Genetics techniques for Thermococcus kodakarensis

Frontiers in Microbiology

Volumn 3, Issue JUN, 2012, Pages

  Hileman, Travis H ^a   Santangelo, Thomas J ^a  

^a OHIO STATE UNIVERSITY   (United States)

Author keywords

Archaea; Genetics; Hyperthermophilic; Recombination; Thermococcales; Transcription

Indexed keywords

EID: 84870695619     PISSN: None     EISSN: 1664302X     Source Type: Journal    
DOI: 10.3389/fmicb.2012.00195     Document Type: Article

References (89)

1. Albers, S. V., and Driessen, A. J. (2008). Conditions for gene disruption by homologous recombination of exogenous DNA into the Sulfolobus solfataricus genome. Archaea 2, 145-149.
2. Atomi, H. (2005). Recent progress towards the application of hyper-thermophiles and their enzymes. Curr. Opin. Chem. Biol. 9, 166-173.
3. Atomi, H., Ezaki, S., and Imanaka, T. (2001). Ribulose-1,5-bisphosphate carboxylase/oxygenase from Ther-mococcus kodakarensis KOD1. Meth. Enzymol. 331, 353-365.
4. Atomi,H.,Matsumi,R.,and Imanaka,T. (2004a). Reverse gyrase is not a pre-requisite for hyperthermophilic life. J. Bacteriol. 186, 4829-4833.
5. Atomi, H., Fukui, T., Kanai, T., Morikawa, M., and Imanaka, T. (2004b). Description of Thermococ-cus kodakarensis sp nov, a well studied hyperthermophilic archaeon previously reported as Pyrococcus sp KOD1. Archaea 1, 263-267.
6. Atomi, H., Sato, T., and Kanai, T. (2011). Application of hyperther-mophiles and their enzymes. Curr. Opin. Biotechnol. 22, 618-626.
7. Bae, H., Kim, K. P., Lee, J. I., Song, J. G., Kil, E. J., Kim, J. S., and Kwon, S. T. (2009). Characterization of DNA polymerase from the hyperthermophilic archaeon Ther-mococcus marinus and its application to PCR. Extremophiles 13, 657-667.
8. Bae, S. S., Kim, T. W., Lee, H. S., Kwon, K. K., Kim, Y. J., Kim, M. S., Lee, J. H., and Kang, S. G. (2012). H2 production from CO, formate or starch using the hyper-thermophilic archaeon, Thermococ-cus onnurineus. Biotechnol. Lett. 34, 75-79.
9. Blumer-Schuette, S. E., Kataeva, I., Westpheling, J., Adams, M. W., and Kelly, R. M. (2008). Extremely thermophilic microorganisms for biomass conversion: status and prospects. Curr. Opin. Biotechnol. 19, 210-217.
10. Borges, N., Matsumi, R., Imanaka, T., Atomi, H., and Santos, H. (2010). Thermococcus kodakarensis mutants deficient in dimyoinositol phos-phate use aspartate to cope with heat stress. J. Bacteriol. 192, 191-197.
11. Bridger, S. L., Clarkson, S. M., Stirrett, K., DeBarry, M. B., Lipscomb, G. L., Schut, G. J., Westpheling, J., Scott, R. A., and Adams, M. W. (2011). Deletion strains reveal metabolic roles for key elemental sulfur-responsive proteins in Pyrococcus furiosus. J. Bacteriol. 193, 6498-6504.
12. Cho, Y., Lee, H. S., Kim, Y. J., Kang, S. G., Kim, S. J., and Lee, J. H. (2007). Characterization of a dUT-Pase from the hyperthermophilic archaeon Thermococcus onnurineus NA1 and its application in poly-merase chain reaction amplification. Mar. Biotechnol. 9, 450-458.
13. Chou, C. J., Jenney, F. E. Jr., Adams, M. W., and Kelly, R. M. (2008). Hydrogenesis in hyperthermophilic microorganisms: implications for biofuels. Metab. Eng. 10, 394-404.
14. Danno,A.,Fukuda,W.,Yoshida,M.,Aki, R., Tanaka, T., Kanai, T., Imanaka, T., and Fujiwara, S. (2008). Expression profiles and physiological roles of two types of prefoldins from the hyperthermophilic archaeon Thermococcus kodakarensis. J. Mol. Biol. 382, 298-311.
15. Davidova, I. A., Duncan, K. E., Perez-Ibarra, B. M., and Suflita, J. M. (2012). Involvement of thermophilic archaea in the bio-corrosion of oil pipelines. Environ. Microbiol. doi: 10.1111/j.1462-2920.2012.02721.x. [Epub ahead of print].
16. De Stefano, L., Vitale, A., Rea, I., Staiano, M., Rotiroti, L., Labella, T., Rendina, I., Aurilia, V., Rossi, M., and D'Auria, S. (2008). Enzymes and proteins from extremophiles as hyperstable probes in nanotechnology: the use of D-trehalose/D-maltose-binding protein from the hyperthermophilic archaeon Ther-mococcus litoralis for sugars monitoring. Extremophiles 12, 69-73.
17. Dev, K., Santangelo, T. J., Rothen-burg, S., Neculai, D., Dey, M., Sicheri, F., Dever, T. E., Reeve, J. N., and Hinnebusch, A. G. (2009). Archaeal aIF2B interacts with eukaryotic translation initiation factors eIF2alpha and eIF2Balpha: implications for aIF2B function and eIF2B regulation. J. Mol. Biol. 392, 701-722.
18. Endoh, T., Kanai, T., and Imanaka, T. (2007). A highly productive system for cell-free protein synthesis using a lysate of the hyperthermophilic archaeon, Thermococcus kodakarensis. Appl. Microbiol. Biotechnol. 74, 1153-1161.
19. Endoh, T., Kanai, T., and Imanaka, T. (2008). Effective approaches for the production of heterolo-gous proteins using the Thermo-coccus kodakarensis-based translation system. J. Biotechnol. 133, 177-182.
20. Endoh, T., Kanai, T., Sato, Y. T., Liu, D. V., Yoshikawa, K., Atomi, H., and Imanaka, T. (2006). Cell-free protein synthesis at high temperatures using the lysate of a hyperthermophile. J. Biotechnol. 126, 186-195.
21. Farkas, J., Chung, D., DeBarry, M., Adams, M. W., and Westpheling, J. (2011). Defining components of the chromosomal origin of replication of the hyperthermophilic archaeon Pyrococcus furiosus needed for construction of a stable replicating shuttle vector. Appl. Environ. Microbiol. 77, 6343-6349.
22. Fujikane, R., Ishino, S., Ishino, Y., and Forterre, P. (2010). Genetic analysis of DNA repair in the hyper-thermophilic archaeon, Thermococ-cus kodakarensis. Genes Genet. Syst. 85, 243-257.
23. Fujiwara, S., Aki, R., Yoshida, M., Higashibata, H., Imanaka, T., and Fukuda, W. (2008). Expression profiles and physiological roles of two types of molecular chaperonins from the hyperthermophilic archaeon Thermococcus kodakarensis. Appl. Environ. Microbiol. 74, 7306-7312.
24. Fujiwara, S., Takagi, M., and Imanaka, T. (1998). Archaeon Pyrococcus kodakaraensis KOD1: application and evolution. Biotechnol. Annu. Rev. 4, 259-284.
25. Fukuda, W., Fukui, T., Atomi, H., and Imanaka, T. (2004). First characterization of an archaeal GTP-dependent phosphoenolpyru-vate carboxykinase from the hyper-thermophilic archaeon Thermococ-cus kodakarensis KOD1. J. Bacteriol. 186, 4620-4627.
26. Fukuda, W., Morimoto, N., Imanaka, T., and Fujiwara, S. (2008). Agma-tine is essential for the cell growth of Thermococcus kodakarensis. FEMS Microbiol. Lett. 287, 113-120.
27. Fukui,T.,Atomi,H.,Kanai, T.,Matsumi, R., Fujiwara, S., and Imanaka, T. (2005). Complete genome sequence of the hyperthermophilic archaeon Thermococcus kodakarensis KOD1 and comparison with Pyrococ-cus genomes. Genome Res. 15, 352-363.
28. Gaidamaviciute, E., Tauraite, D., Gagilas, J., and Lagunavicius, A. (2010). Site-directed chemical modification of archaeal Thermococcus litoralis Sh1B DNA polymerase: acquired ability to read through template-strand uracils. Biochim. Biophys. Acta 1804, 1385-1393.
29. Gonnet, M., Erauso, G., Prieur, D., and Le Romancer, M. (2011). pAMT11, a novel plasmid isolated from a Ther-mococcus sp strain closely related to the virus-like integrated element TKV1 of the Thermococcus kodakarensis genome. Res. Microbiol. 162, 132-143.
30. Gorlas, A., Koonin, E. V., Bienvenu, N., Prieur, D., and Geslin, C. (2012). TPV1, the first virus isolated from the hyperthermophilic genus Ther-mococcus. Environ. Microbiol. 14, 503-516.
31. Griffiths, K., Nayak, S., Park, K., Man-delman, D., Modrell, B., Lee, J., Ng, B., Gibbs, M. D., and Bergquist, P. L. (2007). New high fidelity poly-merases from Thermococcus species. Protein Expr. Purif. 52, 19-30.
32. Hashimoto, H., Nishioka, M., Fujiwara, S., Takagi, M., Imanaka, T., Inoue, T., and Kai, Y. (2001). Crystal structure of DNA polymerase from hyper-thermophilic archaeon Pyrococcus kodakaraensis KOD1. J. Mol. Biol. 306, 469-477.
33. Hirata, A., Kanai, T., Santangelo, T. J., Tajiri, M., Manabe, K., Reeve, J. N., Imanaka, T., and Murakami, K. S. (2008). Archaeal RNA polymerase subunits E and F are not required for transcription in vitro, but a Thermo-coccus kodakarensis mutant lacking subunit F is temperature-sensitive. Mol. Microbiol. 70, 623-633.
34. Hotta, Y., Ezaki, S., Atomi, H., and Imanaka, T. (2002). Extremely stable and versatile carboxylesterase from a hyperthermophilic archaeon. Appl. Environ. Microbiol. 68, 3925-3931.
35. Imanaka, H., Yamatsu, A., Fukui, T., Atomi, H., and Imanaka, T. (2006). Phosphoenolpyruvate syn-thase plays an essential role for gly-colysis in the modified Embden-Meyerhof pathway in Thermococ-cus kodakarensis. Mol. Microbiol. 61, 898-909.
36. Imanaka, T., and Atomi, H. (2002). Catalyzing "hot" reactions: enzymes from hyperthermophilic archaea. Chem. Rec. 2, 149-163.
37. Imanaka, T., Fukui, T., and Fujiwara, S. (2001). Chitinase from Thermococ-cus kodakarensis KOD1. Meth. Enzy-mol. 330, 319-329.
38. Ishino, S., Fujino, S., Tomita, H., Ogino, H., Takao, K., Daiyasu, H., Kanai, T., Atomi, H., and Ishino, Y. (2011). Biochemical and genetical analyses of the three mcm genes from the hyperthermophilic archaeon, Ther-mococcus kodakarensis. Genes Cells 16, 1176-1189.
39. Izumi, M., Fujiwara, S., Shiraki, K., Tak-agi, M., Fukui, K., and Imanaka, T. (2001). Utilization of immobilized archaeal chaperonin for enzyme stabilization. J. Biosci. Bioeng. 91, 316-318.
40. Jarrell, K. F., Walters, A. D., Bochiwal, C., Borgia, J. M., Dickinson, T., and Chong, J. P. (2011). Major players on the microbial stage: why archaea are important. Microbiology 157(Pt 4), 919-936.
41. Kanai, T., Akerboom, J., Takedomi, S., van de Werken, H. J., Blombach, F., van der Oost, J., Murakami, T., Atomi, H., and Imanaka, T. (2007). A global transcriptional regulator in Thermococcus kodakarensis controls the expression levels of both gly-colytic and gluconeogenic enzyme-encoding genes. J. Biol. Chem. 282, 33659-33670.
42. Kanai, T., Imanaka, H., Nakajima, A., Uwamori, K., Omori, Y., Fukui, T., Atomi, H., and Imanaka, T. (2005). Continuous hydrogen production by the hyperthermophilic archaeon, Thermococcus kodakarensis KOD1. J. Biotechnol. 116, 271-282.
43. Kanai, T., Matsuoka, R., Beppu, H., Nakajima, A., Okada, Y., Atomi, H., and Imanaka, T. (2011). Dis-tinct physiological roles of the three [NiFe]-hydrogenase orthologs in the hyperthermophilic archaeon Ther-mococcus kodakarensis. J. Bacteriol. 193, 3109-3116.
44. Kanai, T., Takedomi, S., Fujiwara, S., Atomi, H., and Imanaka, T. (2010). Identification of the Phr-dependent heat shock regulon in the hyper-thermophilic archaeon, Thermococ-cus kodakarensis. J. Biochem. 147, 361-370.
45. Kelly, R. M., Dijkhuizen, L., and Leemhuis, H. (2009). Starch and alpha-glucan acting enzymes, modulating their properties by directed evolution. J. Biotechnol. 140, 184-193.
46. Kim, J. W., and Peeples, T. L. (2006). Screening extremophiles for biocon-version potentials. Biotechnol. Prog. 22, 1720-1724.
47. Kim, Y. J., Lee, H. S., Kim, E. S., Bae, S. S., Lim, J. K., Matsumi, R., Lebedinsky, A. V., Sokolova, T. G., Kozhevnikova, D. A., Cha, S. S., Kim, S. J., Kwon, K. K., Imanaka, T., Atomi, H., Bonch-Osmolovskaya, E. A., Lee, J. H., and Kang, S. G. (2010). Formate-driven growth coupled with H(2) production. Nature 467, 352-355.
48. Kobori, H., Ogino, M., Orita, I., Nakamura, S., Imanaka, T., and Fukui, T. (2010). Characterization of NADH oxidase/NADPH polysul-fide oxidoreductase and its unex-pected participation in oxygen sen-sitivity in an anaerobic hyperther-mophilic archaeon. J. Bacteriol. 192, 5192-5202.
49. Lam, W. L., and Doolittle, W. F. (1992). Mevinolin-resistant mutations identify a promoter and the gene for a eukaryote-like 3-hydroxy-3-methylglutaryl-coenzyme A reductase in the archaebacterium Haloferax volcanii. J. Biol. Chem. 267, 5829-5834.
50. Leigh, J. A., Albers, S. V., Atomi, H., and Allers, T. (2011). Model organ-isms for genetics in the domain archaea: methanogens, halophiles, Thermococcales and Sulfolob-ales. FEMS Microbiol. Rev. 35, 577-608.
51. Li, Z., Pan, M., Santangelo, T. J., Chem-nitz, W., Yuan, W., Edwards, J. L., Hurwitz, J., Reeve, J. N., and Kelman, Z. (2011). A novel DNA nuclease is stimulated by association with the GINS complex. Nucleic Acids Res. 39, 6114-6123.
52. Li, Z., Santangelo, T. J., Cuboňová, L., Reeve, J. N., and Kelman, Z. (2010). Affinity purification of an archaeal DNA replication protein network. MBio 1, e00221-10.
53. Lipscomb, G. L., Stirrett, K., Schut, G. J., Yang, F., Jenney, F. E. Jr., Scott, R. A., Adams, M. W., and West-pheling, J. (2011). Natural com-petence in the hyperthermophilic archaeon Pyrococcus furiosus facil-itates genetic manipulation: con-struction of markerless deletions of genes encoding the two cytoplasmic hydrogenases. Appl. Environ. Micro-biol. 77, 2232-2238.
54. Littlechild, J. A. (2011). Thermophilic archaeal enzymes and applications in biocatalysis. Biochem. Soc. Trans. 39, 155-158.
55. Louvel, H., Kanai, T., Atomi, H., and Reeve, J. N. (2009). The Fur iron regulator-like protein is cryptic in the hyperthermophilic archaeon Thermococcus kodakarensis. FEMS Microbiol. Lett. 295, 117-128.
56. Matsubara, K., Yokooji, Y., Atomi, H., and Imanaka, T. (2011). Biochem-ical and genetic characterization of the three metabolic routes in Thermococcus kodakarensis linking glyceraldehyde 3-phosphate and 3-phosphoglycerate. Mol. Microbiol. 81, 1300-1312.
57. Matsumi, R., Atomi, H., Driessen, A. J., and van der Oost, J. (2011). Isoprenoid biosynthesis in archaea - biochemical and evolutionary implications. Res. Microbiol. 162, 39-52.
58. Matsumi, R., Manabe, K., Fukui, T., Atomi, H., and Imanaka, T. (2007). Disruption of a sugar transporter gene cluster in a hyperthermophilic archaeon using a host-marker sys-tem based on antibiotic resistance. J. Bacteriol. 189, 2683-2691.
59. Morikawa, M., Izawa, Y., Rashid, N., Hoaki, T., and, Imanaka, T. (1994). Purification and characterization of a thermostable thiol protease from a newly isolated hyperthermophilic Pyrococcus sp. Appl. Environ. Micro-biol. 60, 4559-4566.
60. Morimoto, N., Fukuda, W., Naka-jima, N., Masuda, T., Terui, Y., Kanai, T., Oshima, T., Imanaka, T., and Fujiwara, S. (2010). Dual biosynthesis pathway for longer-chain polyamines in the hyper-thermophilic archaeon Thermococ-cus kodakarensis. J. Bacteriol. 192, 4991-5001.
61. Murakami, T., Kanai, T., Takata, H., Kuriki, T., and Imanaka, T. (2006). A novel branching enzyme of the GH-57 family in the hyperther-mophilic archaeon Thermococcus kodakarensis KOD1. J. Bacteriol. 188, 5915-5924.
62. Nunoura, T., Takaki, Y., Kakuta, J., Nishi, S., Sugahara, J., Kazama, H., Chee, G. J., Hattori, M., Kanai, A., Atomi, H., Takai, K., and Takami, H. (2011). Insights into the evolution of Archaea and eukaryotic protein modifier systems revealed by the genome of a novel archaeal group. Nucleic Acids Res. 39, 3204-3223.
63. Orita, I., Sato, T.,Yurimoto, H., Kato, N., Atomi, H., Imanaka, T., and Sakai, Y. (2006). The ribulose monophos-phate pathway substitutes for the missing pentose phosphate path-way in the archaeon Thermococ-cus kodakarensis. J. Bacteriol. 188, 4698-4704.
64. Pan, M., Santangelo, T. J., Li, Z., Reeve, J. N., and Kelman, Z. (2011). Thermococcus kodakarensis encodes three MCM homologs but only one is essential. Nucleic Acids Res. 39, 9671-9680.
65. Rashid, N., Kanai, T., Atomi, H., and Imanaka, T. (2004). Among multiple phosphomannomutase gene ortho-logues, only one gene encodes a pro-tein with phosphoglucomutase and phosphomannomutase activities in Thermococcus kodakarensis. J. Bacte-riol. 186, 6070-6076.
66. Rother, M., and Metcalf, W. W. (2005). Genetic technologies for Archaea. Curr. Opin. Microbiol. 8, 745-751.
67. Santangelo, T. J., and Artsimovitch, I. (2011). Termination and antiter-mination: RNA polymerase runs a stop sign. Nat. Rev. Microbiol. 9, 319-329.
68. Santangelo,T. J.,Cubonová,L.,James,C. L., and Reeve, J. N. (2007). TFB1 or TFB2 is sufficient for Thermococcus kodakarensis viability and for basal transcription in vitro. J. Mol. Biol. 367, 344-357.
69. Santangelo, T. J., Cubonová, L., Mat-sumi, R., Atomi, H., Imanaka, T., and Reeve, J. N. (2008a). Polarity in archaeal operon transcription in Thermococcus kodakarensis. J. Bacte-riol. 190, 2244-2248.
70. Santangelo, T. J., Cubonová, L., and Reeve, J. N. (2008b). Shuttle vec-tor expression in Thermococcus kodakarensis: contributions of cis elements to protein synthesis in a hyperthermophilic archaeon. Appl. Environ. Microbiol. 74, 3099-3104.
71. Santangelo, T. J., Cubonová, L., and Reeve, J. N. (2010). Ther-mococcus kodakarensis genetics: TK1827-encoded beta-glycosidase, new positive-selection protocol, and targeted and repetitive dele-tion technology. Appl. Environ. Microbiol. 76, 1044-1052.
72. Santangelo, T. J., Cubonová, L., and Reeve, J. N. (2011). Deletion of alter-native pathways for reductant recy-cling in Thermococcus kodakarensis increases hydrogen production. Mol. Microbiol. 81, 897-911.
73. Santangelo, T. J., Cubonová, L., Skin-ner, K. M., and Reeve, J. N. (2009). Archaeal intrinsic transcription ter-mination in vivo. J. Bacteriol. 191, 7102-7108.
74. Santangelo, T. J., and Reeve, J. N. (2006). Archaeal RNA polymerase is sensitive to intrinsic termination directed by transcribed and remote sequences. J. Mol. Biol. 355,196-210.
75. Santangelo, T. J., and Reeve, J. N. (2010a). Deletion of switch 3 results in an archaeal RNA polymerase that is defective in transcript elongation. J. Biol. Chem. 285, 23908-23915.
76. Santangelo, T. J., and Reeve, J. N. (2010b). "Genetic tools and manip-ulations of the hyperthermophilic heterotrophic archaeon Thermococ-cus kodakarensis," in Extremophiles Handbook, ed. K. Horikoshi (Tokyo: Springer), 567-582.
77. Sato, T., and Atomi, H. (2011). Novel metabolic pathways in Archaea. Curr. Opin. Microbiol. 14, 307-314.
78. Sato, T., Atomi, H., and Imanaka, T. (2007). Archaeal type III RuBis-COs function in a pathway for AMP metabolism. Science 315, 1003-1006.
79. Sato, T., Fukui, T., Atomi, H., and Imanaka, T. (2003). Targeted gene disruption by homologous recom-bination in the hyperthermophilic archaeon Thermococcus kodakarensis KOD1. J. Bacteriol. 185, 210-220.
80. Sato, T., Fukui, T., Atomi, H., and Imanaka, T. (2005). Improved and versatile transformation system allowing multiple genetic manip-ulations of the hyperthermophilic archaeon Thermococcus kodakaren-sis. Appl. Environ. Microbiol. 71, 3889-3899.
81. Sato, T., Imanaka, H., Rashid, N., Fukui, T., Atomi, H., and Imanaka, T. (2004). Genetic evidence identifying the true gluconeogenic fructose-1,6-bisphosphatase in Thermococ-cus kodakarensis and other hyper-thermophiles. J. Bacteriol. 186, 5799-5807.
82. Shiraki, K., Tsuji, M., Hashimoto, Y., Fujimoto, K., Fujiwara, S., Tak-agi, M., and Imanaka, T. (2003). Genetic, enzymatic, and struc-tural analyses of phenylalanyl-tRNA synthetase from Thermococcus kodakarensis KOD1. J. Biochem. 134, 567-574.
83. Soler, N., Gaudin, M., Marguet, E., and Forterre, P. (2011). Plasmids, viruses and virus-like membrane vesicles from Thermococcales. Biochem. Soc. Trans. 39, 36-44.
84. Soler, N., Justome, A., Quevillon-Cheruel, S., Lorieux, F., Le Cam, E., Marguet, E., and Forterre, P. (2007). The rolling-circle plasmid pTN1 from the hyperthermophilic archaeon Thermococcus nautilus. Mol. Microbiol. 66, 357-370.
85. Takemasa, R., Yokooji, Y., Yamatsu, A., Atomi, H., and Imanaka, T. (2011). Thermococcus kodakarensis as a host for gene expression and protein secretion. Appl. Environ. Microbiol. 77, 2392-2398.
86. Tumbula, D. L., and Whitman, W. B. (1999). Genetics of Methanococcus: possibilities for functional genomics in archaea. Mol. Microbiol. 33, 1-7.
87. Yamaji, K., Kanai, T., Nomura, S. M., Akiyoshi, K., Negishi, M., Chen, Y., Atomi, H., Yoshikawa, K., and Imanaka, T. (2009). Protein syn-thesis in giant liposomes using the in vitro translation system of Thermococcus kodakarensis. IEEE Trans. Nanobioscience 8, 325-331.
88. Yamamoto, T., Matsuda, T., Sakamoto, N., Matsumura, H., Inoue, T., Morikawa, M., Kanaya, S., and Kai, Y. (2003). Crystallization and pre-liminary X-ray analysis of TBP-interacting protein from the hyper-thermophilic archaeon Thermococ-cus kodakaraensis strain KOD1. Acta Crystallogr. D Biol. Crystallogr. 59, 372-374.
89. Yokooji, Y., Tomita, H., Atomi, H., and Imanaka, T. (2009). Pantoate kinase and phosphopantothenate synthetase, two novel enzymes nec-essary for CoA biosynthesis in the archaea. J. Biol. Chem. 284, 28137-28145.