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''Pasteuria ramosa'' is a gram-positive, endospore-forming bacterium in the
Bacillus ''Bacillus'', from Latin "bacillus", meaning "little staff, wand", is a genus of Gram-positive, rod-shaped bacteria, a member of the phylum ''Bacillota'', with 266 named species. The term is also used to describe the shape (rod) of other so-sh ...
/
Clostridia The Clostridia are a highly polyphyletic class of Bacillota, including '' Clostridium'' and other similar genera. They are distinguished from the Bacilli by lacking aerobic respiration. They are obligate anaerobes and oxygen is toxic to them ...
clade within
Bacillota The Bacillota (synonym Firmicutes) are a phylum of bacteria, most of which have Gram-positive cell wall structure. They have round cells, called cocci (singular coccus), or rod-like forms (bacillus). A few Bacillota, such as '' Megasphaera'', ...
. It is an obligate pathogen of
cladocera The Diplostraca or Cladocera, commonly known as water fleas, is a superorder (biology), superorder of small, mostly freshwater crustaceans, most of which feed on microscopic chunks of organic matter, though some forms are predatory. Over 1000 sp ...
n
crustacean Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquatic arthrop ...
s from the genus ''
Daphnia ''Daphnia'' is a genus of small planktonic crustaceans, in length. ''Daphnia'' are members of the Order (biology), order Anomopoda, and are one of the several small aquatic crustaceans commonly called water fleas because their Saltation (gait), ...
''. ''Daphnia'' is a genus of small planktonic crustaceans including '' D. magna'', ''P. ramosa'''s most popular host target''.'' Other hosts include '' D. pulex'', '' D. longispina'', '' D. dentifera'', and '' Moina rectirostris''. An established and widely used
coevolution In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well a ...
ary model of host-pathogen interactions exists with ''P. ramosa'' and '' D. magna''.


Growth and sporulation

''P. ramosa'' is an
obligate {{wiktionary, obligate As an adjective, obligate means "by necessity" (antonym '' facultative'') and is used mainly in biology in phrases such as: * Obligate aerobe, an organism that cannot survive without oxygen * Obligate anaerobe, an organism ...
pathogen and it can only grow inside its host. Transmission between hosts takes place through the
endospore An endospore is a dormant, tough, and non-reproductive structure produced by some bacteria in the phylum Bacillota. The name "endospore" is suggestive of a spore or seed-like form (''endo'' means 'within'), but it is not a true spore (i.e., not ...
stage, and is strictly horizontal. These endospores are highly resistant to different environmental stresses, including freezing temperatures, and can remain in the environment for decades without any deleterious effects. The infection can be explained in 5 steps: (1.) Encounter (2.) Activation (3.) Attachment (4.) Proliferation and (5.) Termination. The process starts when a ''Daphnia'' has ingested a spore of ''P. ramosa'' during filter feeding. The spore receives a signal to begin germination, and attaches to the host esophagus. The pathogen then enters the body cavity of the host by penetrating the esophagus wall. Once inside the body cavity, the bacterium begins to propagate in cauliflower like colonies. Propagation of spores is usually observed in the haemocoel and musculature. After the infection has spread throughout the host, the bacterium begins to sporulate. The spores are shed into the environment from the dead host and can remain in the sediment for decades while maintaining their infectivity. Additionally, these spores may be ingested by their immune hosts and pass through the gut of the ''Daphnia'' unharmed by any immune cells. This makes the spores very difficult to kill.


Pathogenicity

The infection success of ''P. ramosa'' depends on its ability to attach to the host esophagus and to spread into its body cavity where the propagation of the pathogen takes place. Propagation of the spores take place over a period of 10-20 days and ultimately leads to death of the infected host and the release of millions of created spores into the surrounding area. The attachment step of the infection depends on the genotypes of the host and the bacterium, meaning that only certain host genotypes can be infected by certain strains of the bacterium. Although the process through which the genotypic interactions occur is unclear, environmental factors, such as temperature, play a large role in the castration of ''Daphnia''. Studies have shown that female ''Daphnia'' are sterilized at warmer temperatures (), but still have the ability to reproduce at lower temperatures (). This difference in temperatures can be observed in different seasons and can lead to a high amount of variability between ''Daphnia'', a crucial part of its ability to coevolve with ''P. ramosa.'' During ''P. ramosa'' infection, the size of the ''Daphnia'' increases significantly. This phenomenon is known as pathogen-induced
gigantism Gigantism (, ''gígas'', "wiktionary:giant, giant", plural γίγαντες, ''gígantes''), also known as giantism, is a condition characterized by excessive growth and height significantly above average height, average. In humans, this conditi ...
. In addition, the lifespan of the host is significantly reduced.


Coevolutionary model with ''Daphnia magna''

''P. ramosa'' has coevolved with its host ''Daphnia magna''. The mode of
coevolution In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well a ...
in this system fits the model with
negative frequency-dependent selection Frequency-dependent selection is an evolutionary process by which the fitness of a phenotype or genotype depends on the phenotype or genotype composition of a given population. * In positive frequency-dependent selection, the fitness of a phenotyp ...
where the rare genotype is favored since the more common host genotype is more likely to become the target of a specialized pathogen.


Taxonomy

A culture established by James T. Staley, ATCC 27377T, was previously considered to be the
neotype In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally associated. In other words, a type is an example that serves to anchor or centralizes ...
for this species, but has been reassigned to '' Pirellula staleyi'' Schlesner and Hirsch, 1987 because it did not conform to Metchnikoff's original description of ''Pasteuria ramosa''.


References

{{Taxonbar, from=Q23817021 Bacillales Pathogenic bacteria