Pneumocystis: From a doubtful unique entity to a group of highly diversified fungal species

FEMS Yeast Research

Volumn 11, Issue 1, 2011, Pages 2-17

  Chabe M ^a   Aliouat Denis, Cécile Marie ^a   Delhaes, Laurence ^a   Aliouat, El Moukhtar ^a   Viscogliosi, Eric ^a   Dei Cas, Eduardo ^a  

^a UNIV LILLE   (France)

Author keywords

Epidemiology; Life cycle; Phylogenetic species concept; Pneumocystis spp.

Indexed keywords

ASCOMYCETES; DISEASE TRANSMISSION; FUNGAL LIFE CYCLE STAGE; GENOTYPE PHENOTYPE CORRELATION; NONHUMAN; PHYLOGENY; PNEUMOCYSTIS; PNEUMOCYSTIS CARINII; PNEUMOCYSTIS JIROVECI; PNEUMOCYSTIS MURINA; PNEUMOCYSTIS WAKEFIELDIAE; PNEUMOCYSTOSIS; REVIEW;

ANIMALS; HUMANS; LUNG; MAMMALS; PHYLOGENY; PNEUMOCYSTIS; PNEUMOCYSTIS INFECTIONS;

FUNGI; MAMMALIA; PNEUMOCYSTIS; PNEUMOCYSTIS CARINII; PNEUMOCYSTIS JIROVECII; PNEUMOCYSTIS MURINA; PNEUMOCYSTIS WAKEFIELDIAE;

EID: 78651499616     PISSN: 15671356     EISSN: 15671364     Source Type: Journal    
DOI: 10.1111/j.1567-1364.2010.00698.x     Document Type: Review

References (173)

1. Aliouat EM, Dei-Cas E, Ouaissi A, Palluault F, Soulez B & Camus D (1993a) In vitro attachment of Pneumocystis carinii from mouse and rat origin. Biol Cell 77: 209-217.
2. Aliouat EM, Mazars E, Dei-Cas E, Cesbron J & Camus D (1993b) Intranasal inoculation of mouse, rat or rabbit-derived Pneumocystis in SCID mice. J Protozool Res 3: 94-98.
3. Aliouat EM, Mazars E, Dei-Cas E, Delcourt P, Billaut P & Camus D (1994) Pneumocystis cross infection experiments using SCID mice and Nude rats as recipient host, showed strong host-species specificity. J Eukaryot Microbiol 41: S71.
4. Aliouat EM, Dujardin L, Martinez A, Duriez T, Ricard I & Dei-Cas E (1999) Pneumocystis carinii growth kinetics in culture systems and in hosts: involvement of each life cycle parasite stage. J Eukaryot Microbiol 46: S116-S117.
5. Aliouat-Denis CM, Chabé M, Demanche C, Aliouat EM, Viscogliosi E, Guillot J, Delhaes L & Dei-Cas E (2008) Pneumocystis species, co-evolution and pathogenic power. Infect Genet Evol 8: 708-726.
6. Aliouat-Denis CM, Martinez A, Aliouat EM, Pottier M, Gantois N & Dei-Cas E (2009) The Pneumocystis life cycle. Mem I Oswaldo Cruz 104: 419-426.
7. Anderson AC (2005) Targeting DHFR in parasitic protozoa. Drug Discov Today 10: 121-128.
8. Anonymous (1994) Revised nomenclature for Pneumocystis carinii. The Pneumocystis Workshop. J Eukaryot Microbiol 41: S121-S122.
9. Aragao HB (1913) Nota sobre as schizogonias e gametogonidas dos trypanosomas. Bras Med 11: 271.
10. Arcenas RC, Uhl JR, Buckwalter SP, Limper AH, Crino D, Roberts GD & Wengenack NL (2006) A real-time polymerase chain reaction assay for detection of Pneumocystis from bronchoalveolar lavage fluid. Diagn Microbiol Infect Dis 54: 169-175.
11. Atzori C, Agostoni F, Angeli E, Mainini A, Micheli V & Cargnel A (1999) P. carinii host specificity: attempt of cross infections with human derived strains in rats. J Eukaryot Microbiol 46: S112.
12. Baldauf SL, Roger AJ, Wenk-Siefert I & Doolittle WF (2000) A kingdom-level phylogeny of eukaryotes based on combined protein data. Science 290: 972-977.
13. Banerji S, Lugli EB, Miller RF & Wakefield AE (1995) Analysis of genetic diversity at the arom locus in isolates of Pneumocystis carinii. J Eukaryot Microbiol 42: 675-679.
14. Bartlett MS, Lee CH, Lu JJ, Bauer NL, Bettz JF, McLaughlin GL & Smith JW (1994) Pneumocystis carinii detected in air. J Eukaryot Microbiol 41: S75.
15. Bartlett MS, Vermund SH, Jacobs R et al. (1997) Detection of Pneumocystis carinii DNA in air samples: likely environmental risk to susceptible persons. J Clin Microbiol 35: 2511-2513.
16. Barton EG & Campbell WG (1967) Further observations on the ultrastructure of Pneumocystis. Arch Pathol 83: 527-534.
17. Beck JM, Preston AM, Wagner JG, Wilcoxen SE, Hossler P, Meshnick SR & Paine R III (1998) Interaction of rat Pneumocystis carinii and rat alveolar epithelial cells in vitro. Am J Physiol 275: L118-L125.
18. Burgess JW, Kottom TJ & Limper AH (2008) Pneumocystis carinii exhibits a conserved meiotic control pathway. Infect Immun 76: 417-425.
19. Burgess JW, Kottom TJ, Villegas LR, Lamont JD, Baden EM, Ramirez-Alvarado M & Limper AH (2009) The Pneumocystis meiotic PCRan1p kinase exhibits unique temperature-regulated activity. Am J Resp Cell Mol 41: 714-721.
20. Calderón EJ, Regordan C, Medrano FJ, Ollero M & Varela JM (1996) Pneumocystis carinii infection in patients with chronic bronchial disease. Lancet 347: 977.
21. Calderón EJ, Varela JM, Medrano FJ et al. (2004) Epidemiology of Pneumocystis carinii pneumonia in southern Spain. Clin Microbiol Infect 10: 673-676.
22. Calderón EJ, Rivero L, Respaldiza N, Morilla R, Montes-Cano MA, Friaza V, Muñoz-Lobato F, Varela JM, Medrano FJ & De La Horra C (2007) Systemic inflammation in patients with chronic obstructive pulmonary disease who are colonized with Pneumocystis jirovecii. Clin Infect Dis 45: 17-19.
23. Calderón-Sandubete EJ, Varela-Aguilar JM, Medrano-Ortega FJ, Nieto-Guerrer V, Respaldiza-Salas N, de la Horra-Padilla C & Dei-Cas E (2002) Historical perspective on Pneumocystis carinii infection. Protist 153: 303-310.
24. Carini A (1910) Formas de eschizogonia do Trypanosoma lewisi. Bol Soc de Med e Cir de Sao Paulo 18: 204.
25. Casanova-Cardiel L & Leibowitz MJ (1997) Presence of Pneumocystis carinii DNA in pond water. J Eukaryot Microbiol 44: S28.
26. Cavallini-Sanches EM, Pescador C, Rozza D, Spanamberg A, Borba MR, Ravazzolo AP, Driemeier D, Guillot J & Ferreiro L (2007) Detection of Pneumocystis spp. in lung samples from pigs in Brazil. Med Mycol 45: 395-399.
27. Ceré N, Drouet-Viard F, Dei-Cas E, Chanteloup N & Coudert P (1997) In utero transmission of Pneumocystis carinii sp. f. oryctolagi. Parasite 4: 325-330.
28. Chabé M, Dei-Cas E, Creusy C, Fleurisse L, Respaldiza N, Camus D & Durand-Joly I (2004) Immunocompetent hosts as a reservoir of Pneumocystis organisms: histological and RT-PCR data demonstrate active replication. Eur J Clin Microbiol 23: 89-97.
29. Chabé M, Nevez G, Totet A, Fréalle E, Delhaes L, Aliouat EM & Dei-Cas E (2009) Transmission de Pneumocystis. J Mycol Med 19: 276-284.
30. Chabé M, Herbreteau V, Hugot JP, Bouzard N, Deruyter L, Morand S & Dei-Cas E (2010) Pneumocystis carinii and Pneumocystis wakefieldiae in wild Rattus norvegicus trapped in Thailand. J Eukaryot Microbiol 57: 213-217.
31. Chagas C (1909) Nova tripanozomiazaea humana. Mem I Oswaldo Cruz 1: 159-218.
32. Chagas C (1911) Nova entidade morbida do homen: regumo general de estudos etiologicos e clinicos. Mem I Oswaldo Cruz 3: 219-275.
33. Chagas C (1913) Revisao do ciclo evolutivo do Trypanosoma cruzi. Braz Med 11: 225.
34. Chin K, Luttrell TD, Roe JD, Shadzi S, Wyder MA & Kaneshiro ES (1999) Putative Pneumocystis dormant forms outside the mammalian host, and long-term culture derived from them: initial characterizations. J Eukaryot Microbiol 46: S95-S99.
35. Cho SR, Park YG, Moon HN, Lee SH & Hong ST (1999) Karyotypes of Pneumocystis carinii derived from several mammals. Korean J Parasitol 37: 271-275.
36. Cornillot E, Keller B, Cushion MT, Méténier G & Vivarès CP (2002) Fine analysis of the Pneumocystis carinii f. sp. carinii genome by two-dimensional pulsed-field gel electrophoresis. Gene 293: 87-95.
37. Cushion M, Keely S & Stringer J (2004) Molecular and phenotypic description of Pneumocystis wakefieldiae sp.nov., a new species in rats. Mycologia 96: 429-438.
38. Cushion MT (1998) Genetic heterogeneity of rat-derived Pneumocystis. FEMS Immunol Med Mic 22: 51-58.
39. Cushion MT (2004) Pneumocystis: unraveling the cloak of obscurity. Trends Microbiol 12: 243-249.
40. Cushion MT & Beck JM (2001) Summary of Pneumocystis research presented at the 7th International Workshop on Opportunistic Protists. J Eukaryot Microbiol 48: S101-S105.
41. Cushion MT, Zhang J, Kaselis M, Giuntoli D, Stringer SL & Stringer JR (1993) Evidence for two genetic variants of Pneumocystis carinii coinfecting laboratory rats. J Clin Microbiol 31: 1217-1223.
42. Cushion MT, Smulian AG, Slaven BE, Sesterhenn T, Arnold J, Staben C, Porollo A, Adamczak R & Meller J (2007) Transcriptome of Pneumocystis carinii during fulminate infection: carbohydrate metabolism and the concept of a compatible parasite. PLoS ONE 2: e423.
43. Dei-Cas E (2000) Pneumocystis infections: the iceberg? Med Mycol 1: 23-32.
44. Dei-Cas E & Vernes A (1986) Parasitic adaptation of pathogenic fungi to the mammalian hosts. CRC Crit Rev Microbiol 13: 173-218.
45. Dei-Cas E, Jackson H, Palluault F, Aliouat EM, Hancock V, Soulez B & Camus D (1991) Ultrastructural observations on the attachment of Pneumocystis carinii in vitro. J Protozool 38: S205-S207.
46. Dei-Cas E, Mazars E, Ödberg-Ferragut C et al. (1994) Ultrastructural, genomic, isoenzymatic and biological features make it possible to distinguish rabbit Pneumocystis from other mammal Pneumocystis strains. J Eukaryot Microbiol 41: S84.
47. Dei-Cas E, Aliouat EM & Cailliez JC (2004) Cellular structure. Pneumocystis Pneumonia, 3rd edn (Walzer PD & Cushion MT, eds), pp. 61-94. Marcel Dekker, New York.
48. Dei-Cas E, Chabé M, Moukhlis R et al. (2006) Pneumocystis oryctolagi sp. nov., an uncultured fungus causing pneumonia in rabbits at weaning: review of current knowledge, and description of a new taxon on genotypic, phylogenetic and phenotypic bases. FEMS Microbiol Rev 80: 853-871.
49. Delanoë P & Delanoë E (1912) Sur les rapports des kystes de carinii du poumon des rats avec le Trypanosoma lewisi. CR Acad Sci 155: 658-660.
50. Delaporte F (1999) La Maladie de Chagas. Histoire d'un Fléau Continental. Payot & Rivages, Paris.
51. Demanche C, Berthelemy M, Petit T, Polack B, Wakefield AE, Dei-Cas E & Guillot J (2001) Phylogeny of Pneumocystis carinii from 18 primate species confirms host specificity and suggests coevolution. J Clin Microbiol 39: 2126-2133.
52. Demanche C, Petit T, Moisson P, Ollivet F, Rigoulet J, Chermette R, Dei-Cas E, Wakefield AE & Guillot J (2003) Assessment of Pneumocystis species carriage in captive primates. Vet Rec 28: 811-813.
53. De Stefano JA, Cushion MT, Sleught R & Walzer PD (1990) Analysis of Pneumocystis carinii cyst wall. I. Evidence for an outer surface membrane. J Protozool 37: 428-435.
54. Diezmann S, Cox CJ, Sconian G, Vilgalys RJ & Mitchell TG (2004) Phylogeny and evolution of medical species of Candida and related taxa: a multigenic analysis. J Clin Microbiol 42: 5624-5635.
55. Dumoulin A, Mazars E, Seguy N, Gargallo-Viola D, Vargas S, Cailliez JC, Aliouat EM, Wakefield AE & Dei-Cas E (2000) Transmission of Pneumocystis carinii disease from immunocompetent contacts of infected hosts to susceptible hosts. Eur J Clin Microbiol 19: 671-678.
56. Durand-Joly I, Aliouat EM, Recourt C, Guyot K, Francois N, Wauquier M, Camus D & Dei-Cas E (2002) Pneumocystis carinii f. sp. hominis is not infectious for SCID mice. J Clin Microbiol 40: 1862-1865.
57. Durand-Joly I, Soula F, Chabé M, Dalle JH, Lafitte JJ, Senechal M, Pinon A, Camus D & Dei-Cas E (2003) Long-term colonization with Pneumocystis jirovecii in hospital staffs: a challenge to prevent nosocomial pneumocystosis. J Eukaryot Microbiol 50: S614-S615.
58. Edman JC, Kovacs JA, Masur H, Santi DV, Elwood HJ & Sogin ML (1988) Ribosomal RNA sequence shows Pneumocystis carinii to be a member of the fungi. Nature 334: 519-522.
59. Edman JC, Kovacs JA, Masur H, Santi DV, Elwood HJ & Sogin ML (1989a) Ribosomal RNA sequence shows Pneumocystis carinii to be a member of the fungi. J Protozool 36: S18-S20.
60. Edman U, Edman JC, Lundgren B & Santi DV (1989b) Isolation and expression of the Pneumocystis carinii thymidylate synthase gene. P Natl Acad Sci USA 86: 6503-6507.
61. Eriksson OE (1994) Pneumocystis carinii, a parasite in lungs of mammals, referred to a new family and order (Pneumocystidaceae, Pneumocystidales, Ascomycota). Systema Ascomycetum 13: 165-180.
62. Eriksson OE (1999) Outline of Ascomycota - 1999. Myconet 3: 1-88.
63. Eriksson OE & Winka K (1997) Supraordinal taxa of the Ascomycota. Myconet 1: 1-16.
64. Frenkel JK (1976) Pneumocystis jiroveci n. sp. from man: morphology, physiology, and immunology in relation to pathology. Natl Cancer I Monogr 43: 13-30.
65. Frenkel JK (1999) Pneumocystis pneumonia, an immunodeficiency-dependent disease (IDD): a critical historical overview. J Eukaryot Microbiol 46: S89-S92.
66. Furuta T, Fujita M, Mukai R, Sakakibara I, Sata T, Miki K, Hayami M, Kojima S & Yoshikawa Y (1993) Severe pulmonary pneumocystosis in simian acquired immunodeficiency syndrome induced by simian immunodeficiency virus: its characterization by the polymerase-chain-reaction method and failure of experimental transmission to immunodeficient animals. Parasitol Res 79: 624-628.
67. Gigliotti F, Harmsen AG, Haidaris CG & Haidaris PJ (1993) Pneumocystis carinii is not universally transmissible between mammalian species. Infect Immun 61: 2886-2890.
68. Goheen M, Blumershine R, Hull M, Bartlett M & Smith J (1989) Enhancement of the fine structure of Pneumocystis carinii using potassium ferrocyanide and tannic acid. Proceedings of the 47th Annual Meeting of the Electron Microscopy Society of America (Bailey GW, ed), pp. 984-985. San Francisco.
69. Green PJ (1995) Reversible jump Markov chain Monte Carlo computation and Bayesian model determination. Biometrika 82: 711-732.
70. Guillot J, Demanche C, Hugot JP, Berthelemy M, Wakefield AE, Dei-Cas E & Chermette R (2001) Parallel phylogenies of Pneumocystis species and their mammalian hosts. J Eukaryot Microbiol 47: S113-S115.
71. Gustafson MP, Thomas CF Jr, Rusnak F, Limper AH & Leof EB (2001) Differential regulation of growth and checkpoint control mediated by a Cdc25 mitotic phosphatase from Pneumocystis carinii. J Biol Chem 276: 835-843.
72. Haque AU, Plattner SB, Cook RT & Hart MN (1987) Pneumocystis carinii: taxonomy as viewed by electron microscopy. Am J Clin Pathol 87: 504-510.
73. Harigaya Y & Yamamoto M (2007) Molecular mechanisms underlying the mitosis-meiosis decision. Chromosome Res 15: 523-537.
74. Hey J (2001) The mind of the species problem. Trends Ecol Evol 16: 326-329.
75. Hughes WT (1982) Natural mode of acquisition for de novo infection with Pneumocystis carinii. J Infect Dis 145: 842-848.
76. Hugot JP, Demanche C, Barriel V, Dei-Cas E & Guillot J (2003) Phylogenetic systematics and evolution of primate-derived Pneumocystis based on mitochondrial or nuclear DNA sequence comparison. Syst Biol 52: 735-744.
77. Hunter JA & Wakefield AE (1996) Genetic divergence at the mitochondrial small subunit ribosomal RNA gene among isolates of Pneumocystis carinii from five mammalian host species. J Eukaryot Microbiol 43: S24-S25.
78. Itatani CA (1994) Ultrastructural demonstration of a pore in the cyst wall of Pneumocystis carinii. J Parasitol 80: 644-648.
79. Itatani CA (1996) Ultrastructural morphology of intermediate forms and forms suggestive of conjugation in the life cycle of Pneumocystis carinii. J Parasitol 82: 163-171.
80. James TY, Kauff F, Schoch CL et al. (2006) Reconstructing the early evolution of Fungi using a six-gene phylogeny. Nature 443: 818-822.
81. Kaiser K, Rabodonirina M, Mayencon M & Picot S (1999) Cdc2 gene of Pneumocystis carinii hominis and its expression during culture. J Eukaryot Microbiol 46: S130.
82. Kaneshiro ES (1998) The lipids of Pneumocystis carinii. Clin Microbiol Rev 11: 27-41.
83. Kaneshiro ES & Maiorano JN (1996) Survival and infectivity of Pneumocystis carinii outside the mammalian host. J Eukaryot Microbiol 43: S35.
84. Keely S, Pai HJ, Baughman R, Sidman C, Sunkin C, Stringer JR & Stringer SL (1994) Pneumocystis species inferred from analysis of multiple genes. J Eukaryot Microbiol 41: S94.
85. Keely SP & Stringer JR (2005) Nomenclature and genetic variation of Pneumocystis. Pneumocystis Pneumonia, 3rd edn (Walzer PD & Cushion MT, eds), pp. 39-59. Marcel Dekker, New York.
86. Keely SP, Fischer JM & Stringer JR (2003) Evolution and speciation of Pneumocystis. J Eukaryot Microbiol 50: S624-S626.
87. Keely SP, Fischer JM, Cushion MT & Stringer JR (2004) Phylogenetic identification of Pneumocystis murina sp. nov., a new species in laboratory mice. Microbiology 150: 1153-1165.
88. Kottom TJ, Thomas CF Jr, Mubarak KK, Leof EB & Limper AH (2000) Pneumocystis carinii uses a functional cdc13 B-type cyclin complex during its life cycle. Am J Resp Cell Mol 22: 722-731.
89. Kottom TJ, Köhler JR, Thomas CF, Fink GR & Limper AH (2003) Lung epithelial cells and extracellular matrix components induce expression of Pneumocystis carinii STE20, a gene complementing the mating and pseudohyphal growth defects of STE20 mutant yeast. Infect Immun 71: 6463-6471.
90. Kovacs JA, Powell F, Edman JC, Lundgren B, Martinez A, Drew B & Angus CW (1993) Multiple genes encode the major surface glycoprotein of Pneumocystis carinii. J Biol Chem 268: 6034-6040.
91. Krajicek BJ, Kottom TJ, Villegas L & Limper AH (2010) Characterization of the PcCdc42 small G protein from Pneumocystis carinii, which interacts with the PcSte20 life cycle regulatory kinase. Am J Physiol Lung C 298: L252-L260.
92. Krungkrai J, Webster HK & Yuthavong Y (1990) Folate and cobalamin metabolism in Plasmodium falciparum. Parasitol Today 6: 388-391.
93. Kwon KY, Kim SP & Limper AH (1998) Recognition of Pneumocystis carinii antigen on its surface by immunohistochemistry and immunoelectron microscopy. J Korean Med Sci 13: 131-137.
94. Laakkonen J (1998) Pneumocystis carinii in wildlife. Int J Parasitol 28: 241-252.
95. Laakkonen J & Sukura A (1997) Pneumocystis carinii of the common shrew, Sorex araneus, shows a discrete phenotype. J Eukaryot Microbiol 44: 117-121.
96. Landvik S, Eriksson OE & Berbee ML (2001) Neolecta - a fungal dinosaur? Evidence from β-tubulin amino acid sequences. Mycologia 93: 1151-1163.
97. Larsen HH, von Linstow ML, Lundgren B, Hogh B, Westh H & Lundgren JD (2007) Primary Pneumocystis infection in infants hospitalized with acute respiratory tract infection. Emerg Infect Dis 13: 66-72.
98. Latouche S, Olsson M, Polack B, Brun-Pascaud M, Bernard C & Roux P (1997) Detection of Pneumocystis carinii f. sp. in air samples collected in animal rooms. J Eukaryot Microbiol 44: 46-47.
99. Latouche S, Totet A, Lacube P, Bolognini J, Nevez G & Roux P (2001) Development of an RT-PCR on the heat shock protein 70 gene for viability detection of Pneumocystis carinii f. sp. hominis in patients with pneumocystosis and in air sample. J Eukaryot Microbiol 48: S176-S177.
100. Li L, Wright SJ, Krystofova S, Park G & Borkovich KA (2007) Heterotrimeric G protein signaling in filamentous fungi. Annu Rev Microbiol 61: 423-452.
101. Liu Y, Rocourt M, Pan S, Liu C & Leibowitz MJ (1992) Sequence and variability of the 5.8S and 26S rRNA genes of Pneumocystis carinii. Nucleic Acids Res 20: 3763-3772.
102. Liu Y, Leigh JW, Brinkmann H, Cushion MT, Rodriguez-Ezpeleta N, Philippe H & Lang BF (2009) Phylogenomic analyses support the monophyly of Taphrinomycotina, including Schizosaccharomyces fission yeasts. Mol Biol Evol 26: 27-34.
103. Liu YJ, Hodson MC & Hall BD (2006) Loss of flagellum happened only once in the fungal lineage: phylogenetic struture of kingdom Fungi inferred from RNA polymerase II subunit genes. BMC Evol Biol 6: 74.
104. Lu JJ, Bartlett MS, Shaw MM, Queener SF, Smith JW, Ortiz-Rivera M, Leibowitz MJ & Lee CH (1994) Typing of Pneumocystis carinii strains that infect humans based on nucleotide sequence variations of internal transcribed spacers of rRNA genes. J Clin Microbiol 32: 2904-2912.
105. Maher NH, Vermund SH, Welsh DA, Dillon HK, Awooda A & Unnasch TR (2001) Development and characterization of a molecular viability assay for Pneumocystis carinii f sp hominis. J Infect Dis 183: 1825-1827.
106. Martinez A, Aliouat EM, Pottier M, Gantois M, Pincon C, Standaert-Vitse A, Dei-Cas E & Aliouat CM (2009) High-speed cell sorting of infectious trophic and cystic forms of Pneumocystis carinii. J Eukaryot Microbiol 56: 446-453.
107. Matsumoto Y & Yoshida Y (1984) Sporogony in Pneumocystis carinii: synaptonemal complexes and meiotic nuclear divisions observed in precysts. J Protozool 31: 420-428.
108. Mayden RL (1997) A hierarchy of species concepts: the denouement in the saga of the species problem. Species: The Units of Biodiversity (Claridge MF, Dawah HA & Wilson MR, eds), pp. 381-424. Chapman & Hall, London.
109. Mayr E (1963) Animal Species and Evolution. Harvard University Press, Cambridge, MA.
110. Mazars E, Ödberg-Ferragut C, Dei-Cas E, Fourmaux MN, Aliouat EM, Brun-Pascaud M, Mougeot G & Camus D (1995) Polymorphism of Pneumocystis carinii from different host species. J Eukaryot Microbiol 42: 26-32.
111. Mazars E, Guyot K, Durand I, Dei-Cas E, Boucher S, Ben Abderrazak S, Banuls AL, Tibayrenc M & Camus D (1997) Isoenzyme diversity in Pneumocystis carinii from rats, mice and rabbits. J Infect Dis 175: 655-660.
112. Miller RF, Ambrose HE & Wakefield AE (2001) Pneumocystis carinii f. sp. hominis DNA in immunocompetent health care workers in contact with patients with P. carinii pneumonia. J Clin Microbiol 39: 3877-3882.
113. Montes-Cano MA, Chabé M, Fontillon-Alberdi M, de-Lahorra C, Respaldiza N, Medrano FJ, Varela JM, Dei-Cas E & Calderon EJ (2009) Vertical transmission of Pneumocystis jirovecii in humans. Emerg Infect Dis 15: 125-127.
114. Nei M, Xu P & Glazko G (2001) Estimation of divergence times from multiprotein sequences for a few mammalian species and several distantly related organisms. P Natl Acad Sci USA 98: 2497-2502.
115. Nielsen MH, Settnes OP, Aliouat EM, Cailliez JC & Dei-Cas E (1998) Different ultrastructural morphology of Pneumocystis carinii derived from mice, rats, and rabbits. APMIS 106: 771-779.
116. Nishida H & Sugiyama J (1993) Phylogenetic relationships among Taphrina, Saitoella, and other higher fungi. Mol Biol Evol 10: 431-436.
117. Nishida H & Sugiyama J (1994) Archiascomycetes: detection of a major new lineage within the Ascomycota. Mycoscience 35: 361-366.
118. Olsson M, Sukura A, Lindberg LA & Linder E (1996) Detection of Pneumocystis carinii DNA by filtration of air. Scand J Infect Dis 28: 279-282.
119. Olsson M, Lidman C, Latouche S, Bjorkman A, Roux P, Linder E & Wahlgren M (1998) Identification of Pneumocystis carinii f. sp. hominis gene sequences in filtered air in hospital environments. J Clin Microbiol 36: 1737-1740.
120. Ortiz-Rivera M, Liu Y, Felder R & Leibowitz MJ (1995) Comparison of coding and spacer region sequences of chromosomal rRNA-coding genes of two sequevars of Pneumocystis carinii. J Eukaryot Microbiol 42: 44-49.
121. Palluault F, Dei-Cas E, Slomianny C, Soulez B & Camus D (1990) Golgi complex and lysosomes in rabbit-derived Pneumocystis carinii. Biol Cell 70: 73-82.
122. Palluault F, Pietrzyck B, Dei-Cas E, Slomianny C, Soulez B & Camus D (1991a) Three-dimensional reconstruction of rabbit-derived Pneumocystis carinii. I. Trophozoite. J Protozool 38: 402-407.
123. Palluault F, Pietrzyck B, Dei-Cas E, Slomianny C, Soulez B & Camus D (1991b) Three-dimensional reconstruction of rabbit-derived Pneumocystis carinii. II. Intermediate precyst. J Protozool 38: 407-411.
124. Palluault F, Slomianny C, Soulez B, Dei-Cas E & Camus D (1992a) High osmotic pressure enables fine ultrastructural and cytochemical studies on Pneumocystis carinii. I: epon embedding. Parasitol Res 78: 437-444.
125. Palluault F, Soulez B, Slomianny C, Dei-Cas E, Cesbron JY & Camus D (1992b) High osmotic pressure for Pneumocystis carinii London Resin White embedding enables fine immunocytochemistry studies: I. Golgi complex and cell-wall synthesis. Parasitol Res 78: 482-488.
126. Palmer RJ, Settnes OP, Lodal J & Wakefield AE (2000) Population structure of rat-derived Pneumocystis carinii in Danish wild rats. Appl Environ Microb 66: 4954-4961.
127. Peters SE, English K, Rana A, Akter S, Malik S, Warburton NC & Duckett JG (2001) Synaptonemal complexes in the pre-cyst of Pneumocystis carinii. J Eukaryot Microbiol 48: S134.
128. Peterson JC & Cushion MT (2005) Pneumocystis: not just pneumonia. Curr Opin Microbiol 8: 393-398.
129. Pixley FJ, Wakefield AE, Banerji S & Hopkin JM (1991) Mitochondrial gene sequences show fungal homology for Pneumocystis carinii. Mol Microbiol 5: 1347-1351.
130. Rabodonirina M (2001) La pneumocystose est-elle une maladie contagieuse? Lettre l'Infectiol 16: 110-112.
131. Rajagopalan-Levasseur P, Allaert A, Dridba M, Ödberg-Ferragut C, Jouault T, Creusy C, Camus D & Dei-Cas E (1998) Response to Pneumocystis infection in an immunocompetent host. FEMS Immunol Med Mic 22: 107-121.
132. Redhead SA, Cushion MT, Frenkel JK & Stringer JR (2006) Pneumocystis and Trypanosoma cruzi: nomenclature and typifications. J Eukaryot Microbiol 53: 2-11.
133. Ronquist F & Huelsenbeck JP (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19: 1572-1574.
134. Sanchez CA, Chabé M, Aliouat el M, Durand-Joly I, Gantois N, Conseil V, López C, Duriez T, Dei-Cas E & Vargas SL (2007) Exploring transplacental transmission of Pneumocystis oryctolagi in first-time pregnant and multiparous rabbit does. Med Mycol 45: 701-707.
135. Settnes OP & Nielsen MJ (1991) Host-parasite relationship in Pneumocystis carinii infection: activation of the plasmalemmal vesicular system in type I alveolar epithelial cells. J Protozool 38: S174-S176.
136. Sexton AC & Howlett BJ (2006) Parallels in fungal pathogenesis on plant and animal hosts. Eukaryot Cell 5: 1941-1949.
137. Sienkiewicz N, Jaroslawski S, Wyllie S & Fairlamb AH (2008) Chemical and genetic validation of dihydrofolate reductase-thymidylate synthase as a drug target in African trypanosomes. Mol Microbiol 69: 520-533.
138. Sinclair K, Wakefield AE, Banerji S & Hopkin JM (1991) Pneumocystis carinii organisms derived from rat and human hosts are genetically distinct. Mol Biochem Parasitol 45: 183-184.
139. Sing A, Wonhas C, Bader L, Luther M & Heesemann J (1999) Detection of Pneumocystis carinii DNA in the air filter of a ventilated patient with AIDS. Clin Infect Dis 29: 952-953.
140. Smulian AG, Sesterhenn T, Tanaka R & Cushion MT (2001) The ste3 pheromone receptor gene of Pneumocystis carinii is surrounded by a cluster of signal transduction genes. Genetics 157: 991-1002.
141. Soulez B, Palluault F, Cesbron JY, Dei-Cas E, Capron A & Camus D (1991) Introduction of Pneumocystis carinii in a colony of SCID mice. J Protozool 38: S123-S125.
142. Spatafora JW, Sung GH, Johnson D et al. (2006) A five-gene phylogeny of Pezizomycotina. Mycologia 98: 1018-1028.
143. Stagno S, Pifer LL, Hughes WT, Brasfield DM & Tiller RE (1980) Pneumocystis carinii pneumonitis in young immunocompetent infants. Pediatrics 66: 56-62.
144. Stedman TT, Butler DR & Buck GA (1998) The HSP70 gene family in Pneumocystis carinii: molecular and phylogenetic characterization of cytoplasmic members. J Eukaryot Microbiol 45: 589-599.
145. Stewart WC, Pollock KG, Browning LM, Young D, Smith-Palmer A & Reilly WJ (2005) Survey of zoonoses recorded in Scotland between 1993 and 2002. Vet Rec 157: 697-702.
146. Stringer JR & Cushion MT (1998) The genome of Pneumocystis carinii. FEMS Immunol Med Mic 22: 15-26.
147. Stringer JR, Cushion MT & Wakefield AE (2001) New nomenclature for the genus Pneumocystis. J Eukaryot Microbiol 48: 184-189.
148. Stringer SL, Stringer JR, Blase MA, Walzer PD & Cushion MT (1989) Pneumocystis carinii: sequence from ribosomal RNA implies a close relationship with fungi. Exp Parasitol 68: 450-461.
149. Sugiyama J, Hosaka K & Suh SO (2006) Early diverging Ascomycota: phylogenetic divergence and related evolutionary enigmas. Mycologia 298: 996-1005.
150. Tan WC & Ng TP (2008) COPD in Asia: where east meets west. Chest 133: 517-527.
151. Taylor JW, Jacobson DJ, Kroken S, Kasuga T, Geiser DM, Hibbett DS & Fisher MC (2000) Phylogenetic species recognition and species concepts in fungi. Fungal Genet Biol 31: 21-32.
152. Thomas CF, Anders RA, Gustafson MP, Leof EB & Limper AH (1998a) Pneumocystis carinii contains a functional cell-division-cycle Cdc2 homologue. Am J Resp Cell Mol 18: 297-306.
153. Thomas CF Jr, Kottom TJ, Leof EB & Limper AH (1998b) Characterization of a mitogen-activated protein kinase from Pneumocystis carinii. Am J Physiol 275: L193-L199.
154. Totet A, Latouche S, Lacube P, Pautard JC, Jounieaux V, Raccurt C, Roux P & Nevez G (2004) Pneumocystis jirovecii dihydropteroate synthase genotypes in immunocompetent infants and immunosuppressed adults, Amiens, France. Emerg Infect Dis 10: 667-673.
155. Vargas SL, Ponce CA, Gigliotti F, Ulloa AV, Prieto S, Muñoz MP & Hughes WT (2000) Transmission of Pneumocystis carinii DNA from a patient with P. carinii pneumonia to immunocompetent contact health care workers. J Clin Microbiol 38: 1536-1538.
156. Vavra J & Kucera K (1970) Pneumocystis carinii delanoë, its ultrastructure and ultrastructural affinities. J Protozool 17: 463-483.
157. Villegas LR, Kottom TJ & Limper AH (2010) Characterization of PCEng2 a {beta}-1, 3-endoglucanase homologue in Pneumocystis carinii with activity in cell wall regulation. Am J Resp Cell Mol 43: 192-200.
158. Vohra PK, Puri V, Kottom TJ, Limper AH & Thomas CF Jr (2003a) Pneumocystis carinii STE11, an HMG-box protein, is phosphorylated by the mitogen activated protein kinase PCM. Gene 312: 173-179.
159. Vohra PK, Puri V & Thomas CF Jr (2003b) Complementation and characterization of the Pneumocystis carinii MAPK, PCM. FEBS Lett 551: 139-146.
160. Vohra PK, Park JG, Sanyal B & Thomas CF Jr (2004) Expression analysis of PCSTE3, a putative pheromone receptor from the lung pathogenic fungus Pneumocystis carinii. Biochem Bioph Res Co 319: 193-199.
161. Wakefield AE (1996) DNA sequences identical to Pneumocystis carinii f. sp. carinii and Pneumocystis carinii f. sp. hominis in samples of air spora. J Clin Microbiol 34: 1754-1759.
162. Wakefield AE (1998) Genetic heterogeneity in Pneumocystis carinii: an introduction. FEMS Immunol Med Mic 22: 97-101.
163. Wakefield AE, Peters SE, Banerji S, Bridge PD, Hall GS, Hawksworth DL, Guiver LA, Allen AG & Hopkin JM (1992) Pneumocystis carinii shows DNA homology with the Ustomycetous red yeast fungi. Mol Microbiol 6: 1903-1911.
164. Walzer PD, Schnelle V, Armstrong D & Rosen PP (1977) Nude mouse: a new experimental model for Pneumocystis carinii infection. Science 197: 177-179.
165. Wyder MA, Rasch EM & Kaneshiro ES (1994) Assessment of Pneumocystis carinii DNA content. J Eukaryot Microbiol 41: S120.
166. Wyder MA, Rasch EM & Kaneshiro ES (1998) Quantitation of absolute Pneumocystis carinii nuclear DNA content. Trophic and cystic forms isolated from infected rat lungs are haploid organisms. J Eukaryot Microbiol 45: 233-239.
167. Yamada M, Matsumoto Y, Hamada S, Fujita S & Yoshida Y (1986) Demonstration and determination of DNA in Pneumocystis carinii by fluorescence microscopy with 4', 6-diamidino-2-phenylindole (DAPI). Zbl Bakt Mik Hyg A 262: 240-246.
168. Yoshida Y (1989) Ultrastructural studies of Pneumocystis carinii. J Protozool 36: 53-60.
169. Yoshida Y, Matsumoto Y, Yamada M, Okabayashi K, Yoshikawa H & Nakazawa M (1984) Pneumocystis carinii: electron microscopic investigation on the interaction of trophozoite and alveolar lining cell. Zbl Bakt Mik Hyg A 256: 390-399.
170. Yoshikawa H & Yoshida Y (1987) Localization of silver deposits on Pneumocystis carinii treated with Gomori's methenamine silver nitrate stain. Zbl Bakt Mik Hyg A 264: 363-372.
171. Yoshikawa H, Morioka H & Yoshida Y (1988) Ultrastructural detection of carbohydrates in the pellicle of Pneumocystis carinii. Parasitol Res 74: 537-543.
172. Youn H (2009) Review of zoonotic parasites in medical and veterinary fields in the Republic of Korea. Korean J Parasitol 47: S133-S141.
173. Ypma-Wong MF, Fonzi WA & Sypherd PS (1992) Fungus-specific translation elongation factor 3 gene present in Pneumocystis carinii. Infect Immun 60: 4140-4145.