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. 1996 Oct;9(4):489–498. doi: 10.1128/cmr.9.4.489

Pneumocystis carinii: what is it, exactly?

J R Stringer 1
PMCID: PMC172906  PMID: 8894348

Abstract

The identity of Pneumocystis carinii has been uncertain for many years. Until recently, it was widely regarded to be a protozoan because it does not grow in culture and is not susceptible to antifungal drugs. Although these and a number of other phenotypic characteristics of P. carinii differ from those of typical fungi, analysis of DNA sequences has shown that P. carinii is a member of the fungal lineage of eukaryotes. However, a close phylogenetic relative of P. carinii has not yet been found. Analysis of gene sequences has also revealed that P. carinii is not a single entity but that the genus Pneumocystis contains a complex group of organisms. P. carinii organisms from one host species do not grow when introduced into another host species, and P. carinii isolates from different host species are more genetically divergent from one another than might be expected for members of the same species. Genetic variation of a lesser degree also occurs among P. carinii organisms from the same host species, suggesting that it may be possible to identify strains and to conduct transmission and epidemiological studies. Results of early studies exploiting genetic variation among P. carinii isolates from patients have suggested that recurrent P. carinii pneumonia may not always be caused by reactivation of latent organisms, as is commonly believed. However, other features of P. carinii suggest that this microbe has established a long-term relationship with its host. A striking new development in this regard is the discovery of a genetic system that is designed to allow variation in the structure of a major antigen on the surface of P. carinii.

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Selected References

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  1. Aliouat E. M., Mazars E., Dei-Cas E., Delcourt P., Billaut P., Camus D. Pneumocystis cross infection experiments using SCID mice and nude rats as recipient host, showed strong host-species specificity. J Eukaryot Microbiol. 1994 Sep-Oct;41(5):71S–71S. [PubMed] [Google Scholar]
  2. Andrews R. P., Theus S. A., Cushion M. T., Walzer P. D. Comparison of the antigenic recognition of the major surface glycoprotein of Pneumocystis carinii isolated from different rat strains. J Eukaryot Microbiol. 1994 Sep-Oct;41(5):72S–72S. [PubMed] [Google Scholar]
  3. Baldauf S. L., Palmer J. D. Animals and fungi are each other's closest relatives: congruent evidence from multiple proteins. Proc Natl Acad Sci U S A. 1993 Dec 15;90(24):11558–11562. doi: 10.1073/pnas.90.24.11558. [DOI] [PMC free article] [PubMed] [Google Scholar]
  4. Banerji S., Lugli E. B., Miller R. F., Wakefield A. E. Analysis of genetic diversity at the arom locus in isolates of Pneumocystis carinii. J Eukaryot Microbiol. 1995 Nov-Dec;42(6):675–679. doi: 10.1111/j.1550-7408.1995.tb01614.x. [DOI] [PubMed] [Google Scholar]
  5. Banerji S., Lugli E. B., Wakefield A. E. Identification of two genetically distinct strains of Pneumocystis carinii in infected ferret lungs. J Eukaryot Microbiol. 1994 Sep-Oct;41(5):73S–73S. [PubMed] [Google Scholar]
  6. Banerji S., Wakefield A. E., Allen A. G., Maskell D. J., Peters S. E., Hopkin J. M. The cloning and characterization of the arom gene of Pneumocystis carinii. J Gen Microbiol. 1993 Dec;139(12):2901–2914. doi: 10.1099/00221287-139-12-2901. [DOI] [PubMed] [Google Scholar]
  7. Bartlett M. S., Lee C. H., Lu J. J., Bauer N. L., Bettz J. F., McLaughlin G. L., Smith J. W. Pneumocystis carinii detected in air. J Eukaryot Microbiol. 1994 Sep-Oct;41(5):75S–75S. [PubMed] [Google Scholar]
  8. Bauer N. L., Paulsrud J. R., Bartlett M. S., Smith J. W., Wilde C. E., 3rd Pneumocystis carinii organisms obtained from rats, ferrets, and mice are antigenically different. Infect Immun. 1993 Apr;61(4):1315–1319. doi: 10.1128/iai.61.4.1315-1319.1993. [DOI] [PMC free article] [PubMed] [Google Scholar]
  9. Bruns T. D., Vilgalys R., Barns S. M., Gonzalez D., Hibbett D. S., Lane D. J., Simon L., Stickel S., Szaro T. M., Weisburg W. G. Evolutionary relationships within the fungi: analyses of nuclear small subunit rRNA sequences. Mol Phylogenet Evol. 1992 Sep;1(3):231–241. doi: 10.1016/1055-7903(92)90020-h. [DOI] [PubMed] [Google Scholar]
  10. Chave J. P., David S., Wauters J. P., Van Melle G., Francioli P. Transmission of Pneumocystis carinii from AIDS patients to other immunosuppressed patients: a cluster of Pneumocystis carinii pneumonia in renal transplant recipients. AIDS. 1991 Aug;5(8):927–932. doi: 10.1097/00002030-199108000-00002. [DOI] [PubMed] [Google Scholar]
  11. Chen W., Gigliotti F., Harmsen A. G. Latency is not an inevitable outcome of infection with Pneumocystis carinii. Infect Immun. 1993 Dec;61(12):5406–5409. doi: 10.1128/iai.61.12.5406-5409.1993. [DOI] [PMC free article] [PubMed] [Google Scholar]
  12. Cushion M. T., DeStefano J. A., Walzer P. D. Pneumocystis carinii: surface reactive carbohydrates detected by lectin probes. Exp Parasitol. 1988 Dec;67(2):137–147. doi: 10.1016/0014-4894(88)90061-6. [DOI] [PubMed] [Google Scholar]
  13. Cushion M. T., Kaselis M., Stringer S. L., Stringer J. R. Genetic stability and diversity of Pneumocystis carinii infecting rat colonies. Infect Immun. 1993 Nov;61(11):4801–4813. doi: 10.1128/iai.61.11.4801-4813.1993. [DOI] [PMC free article] [PubMed] [Google Scholar]
  14. Cushion M. T., Zhang J., Kaselis M., Giuntoli D., Stringer S. L., Stringer J. R. Evidence for two genetic variants of Pneumocystis carinii coinfecting laboratory rats. J Clin Microbiol. 1993 May;31(5):1217–1223. doi: 10.1128/jcm.31.5.1217-1223.1993. [DOI] [PMC free article] [PubMed] [Google Scholar]
  15. Dei-Cas E., Mazars E., Ferragut C. O., Durand I., Aliouat E. M., Dridba M., Palluault F., Cailliez J. C., Seguy N., Tibayrenc M. Ultrastructural, genomic, isoenzymatic and biological features make it possible to distinguish rabbit Pneumocystis from other mammal Pneumocystis strains. J Eukaryot Microbiol. 1994 Sep-Oct;41(5):84S–84S. [PubMed] [Google Scholar]
  16. Dyer M., Volpe F., Delves C. J., Somia N., Burns S., Scaife J. G. Cloning and sequence of a beta-tubulin cDNA from Pneumocystis carinii: possible implications for drug therapy. Mol Microbiol. 1992 Apr;6(8):991–1001. doi: 10.1111/j.1365-2958.1992.tb02165.x. [DOI] [PubMed] [Google Scholar]
  17. Edlind T. D., Bartlett M. S., Weinberg G. A., Prah G. N., Smith J. W. The beta-tubulin gene from rat and human isolates of Pneumocystis carinii. Mol Microbiol. 1992 Nov;6(22):3365–3373. doi: 10.1111/j.1365-2958.1992.tb02204.x. [DOI] [PubMed] [Google Scholar]
  18. Edman J. C., Edman U., Cao M., Lundgren B., Kovacs J. A., Santi D. V. Isolation and expression of the Pneumocystis carinii dihydrofolate reductase gene. Proc Natl Acad Sci U S A. 1989 Nov;86(22):8625–8629. doi: 10.1073/pnas.86.22.8625. [DOI] [PMC free article] [PubMed] [Google Scholar]
  19. Edman J. C., Kovacs J. A., Masur H., Santi D. V., Elwood H. J., Sogin M. L. Ribosomal RNA genes of Pneumocystis carinii. J Protozool. 1989 Jan-Feb;36(1):18S–20S. doi: 10.1111/j.1550-7408.1989.tb02672.x. [DOI] [PubMed] [Google Scholar]
  20. Edman J. C., Kovacs J. A., Masur H., Santi D. V., Elwood H. J., Sogin M. L. Ribosomal RNA sequence shows Pneumocystis carinii to be a member of the fungi. Nature. 1988 Aug 11;334(6182):519–522. doi: 10.1038/334519a0. [DOI] [PubMed] [Google Scholar]
  21. Edman U., Edman J. C., Lundgren B., Santi D. V. Isolation and expression of the Pneumocystis carinii thymidylate synthase gene. Proc Natl Acad Sci U S A. 1989 Sep;86(17):6503–6507. doi: 10.1073/pnas.86.17.6503. [DOI] [PMC free article] [PubMed] [Google Scholar]
  22. Fletcher L. D., McDowell J. M., Tidwell R. R., Meagher R. B., Dykstra C. C. Structure, expression and phylogenetic analysis of the gene encoding actin I in Pneumocystis carinii. Genetics. 1994 Jul;137(3):743–750. doi: 10.1093/genetics/137.3.743. [DOI] [PMC free article] [PubMed] [Google Scholar]
  23. Frenkel J. K. Pneumocystis jiroveci n. sp. from man: morphology, physiology, and immunology in relation to pathology. Natl Cancer Inst Monogr. 1976 Oct;43:13–30. [PubMed] [Google Scholar]
  24. Furuta T., Fujita M., Mukai R., Sakakibara I., Sata T., Miki K., Hayami M., Kojima S., Yoshikawa Y. 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. 1993;79(8):624–628. doi: 10.1007/BF00932502. [DOI] [PubMed] [Google Scholar]
  25. Garbe T. R., Stringer J. R. Molecular characterization of clustered variants of genes encoding major surface antigens of human Pneumocystis carinii. Infect Immun. 1994 Aug;62(8):3092–3101. doi: 10.1128/iai.62.8.3092-3101.1994. [DOI] [PMC free article] [PubMed] [Google Scholar]
  26. Garner R. E., Walker A. N., Horst M. N. Morphologic and biochemical studies of chitin expression in Pneumocystis carinii. J Protozool. 1991 Nov-Dec;38(6):12S–14S. [PubMed] [Google Scholar]
  27. Gigliotti F., Ballou L. R., Hughes W. T., Mosley B. D. Purification and initial characterization of a ferret Pneumocystis carinii surface antigen. J Infect Dis. 1988 Oct;158(4):848–854. doi: 10.1093/infdis/158.4.848. [DOI] [PubMed] [Google Scholar]
  28. Gigliotti F., Harmsen A. G., Haidaris C. G., Haidaris P. J. Pneumocystis carinii is not universally transmissible between mammalian species. Infect Immun. 1993 Jul;61(7):2886–2890. doi: 10.1128/iai.61.7.2886-2890.1993. [DOI] [PMC free article] [PubMed] [Google Scholar]
  29. Gigliotti F. Host species-specific antigenic variation of a mannosylated surface glycoprotein of Pneumocystis carinii. J Infect Dis. 1992 Feb;165(2):329–336. doi: 10.1093/infdis/165.2.329. [DOI] [PubMed] [Google Scholar]
  30. Gigliotti F., Stokes D. C., Cheatham A. B., Davis D. S., Hughes W. T. Development of murine monoclonal antibodies to Pneumocystis carinii. J Infect Dis. 1986 Aug;154(2):315–322. doi: 10.1093/infdis/154.2.315. [DOI] [PubMed] [Google Scholar]
  31. Giuntoli D., Stringer S. L., Stringer J. R. Extraordinarily low number of ribosomal RNA genes in P. carinii. J Eukaryot Microbiol. 1994 Sep-Oct;41(5):88S–88S. [PubMed] [Google Scholar]
  32. Graves D. C., McNabb S. J., Worley M. A., Downs T. D., Ivey M. H. Analyses of rat Pneumocystis carinii antigens recognized by human and rat antibodies by using western immunoblotting. Infect Immun. 1986 Oct;54(1):96–103. doi: 10.1128/iai.54.1.96-103.1986. [DOI] [PMC free article] [PubMed] [Google Scholar]
  33. Haidaris C. G., Wright T. W., Gigliotti F., Haidaris P. J. Molecular cloning and characterization of ferret Pneumocystis carinii gp120. J Protozool. 1991 Nov-Dec;38(6):5S–6S. [PubMed] [Google Scholar]
  34. Haidaris P. J., Wright T. W., Gigliotti F., Haidaris C. G. Expression and characterization of a cDNA clone encoding an immunodominant surface glycoprotein of Pneumocystis carinii. J Infect Dis. 1992 Nov;166(5):1113–1123. doi: 10.1093/infdis/166.5.1113. [DOI] [PubMed] [Google Scholar]
  35. Hong S. T., Steele P. E., Cushion M. T., Walzer P. D., Stringer S. L., Stringer J. R. Pneumocystis carinii karyotypes. J Clin Microbiol. 1990 Aug;28(8):1785–1795. doi: 10.1128/jcm.28.8.1785-1795.1990. [DOI] [PMC free article] [PubMed] [Google Scholar]
  36. Hughes W. T., Gigliotti F. Nomenclature for Pneumocystis carinii. J Infect Dis. 1988 Mar;157(3):432–433. doi: 10.1093/infdis/157.3.432. [DOI] [PubMed] [Google Scholar]
  37. Kaneshiro E. S., Ellis J. E., Jayasimhulu K., Beach D. H. Evidence for the presence of "metabolic sterols" in Pneumocystis: identification and initial characterization of Pneumocystis carinii sterols. J Eukaryot Microbiol. 1994 Jan-Feb;41(1):78–85. doi: 10.1111/j.1550-7408.1994.tb05938.x. [DOI] [PubMed] [Google Scholar]
  38. Keely S. P., Stringer J. R., Baughman R. P., Linke M. J., Walzer P. D., Smulian A. G. Genetic variation among Pneumocystis carinii hominis isolates in recurrent pneumocystosis. J Infect Dis. 1995 Aug;172(2):595–598. doi: 10.1093/infdis/172.2.595. [DOI] [PubMed] [Google Scholar]
  39. Keely S., Pai H. J., Baughman R., Sidman C., Sunkin S. M., Stringer J. R., Stringer S. L. Pneumocystis species inferred from analysis of multiple genes. J Eukaryot Microbiol. 1994 Sep-Oct;41(5):94S–94S. [PubMed] [Google Scholar]
  40. Kovacs J. A., Halpern J. L., Lundgren B., Swan J. C., Parrillo J. E., Masur H. Monoclonal antibodies to Pneumocystis carinii: identification of specific antigens and characterization of antigenic differences between rat and human isolates. J Infect Dis. 1989 Jan;159(1):60–70. doi: 10.1093/infdis/159.1.60. [DOI] [PubMed] [Google Scholar]
  41. Kovacs J. A., Halpern J. L., Swan J. C., Moss J., Parrillo J. E., Masur H. Identification of antigens and antibodies specific for Pneumocystis carinii. J Immunol. 1988 Mar 15;140(6):2023–2031. [PubMed] [Google Scholar]
  42. Kovacs J. A., Powell F., Edman J. C., Lundgren B., Martinez A., Drew B., Angus C. W. Multiple genes encode the major surface glycoprotein of Pneumocystis carinii. J Biol Chem. 1993 Mar 15;268(8):6034–6040. [PubMed] [Google Scholar]
  43. Latouche S., Roux P., Poirot J. L., Lavrard I., Hermelin B., Bertrand V. Preliminary results of Pneumocystis carinii strain differentiation by using molecular biology. J Clin Microbiol. 1994 Dec;32(12):3052–3053. doi: 10.1128/jcm.32.12.3052-3053.1994. [DOI] [PMC free article] [PubMed] [Google Scholar]
  44. Lee C. H., Lu J. J., Bartlett M. S., Durkin M. M., Liu T. H., Wang J., Jiang B., Smith J. W. Nucleotide sequence variation in Pneumocystis carinii strains that infect humans. J Clin Microbiol. 1993 Mar;31(3):754–757. doi: 10.1128/jcm.31.3.754-757.1993. [DOI] [PMC free article] [PubMed] [Google Scholar]
  45. Li J., Edlind T. Phylogeny of Pneumocystis carinii based on beta-tubulin sequence. J Eukaryot Microbiol. 1994 Sep-Oct;41(5):97S–97S. [PubMed] [Google Scholar]
  46. Lin H., Niu M. T., Yoganathan T., Buck G. A. Characterization of the rRNA-encoding genes and transcripts, and a group-I self-splicing intron in Pneumocystis carinii. Gene. 1992 Oct 1;119(2):163–173. doi: 10.1016/0378-1119(92)90268-t. [DOI] [PubMed] [Google Scholar]
  47. Linke M. J., Cushion M. T., Walzer P. D. Properties of the major antigens of rat and human Pneumocystis carinii. Infect Immun. 1989 May;57(5):1547–1555. doi: 10.1128/iai.57.5.1547-1555.1989. [DOI] [PMC free article] [PubMed] [Google Scholar]
  48. Linke M. J., Smulian A. G., Yoshihara P., Walzer P. D. Production and characterization of monoclonal antibodies specific for the major surface glycoprotein of Pneumocystis carinii. J Eukaryot Microbiol. 1994 Sep-Oct;41(5):99S–100S. [PubMed] [Google Scholar]
  49. Liu Y., Leibowitz M. J. Variation and in vitro splicing of group I introns in rRNA genes of Pneumocystis carinii. Nucleic Acids Res. 1993 May 25;21(10):2415–2421. doi: 10.1093/nar/21.10.2415. [DOI] [PMC free article] [PubMed] [Google Scholar]
  50. Liu Y., Rocourt M., Pan S., Liu C., Leibowitz M. J. Sequence and variability of the 5.8S and 26S rRNA genes of Pneumocystis carinii. Nucleic Acids Res. 1992 Jul 25;20(14):3763–3772. doi: 10.1093/nar/20.14.3763. [DOI] [PMC free article] [PubMed] [Google Scholar]
  51. Lu J. J., Bartlett M. S., Shaw M. M., Queener S. F., Smith J. W., Ortiz-Rivera M., Leibowitz M. J., Lee C. H. Typing of Pneumocystis carinii strains that infect humans based on nucleotide sequence variations of internal transcribed spacers of rRNA genes. J Clin Microbiol. 1994 Dec;32(12):2904–2912. doi: 10.1128/jcm.32.12.2904-2912.1994. [DOI] [PMC free article] [PubMed] [Google Scholar]
  52. Lu J. J., Bartlett M. S., Shaw M. M., Smith J. W., Lee C. H. Typing of Pneumocystis carinii strains that infect humans based on nucleotide sequence variations of internal transcribed spacers of rRNA genes. J Eukaryot Microbiol. 1994 Sep-Oct;41(5):102S–102S. [PubMed] [Google Scholar]
  53. Lu J. J., Bartlett M. S., Smith J. W., Lee C. H. Typing of Pneumocystis carinii strains with type-specific oligonucleotide probes derived from nucleotide sequences of internal transcribed spacers of rRNA genes. J Clin Microbiol. 1995 Nov;33(11):2973–2977. doi: 10.1128/jcm.33.11.2973-2977.1995. [DOI] [PMC free article] [PubMed] [Google Scholar]
  54. Lundgren B., Cotton R., Lundgren J. D., Edman J. C., Kovacs J. A. Identification of Pneumocystis carinii chromosomes and mapping of five genes. Infect Immun. 1990 Jun;58(6):1705–1710. doi: 10.1128/iai.58.6.1705-1710.1990. [DOI] [PMC free article] [PubMed] [Google Scholar]
  55. Lundgren B., Koch C., Mathiesen L., Nielsen J. O., Hansen J. E. Glycosylation of the major human Pneumocystis carinii surface antigen. APMIS. 1993 Mar;101(3):194–200. [PubMed] [Google Scholar]
  56. Matsumoto Y., Yamada M., Amagai T. Yeast glucan of Pneumocystis carinii cyst wall: an excellent target for chemotherapy. J Protozool. 1991 Nov-Dec;38(6):6S–7S. [PubMed] [Google Scholar]
  57. Matsumoto Y., Yoshida Y. Sporogony in Pneumocystis carinii: synaptonemal complexes and meiotic nuclear divisions observed in precysts. J Protozool. 1984 Aug;31(3):420–428. doi: 10.1111/j.1550-7408.1984.tb02989.x. [DOI] [PubMed] [Google Scholar]
  58. Matusiewicz S. P., Fergusson R. J., Greening A. P., Crompton G. K., Burns S. M. Pneumocystis carinii in bronchoalveolar lavage fluid and bronchial washings. BMJ. 1994 May 7;308(6938):1206–1207. doi: 10.1136/bmj.308.6938.1206. [DOI] [PMC free article] [PubMed] [Google Scholar]
  59. Mazars E., Odberg-Ferragut C., Dei-Cas E., Fourmaux M. N., Aliouat E. M., Brun-Pascaud M., Mougeot G., Camus D. Polymorphism of the thymidylate synthase gene of Pneumocystis carinii from different host species. J Eukaryot Microbiol. 1995 Jan-Feb;42(1):26–32. doi: 10.1111/j.1550-7408.1995.tb01536.x. [DOI] [PubMed] [Google Scholar]
  60. Mazars E., Odberg-Ferragut C., Durand I., Tibayrenc M., Dei-Cas E., Camus D. Genomic and isoenzymatic markers of Pneumocystis from different host species. J Eukaryot Microbiol. 1994 Sep-Oct;41(5):104S–104S. [PubMed] [Google Scholar]
  61. Meade J. C., Stringer J. R. Cloning and characterization of an ATPase gene from Pneumocystis carinii which closely resembles fungal H+ ATPases. J Eukaryot Microbiol. 1995 May-Jun;42(3):298–307. doi: 10.1111/j.1550-7408.1995.tb01584.x. [DOI] [PubMed] [Google Scholar]
  62. Meade J. C., Stringer J. R. PCR amplification of DNA sequences from the transcription factor IID and cation transporting ATPase genes in Pneumocystis carinii. J Protozool. 1991 Nov-Dec;38(6):66S–68S. [PubMed] [Google Scholar]
  63. Millard P. R., Heryet A. R. Observations favouring Pneumocystis carinii pneumonia as a primary infection: a monoclonal antibody study on paraffin sections. J Pathol. 1988 Apr;154(4):365–370. doi: 10.1002/path.1711540413. [DOI] [PubMed] [Google Scholar]
  64. Ortiz-Rivera M., Liu Y., Felder R., Leibowitz M. J. Comparison of coding and spacer region sequences of chromosomal rRNA-coding genes of two sequevars of Pneumocystis carinii. J Eukaryot Microbiol. 1995 Jan-Feb;42(1):44–49. doi: 10.1111/j.1550-7408.1995.tb01538.x. [DOI] [PubMed] [Google Scholar]
  65. Peglow S. L., Smulian A. G., Linke M. J., Pogue C. L., Nurre S., Crisler J., Phair J., Gold J. W., Armstrong D., Walzer P. D. Serologic responses to Pneumocystis carinii antigens in health and disease. J Infect Dis. 1990 Feb;161(2):296–306. doi: 10.1093/infdis/161.2.296. [DOI] [PubMed] [Google Scholar]
  66. Pesanti E. L., Shanley J. D. Glycoproteins of Pneumocystis carinii: characterization by electrophoresis and microscopy. J Infect Dis. 1988 Dec;158(6):1353–1359. doi: 10.1093/infdis/158.6.1353. [DOI] [PubMed] [Google Scholar]
  67. Peters S. E., English K., Laakkonen J., Gurnell J. DNA analysis of Pneumocystis carinii infecting Finnish and English shrews. J Eukaryot Microbiol. 1994 Sep-Oct;41(5):108S–108S. [PubMed] [Google Scholar]
  68. Peters S. E., Wakefield A. E., Sinclair K., Millard P. R., Hopkin J. M. A search for Pneumocystis carinii in post-mortem lungs by DNA amplification. J Pathol. 1992 Feb;166(2):195–198. doi: 10.1002/path.1711660217. [DOI] [PubMed] [Google Scholar]
  69. Peters S. E., Wakefield A. E., Whitwell K. E., Hopkin J. M. Pneumocystis carinii pneumonia in thoroughbred foals: identification of a genetically distinct organism by DNA amplification. J Clin Microbiol. 1994 Jan;32(1):213–216. doi: 10.1128/jcm.32.1.213-216.1994. [DOI] [PMC free article] [PubMed] [Google Scholar]
  70. Pixley F. J., Wakefield A. E., Banerji S., Hopkin J. M. Mitochondrial gene sequences show fungal homology for Pneumocystis carinii. Mol Microbiol. 1991 Jun;5(6):1347–1351. doi: 10.1111/j.1365-2958.1991.tb00781.x. [DOI] [PubMed] [Google Scholar]
  71. Powles M. A., McFadden D. C., Liberator P. A., Anderson J. W., Vadas E. B., Meisner D., Schmatz D. M. Aerosolized L-693,989 for Pneumocystis carinii prophylaxis in rats. Antimicrob Agents Chemother. 1994 Jun;38(6):1397–1401. doi: 10.1128/aac.38.6.1397. [DOI] [PMC free article] [PubMed] [Google Scholar]
  72. Päbo S., Irwin D. M., Wilson A. C. DNA damage promotes jumping between templates during enzymatic amplification. J Biol Chem. 1990 Mar 15;265(8):4718–4721. [PubMed] [Google Scholar]
  73. Schmatz D. M., Romancheck M. A., Pittarelli L. A., Schwartz R. E., Fromtling R. A., Nollstadt K. H., Vanmiddlesworth F. L., Wilson K. E., Turner M. J. Treatment of Pneumocystis carinii pneumonia with 1,3-beta-glucan synthesis inhibitors. Proc Natl Acad Sci U S A. 1990 Aug;87(15):5950–5954. doi: 10.1073/pnas.87.15.5950. [DOI] [PMC free article] [PubMed] [Google Scholar]
  74. Sethi K. K. Multiplication of human-derived Pneumocystis carinii in severe combined immunodeficient (SCID) mice. Experientia. 1992 Jan 15;48(1):63–66. doi: 10.1007/BF01923610. [DOI] [PubMed] [Google Scholar]
  75. Sinclair K., Wakefield A. E., Banerji S., Hopkin J. M. Pneumocystis carinii organisms derived from rat and human hosts are genetically distinct. Mol Biochem Parasitol. 1991 Mar;45(1):183–184. doi: 10.1016/0166-6851(91)90042-5. [DOI] [PubMed] [Google Scholar]
  76. Smulian A. G., Sullivan D. W., Linke M. J., Halsey N. A., Quinn T. C., MacPhail A. P., Hernandez-Avila M. A., Hong S. T., Walzer P. D. Geographic variation in the humoral response to Pneumocystis carinii. J Infect Dis. 1993 May;167(5):1243–1247. doi: 10.1093/infdis/167.5.1243. [DOI] [PubMed] [Google Scholar]
  77. Stringer J. R., Stringer S. L., Zhang J., Baughman R., Smulian A. G., Cushion M. T. Molecular genetic distinction of Pneumocystis carinii from rats and humans. J Eukaryot Microbiol. 1993 Nov-Dec;40(6):733–741. doi: 10.1111/j.1550-7408.1993.tb04468.x. [DOI] [PubMed] [Google Scholar]
  78. Stringer J. R. The identity of Pneumocystis carinii: not a single protozoan, but a diverse group of exotic fungi. Infect Agents Dis. 1993 Jun;2(3):109–117. [PubMed] [Google Scholar]
  79. Stringer S. L., Garbe T., Sunkin S. M., Stringer J. R. Genes encoding antigenic surface glycoproteins in Pneumocystis from humans. J Eukaryot Microbiol. 1993 Nov-Dec;40(6):821–826. doi: 10.1111/j.1550-7408.1993.tb04481.x. [DOI] [PubMed] [Google Scholar]
  80. Stringer S. L., Hong S. T., Giuntoli D., Stringer J. R. Repeated DNA in Pneumocystis carinii. J Clin Microbiol. 1991 Jun;29(6):1194–1201. doi: 10.1128/jcm.29.6.1194-1201.1991. [DOI] [PMC free article] [PubMed] [Google Scholar]
  81. Stringer S. L., Hudson K., Blase M. A., Walzer P. D., Cushion M. T., Stringer J. R. Sequence from ribosomal RNA of Pneumocystis carinii compared to those of four fungi suggests an ascomycetous affinity. J Protozool. 1989 Jan-Feb;36(1):14S–16S. doi: 10.1111/j.1550-7408.1989.tb02670.x. [DOI] [PubMed] [Google Scholar]
  82. Stringer S. L., Stringer J. R., Blase M. A., Walzer P. D., Cushion M. T. Pneumocystis carinii: sequence from ribosomal RNA implies a close relationship with fungi. Exp Parasitol. 1989 May;68(4):450–461. doi: 10.1016/0014-4894(89)90130-6. [DOI] [PubMed] [Google Scholar]
  83. Sunkin S. M., Stringer J. R. Transcription factor genes from rat Pneumocystis carinii. J Eukaryot Microbiol. 1995 Jan-Feb;42(1):12–19. doi: 10.1111/j.1550-7408.1995.tb01534.x. [DOI] [PubMed] [Google Scholar]
  84. Sunkin S. M., Stringer J. R. Translocation of surface antigen genes to a unique telomeric expression site in Pneumocystis carinii. Mol Microbiol. 1996 Jan;19(2):283–295. doi: 10.1046/j.1365-2958.1996.375905.x. [DOI] [PubMed] [Google Scholar]
  85. Sunkin S. M., Stringer S. L., Stringer J. R. A tandem repeat of rat-derived Pneumocystis carinii genes encoding the major surface glycoprotein. J Eukaryot Microbiol. 1994 May-Jun;41(3):292–300. doi: 10.1111/j.1550-7408.1994.tb01509.x. [DOI] [PubMed] [Google Scholar]
  86. Tibayrenc M., Kjellberg F., Arnaud J., Oury B., Brenière S. F., Dardé M. L., Ayala F. J. Are eukaryotic microorganisms clonal or sexual? A population genetics vantage. Proc Natl Acad Sci U S A. 1991 Jun 15;88(12):5129–5133. doi: 10.1073/pnas.88.12.5129. [DOI] [PMC free article] [PubMed] [Google Scholar]
  87. Van der Ploeg L. H., Gottesdiener K., Lee M. G. Antigenic variation in African trypanosomes. Trends Genet. 1992 Dec;8(12):452–457. doi: 10.1016/0168-9525(92)90330-7. [DOI] [PubMed] [Google Scholar]
  88. Vargas S. L., Hughes W. T., Wakefield A. E., Oz H. S. Limited persistence in and subsequent elimination of Pneumocystis carinii from the lungs after P. carinii pneumonia. J Infect Dis. 1995 Aug;172(2):506–510. doi: 10.1093/infdis/172.2.506. [DOI] [PubMed] [Google Scholar]
  89. Vasquez J., Smulian A. G., Linke M. J., Cushion M. T. Antigenic differences among genetically distinct types of rat-derived Pneumocystis carinii. J Eukaryot Microbiol. 1994 Sep-Oct;41(5):114S–114S. [PubMed] [Google Scholar]
  90. Vavra J., Kucera K. Pneumocystis carinii delanoë, its ultrastructure and ultrastructural affinities. J Protozool. 1970 Aug;17(3):463–483. doi: 10.1111/j.1550-7408.1970.tb04715.x. [DOI] [PubMed] [Google Scholar]
  91. Wada M., Kitada K., Saito M., Egawa K., Nakamura Y. cDNA sequence diversity and genomic clusters of major surface glycoprotein genes of Pneumocystis carinii. J Infect Dis. 1993 Oct;168(4):979–985. doi: 10.1093/infdis/168.4.979. [DOI] [PubMed] [Google Scholar]
  92. Wada M., Sunkin S. M., Stringer J. R., Nakamura Y. Antigenic variation by positional control of major surface glycoprotein gene expression in Pneumocystis carinii. J Infect Dis. 1995 Jun;171(6):1563–1568. doi: 10.1093/infdis/171.6.1563. [DOI] [PubMed] [Google Scholar]
  93. Wakefield A. E. Detection of DNA sequences identical to Pneumocystis carinii in samples of ambient air. J Eukaryot Microbiol. 1994 Sep-Oct;41(5):116S–116S. [PubMed] [Google Scholar]
  94. Wakefield A. E., Fritscher C. C., Malin A. S., Gwanzura L., Hughes W. T., Miller R. F. Genetic diversity in human-derived Pneumocystis carinii isolates from four geographical locations shown by analysis of mitochondrial rRNA gene sequences. J Clin Microbiol. 1994 Dec;32(12):2959–2961. doi: 10.1128/jcm.32.12.2959-2961.1994. [DOI] [PMC free article] [PubMed] [Google Scholar]
  95. Wakefield A. E., Pixley F. J., Banerji S., Sinclair K., Miller R. F., Moxon E. R., Hopkin J. M. Amplification of mitochondrial ribosomal RNA sequences from Pneumocystis carinii DNA of rat and human origin. Mol Biochem Parasitol. 1990 Nov;43(1):69–76. doi: 10.1016/0166-6851(90)90131-5. [DOI] [PubMed] [Google Scholar]
  96. Wakefield A. E., Pixley F. J., Banerji S., Sinclair K., Miller R. F., Moxon E. R., Hopkin J. M. Detection of Pneumocystis carinii with DNA amplification. Lancet. 1990 Aug 25;336(8713):451–453. doi: 10.1016/0140-6736(90)92008-6. [DOI] [PubMed] [Google Scholar]
  97. Walker A. N., Garner R. E., Horst M. N. Immunocytochemical detection of chitin in Pneumocystis carinii. Infect Immun. 1990 Feb;58(2):412–415. doi: 10.1128/iai.58.2.412-415.1990. [DOI] [PMC free article] [PubMed] [Google Scholar]
  98. Walzer P. D., Linke M. J. A comparison of the antigenic characteristics of rat and human Pneumocystis carinii by immunoblotting. J Immunol. 1987 Apr 1;138(7):2257–2265. [PubMed] [Google Scholar]
  99. Walzer P. D. Pneumocystis carinii--new clinical spectrum? N Engl J Med. 1991 Jan 24;324(4):263–265. doi: 10.1056/NEJM199101243240411. [DOI] [PubMed] [Google Scholar]
  100. Watanabe J., Hori H., Tanabe K., Nakamura Y. Phylogenetic association of Pneumocystis carinii with the 'Rhizopoda/Myxomycota/Zygomycota group' indicated by comparison of 5S ribosomal RNA sequences. Mol Biochem Parasitol. 1989 Jan 15;32(2-3):163–167. doi: 10.1016/0166-6851(89)90067-4. [DOI] [PubMed] [Google Scholar]
  101. Weinberg G. A., Durant P. J. Genetic diversity of Pneumocystis carinii derived from infected rats, mice, ferrets, and cell cultures. J Eukaryot Microbiol. 1994 May-Jun;41(3):223–228. doi: 10.1111/j.1550-7408.1994.tb01501.x. [DOI] [PubMed] [Google Scholar]
  102. Williams D. J., Radding J. A., Dell A., Khoo K. H., Rogers M. E., Richards F. F., Armstrong M. Y. Glucan synthesis in Pneumocystis carinii. J Protozool. 1991 Jul-Aug;38(4):427–437. doi: 10.1111/j.1550-7408.1991.tb01382.x. [DOI] [PubMed] [Google Scholar]
  103. Wright T. W., Simpson-Haidaris P. J., Gigliotti F., Harmsen A. G., Haidaris C. G. Conserved sequence homology of cysteine-rich regions in genes encoding glycoprotein A in Pneumocystis carinii derived from different host species. Infect Immun. 1994 May;62(5):1513–1519. doi: 10.1128/iai.62.5.1513-1519.1994. [DOI] [PMC free article] [PubMed] [Google Scholar]
  104. Wyder M. A., Rasch E. M., Kaneshiro E. S. Assessment of Pneumocystis carinii DNA content. J Eukaryot Microbiol. 1994 Sep-Oct;41(5):120S–120S. [PubMed] [Google Scholar]
  105. Yoshikawa H., Morioka H., Yoshida Y. Ultrastructural detection of carbohydrates in the pellicle of Pneumocystis carinii. Parasitol Res. 1988;74(6):537–543. doi: 10.1007/BF00531631. [DOI] [PubMed] [Google Scholar]
  106. Yoshikawa H., Tegoshi T., Yoshida Y. Detection of surface carbohydrates on Pneumocystis carinii by fluorescein-conjugated lectins. Parasitol Res. 1987;74(1):43–49. doi: 10.1007/BF00534931. [DOI] [PubMed] [Google Scholar]
  107. Ypma-Wong M. F., Fonzi W. A., Sypherd P. S. Fungus-specific translation elongation factor 3 gene present in Pneumocystis carinii. Infect Immun. 1992 Oct;60(10):4140–4145. doi: 10.1128/iai.60.10.4140-4145.1992. [DOI] [PMC free article] [PubMed] [Google Scholar]
  108. Zhang J., Cushion M. T., Stringer J. R. Molecular characterization of a novel repetitive element from Pneumocystis carinii from rats. J Clin Microbiol. 1993 Feb;31(2):244–248. doi: 10.1128/jcm.31.2.244-248.1993. [DOI] [PMC free article] [PubMed] [Google Scholar]
  109. Zhang J., Stringer J. R. Cloning and characterization of an alpha-tubulin-encoding gene from rat-derived Pneumocystis carinii. Gene. 1993 Jan 15;123(1):137–141. doi: 10.1016/0378-1119(93)90553-f. [DOI] [PubMed] [Google Scholar]

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