[Frontiers in Bioscience 3, d376-398, March 26, 98]
Reprints
PubMed
CAVEAT LECTOR




Table of Conents
 Previous Section   Next Section

TRANSCRIPTION BY RNA POLYMERASE I

Katherine M. Hannan, Ross D. Hannan and Lawrence I. Rothblum

Henry Hood Research Program, Department of Molecular and Cellular Physiology, Penn State College of Medicine, Weis Centre for Research, 100 N. Academy Avenue, Danville, PA 17822-2618

Received 3/18/98 Accepted 3/23/98

6. REFERENCES

1. Hannan, R.D, L. Taylor, A. Cavanaugh, K. Hannan, & L. Rothblum: The role of UBF in regulating rDNA transcription. (1997) In: RNA Polymerase I: Transcription of Eukaryotic Ribosomal RNA. Eds: Paule, M., in press.

2. Moss, T., & V.Y. Stefanovsky: Promotion and regulation of ribosomal transcription in eukaryotes by RNA polymerase I. Prog. Nucleic Acid Res. Mol. Biol. 50, 25-66 (1995)

3. Paule, M.R: Transcription of ribosomal RNA by eukaryotic RNA polymerase I. In: Transcription: Mechanisms and regulation. Eds: Conaway R.C., Conway J.W., Raven Press, NY (1994)

4. Reeder, R.H: Regulation of transcription by RNA polymerase I. In: Transcription Regulation. Eds: McKnight S.L., Yamamoto K.R., Cold Spring Harbor Lab. Press. (1992)

5. Hannan, R.D., & L.I. Rothblum:. Regulation of ribosomal DNA transcription during neonatal cardiomyocyte hypertrophy. Cardiovas. Res. 30, 501-510 (1995)

6. Larson, D.E., P. Zahradka, & B.H. Sells: Control points in eucaryotic ribosome biogenesis. Biochem. Cell Biol. 69, 5-22 (1991)

7. Paule, M.R., C.T. Iida, P.J. Perna, G.H. Harris, D.A., Knoll, & J.M. D’Alessio: In vitro evidence that eukaryotic ribosomal RNA transcripiton is regulated by modification of RNA polymerase I. Nucleic Acids Res. 12, 8161-8180 (1984)

8. Hannan, R.D., J. Luyken, & L.I. Rothblum: Regulation of rDNA transcription factors during cardiomyocyte hypertrophy induced by adrenergic agents. J. Biol. Chem. 270, 8290-8297 (1995)

9. Hannan, R.D., J. Luyken, & L.I. Rothblum: Regulation of ribosomal DNA transcription during contraction-induced hypertrophy of neonatal cardiomyocytes. J. Biol. Chem. 271, 3213-3220 (1996)

10. Cavanaugh, A.H., W.M. Hemple, L.J. Taylor, V. Rogaisky, G. Todorov, & L.I. Rothblum: Activity of RNA polymerase I transcription factor UBF blocked by Rb gene product. Nature 374, 177-180 (1995)

11. Larson, D., W. Xie, M. Glibetic, D. O’Mahony, B.H.. Sells, & L.I. Rothblum: Coordinated decreases in rRNA gene transcription factors and rRNA synthesis during muscle cell differentiation. Proc. Natl. Acad. Sci. 90, 1-4 (1993)

12. Glibetic, M., L. Taylor, D. Larson, R.D. Hannan, B.H. Sells, & L.I. Rothblum: The RNA polymerase I transcription factor UBF is the product of a primary response gene. J. Biol. Chem. 270, 4209-4212 (1995)

13. O’Mahony, D., W. Xie, S.D.. Smith, H.A. Singer, & L.I. Rothblum: Differential phosphorylation and localization of the transcription factor UBF in vivo in response to serum deprivation. J. Biol. Chem. 267, 35-38 (1992)

14. Antonetti, D.A., S.R. Kimball, R.L. Horetsky, & L.S. Jefferson: Regulation of rDNA transcription by insulin in primary cultures of rat heptocytes. J. Biol. Chem. 268, 25277-25284 (1993)

15. Gokal, P.K., A.H. Cavanaugh, & E.A. Thompson: The effects of cyclohexamide upon transcription of rRNA, 5S RNA, and tRNA genes. J. Biol. Chem. 261, 2536-2541 (1986)

16. Kermekchiev, M., & M. Muramatsu: Presence of an inhibitor of RNA polymerase I mediated transcription in extracts from growth arrested mouse cells. Nucleic Acids Res. 21, 447-453 (1993)

17. Jacob, S.T: Regulation of ribosomal gene transcription. Biochem. J. 306, 617-626 (1995)

18. Reeder, R.H. rRNA synthesis in the nucleolus. Trends Genet. 6, 390-395 (1990)

19. Roussel, P., C. Andre, L. Comai, & D. Hernandez-Verdun: The rDNA transcription machinery is assembled during mitosis in active NORs and absent in inactive NORs. J. Cell Biol. 133, 235-246 (1996)

20. Schwarzacher, H.G., & F. Wachtler: The nucleolus. Anat. Embryol. 188, 515-536 (1993)

21. Jordan, P., M. Mannervik, L. Tora, & M. Carmo-Fonseca: In vivo evidence that TATA-binding protein/SL-1 colocalizes with UBF and RNA polymerase I when rRNA synthesis is either active or inactive. J. Cell Biol. 133, 225-234 (1996)

22. Roussel, P., C. Andre, C. Masson, G. Geraud, & D. Hernandez-Verdun: Localization of the RNA polymerase I transcription factor hUBF during the cell cycle. J. Cell Sci. 104, 327-337 (1993)

23. Junera, H.R., C. Masson, G. Geraud, J. Suja, & D. Hernandez-Verdun: Involvement of in situ conformation of ribosomal genes and selective distribution of upstream binding factor in rRNA transcription. Mol. Biol. Cell 8, 145-156 (1997)

24. Matera, A.G., K.T. Tycowski, J.A. Steitz, & D.C. Ward: Organization of small nucleolar ribonucleoproteins (snoRNPs) by fluorescence in situ hybridization and immunocytochemistry. Mol. Biol. Cell 5, 1289-1299 (1994)

25. Peter, M., J. Nakagawa, M. Doree, J.C. Labbe, & E.A. Nigg: Identification of major nucleolar proteins as candidate mitotic substrates of cdc2 kinase. Cell 60, 791-801 (1990)

26. Weisenberger, D., & U. Scheer: A possible mechanism for the inhibition of ribosomal RNA gene transcription during mitosis. J. Cell Biol. 129, 561-575 (1995)

27. Culotta, V., & B. Sollner-Webb: Sites of topoisomerase I action on X. laevis ribosomal chromatin: Transcriptionally active rDNA has an ~200 bp repeating structure. Cell 52, 585-597 (1988)

28. Labhart, P., & T. Koller: Structure of the active nucleolar chromatin of Xenopus laevis oocytes. Cell 28, 279-292 (1982)

29. Langst, G., T. Schatz, J. Langowski, & I. Grummt: Structural analysis of mouse rDNA: coincidence between nuclease hypersensitive sites, DNA curvature and regulatory elements in the intergenic spacer. Nucleic Acids Res. 25, 511-517 (1997)

30. Mutskov, V.J., V.R. Russanova, S.I. Dimitrov, & I.G. Pashev. Histones associated with non-nucleosomal rat ribosomal genes are acetylated while those bound to nucleosome-organized gene copies are not. J. Biol. Chem. 271, 11852-11857 (1996).

31. Dimitrov, S.I., H.N. Tateossyan, V.Y. Stepanovsky, V.R. Russanova, L. Karagyozov, & I.G. Pashev: Binding of histones to Xenopus laevis ribosomal genes with different levels of expression. Eur. J. Biochem. 204, 977-981 (1992)

32. George, P., B. Demeler, C. Terpening, M.R. Paule, & K.E. van Holde: Binding of the RNA polymerase I transcription complex to its promoter can modify positioning of downstream nucleosomes assembled in vitro. J. Biol. Chem. 268, 1947-1954 (1993)

33. Finley, D., B. Bartel, & A. Vershavsky: The tails of ubiquitin precursors are ribosomal proteins whose fusion to ubiquitin facilitates ribosome biogenesis. Nature 338, 394-401 (1989)

34. Belenguer, P., V. Baldin, C. Mathieu, H. Prats, M. Bensaid, G. Bouche, & F. Amalric: Protein kinase NII and the regulation of rDNA transcription in mammalian cells. Nucleic Acids Res. 17, 6625-6636 (1989)

35. Csermely, P., T. Schnaider, B. Cheatham, M.O.J. Olson, & C.R. Kahn: Insulin induces the phosphorylation of nucleolin. A possible mechanism of insulin-induced RNA efflux from nuclei. J. Biol. Chem. 268, 9747-9752 (1993)

36. Caizergues-Ferrer, M., P. Belenguer, B. Lapeyre, F. Amalric, M.O. Wallace, & M.O.J Olson: Phosphorylation of nucleolin by a nucleolar type NII protein kinase. Biochemistry 26, 7876-7883 (1987)

37. Maridor, G., & E.A. Nigg: cDNA sequences of chicken nuclolin/C23 and NO38/B23, two major nucleolar proteins. Nucleic Acids Res. 18, 1286 (1990)

38. Miller, K.G., J. Tower, & B. Sollner-Webb: A complex control region of the mouse rRNA gene directs accurate initiation by RNA polymerase I. Mol. Cell. Biol. 5, 554-562(1985)

39. Cassidy, B., R. Haglund, & L.I. Rothblum: Regions upstream from the core promoter of the rat ribosomal gene are required for the formation of a stable transcription initiation complex by RNA polymerase I in vitro. Biochem. Biophys. Acta 9, 133-144 (1987)

40. Smith, S., S. Hong, H-F. Yang-Yen, E. Oriahi, D. Lowe, & L.I. Rothblum: In: Mechanisms of control of gene expression. Eds: Cullen B., et al., eds. UCLA Symposia on Molecular and Cellular Biology, New Series, 95, NY (1989)

41. Xie, W.Q., D.J. O’Mahony, S.D. Smith, S.D., & L.I. Rothblum: Complementary in vivo and in vitro analyses of the interactions between cis-acting elements of the rat rDNA promoter. Mol. Cell. Biochem. 104, 127-135 (1991)

42. Pape, L., J. Windle, J. & B. Sollner-Webb: Half helical turn spacing changes convert a frog into a mouse rDNA promoter: a distant upstream domain determines the helix face of the initiation site. Genes Dev. 4,52-62, 1990

43. Leblanc, B., C. Read, & T. Moss: Recognition of the Xenopus ribosomal core promoter by the transcription factor xUBF involves multiple HMG box domains and leads to an xUBF interdomain interaction. EMBO J. 12, 513-525 (1993)

44. Smith, S.D., E. Oriahi, D. Lowe, H.F. Yang-Yen, D. O’Mahony, K. Rose, K. Chen, K., & L.I. Rothblum: Characterization of factors that direct transcription of rat ribosomal DNA. Mol. Cell. Biol. 10, 3105-3116 (1990)

45. Smith, S.D., E. Oriahi, H.F. Yang-Yen, W.Q. Xie, C. Chen, & L.I. Rothblum: Interaction of RNA polymerase I transcription factors with a promoter in the nontranscribed spacer of rat ribosomal DNA. Nucleic Acids Res. 18, 1677-85 (1990)

46. Xie, W., D.J. O’Mahony, S.D. Smith, D. Lowe, & L.I. Rothblum: Analysis of the rat ribosomal DNA promoter: characterization of linker-scanning mutants and of the binding of UBF. Nucleic Acids Res. 20, 1587-92 (1992).

47. Putnam, C.D., & C.S. Pikaard: Cooperative binding of the Xenopus RNA polymerase I transcription factor xUBF to repetitive ribosomal gene enhancers. Mol. Cell. Biol. 12, 4970-4980 (1992)

48. Reeder, R. Enhancers and ribosomal gene spacers. Cell 38, 349-351 (1984)

49. De Winter, R.F.J. & T. Moss: Spacer promoters are essential for efficient enhancement of X. laevis ribosomal transcription. Cell 44, 313-318 (1965)

50. Pikaard, C.S., L.K. Pape, S.L. Henderson, K. Ryan, M.H. Paalman, M.A. Lopata, R.H. Reeder, & B. Sollner-Webb: Enhancers for RNA polymerase I in mouse ribosomal DNA. Mol. Cell. Biol. 10, 4816-4825 (1990)

51. Mason, S.W., E.E. Sander, & I. Grummt: Identification of a transcript release activity acting on ternary transcription complexes containing murine RNA polymerase I. EMBO J 16, 163-172 (1997)

52. Smid, A., M. Finsterer, & I. Grummt: Limited proteolysis unmasks specific DNA-binding of the murine RNA polymerase I-specific transcription termination factor TTFI. J. Mol. Biol. 227, 635-647 (1992)

53. Evers, R., A. Smid, U., Rudloff, F., Lottspeich, & I. Grummt: Different domains of the murine RNA polymerase I-specific termination factor mTTF-I serve distinct functions in transcription termination. EMBO J. 14: 1248-1256 (1995)

54. Labhart, P., & R.H. Reeder: A point mutation uncouples RNA 3’-end formation and termination during ribosomal gene transcription in Xenopus laevis. Genes Dev. 4, 269-276 (1990)

55. McStay, B., & R.H. Reeder: A DNA-binding protein is required for termination of transcription by RNA polymerase I in Xenopus laevis. Mol. Cell. Biol. 10, 2793-2800 (1990)

56. Lang, W.H., B.E. Morrow, Q. Ju, J.R. Warner, & R.H. Reeder: A model for transcription termination by RNA polymerase I. Cell 79, 527-534 (1994)

57. Lang, W.H., & R.H. Reeder: The REB1 site is an essential component of a terminator for RNA polymerase I in Saccharomyces cerevisiae. Mol. Cell. Biol. 13, 649-658 (1993)

58. Lang, W.H., & R.H. Reeder: Transcription termination of RNA polymerase I due to a T-rich element interacting with Reb1p. Proc. Natl. Acad. Sci. 92, 9781-5 (1995)

59. Jeong, S.W., W.H. Lang, & R.H. Reeder, R.H: The release element of the yeast polymerase I transcription terminator can function independently of Reb1p. Mol. Cell. Biol. 15, 5929-36 (1995)

60. Langst, G., T.A. Blank, P.B. Becker, & I. Grummt: RNA polymerase I transcription on nucleosomal templates: the transcription termination factor TTF-I induces chromatin remodeling a relieves transcriptional repression. EMBO J. 16, 760-768 (1997)

61. Gerber, J.K., E. Gogel, C. Berger, M. Wallisch, F. Muller, I. Grummt, I., & F. Grummt: Termination of mammalian rDNA replication: polar arrest of replication fork movement by transcription termination factor TTF-I. Cell. 90,559-67 (1997)

62. Sentenac, A., M. Riva, P. Thuriaux, J-M. Buhler, I. Treich, C. Carles, M. Werner, A. Ruet, J. Huet, C. Mann, N. Chiannikulchai, S. Stettler, & S. Mariotte: Yeast RNA polymerase subunits and genes. In: Transcription Regulation. Eds: McKnight S.L., Yamamoto K.R., Cold Spring Harbor Lab. Press. (1992)

63. Goldberg, M.I., J-C. Perriard, & W.J. Rutter:Purification of rat liver and mouse ascites DNA-dependent RNA polymerase I. Biochemistry 16, 1655-1665 (1977)

64. Song, C.Z., K-I. Hanada, K-I. Yano, Y. Maeda, K. Yamamoto, & M. Muramatsu: High conservation of subunit composition of RNA polymerase I(A) between yeast and mouse and the molecular cloning of mouse RNA polymerase I 40-kDa subunit RPA40. J. Biol. Chem. 269, 26976-26981 (1994)

65. Schwartz, L.B., & R.G. Roeder: Purification and subunit structure of deoxyribonucleic acid-dependent ribonucleic acid polymerase I from the mouse myeloma, MOPC 315. J. Biol. Chem. 249, 5898-5906 (1974)

66. Matsui, T., T. Onishi, & M. Muramatsu: Nucleolar DNA-dependent RNA polymerase from rat liver. 1. Purification and subunit structure. Eur. J. Biochem. 71, 351-360 (1976)

67. Hannan, R.D., W.M. Hempel, A. Cavanaugh, T. Arino, S.I. Dimitrov, T. Moss, & L.I. Rothblum: Affinity purification of mammalian RNA polymerase I. Identification of an associated kinase. J. Biol. Chem. 273, 1257-1267 (1998)

68. Hanada, K-I., C.Z. Song, K. Yamamoto, K-I. Yano, Y. Meada, K. Yamaguchi, & M. Muramatsu: RNA polymerase I associated factor 53 binds to the nucleolar factor UBF and functions in specific rDNA transcription. EMBO J. 15, 2217-2226 (1996)

69. Gadal, O., S. Mariotte-Labarre, S. Chedin, E. Quemeneur, C. Carles, A. Sentenac, & P. Thuriaux: A34.5, a nonessential component of yeast RNA polymerase I, cooperates with subunit A14 and DNA topoisomerase I to produce a functional rRNA synthesis machine. Mol. Cell. Biol. 17, 1787-95 (1997)

69. Yao, Y., K. Yamamoto, Y. Nishi, Y. Nogi, & M. Muramatsu: Mouse RNA polymerase I 16-kDa subunit able to associate with 40-kDa subunit is a homolog of yeast AC19 subunit of RNA polymerase I and III. J. Biol .Chem. 271, 32881-5 (1996)

70. Riva, M., A.R. Schaffner, A. Sentenac, G.R. Hartmann, A.A., Mustaev, E.F. Zaychikov, & M.A. Grachev: Active site labeling of the RNA polymerases A, B, and C from yeast. J. Biol. Chem. 262, 14377-14380 (1987)

71. Sawadogo, M., & A. Sentenac: RNA polymerase B (II) and general transcirption factors. Annu. Rev. Biochem. 59, 711-754 (1990)

72. Tower, J., & B. Sollner-Webb: Transcription of mouse rDNA is regulated by an activated subform of RNA polymerase I. Cell 50, 873-883 (1987)

73. Webb, M.L., J.F. Mealey-Cavender, & S.T. Jacob: Glucocorticoid-induced stimulation of ribosomal gene transcription in rat hepatoma cells is mediated by modification of RNA polymerase I or an associated factor. Mol. Endo. 3, 1861-1868 (1989)

74. Schnapp, A., C. Pfleiderer, R. Rosenbauer, & I. Grummt: A growth-dependent transcription initiation factor (TIF-IA) interacting with RNA polymerase I regulates mouse ribosomal RNA synthesis. EMBO J. 9, 2857-2863 (1990)

75. Schnapp, G., F. Santori, C. Carles, M. Riva, & I. Grummt: The HMG box-containing nucleolar transcription factor UBF interacts with a specific subunit of RNA polymerase I. EMBO J. 13, 190-199 (1994)

76. Seither, P., O. Zatsepina, M. Hoffmann, & I. Grummt: Constitutive and strong association of PAF53 with RNA polymerase I. Chromosoma 106, 216-225 (1997)

77. Hempel, W.M., A.H. Cavanaugh, R.D. Hannan, L. Taylor, & L.I. Rothblum: The species-specific RNA polymerase I transcription factor SL-1 binds to upstream binding factor. Mol. Cell. Biol. 16, 557-563 (1996)

78. Schnapp, A., G. Schnapp, B. Erny, & I. Grummt: Function of the growth-regulated transcription initiation factor TIF-IA in initiation complex formation at the murine ribosomal gene promoter. Mol. Cell. Biol. 13, 6723-6732 (1993)

79. Brun, R.P., K. Ryan, & B. Sollner-Webb: Factor C*, the specific initiation component of the mouse RNA polymerase I holoenzyme, is inactivated early in the transcription process. Mol. Cell. Biol. 14, 5010-5021 (1994)

80. Mahajan, P.B., P.K. Gokal, & E.A. Thompson: Hormonal regulation of transcription of rDNA: The role of TFIC in formation of initiation complexes. J. Biol. Chem. 265, 16244-16247 (1990)

81. Mahajan, P.B., & E.A. Thompson: Hormonal regulation of transcription of rDNA: Purification and characterization of the hormone-regulated transcription factor IC. J. Biol. Chem. 265, 16225-16233 (1990)

82. Yamamoto, R.T., Y. Nogi, J.A. Dodd, & M. Nomura: Rrn3 gene of Saccharomyces cerevisiae encodes an essential RNA polymerase I transcription factor which interacts with the polymerase independently of DNA template. EMBO J. 15, 3964-3973 (1996)

83. Cavanaugh, A.H., & E.A. Thompson: Hormonal regulation of transcription of rDNA: Initiation of transcription by RNA polymerase I in vitro. J. Biol. Chem. 261, 12738-12744 (1986)

84. Schnapp, G., A. Schnapp, H. Rosenbauer, & I. Grummt: TIF-IC, a factor involved in both transcription initiation and elongation of RNA polymerase I. EMBO J. 13, 4028-4035 (1994)

85. Jantzen, H.M., A.M. Chow, D.S., King, & R. Tjian: Multiple domains of the RNA polymerase I activator hUBF interact with the TATA-binding protein complex hSL-1 to mediate transcirption. Genes Dev. 6, 1950-1963 (1992)

86. Radebaugh, C.A., J.L. Matthews, G.K. Geiss, F., Liu, J-M. Wong, E. Bateman, S. Camier, A. Sentenac, & M.R. Paule: TATA box-binding protein (TBP) is a constituent of the polymerase I-specific transcription initiation factor TIF-IB (SL-1) bound to the rRNA promoter and shows differential sensitivity to TBP-directed reagents in Polymerase I, II, and III transcription factors. Mol. Cell. Biol. 14, 597-605 (1994)

87. Smith, S.D., D.J. O’Mahony, T. Kinsella, & L.I. Rothblum: Transcription from the rat 45S ribosomal DNA promoter does not require the factor UBF. Gene Expr. 3, 229-236 (1993)

88. Voit, R., A. Schnapp, A. Kuhn, H., Rosenbauer, P. Hirshmann, H.G. Stunnernberg, & I. Grummt: The nucleolar transcription factor mUBF is phosphorylated by casein kinase II in the C-terminal hyperacidic tail which is essential for transactivation. EMBO J. 11, 2211-2218 (1992)

89. Jantzen, H-M., A. Admon, S.P. Bell, & R. Tjian: Nucleolar transcription factor hUBF contains a DNA-binding motif with homology to HMG proteins. Nature 344, 830-836 (1990)

90. Hannan, R.D., V. Stefanovsky, L. Taylor, T. Moss, & L.I. Rothblum: Overexpression of the transcription factor UBF1 is sufficient to increase ribosomal DNA transcription in neonatal cardiomyocytes: Implications for cardiac hypertrophy. Proc. Natl. Acad.Sci. 93, 8750-8755 (1996)

91. Colbert, T., & S. Hahn: A yeast TFIIB-related factor involved in RNA polymerase III transcription. Genes Dev. 61, 1940-9 (1992)

92. Huet, J., & A. Sentenac: The TATA-binding protein participates in TFIIIB assembly on tRNA genes. Nucleic Acids Res. 20, 6451-4 (1992)

93. Kassavetis, G.A., C.A. Joazeiro, M. Pisano, E.P. Geiduschek ,T. Colbert, S. Hahn, & J.A. Blanco: The role of the TATA-binding protein in the assembly and function of the multisubunit yeast RNA polymerase III transcription factor, TFIIIB. Cell. 71, 1055-64 (1992)

94. Schultz, M.C., R.H. Reeder, & S. Hahn: Variants of the TATA-binding protein can distinguish subsets of RNA polymerase I, II, and III promoters. Cell 69, 697-702 (1992)

95. Cormack, B.P. & K. Struhl: The TATA-binding protein is required for transcription by all three nuclear RNA polymerases in yeast cells. Cell 69, 685-96, (1992)

96. Lalo, D., J.S. Steffan, J.A. Dodd, & M. Nomura: RRN11 encodes the third subunit of the complex containing Rrn6p and Rrn7p that is essential for the initiation of rDNA transcription by yeast RNA polymerase I. J. Biol. Chem. 271, 21062-7 (1996)

97. Keys, D.A., L. Vu, J.S. Steffan, J.A. Dodd, R.T., Yamamoto, Y. Nogi, & M. Nomura: RRN6 and RRN7 encode subunits of a multiprotein complex essential for the initiation of rDNA transcription by RNA polymerase I in Saccharomyces cerevisiae. Genes Dev. 8, 2349-62 (1994)

98. Keys, D.A., B.S. Lee, J.A. Dodd, T.T., Nguyen, L. Vu, E., Fantino, L.M. Burson, Y. Nogi, & M. Nomura: Multiprotein transcription factor UAF interacts with the upstream element of the yeast RNA polymerase I promoter and forms a stable preinitiation complex. Genes Dev. 10, 887-903 (1996)

99. Lin, C.W., B. Moorefield, J. Payne, P. Aprikian, K. Mitomo, & R.H. Reeder: A novel 66-kilodalton protein complexes with Rrn6, Rrn7, and TATA-binding protein to promote polymerase I transcription initiation in Saccharomyces cerevisiae. Mol. Cell. Biol. 16, 6436-43 (1996)

100. Steffan, J.S., D.A. Keys, J.A. Dodd, & M. Nomura: The role of TBP in rDNA transcription by RNA polymerase I in Saccharomyces cerevisiae: TBP is required for upstream activation factor-dependent recruitment of core factor. Genes Dev. 10, 2551-63 (1996)

101. Schnapp, A., J. Clos, W. Hadelt, R. Schreck, A. Cveki, & I. Grummt:. Isolation and functional characterization of TIF-IB, a factor that confers promoter specificity to mouse RNA polymerase I. Nucleic Acids Res. 18, 1385-1393 (1990)

102. Gong, X., C.A. Radebaugh, G.K., Geiss, M.N., Simon, & M.R Paule: Site-directed photo-cross-linking of rRNA transcription initiation complexes. Mol. Cell. Biol. 15, 4956-4963 (1995)

103. Geiss, G., C. Radebaugh, & M. Paule: The Fundamental Ribosomal RNA Transcription Initiation Factor TIF-IB (TIF-IB, SL1, Factor D) Binds to the rRNA Core Promoter Primarily by Minor Groove Contacts. J. Biol. Chem. 272, 29243-29254 (1997)

104. Radebaugh, C., X. Gong, B. Bartholomew, & M. Paule: Identification of previously unrecognized common elements in eukaryotic promoters. A ribosomal RNA gene initiator element for RNA polymerase I. J. Biol. Chem. 272, 3141-3144 (1997)

105. Perna, P., G. Harris, C. Iida, P. Kownin, S. Bugren, & M. Paule:. The start site of the Acanthamoeba castellanii ribosomal RNA transcription unit. Gene Exp. 2, 71-78 (1992)

106. Comai, L., J.C.B.M. Zomerdijk, H. Beckmann, S. Zhou, A. Admon, & R. Tjian: Reconstitution of transcription factor SL-1: Exclusive binding of TBP by SL-1 or TFIID subunits. Science 266, 1966-1972 (1994)

107. Comai, L., N. Tanese, & R. Tjian: The TATA-binding protein and associated factors are integral components of the RNA polymerase I transcription factor, SL-1. Cell 68, 965-976 (1992)

108. Eberhard, D., L. Tora J-M. Egly, & I. Grummt: A TBP-containing multiprotein complex (TIF-IB) mediates transcription specifically of murine RNA polymerase I. Nucleic Acids Res. 21, 4180-4186 (1993)

109. Zomerdijk, J.C.B.M., H. Beckmann, L. Comai, & R. Tjian: Assembly of transcriptionally active RNA polymerase I initiation factor SL-1 from recombinant subunits. Science 266, 2015-2018(1994)

110. Helix, J., J.C.B.M. Zomerdijk, A. Ravanpay, R. Tjian, & I. Grummt: Cloning of murine RNA polymerase I-specific TAF factors: Conserved interactions between subunits of the species-specific transcription factor TIF-IB/SL-1. Proc. Natl. Acad. Sci. 94, 1733-1738 (1997)

111. McStay, B., C.H. Hu, C.S. Pikaard, & R.H. Reeder: xUBF and Rib1 are both required for formation of a stable polymerase I promoter complex with X.laevis. EMBO J. 10, 2297-2303 (1991)

112. Rudloff, U., D. Eberhard, & I. Grummt: The conserved core domain of the human TATA binding protein is sufficient to assemble the multisubunit RNA polymerase I-specific transcription factor SL-1. Proc. Natl. Acad. Sci. 91, 8229-8233 (1994)

113. Learned, R.M., T.K. Learned, M.M. Haltiner, & R. Tjian: Human rRNA transcription is modulated by the coordinant binding of two factors to an upstream control element. Cell 45, 847-857 (1986)

114. Clos, J., D. Buttgereit, & I. Grummt: A purified transcription factor (TIF-IB) binds to essential sequences of the muse rDNA promoter. Proc. Natl. Acad. Sci. 83, 604-608 (1986)

115. Bell S.P., R.M. Learned, H-M, Jantzen, & R. Tjian: Functional cooperativity between transcription factors UBF1 and SL-1 mediates human ribosomal RNA synthesis. Science 241, 1192-1197 (1988)

116. Bell, S.P., H-M. Jantzen, & R. Tjian:. Assembly of alternative multiprotein complexes directs rRNA promoter selectivity. Genes Dev. 4, 943-954 (1990)

117. Beckmann, H., J-L. Chen, T. O’Brien, & R. Tjian: Coactivator and promoter-selective properties of RNA polymerase I TAFs. Science 270, 1506-1509 (1995)

118. Rudloff, U., D. Eberhard, L. Tora, H. Stunnenberg, & I. Grummt: TBP-associated factors interact with DNA and govern species specificity of RNA polymerase I transcription. EMBO J. 13, 2611-2616 (1994)

119. Yang ,Q., C.A. Radebaugh , W. Kubaska, G.K. Geiss, & M.R. Paule. Acanthamoeba castellanii contains a ribosomal RNA enhancer binding protein which stimulates TIF-IB binding and transcription under stringent conditions. Nucleic Acids Res. 23, 4345-52 (1995)

120. Kwon, H., & M.R. Green: The RNA polymerase I transcription factor, upstream binding factor, interacts directly with the TATA box-binding protein. J. Biol. Chem. 269, 30140-30146 (1994)

121. Mishima, Y., I. Financsek, R. Kominami, & M. Muramatsu: Fractionation and reconstitution of factors required for accurate transcription of mammalian ribosomal RNA genes: identification of a species-dependent initiation factor. Nucleic Acids Res. 10, 6659-6670 (1982)

122. Grummt, I., E. Roth, & M.R. Paule: Ribosomal RNA transcription in vitro is species specific. Nature 296, 173-174 (1982)

123. Pikaard, G.S., D.S. Smith, R.H. Reeder, & L.I. Rothblum: rUBF, an RNA Polymerase I transcription factor from rats, produces Dnase I footprints identical to those produced by xUBF, its homolog from frogs. Mol. Cell. Biol. 10, 3810-3812 (1990)

124. Learned, R.M., S. Cordes, & R. Tjian: Purification and characterization of a transcription factor that confers promoter specificity to human RNA polymerase I. Mol. Cell. Biol. 5, 1358-1369 (1985)

125. Bell, S.P., C.S. Pikaard, R.H. Reeder, & R. Tjian: Molecular mechanisms governing species-specific transcription of ribosomal RNA. Cell 59, 489-497 (1989)

126. Culotta, V.C., J.K. Wilkinson, & B. Sollner-Webb: Mouse and frog violate the paradigm of species -specific transcription of ribosomal RNA genes. Proc. Natl. Acad. Sci. 84, 7498-7502 (1987)

127. Ghosh, A.K., H. Niu, & S.T. Jacob: Rat ribosomal RNA gene can utilize primate RNA polymerase I transcription machinery: Lack of absolute species specificity in rDNA transcription. Biochem. Biophys. Res. Commun. 225, 890-895 (1996)

128. Histake, K., T. Nishimura, Y. Maeda, K-I. Hanada, C-Z., Song, & M. Muramatsu: Cloning and structural analysis of cDNA and the gene for mouse transcription factor UBF. Nucleic Acids Res. 19, 4631-4637 (1991)

129. O’Mahony, D.J., & L.I. Rothblum: Identification of two forms of the RNA polymerase I transcription factor UBF. Proc. Natl. Acad. Sci. 88, 3180-3184 (1991)

130. Bachvarov, D., M. Normandaeu, & T. Moss: Heterogeneity in the Xenopus ribosomal transcription factor xUBF has a molecular basis distinct from that in mammals. FEBS 288, 55-59 (1991)

131. Bachvarov, D., & T. Moss: The RNA polymerase I transcription factor xUBF contains 5 tandemly repeated HMG homology boxes. Nucleic Acids Res. 19: 2331-2335 (1991)

132. Kuhn, A., R. Voit, V. Stefanovsky, R. Evers, M. Bianchi, & I. Grummt: Functional differences between the two splice variants of the nucleolar transcription factor UBF: the second HMG box determines specificity of DNA binding and transcriptional activity. EMBO J. 13, 416-424 (1994)

133. McStay, B., G.J. Sullivan, & C. Cairns: The Xenopus RNA polymerase I transcirption factor, UBF, has a role in transcirptional enhancement distinct from that at the promoter. EMBO J. 16,396-405 (1997)

134. Nishimura, T., K-I. Hanada, Y. Maeda, C.Z.. Song, K. Hisatake, & M. Muramatsu: Regulation of mouse UBF gene by multiple growth-related control elements. Biochem. Biophysic. Res. Commun. 205, 1217-1225 (1994)

135. Cairns, C., & B. McStay: HMG box 4 is the principal determinant of specifies specificity in the RNA polymerase I transcription factor UBF. Nucleic Acids Res. 23, 4583-4590 (1995)

136. Hu, C.H., B. McStay, S-W. Jeong, & R.H. Reeder: xUBF, an RNA polymerase I transcription factor, binds crossover DNA with low sequence specificity. Mol. Cell. Biol. 14, 2871-2882 (1994)

137. McStay, B., M.W. Frazier, & R.H. Reeder: xUBF contains a novel dimerization domain essential for RNA polymerase I transcription. Genes Dev. 5, 1957-1968 (1991)

138. Maeda, Y., K. Histake, T. Kondo, K. Hanada, C.Z. Song, T. Nishimura, & M. Muramatsu: Mouse rRNA gene transcription factor mUBF requires both HMG-box1 and an acidic tail for nucleolar accumulation:molecular analysis of the nucleolar targetting mechansim. EMBO J 11, 3695-704 (1992)

139. O’Mahony, D.J., S.D. Smith, W. Xie, & L.I. Rothblum: Analysis of the phosphorylation, DNA-binding and dimerization properties of the RNA polymerase I transcription factors UBF1 and UBF2. Nucleic Acids Res. 20, 1301-8 (1992)

140. Copenhaver, G.P., C.D. Putnam, M.L. Denton, & C.S. Pikaard: The RNA polymerase I transcription factor UBF is a sequence-tolerant HMG-box protein that can recognize structural nucleic acids. Nucleic Acids Res. 22, 2651-2657 (1994)

141. Bazett-Jones, D.P., B. Leblanc, M. Herfort, & T. Moss: Short-range DNA looping by the Xenopus HMG-box transcription factor, xUBF. Science 264, 1134-1137 (1994)

142. Putnam, C.D., G.P. Copenhaver, M.L. Denton, & C.S. Pikaard: The RNA polymerase I transactivator upstream binding factor requires its dimerization domain and high-mobility-group (HMG) box 1 to bend, wrap, and positively supercoil enhancer DNA. Mol. Cell. Biochem. 14, 6476-6488 (1994)

143. Luyken, J., R.D. Hannan, J.Y. Cheung, & L.I. Rothblum: Regulation of rDNA transcription during endothelin-1-induced hypertrophy of neonatal cardiomyocytes. Hyperphosphorylation of upstream binding factor, an rDNA transcription factor. Circ. Res. 78, 354-361 (1996)

144. White, R.J. Regulation of RNA polymerase I and III by the retinoblastoma protein: a mechanism for growth control? Trends in Biol. Sci. 22, 77-80 (1997)

145. Zhang, J., & S.T. Jacob: Purification and characterization of a novel factor which stimulates rat ribosomal gene transcription in vitro by interacting with enhancer and core promoter elements. Mol. Cell. Biol. 10, 5177-5186 (1990)

146. Niu, H., J. Zhang, & S.T. Jacob: E1BF/Ku interacts physically and functionally with the core promoter binding factor CPBP and promotes the basal transcription of rat and human ribosomal RNA genes. Gene Exp. 4, 111-124 (1995)

147. Kuhn, A., V. Stefanovsky, & I. Grummt: The nucleolar transcription activator UBF relieves Ku antigen-mediated repression of mouse ribosomal gene transcription. Nucleic Acids Res. 21, 2057-2063 (1993)

148. Hoff, C.M., & S.T. Jacob: Characterization of the factor E1BF from a rat hepatoma that modulates ribosomal RNA gene transcription and its relationship to the human Ku autoantigen. Biochem. Biophys. Res. Commun. 190, 747-753 (1993)

149. Ghosh, A.K., C.M. Hoff, & S.T. Jacob: Characterization of the 130-bp repeat enhancer element of the rat ribosomal gene: functional interaction with transcription factor E1BF. Gene 125, 217-222 (1993)

150. Labhart, P. DNA-dependent protein kinase specifically represses promoter-directed transcription initiation by RNA polymerase I. Proc. Natl. Acad. Sci. 92, 2934-2938 (1995)

151. Kuhn, A., T.M. Gottlieb, S.P. Jackson, & I. Grummt: DNA-dependent protein kinase: a potent inhibitor of transcription by RNA polymerase I. Genes Dev. 9, 193-203 (1995)

152. Liu, Z., & S.T. Jacob: Characterization of a protein that interacts with the rat ribosomal gene promoter and modulates RNA polymerase I transcription. J. Biol. Chem. 269, 16618-16626 (1994)

153. Datta, P.K., A.K. Ghosh, & S.T. Jacob: The RNA polymerase I promoter-activating factor CPBP is functionally and immunologically related to the basic helix-loop-helix-zipper DNA-binding protein USF. J. Biol. Chem. 270: 8637-8641 (1995)

154. Ghosh, A.K., P.K. Datta, & S.T. Jacob: The dual role of helix-loop-helix-zipper protein USF in ribosomal RNA gene transcription in vivo. Oncogene 14, 589-594 (1997)

155. Wang, J.C. DNA topoisomerases. Ann. Rev. Biochem. 54, 665-97 (1985)

156. Brill, S.J., S. DiNardo, K., Voelkel-Meiman, & R. Sternglanz: Need for DNA topoisomerase activity as a swivel for DNA replication for transcription of ribosomal RNA. Nature 326, 414-416 (1987)

157. Zini, N., S. Santi, A. Ognibene, A. Bavelloni, L.M. Neri, A. Valmori, E. Mariani, C. Negri, G.C.B. Astaldi-Ricotti, & N.M. Maraldi: Discrete localization of different DNA topoisomerases in HeLa and K562 cell nuclei and subnuclear fractions. Exp. Cell Res. 210, 336-348 (1994)

158. Brou, C., A. Kuhn, A. Staub, S. Chaudhary, I. Grummt, I. Davidson, & L. Tora: Sequence-specific transactivators counteract topoisomerase II-mediated inhibition of in vitro transcription by RNA polymerases I and II. Nucleic Acids Res. 21, 4011-4018 (1993)

159. Muller, M.T., W.P. Pfund, V.B., Mehta, & D.K. Trask: Eukaryotic type I topoisomerase is enriched in the nucleolus and catalytically active on ribosomal DNA. EMBO J. 4, 1237-1243 (1985)

160. Fleischmann, G., G. Pflugfelder, E.K. Steiner, K. Javaherian, G.C. Howard, J.C. Wang, & S.C.R. Elgin: Drosophila DNA topoisomerase I is associated with transcriptionally active regions of the genome. Proc. Natl. Acad. Sci. 6958-6962 (1984)

161. Schultz, M.C., S.J. Brill, Q. Ju, R. Sternglanz, & R.H. Reeder: Topoisomerases and yeast rRNA transcription: negative supercoiling stimulates initiation and topoisomerase activity is required for elongation. Genes Dev. 6, 1332-41 (1992)

162. Yang-Yen, H-F., & L.I. Rothblum: Purification and characterization of a high-mobility-group-like DNA-binding protein that stimulates rRNA synthesis in vitro. Mol. Cell. Biol. 8: 3406-3414 (1988)

163. Bateman, E., & M.R. Paule: Events during eucaryotic rRNA transcription initiation and elongation: Conversion from the closed to the open promoter complex requires nucleotide substrates. Mol. Cell. Biol. 8, 1940-1946 (1988)

164. Paule, M.R: Regulation of ribosomal RNA transcription during differentiation of Acanthamoeba castellanii: a review. J. Protozoology 30, 211-214 (1983)

165. Kownin, P., E. Bateman, & M.R. Paule: Eukaryotic RNA polymerase I promoter binding is directed by protein contacts with transcription initiation factor and is DNA sequence-independent. Cell 50, 693-699 (1987)

166. Kato, H., M. Nagamine, R. Kominami, & M. Muramatsu:. Formation of the transcription initiation complex on mammalian rDNA. Mol. Cell. Biol. 6, 3418-3427 (1986)

167. Schnapp, A., & I. Grummt: Transcription complex formation at the mouse rDNA promoter involves the stepwise association of four transcription factors and RNA polymerase I. J. Biol. Chem. 266, 24588-24595 (1991)

168. Kuhn, A., & I. Grummt: Dual role of the nucleolar transcription factor UBF: Trans-activator and antirepressor. Proc. Natl. Acad. Sci. 89, 7340-7344 (1992)

169. Derenzini, M., D. Hernandez-Verdun, F. Farabegoli, A. Pession, & F. Novello: Structure of ribosomal genes of mammalian cells in situ. Chromosoma 95, 63-70 (1987)

170. Gebrane-Younes, J., N. Fomproix, & D. Hernandez-Verdun: When rDNA transcription is arrested during mitosis, UBF is still associated with non-condensed rDNA. J. Cell Science 110, 2429-40 (1997)

171. Bateman, E., & M.R. Paule: Regulation of eukaryotic ribosomal RNA transcription by RNA polymerase modification. Cell 47: 445-450 (1986)

172. Buttergeit, D., G. Pflugfelder, & I. Grummt: Growth-dependent regulation of rRNA synthesis is mediated by a transcription factor (TIF-IA). Nucleic Acids Res. 13, 8165-8180 (1985)

173. Zhai, W., J.A. Tuan, & L. Comai: SV40 large T antigen binds to the TBP-TAF(I) complex SL-1 and coactivates ribosomal RNA transcription. Genes Dev. 11,1605-17 (1997)

174. Kabler, R.L., A. Srinivasan, L.J. Taylor, J. Mowad, L.I. Rothblum, L.I., & A.H. Cavanaugh: Androgen regulation of ribosomal RNA synthesis in LNCaP cells and rat prostate. J. Steroid Biochem. Mol. Biol. 59, 431-439 (1996)

175. Hershey, J.C., M. Hautmann, M.M. Thompson, L.I. Rothblum, T. A.J. Haystead, T.A.J., & G.K. Owens: Angiotensin II-induced hypertrophy of rat vascular smooth muscle is associated with increased 18S rRNA synthesis and phosphorylation of the rRNA transcription factor, upstream binding factor. J. Biol. Chem. 270, 25096-25101 (1995)

176. Levine, A.J. The tumor suppressor genes. Ann. Rev. Biochem. 62, 623-651 (1993)

177. Voit, R., K. Schafer, & I. Grummt: Mechanism of repression of RNA polymerase I transcription by the retinoblastoma protein. Mol. Cell. Biol. 17, 4230-4237 (1997)

178. Hoff, C.M., A.K. Ghosh, B.S. Prabhakar, & S.T. Jacob: Enhancer 1 binding factor, a Ku-related protein, is a positive regulator of RNA polymerase I transcription initiation. Proc. Natl. Acad. Sci. 91, 762-766 (1994)

179. Niu, H., & S.T. Jacob: Enhancer 1 binding factor, a Ku-related protein, is a growth-regulated RNA polymerase I transcription factor: Association of a repressor activity with purified E1BF from serum-deprived cells. Proc. Natl. Acad. Sci. 91, 9101-9105 (1994)

180. Fewell, J.W., & E.L. Kuff: Intracellular redistribution of Ku immunoreactivity in response to cell-cell contact and growth modulating components in the medium. J. Cell Sci. 109, 1937-1946 (1996)

181. Datta, P.K., S. Budhiraja, R.R. Reichel, & S.T. Jacob: Regulation of ribosomal RNA gene transcription during retinoic acid-induced differentiation of mouse teratocarcinoma cells. Exp. Cell Res. 231,198-205 (1997)

182. Ghoshal, K., & S.T. Jacob: Heat shock selectivity inhibits ribosomal RNA gene transcription and down-regulates E1BF/Ku in mouse lymphosarcoma cells. Biochem. J 317, 689-95 (1996)

183. Zhang, H., J.C. Wang, & L.F. Liu: Involvement of DNA topoisomerase I in transcription of human ribosomal RNA genes. Proc. Natl. Acad. Sci. 85, 1060-1064 (1988)

184. Kretzschmar, M., M. Meisterernst, & R.G. Roeder: Identification of human DNA topoisomerase I as a cofactor for activator-dependent transcription by RNA polymerase II. Proc. Natl. Acad. Sci. 90, 11508-12 (1993)

185. Rose, K.M., J. Szopa, F.S. Han, Y.C. Cheng, A. Richter, & U. Scheer: Association of DNA topoisomerase I and RNA polymerase I: a possible role for topoisomerase I in ribosomal gene transcription. Chromosoma 96, 411-6 (1988)

186. Rose, K.M. DNA topoisomerases as targets for chemotherapy. FASEB 2, 2474-8 (1988)