Pasteuria ramosa

File:Pasteuria ramosa endospores.jpg

P. ramosa is an obligate pathogen and it can only grow inside its host. Transmission between hosts takes place through the endospore 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.{{cite journal|last1=Duneau |first1=D. |first2=P. |last2=Luijckx |first3=F. |last3=Ben-Ami |first4=C. |last4=Laforsch |first5=D. |last5=Ebert |date=2011 |title=Resolving the infection process reveals striking differences in the contribution of environment, genetics and phylogeny to host-parasite interactions |journal=BMC Biology |volume=9 |page=11 |doi=10.1186/1741-7007-9-11 |pmid=21342515 |pmc=3052238 |doi-access=free }} 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.King, K. C., Stuart, K. J. R. A., Wilson, P. J., James, J., & Little, T. J. (2013). "The bacterial parasite Pasteuria ramosa is not killed if it fails to infect: Implications for coevolution." Ecology and Evolution, 3(2), 197–203. {{doi|10.1002/ece3.438}}.

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.Auld, S. K. J. R., Graham, A. L., Wilson, P. J., & Little, T. J. (2012). "Elevated haemocyte number is associated with infection and low fitness potential in wild Daphnia magna." Functional Ecology. 26(2):434–440. {{doi|10.1111/j.1365-2435.2011.01959.x}}. 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.{{cite journal|last1=Luijckx |first1=P. |first2=F. |last2=Ben-Ami |first3=L. |last3=Mouton |first4=L. |last4=Du Pasquier |first5=D. |last5=Ebert |date=2011 |title=Cloning of the unculturable parasite Pasteuria ramosa and its Daphnia host reveals extreme genotype-genotype interactions |journal=Ecology Letters |volume=14 |issue=2 |pages=125–131 |doi=10.1111/j.1461-0248.2010.01561.x|pmid=21091597 }} 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 ({{convert|20|-|25|C|F}}), but still have the ability to reproduce at lower temperatures ({{convert|10|-|15|C|F}}).{{cite journal|last1=Mitchell|first1=S.E.|last2=Rogers|first2=E.S.|last3=Little|first3=T.J.|last4=Read|first4=A.F.|year=2005|title=Host-parasite and genotype-by-environment interactions: Temperature modifies potential for selection by a sterilizing pathogen|journal=Evolution|volume=59|issue=1|pages=70–80|doi=10.1111/j.0014-3820.2005.tb00895.x|doi-access=free|pmid=15792228}} 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. In addition, the lifespan of the host is significantly reduced.

P. ramosa has coevolved with its host Daphnia magna. The mode of coevolution in this system fits the model with negative frequency-dependent selection where the rare genotype is favored since the more common host genotype is more likely to become the target of a specialized pathogen.{{Cite journal|last1=Decaestecker |first1=Ellen |last2=Vergote |first2=Adelien |last3=Ebert |first3=Dieter |last4=De Meester |first4=Luc |date=2003-04-01 |title=Evidence for strong host clone-parasite species interactions in the Daphnia microparasite system |journal=Evolution; International Journal of Organic Evolution |volume=57 |issue=4 |pages=784–792 |issn=0014-3820 |pmid=12778548 |doi=10.1111/j.0014-3820.2003.tb00290.x|s2cid=1626602 |doi-access=free }}

A culture established by James T. Staley, ATCC 27377^T, was previously considered to be the neotype for this species, but has been reassigned to Pirellula staleyi Schlesner and Hirsch, 1987{{cite journal|vauthors=Clum A, Tindall BJ, Sikorski J, Ivanova N, Mavrommatis K, Lucas S, Del Rio TJ, Nolan M, Chen F, Tice H, Pitluck S, Cheng JF, Chertkov O, Brettin T, Han C, Detter JC, Kuske C, Bruce D, Goodwin L, Ovchinikova G, Pati A, Mikhailova N, Chen A, Palaniappan K, Land M, Hauser L, Chang YJ, Jeffries CD, Chain P, Rohde M, Göker M, Bristow J, Eisen JA, Markowitz V, Hugenholtz P, Kyrpides NC, Klenk HP, Lapidus A|display-authors=6|title=Complete genome sequence of Pirellula staleyi type strain (ATCC 27377^T)|journal=Standards in Genomic Sciences|volume=1|year=2009|issue=3|doi=10.4056/sigs.51657|doi-access=free|pages=308–316|pmid=21304671|pmc=3035237}} because it did not conform to Metchnikoff's original description of Pasteuria ramosa.{{cite journal|author=Judicial Commission of the International Committee on Systematic Bacteriology|year=1986|title=Opinion 61: Rejection of the Type Strain of Pasteuria ramosa (ATCC 27377) and Conservation of the Species Pasteuria ramosa Metchnikoff 1888 on the Basis of the Type Descriptive Material|volume=36|issue=1|journal=International Journal of Systematic and Evolutionary Microbiology|doi=10.1099/00207713-36-1-119|doi-access=free|page=119}}

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Category:Bacillales

Category:Pathogenic bacteria