Stigmatella Aurantiaca
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''Stigmatella aurantiaca'' is a member of
myxobacteria The myxobacteria ("slime bacteria") are a group of bacteria that predominantly live in the soil and feed on insoluble organic substances. The myxobacteria have very large genomes relative to other bacteria, e.g. 9–10 million nucleotides except ...
, a group of
gram-negative bacteria Gram-negative bacteria are bacteria that do not retain the crystal violet stain used in the Gram staining method of bacterial differentiation. They are characterized by their cell envelopes, which are composed of a thin peptidoglycan cell wall ...
with a complex developmental life cycle.


Classification

The bacterial nature of this organism was recognized by Thaxter in 1892, who grouped it among the ''Chrondromyces''. It had been described several times before, but had been misclassified as a member of the ''
fungi imperfecti The fungi imperfecti or imperfect fungi, are fungi which do not fit into the commonly established taxonomic classifications of fungi that are based on biological species concepts or morphological characteristics of sexual structures because thei ...
''. More recent investigations have shown that, contrary to Thaxter's classification, this organism is not closely related to ''Chrondromyces'', and ''Stigmatella'' is currently recognized as a separate genus. Of the three major subgroups of the myxobacteria, Myxococcus, Nannocystis, and Chrondromyces, ''Stigmatella'' is most closely aligned with Myxococcus.


Life cycle

''S. aurantiaca'', like other myxobacterial species, has a complex life cycle including social gliding (swarming), fruiting body formation, and predatory feeding behaviors. The bacteria do not swim, but glide on surfaces leaving slime trails, forming a mobile
biofilm A biofilm comprises any syntrophic consortium of microorganisms in which cells stick to each other and often also to a surface. These adherent cells become embedded within a slimy extracellular matrix that is composed of extracellular ...
. It commonly grows on the surface of rotting soft woods or fungi, where it may form bright orange patches. During the vegetative portion of their life cycles, swarming enables coordinated masses of myxobacteria to pool their secretions of extracellular digestive enzymes which are used to kill and consume prey microorganisms, a bacterial "wolfpack" effect. The best studied of the myxobacteria, ''
Myxococcus xanthus ''Myxococcus xanthus'' is a gram-negative, rod-shaped species of myxobacteria that exhibits various forms of self-organizing behavior in response to environmental cues. Under normal conditions with abundant food, it exists as a predatory, saproph ...
'', has been shown to actively surround prey organisms, trapping them in pockets where they can be consumed. Roaming flares of ''M. xanthus'' can detect clumps of prey bacteria at a distance, making turns towards the clumps and moving directly towards them. Like other myxobacterial species, ''S. aurantiaca'' survives periods of starvation by undergoing a developmental process whereby the individuals of a swarm aggregate to form ''fruiting bodies'' (not to be confused with those in fungi). Within the fruiting bodies, a certain fraction of the cells differentiate into myxospores, which are dormant cells resistant to drying and temperatures up to 90 °C. Differentiation into fruiting bodies appears to be mediated by contact-mediated signaling. Under laboratory growth conditions, the ability to undergo differentiation to form fruiting bodies is rapidly lost unless the cultures are regularly forced to fruit by transferring to starvation media. Shaker cultures of ''S. aurantiaca'' permanently lose the ability to fruit. The complex life cycle of myxobacteria is reminiscent of the life cycle of eukaryotic cellular
slime mold Slime mold or slime mould is an informal name given to several kinds of unrelated eukaryotic organisms with a life cycle that includes a free-living single-celled stage and the formation of spores. Spores are often produced in macroscopic mu ...
s.


Genome structure

Taxonomic identifier
378806
See also
NCBIUniProtKB
''S. aurantiaca'' DW4/3-1, a common laboratory strain, has been completely sequenced (See NCBI record link given above). Its circular DNA chromosome consists of 10.26 million base pairs and has a GC content of 67.5%. 8407 genes have been identified, coding for 8352 proteins.


Cell structure

The vegetative cells of ''S. aurantiaca'' are elongated rods typically measuring about 5–8 μm long and 0.7–0.8 μm wide. The fine structure resembles that of other gram negative bacteria. The cell surface consists of a
cytoplasmic membrane The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment ( ...
with a typical triple layered organization and a
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mech ...
. The cell wall consists of an outer triple layer and third dense monolayer in the periplasm. The myxospores are short, optically refractile rods measuring about 2.6–3.5 μm by 0.9–1.2 μm. The brightly colored, red or orange fruiting bodies comprise 1 to 20 spherical or ovoid cysts measuring 40–60 μm by 25–45 μm on top of a stalk measuring 60 to 140 μ high. Each red-brown cyst contains thousands of myxospores surrounded by thick, fibrous capsules. Dispersal of cysts is thought to benefit myxobacteria by ensuring that
cell growth Cell growth refers to an increase in the total mass of a cell, including both cytoplasmic, nuclear and organelle volume. Cell growth occurs when the overall rate of cellular biosynthesis (production of biomolecules or anabolism) is greater than ...
is resumed with a group (swarm) of myxobacteria, rather than as isolated cells. The stalks consist mostly of tubules which may represent the debris of lysed swarm cells, as well as some unlysed cells; very little fibrous material interpretable as slime is seen.


Ecology

''S. aurantiaca'' is found on rotting wood or fungi and is only rarely found in soil samples. Secreted and non-secreted proteins involved in their feeding behaviors, either identified directly or speculatively identified on the basis of
proteome The proteome is the entire set of proteins that is, or can be, expressed by a genome, cell, tissue, or organism at a certain time. It is the set of expressed proteins in a given type of cell or organism, at a given time, under defined conditions. ...
analysis, include enzymes capable of breaking down a wide selection of peptidoglycans, polysaccharides, proteins and other cellular detritus. Various other secreted compounds possibly involved in predation include antibiotics such as stigmatellin, which is toxic for yeast and filamentous fungi but not most bacteria, and aurafuron A and B, which inhibits the growth of various filamentous fungi. ''Stigmatella'' species hence appear in nature to help decompose otherwise insoluble biological debris. It is only distantly related to the cellulolytic myxobacteria, does not produce cellulases, and is strongly bacteriolytic. Therefore, ''Stigmatella'' consumes organisms that feed on wood rather that feeding on wood directly. Besides bacteria, its production of antifungal antibiotics suggests that ''Stigmatella'' species may feed on yeasts and fungi as well, or alternatively, may suggest that ''Stigmatella'' competes with fungi for shared resources. By producing antimicrobial compounds, ''Stigmatella'' may play a role in maintaining the balance of the microbial population in its habitat.


Current Research


Model system for development

Myxobacteria are distinguished from most bacteria by their remarkable range of social behaviors, and as a result, multiple laboratories have taken up studies of these bacteria as a prokaryotic paradigm for differentiation processes and
signal transduction Signal transduction is the process by which a chemical or physical signal is transmitted through a cell as a series of molecular events, most commonly protein phosphorylation catalyzed by protein kinases, which ultimately results in a cellula ...
. Most studies on social behavior in the myxobacteria have focused on ''M. xanthus'', which has provided an excellent system amenable to many classical genetic experimental approaches. The fruiting bodies of ''M. xanthus'' are relatively simple mounds, and the considerably more elaborate fruiting structures produced by ''S. aurantiaca'' has led to ''S. aurantiaca'' being considered an excellent complementary system to ''M. xanthus'', especially given the advent of contemporary means of genomic analysis. Most of the 95 known ''M. xanthus'' development-specific genes are highly conserved in ''S. aurantiaca''. Genes for entire signal transduction pathways important for fruiting body formation in ''M. xanthus'' are conserved in ''S. aurantiaca'', whereas only a few are conserved in ''Anaeromyxobacter dehalogenans'', a non-fruiting member of the order Myxococcales. Various genes have been identified in ''S. aurantiaca'' involved in the process of fruiting body formation, including ''fbfA'', which encodes a polypeptide homologous to chitin synthases, ''fbfB'', a gene encoding a putative galactose oxidase, various genes including those encoding tRNAAsp and tRNAVal located at the ''attB'' locus (a phage attachment site), and so forth. These genes play a variety of roles in the developmental cycle. For example, in experiments where the ''fbFA'' gene was deactivated, the bacterium formed structured clumps instead of fruiting bodies. To control formation of the elaborate and spatially complex multicellular structure which is the fruiting body, the cells must exchange signals during the entire process. In ''M. xanthus'', various signal molecules involved in this process have been identified. In ''S. aurantiaca'', Stevens (1982) identified an extracellular, diffusible signaling molecule (
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
) that could substitute for light in stimulating fruiting body maturation. A few years later, the structure of this molecule, a hydroxy ketone named stigmolone, was determined by NMR and mass spectroscopy. Besides signaling via exchange of diffusible substances, cell-cell signaling can be mediated by contact through the activity of surface located macromolecules. An example of this in ''S. aurantiaca'' would be the ''csgA'' homolog to the ''M. xanthus'' gene, which is bound to the cell envelope. The ''csgA'' gene product helps the cells to stay together during development and regulates motility of the cells.
Pxr sRNA Pxr sRNA is a regulatory RNA which downregulates genes responsible for the formation of fruiting bodies in ''Myxococcus xanthus''. Fruiting bodies are aggregations of myxobacteria formed when nutrients are scarce, the fruiting bodies permit a small ...
is a
regulatory RNA RNA interference (RNAi) is a biological process in which RNA molecules are involved in sequence-specific suppression of gene expression by double-stranded RNA, through translational or transcriptional repression. Historically, RNAi was known by o ...
which downregulates genes responsible for the formation of fruiting bodies in ''M. xanthus'', and a homolog has been noted in ''S. aurantiaca''. Another short nucleic acid that has been speculatively linked to
cell–cell recognition Cell–cell recognition is a cell's ability to distinguish one type of neighboring cell from another.Campbell, et al., Biology, ''Eighth Edition'', 2008 Pearson Education Inc. This phenomenon occurs when complementary molecules on opposing cell s ...
is
multicopy single-stranded DNA Multicopy single-stranded DNA (msDNA) is a type of extrachromosomal satellite DNA that consists of a single-stranded DNA molecule covalently linked via a 2'-5'phosphodiester bond to an internal guanosine of an RNA molecule. The resultant DNA/RNA ch ...
(msDNA). Sequence comparison of msDNAs from ''M. xanthus'', ''S. aurantiaca'', and other bacteria reveal conserved and hypervariable domains reminiscent of conserved and hypervariable sequences found in
allorecognition Allorecognition is the ability of an individual organism to distinguish its own tissues from those of another. It manifests itself in the recognition of antigens expressed on the surface of cells of non-self origin. Allorecognition has been descri ...
molecules. Another means for intercellular signaling could be via the exchange of outer membrane vesicles (OMVs). These vesicles are produced from the outer membrane of myxobacterial cells and are found in large quantities within bacterial biofilms. OMVs appear to play a variety of roles in myxobacterial swarming, predation, and development.


Natural secondary metabolites

Natural product A natural product is a natural compound or substance produced by a living organism—that is, found in nature. In the broadest sense, natural products include any substance produced by life. Natural products can also be prepared by chemical syn ...
s have been the source of most of the active ingredients in medicine, and continue to be an important source despite the advent of automated
high-throughput screening High-throughput screening (HTS) is a method for scientific experimentation especially used in drug discovery and relevant to the fields of biology, materials science and chemistry. Using robotics, data processing/control software, liquid handlin ...
methods for drug discovery in synthetic compounds. Historically,
actinomycetes The Actinomycetales is an order of Actinomycetota. A member of the order is often called an actinomycete. Actinomycetales are generally gram-positive and anaerobic and have mycelia in a filamentous and branching growth pattern. Some actinomycete ...
and fungi have been the major source of microbial secondary metabolites found useful as starting points for the development of new drugs, but the last several decades have seen myxobacteria come to the forefront of drug research. The pharmaceutical interest in these organisms comes from their production of a wide variety of structurally unique metabolites with interesting biological activities. The epothilones, derived from the myxobacterium ''Sporangium cellulosum'', represent a new, recently approved class of cancer drugs. Other myxobacterial compounds of potential pharmaceutical interest include disorazol, tubulysin, rhizopodin, chondramid, the aurafurons, tuscolid, tuscuron, and dawenol, chivosazol, soraphen, myxochelin, and the leupyrrins. ''S. aurantiaca'' has been the source of several of these bioactive compounds, including myxothiazol, an inhibitor of the
electron transport chain An electron transport chain (ETC) is a series of protein complexes and other molecules that transfer electrons from electron donors to electron acceptors via redox reactions (both reduction and oxidation occurring simultaneously) and couples th ...
, dawenol, a polyene metabolite, stigmatellin, an antifungal agent, the antifungals aurafuron A and B, and the iron
siderophores Siderophores (Greek: "iron carrier") are small, high-affinity iron-chelating compounds that are secreted by microorganisms such as bacteria and fungi. They help the organism accumulate iron. Although a widening range of siderophore functions is no ...
myxochelin A and B.


References


External links


The Myxobacteria Web PageVideo: Schwarmentwicklung und Morphogenese bei MyxobakterienVideo: Myxobacteria form Fruiting Bodies

Video: Myxococcus xanthus preying on an E. coli colonyType strain of ''Stigmatella aurantiaca'' at Bac''Dive'' - the Bacterial Diversity Metadatabase
{{Taxonbar, from=Q7616495 Myxococcota Bacteria described in 1875