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====== Mosquito-associated spiroplasmas ======

===== Summary =====
  * To date five described //Spiroplasma// species were initially isolated from mosquitoes. 
    * //S. culicicola// from the salt marsh mosquito //Aedes sollicitans// collected in New Jersey, USA (Hung et al. 1987)
    * //S. sabaudiense// from a mixed pool of //Aedes sticticus// and //Aedes vexans// collected in the French northern Alps (Abalain-Colloc et al. 1987). Ecological survey (Legoff et al. 1990), found in May-Jun, disappeared in Aug/Sep.
    * //S. taiwanense// from //Culex tritaeniorhynchus// in Taiwan (Abalain-Colloc et al. 1988).
    * //S. cantharicola// (Whitcomb et al. 1993). closely related to S. diminutum (16S 99% id)
    * //S. diminutum// from //Culex annulus// and //Culex tritaeniorhynchus// in Taiwan (Williamson et al. 1996). 
  * These species are divergent in their serotypes and phylogenetic placements (Williamson et al. 1998; Gasparich et al. 2004), suggesting multiple independent origins of spiroplasma-mosquito association.
  * Artificial infection experiments:
    * S. taiwanense: infection of the yellow fever mosquito Aedes aegypti by S. taiwanense significantly reduces the survival of larvae (Humphery-Smith et al. 1991 MVE) and the lifespan of adult females (Humphery-Smith et al. 1991 JME; Vazeille-Falcoz et al. 1994). A histopathological study that used Anopheles stephensi as the host has shown that S. taiwanense can replicate both extra- and intra-cellularly in the host hemolymph, hemocytes, thoracic flight muscles, neural system, and other tissues (Phillips and Humphery-Smith 1995). Moreover, the infected mosquitoes exhibit loss of flight ability and reduced mobility, which are linked to extensive cell lysis and polysaccharide depletion in the thoracic flight muscles. Finally, cytadsorption of S. taiwanense was associated with the swelling and subsequent lysis of Aedes albopictus C6/36 cells in vitro (Chastel and Humphery-Smith 1991).
    * S. culicicola: infection of adult female Aedes aegypti increases mortality and reduces fecundity (Vazeille-Falcoz et al. 1994).
    * S. sabaudiense: no known effect (Chastel and Humphery-Smith 1991).
    * S. diminutum: can replicate inside Aedes albopictus but does not reduce the host lifespan (Vorms-Le Morvan et al. 1991).
    * Important note: in (Vazeille-Falcoz et al. 1994), the abstract describe that "the progeny of Ae. aegypti females inoculated with S. sabaudiense had a significantly reduced emergence rate and a significant distortion of the sex ratio." However, in the text (Table 2), those results are based on S. taiwanense infection.
  * The mosquito-pathogenic spiroplasmas may be developed into biocontrol agents (Humphery-Smith et al. 1993; Anbutsu and Fukatsu 2011).
  * Comparative genomic analysis between the pathogenic S. taiwanense and the non-pathogenic S. diminutum (Lo et al. 2013 GBE). A gene encoding for glycerol-3-phosphate oxidase (glpO) was found to be S. taiwanense-specific. This gene is involved in reactive oxygen species (ROS) and hydrogen peroxide (H2O2) production, has been experimentally verified as a primary virulence factor in Mycoplasma mycoides (Pilo et al. 2005; Pilo et al. 2007). May explain the tissue damage (Phillips and Humphery-Smith 1995) and higher mortality rates (Humphery-Smith et al. 1991 JME; Humphery-Smith et al. 1991 MVE; Vazeille-Falcoz et al. 1994) observed in S. taiwanense-infected mosquitoes.

===== References =====
  * Abalain-Colloc ML, Chastel C, Tully JG, Bove JM, Whitcomb RF, Gilot B, Williamson DL. 1987. Spiroplasma sabaudiense sp. nov. from mosquitoes collected in France. Int J Syst Microbiol 37:260 –265.
  * Abalain-Colloc ML, Rosen L, Tully JG, Bove JM, Chastel C, Williamson DL. 1988. Spiroplasma taiwanense sp. nov. from Culex tritaeniorhynchus mosquitoes collected in Taiwan. Int J Syst Microbiol 38:103 –107.
  * Chastel C, Gilot B, Legoff F, Gruffaz R, Abalaincolloc M. 1985. Isolation of Spiroplasma in France (savoie, North Alps) from Mosquitos. Comptes Rendus Acad. Sci. Ser. III-Sci. Vie-Life Sci. 300:261–&.
  * Chastel C, Humphery-Smith I. 1991. Mosquito Spiroplasmas, p. 149–206. In Harris, KF (ed.), Advances in Disease Vector Research. Springer New York.
  * Humphery-Smith I, Legoff F, Robaux P, Chastel C. 1993. Biosafety of an Experimentally Proven Mosquito Vector Pathogen, Spiroplasma-Taiwanense. Biocontrol Sci. Technol. 3:73–78.
  * Humphery-Smith I, Grulet O, Le Goff F, Chastel C. 1991. Spiroplasma (Mollicutes: Spiroplasmataceae) pathogenic for Aedes aegypti and Anopheles stephensi (Diptera: Culicidae). J. Med. Entomol. 28:219–222.
  * Humphery-Smith I, Grulet O, Chastel C. 1991. Pathogenicity of Spiroplasma taiwanense for larval Aedes aegypti mosquitoes. Med Vet Entomol 5:229–232.
  * Hung SHY, Chen TA, Whitcomb RF, Tully JG, Chen YX. 1987. Spiroplasma culicicola sp. nov. from the salt marsh mosquito Aedes sollicitans. Int J Syst Bacteriol 37:365 –370.
  * Legoff F, Marjolet M, Guilloteau J, Humphery-Smith I, Chastel C. 1990. Characterization and Ecology of Mosquito Spiroplasmas from Atlantic Biotopes in France. Annales De Parasitologie Humaine Et Comparee 65:107–110.
  * Lo W-S, Ku C, Chen L-L, Chang T-H, Kuo C-H. 2013. Comparison of Metabolic Capacities and Inference of Gene Content Evolution in Mosquito-Associated Spiroplasma diminutum and S. taiwanense. Genome Biol Evol 5:1512–1523.
  * Morvan J, Vazeillefalcoz M, Rodhain F. 1991. Experimental-Infection of Aedes-Albopictus Mosquitos by a Spiroplasma Strain Isolated from Culex-Annulus in Taiwan. Bull. Soc. Pathol. Exot. 84:15–24.
  * Phillips RN, Humphery-Smith I. 1995. The histopathology of experimentally induced infections of Spiroplasma taiwanense (class: Mollicutes) in Anopheles stephensi mosquitoes. J Invertebr Pathol 66:185–195.
  * Pilo P, Frey J, Vilei EM. 2007. Molecular mechanisms of pathogenicity of Mycoplasma mycoides subsp. mycoides SC. Vet J 174:513–521.
  * Pilo P, Vilei EM, Peterhans E, Bonvin-Klotz L, Stoffel MH, Dobbelaere D, Frey J. 2005. A metabolic enzyme as a primary virulence factor of Mycoplasma mycoides subsp. mycoides Small Colony. J. Bacteriol. 187:6824–6831.
  * Shaikh A, Johnson W, Stevens C, Tang A. 1987. The Isolation of Spiroplasmas from Mosquitos in Macon County, Alabama. J. Am. Mosq. Control Assoc. 3:289–295.
  * Vazeille-Falcoz M, Perchec-Merien A-M, Rodhain F. 1994. Experimental infection of Aedes aegypti mosquitoes, suckling mice, and rats with four mosquito spiroplasmas. J Invertebr Pathol 63:37–42.
  * Vorms-Le Morvan J, Vazeille-Falcoz M-C, Rodhain F, Chastel C. 1991. Infection expérimentale de moustiques Aedes albopictus par une souche de spiroplasmes isolée de Culex annulus a Taiwan. Bull Soc Pathol Exot 84:15–24.
  * Whitcomb RF et al. 1993. Spiroplasma cantharicola sp. nov., from Cantharid Beetles (Coleoptera: Cantharidae). Int. J. Syst. Bacteriol. 43:421 –424.
  * Williamson DL, Tully JG, Rosen L, Rose DL, Whitcomb RF, Abalain-Colloc ML, Carle P, Bové JM, Smyth J. 1996. Spiroplasma diminutum sp. nov., from Culex annulus mosquitoes collected in Taiwan. Int. J. Syst. Bacteriol. 46:229–233.