family Acetobacteraceae, e.g. Rhodopila globiformis
* Order Hyphomicrobiales
** Family Nitrobacteraceae, e.g. Rhodopseudomonas palustris
** Family Hyphomicrobiaceae, e.g. Rhodomicrobium
** Family Rhodobiaceae, e.g. Rhodobium
* Order Rhodobacterales, family Rhodobacteraceae
Class Betaproteobacteria
* Family Rhodocyclaceae, e.g. Rhodocyclus
* Family Comamonadaceae, e.g. Rhodoferax

Purple sulfur bacteria are named for the ability to produce elemental sulfur. They are included in the class Gammaproteobacteria, in the two families Chromatiaceae and Ectothiorhodospiraceae. While the former family stores the produced sulfur inside the cell, the latter sends the sulfur outside the cell. According to a 1985 phylogeny, Gammaproteobacteria is divided into three sub-lineages, with both families falling into the first along with non-photosynthetic species such as Nitrosococcus oceani.
The similarity between the photosynthetic machinery in these different lines indicates that it had a common origin, either from some common ancestor or passed by lateral transfer. Purple sulfur bacteria and purple nonsulfur bacteria were distinguished on the basis of physiological factors of their tolerance and utilization of sulfide: was considered that purple sulfur bacteria tolerate millimolar levels of sulfide and oxidize sulfide to sulfur globules stored intracellulary while purple nonsulfur bacteria species did neither. This kind of classification was not absoluted. It was refuted with classic chemostat experiments by Hansen and Van Gemerden that demonstrate the growing of many purple nonsulfur bacteria species at low levels of sulfide and in so doing, oxidize sulfide to S⁰,, or. The important distinction that remains from these two different metabolisms is that: any S⁰ formed by purple nonsulfur bacteria is not stored intracellularly but is deposited outside the cell. So if grown on sulfide it is easy to differentiate purple sulfur bacteria from purple non-sulfur bacteria because the microscopically globules of S⁰ are formed.

Metabolism

Purple bacteria are able to perform different metabolic pathways that allow them to adapt to different and even extreme environmental conditions. They are mainly photoautotrophs, but are also known to be chemoautotrophic and photoheterotrophic. Since pigment synthesis does not take place in presence of oxygen, phototrophic growth only occurs in anoxic and light conditions. However purple bacteria can also grow in dark and oxic environments. In fact they can be mixotrophs, capable of anaerobic and aerobic respiration or fermentation basing on the concentration of oxygen and availability of light.

Photosynthesis

Photosynthetic unit

Purple bacteria use bacteriochlorophyll and carotenoids to obtain the light energy for photosynthesis. Electron transfer and photosynthetic reactions occur at the cell membrane in the photosynthetic unit which is composed by the light-harvesting complexes LHI and LHII and the photosynthetic reaction centre where the charge separation reaction occurs. These structures are located in the intracytoplasmic membrane, areas of the cytoplasmic membrane invaginated to form vesicle sacs, tubules, or single-paired or stacked lamellar sheets which have increased surface to maximize light absorption. Light-harvesting complexes are involved in the energy transfer to the reaction centre. These are integral membrane protein complexes consisting of monomers of α- and β-apoproteins, each one binding molecules of bacteriochlorophyll and carotenoids non-covalently. LHI is directly associated with the reaction centre forming a polymeric ring-like structure around it. LHI has an absorption maximum at 870 nm and it contains most of the bacteriochlorophyll of the photosynthetic unit. LHII contains less bacteriochlorophylls, has lower absorption maximum and is not present in all purple bacteria. Moreover, the photosynthetic unit in Purple Bacteria shows great plasticity, being able to adapt to the constantly changing light conditions. In fact these microorganisms are able to rearrange the composition and the concentration of the pigments, and consequently the absorption spectrum, in response to light variation.

Mechanism

Purple bacteria use cyclic electron transport driven by a series of redox reactions. Light-harvesting complexes surrounding a reaction centre harvest photons in the form of resonance energy, exciting chlorophyll pigments P870 or P960 located in the RC. Excited electrons are cycled from P870 to quinones Q_A and Q_B, then passed to cytochrome bc₁, cytochrome c₂, and back to P870. The reduced quinone Q_B attracts two cytoplasmic protons and becomes QH₂, eventually being oxidized and releasing the protons to be pumped into the periplasm by the cytochrome bc₁ complex. The resulting charge separation between the cytoplasm and periplasm generates a proton motive force used by ATP synthase to produce ATP energy.

Electron donors for anabolism

Purple bacteria are anoxygenic because they do not use water as electron donor to produce oxygen. Purple sulfur bacteria, use sulfide, sulfur, thiosulfate or hydrogen as electron donors. In addition, some species use ferrous iron as electron donor and one strain of Thiocapsa can use nitrite. Finally, even if the purple sulfur bacteria are typically photoautotrophic, some of them are photoheterotrophic and use different carbon sources and electron donors such as organic acids. Purple nonsulfur bacteria typically use hydrogen as an electron donor, but can also use sulfide at lower concentrations compared to PSB and some species can use thiosulfate or ferrous iron as electron donor. In contrast to the purple sulfur bacteria, the purple nonsulfur bacteria are mostly photoheterotrophic and can use a variety of organic compounds as both electron donor and carbon source, such as sugars, amino acids, organic acids, and aromatic compounds like toluene or benzoate.
Purple bacteria lack external electron carriers to spontaneously reduce NAD⁺ to NADH, so they must use their reduced quinones to endergonically reduce NAD⁺. This process is driven by the proton motive force and is called reverse electron flow.

Ecology

Distribution

Purple bacteria inhabit illuminated anoxic aquatic and terrestrial environments. Even if sometimes the two major groups of purple bacteria, purple sulfur bacteria and purple nonsulfur bacteria, coexist in the same habitat, they occupy different niches. Purple sulfur bacteria are strong photoautotrophs and are not adapted to an efficient metabolism and growth in the dark. On the other hand, purple nonsulfur bacteria are strong photoheterotrophs, even if they are capable of photoautotrophy, and are equipped for living in dark environments. Purple sulfur bacteria can be found in different ecosystems with enough sulfate and light, for example shallow lagoons polluted by sewage or deep waters of lakes, in which they could even bloom. They can also be found in microbial mats where the lower layer decomposes and sulfate reduction occurs.
Purple nonsulfur bacteria can be found in both illuminated and dark environments with lack of sulfide. However, they hardly form blooms with sufficiently high concentration to be visible without enrichment techniques.
Purple bacteria have evolved effective strategies for photosynthesis in extreme environments, and are quite successful in harsh habitats. In the 1960s the first halophiles and acidophiles of the genus Ectothiorhodospira were discovered. In the 1980s Thermochromatium tepidum, a thermophilic purple bacterium that can be found in North American hot springs, was isolated for the first time.

Biogeochemical cycles

Purple bacteria are involved in the biogeochemical cycles of different nutrients. In fact they are able to photoautotrophically fix carbon, or to consume it photoheterotrophically; in both cases in anoxic conditions. However the most important role is played by consuming hydrogen sulfide: a highly toxic substance for plants, animals and other bacteria. The oxidation of hydrogen sulfide by purple bacteria produces non-toxic forms of sulfur, such as elemental sulfur and sulfate.
In addition, almost all non-sulfur purple bacteria are able to fix nitrogen, and Rhodopseudomonas sphaeroides, an alpha proteobacter, is capable of reducing nitrate to molecular nitrogen by denitrification.