Journal of Biological Chemistry · 2000 · 79 citations · 57 references
In order for RNA polymerase (pol) III to produce a sufficient quantity of RNAs of appropriate structure, initiation, termination, and reinitiation must be accurate and efficient. Termination-associated factors have been shown to facilitate reinitiation and regulate transcription in some species. Suppressor tRNA genes that differ in the dT(n) termination signal were examined for function in Schizosaccharomyces pombe. We also developed an S. pombe extract that is active for tRNA transcription that is described here for the first time. The ability of this tRNA gene to be transcribed in extracts from different species allowed us to compare termination in three model systems. Although human pol III terminates efficiently at 4 dTs and S. pombeat 5 dTs, Saccharomyces cerevisiae pol III requires 6 dTs to direct comparable but lower termination efficiency and also appears qualitatively distinct. Interestingly, this pattern of sensitivity to a minimal dT(n) termination signal was found to correlate with the sensitivity to α-amanitin, as S. pombe was intermediate between human and S. cerevisiae pols III. The results establish that the pols III of S. cerevisiae,S. pombe, and human exhibit distinctive properties and that termination occurs in S. pombe in a manner that is functionally more similar to human than is S. cerevisiae. In order for RNA polymerase (pol) III to produce a sufficient quantity of RNAs of appropriate structure, initiation, termination, and reinitiation must be accurate and efficient. Termination-associated factors have been shown to facilitate reinitiation and regulate transcription in some species. Suppressor tRNA genes that differ in the dT(n) termination signal were examined for function in Schizosaccharomyces pombe. We also developed an S. pombe extract that is active for tRNA transcription that is described here for the first time. The ability of this tRNA gene to be transcribed in extracts from different species allowed us to compare termination in three model systems. Although human pol III terminates efficiently at 4 dTs and S. pombeat 5 dTs, Saccharomyces cerevisiae pol III requires 6 dTs to direct comparable but lower termination efficiency and also appears qualitatively distinct. Interestingly, this pattern of sensitivity to a minimal dT(n) termination signal was found to correlate with the sensitivity to α-amanitin, as S. pombe was intermediate between human and S. cerevisiae pols III. The results establish that the pols III of S. cerevisiae,S. pombe, and human exhibit distinctive properties and that termination occurs in S. pombe in a manner that is functionally more similar to human than is S. cerevisiae. polymerase transcription factors complementary readthrough wild type RNA polymerase (pol)1III is a multisubunit enzyme that is directed to initiate RNA synthesis by transcription factors (TF) that bind to gene promoter elements. pol III transcripts comprise a large variety of small nuclear and cytoplasmic RNAs (1Willis I.M. Eur. J. Biochem. 1993; 212: 1-11Crossref PubMed Scopus (191) Google Scholar). Although there is diversity in the promoter structures of pol III-transcribed genes, three classes are responsible for the synthesis of most cellular pol III transcripts, tRNAs, 5 S rRNA, and U6 small nuclear RNA (2Baserga S.J. Steitz J.A. Gesteland R.F. Atkins J.F. The RNA World. Cold Spring Harbor Laboratory, Cold Spring Harbor, NY1993: 359-381Google Scholar). Each of these represent a gene class that utilizes a characteristic promoter structure and specific set of TFs (3Kassavetis G.A. Bardeleben C. Bartholomew B. Braun B.R. Joazeiro C.A.P. Pisano M. Geiduschek E.P. Conaway R.C. Conaway J.W. Transcription: Mechanisms and Regulation. Raven Press, Ltd., New York1994: 107-126Google Scholar). 5 S rRNA genes comprise class I and contain a principal internal promoter that is recognized by TFIIIA. Class 3 genes utilize upstream TATA elements and in metazoans an upstream element recognized by a distinct multisubunit TF (4Lobo S.M. Hernandez N.T. Conaway R.C. Conaway J.W. Transcription: Mechanisms and Regulation. Raven Press, Ltd., New York1994: 127-159Google Scholar). Class 2 genes are represented by tRNA genes, which use an internal promoter comprised of proximal box A and distal box B elements. Distinct subunits of TFIIIC bind to the A box, B box, and terminator element of the class 2 genes and facilitate the assembly of this class of preinitiation complexes (3Kassavetis G.A. Bardeleben C. Bartholomew B. Braun B.R. Joazeiro C.A.P. Pisano M. Geiduschek E.P. Conaway R.C. Conaway J.W. Transcription: Mechanisms and Regulation. Raven Press, Ltd., New York1994: 107-126Google Scholar, 5Arrebola A. Manaud N. Rozenfeld S. Marsolier M.-C. Lefebvre O. Carles C. Thuriaux P. Conesa C. Sentenac A. Mol. Cell. Biol. 1998; 18: 1-9Crossref PubMed Google Scholar). For each gene class, TFIIIB (or related activity) binds just upstream of the start site of transcription, and this in turn serves as the initiation factor proper as it recruits pol III (Refs. 6Kassavetis G.A. Braun B.R. Nguyen L.H. Geiduschek E.P. Cell. 1990; 60: 235-245Abstract Full Text PDF PubMed Scopus (359) Google Scholar, 7Hernandez N. McKnight S.L. Yamamoto K.R. Transcriptional Regulation. Cold Spring Harbor Laboratory, Cold Spring Harbor, NY1992: 281-313Google Scholar, 8McBryant S.J. Baird E.E. Trauger J.W. Dervan P.B. Gottesfeld J.M. J. Mol. Biol. 1999; 286: 973-981Crossref PubMed Scopus (21) Google Scholar and references therein). Subunits of TFIIIB as well as pol III have been conserved from Saccharomyces cerevisiae to human, as have two TFIIIC subunits that localize near the start site of transcription (9Taggart A.K. Fisher T.S. Pugh B.F. Cell. 1992; 71: 1015-1028Abstract Full Text PDF PubMed Scopus (112) Google Scholar, 10Lopez-De-Leon A. Librizzi M. Puglia K. Willis I.M. Cell. 1992; 71: 211-220Abstract Full Text PDF PubMed Scopus (108) Google Scholar, 11Wang Z. Roeder R.G. Proc. Natl. Acad. Sci. U. S. A. 1995; 89: 7026-7030Crossref Scopus (109) Google Scholar, 12Hsieh Y.J. Wang Z. Kovelman R. Roeder R.G. Mol. Cell. Biol. 1999; 19: 4944-4952Crossref PubMed Scopus (53) Google Scholar, 13Colbert T. Hahn S. Genes Dev. 1992; 6: 1940-1949Crossref PubMed Scopus (128) Google Scholar, 14Wang Z. Roeder R.G. Genes Dev. 1997; 11: 1315-1326Crossref PubMed Scopus (127) Google Scholar). By contrast, the downstream TFIIIC subunits in these organisms reveal no recognizable sequence homology (12Hsieh Y.J. Wang Z. Kovelman R. Roeder R.G. Mol. Cell. Biol. 1999; 19: 4944-4952Crossref PubMed Scopus (53) Google Scholar, 15Lagna G. Kovelman R. Sukegawa J. Roeder R.G. Mol. Cell. Biol. 1994; 14: 3053-3064Crossref PubMed Scopus (44) Google Scholar, 16Sinn E. Wang Z. Kovelman R. Roeder R.G. Genes Dev. 1995; 9: 675-685Crossref PubMed Scopus (56) Google Scholar). Some evidence suggests that efficient transcription requires termination and associated activities that promote pol III recycling (17Dieci G. Sentenac A. Cell. 1996; 84: 245-252Abstract Full Text Full Text PDF PubMed Scopus (147) Google Scholar, 18Fan H. Sakulich A.L. Goodier J.L. Zhang X. Qin J. Maraia R.J. Cell. 1997; 88: 707-715Abstract Full Text Full Text PDF PubMed Scopus (94) Google Scholar, 19Goodier J.L. Maraia R.J. J. Biol. Chem. 1998; 273: 26110-26116Abstract Full Text Full Text PDF PubMed Scopus (25) Google Scholar, 20Maraia R.J. Proc. Natl. Acad. Sci. U. S. A. 1996; 93: 3383-3387Crossref PubMed Scopus (93) Google Scholar, 21Maraia R.J. Kenan D.J. Keene J.D. Mol. Cell. Biol. 1994; 14: 2147-2158Crossref PubMed Scopus (135) Google Scholar, 22Wang Z. Roeder R.G. Mol. Cell. 1998; 1: 749-757Abstract Full Text Full Text PDF PubMed Scopus (67) Google Scholar). Upon encountering the dT(n) tract that comprises the pol III termination signal, by pol III itself, the enzyme pauses and releases the transcript and template (23Bogenhagen D.F. Brown D.D. Cell. 1981; 24: 261-270Abstract Full Text PDF PubMed Scopus (321) Google Scholar, 24Geiduschek E.P. Tocchini-Valentini G.P. Annu. Rev. Biochem. 1988; 57: 873-914Crossref PubMed Scopus (446) Google Scholar, 25Campbell F.E. Setzer D.R. Mol. Cell. Biol. 1992; 12: 2260-2272Crossref PubMed Scopus (61) Google Scholar, 26Shaaban S.A. Bobkova E.V. Chudzik D.M. Hall B.D. Mol. Cell. Biol. 1996; 16: 6468-6476Crossref PubMed Scopus (26) Google Scholar). Multiple subunits of pol III affect termination (Refs. 26Shaaban S.A. Bobkova E.V. Chudzik D.M. Hall B.D. Mol. Cell. Biol. 1996; 16: 6468-6476Crossref PubMed Scopus (26) Google Scholar and 27Chedin S. Riva M. Schultz P. Sentenac A. Carles C. Genes Dev. 1998; 12: 3857-3871Crossref PubMed Scopus (150) Google Scholar and also see Ref. 28Thuillier V. Brun I. Sentenac A. Werner M. EMBO J. 1996; 15: 618-629Crossref PubMed Scopus (37) Google Scholar). Evidence that TFIIIC and/or associated factors, as well as the -UUU-OH terminus binding protein, La, may facilitate pol III termination and reinitiation, exists for human-derived systems (19Goodier J.L. Maraia R.J. J. Biol. Chem. 1998; 273: 26110-26116Abstract Full Text Full Text PDF PubMed Scopus (25) Google Scholar, 21Maraia R.J. Kenan D.J. Keene J.D. Mol. Cell. Biol. 1994; 14: 2147-2158Crossref PubMed Scopus (135) Google Scholar,22Wang Z. Roeder R.G. Mol. Cell. 1998; 1: 749-757Abstract Full Text Full Text PDF PubMed Scopus (67) Google Scholar, 29Gottlieb E. Steitz J.A. EMBO J. 1989; 8: 851-861Crossref PubMed Scopus (298) Google Scholar, 30Chu W.M. Ballard R.E. Schmid C.W. Nucleic Acids Res. 1997; 25: 2077-2082Crossref PubMed Scopus (18) Google Scholar). We modified a suppressor tRNA gene, sup3-e (31Kohli J. Munz P. Soll D. Nasim A. Young P. Johnson B.F. Molecular Biology of the Fission Yeast. Academic Press, Inc., San Diego1989: 75-96Crossref Google Scholar, 32Willis I. Nichols M. Chisholm V. Soll D. Heyer W.D. Szankasi P. Amstutz H. Munz P. Kohli J. Proc. Natl. Acad. Sci. U. S. A. 1986; 83: 7860-7864Crossref PubMed Scopus (14) Google Scholar), to serve as a reporter of pol III termination in the fission yeastSchizosaccharomyces pombe. To complement the results obtained, we developed an S. pombe-derived extract that is active for tRNA transcription. We took advantage of the extraordinary conservation of the tRNA gene promoter and the ability of the pol III machinery from different species to initiate transcription on it. Because sequence context can affect termination and, in some cases, associated activities of apparent regulatory significance, we also examined different pols III on the same template (17Dieci G. Sentenac A. Cell. 1996; 84: 245-252Abstract Full Text Full Text PDF PubMed Scopus (147) Google Scholar, 18Fan H. Sakulich A.L. Goodier J.L. Zhang X. Qin J. Maraia R.J. Cell. 1997; 88: 707-715Abstract Full Text Full Text PDF PubMed Scopus (94) Google Scholar, 19Goodier J.L. Maraia R.J. J. Biol. Chem. 1998; 273: 26110-26116Abstract Full Text Full Text PDF PubMed Scopus (25) Google Scholar, 21Maraia R.J. Kenan D.J. Keene J.D. Mol. Cell. Biol. 1994; 14: 2147-2158Crossref PubMed Scopus (135) Google Scholar, 27Chedin S. Riva M. Schultz P. Sentenac A. Carles C. Genes Dev. 1998; 12: 3857-3871Crossref PubMed Scopus (150) Google Scholar, 30Chu W.M. Ballard R.E. Schmid C.W. Nucleic Acids Res. 1997; 25: 2077-2082Crossref PubMed Scopus (18) Google Scholar,33Maraia R.J. Chang D.Y. Wolffe A.P. Vorce R.L. Hsu K. Mol. Cell. Biol. 1992; 12: 1500-1506Crossref PubMed Scopus (36) Google Scholar, 34Gunnery S. Ma Y. Mathews M.B. J. Mol. Biol. 1999; 286: 745-757Crossref PubMed Scopus (48) Google Scholar). By this approach, a clear difference between S. cerevisiae and human pol III termination signal recognition was demonstrated with S. pombe pol III termination at an intermediate dT(n) length. Another difference supported the idea that S. pombe pol III termination may be mechanistically more similar to human than is S. cerevisiae. Finally, we examined transcription in the presence of α-amanitin, a small toxin that is known to inhibit certain eukaryotic RNA polymerases. This revealed a correlation between the response to a minimal dT(n) signal and sensitivity to α-amanitin by these pols III on the same template and further distinguished the enzymes. A gene H. D. V. T. Soll D. Mol. PubMed Scopus Google was from K. 1993; PubMed Scopus Google and for polymerase and a was the and of J.D. 1994; PubMed Google Scholar). The sequence the in the genes is The the were in as and to Ref. J. Munz P. Soll D. Nasim A. Young P. Johnson B.F. Molecular Biology of the Fission Yeast. Academic Press, Inc., San Diego1989: 75-96Crossref Google in the and in the each a to a the in the of the (31Kohli J. Munz P. Soll D. Nasim A. Young P. Johnson B.F. Molecular Biology of the Fission Yeast. Academic Press, Inc., San Diego1989: 75-96Crossref Google Scholar). In the these between the A and B box promoter in a that is that affect the transcription of tRNA genes H. A. Cell. 1981; 24: Full Text PDF PubMed Scopus Google Scholar, C.A.P. G.A. Geiduschek E.P. Genes Dev. 1996; PubMed Scopus Google Scholar, F.E. Nucleic Acids Res. 11: PubMed Scopus Google Scholar). The of the were with and to from as T. D. Mol. Cell. Biol. 1994; 14: PubMed Scopus Google Scholar). for were in RNA was with and on and with R. Nucleic Acids Res. 19: PubMed Scopus Google Scholar). The is complementary to of in and species. The is directed to the of (31Kohli J. Munz P. Soll D. Nasim A. Young P. Johnson B.F. Molecular Biology of the Fission Yeast. Academic Press, Inc., San Diego1989: 75-96Crossref Google Scholar, J. O. Soll D. Cell. Full Text PDF PubMed Scopus Google Scholar). The was described D.J. D. H. S.L. 1997; Google Scholar). S. pombe and S. cerevisiae extracts for in were to Nichols M. Willis I. Soll D. 1990; PubMed Scopus Google with the for S. pombe. with the was by the of 5 The and from Molecular nuclear extract was by a J.D. Roeder R.G. Nucleic Acids Res. 11: PubMed Scopus Google Scholar). of of and of was from was on a To the elements for termination, we the sequence of of of in of the terminator that readthrough transcripts were at S. M. Nucleic Acids Res. 12: PubMed Scopus Google Scholar, W.M. Mol. Cell. Biol. PubMed Scopus Google Scholar, R. M. Hall B.D. 1992; PubMed Scopus (18) Google tRNA structure appears to be a of recognition by J.A. M. J. Biol. Chem. Full Text PDF PubMed Google Scholar, V. V. R. A. S. J. Biol. Chem. 1995; Full Text Full Text PDF PubMed Scopus (37) Google Scholar), and certain downstream of an terminator can with S. M. Nucleic Acids Res. 12: PubMed Scopus Google Scholar, R. M. Hall B.D. 1992; PubMed Scopus (18) Google Scholar). To that tRNA be on accurate termination, we a sequence downstream of the terminator to with tRNA pol III was to the This sequence is to as the complementary readthrough and can with the of the to an that is recognizable as a tRNA the the the Upon accurate termination at the this gene, be to produce a of To from and we three (31Kohli J. Munz P. Soll D. Nasim A. Young P. Johnson B.F. Molecular Biology of the Fission Yeast. Academic Press, Inc., San Diego1989: 75-96Crossref Google to specific by The gene was between the and box in a of tRNA genes that is that be affect transcription (Refs. J. Munz P. Soll D. Nasim A. Young P. Johnson B.F. Molecular Biology of the Fission Yeast. Academic Press, Inc., San Diego1989: 75-96Crossref Google Scholar and H. A. Cell. 1981; 24: Full Text PDF PubMed Scopus Google Scholar, C.A.P. G.A. Geiduschek E.P. Genes Dev. 1996; PubMed Scopus Google Scholar, F.E. Nucleic Acids Res. 11: PubMed Scopus Google S. pombe the in but as is efficiently by (31Kohli J. Munz P. Soll D. Nasim A. Young P. Johnson B.F. Molecular Biology of the Fission Yeast. Academic Press, Inc., San Diego1989: 75-96Crossref Google Scholar). was with in a that at the J.D. 1994; PubMed Google Scholar). were for and the of was was in with the gene 2 in terminator This that readthrough transcription the terminator to The gene a terminator in a of and 2 The terminator supported more than the terminator but than Suppressor from the and of that have at the as by polymerase for of the that no in this Multiple can also to shown J.D. 1994; PubMed Google these the in the is the of the tract of the the that a and in termination between 3 and 4 of 5 suppressor this we that efficient termination to at 5 transcripts can be in are by The ability to an in and RNA from the same gene us to on for the of this was the terminator at 5 2 This gene suppressor with an as be shown efficient Although a of 5 efficient to a in that the of the gene was to termination efficiency and that a large in termination between and with at of RNAs from is shown in 3 A. signal in and revealed of with tRNA species with of with in to transcripts as 3 A specific for that the transcripts represented readthrough the terminator the may of the with 5 more with with The and with directed to the and of the transcript that be from the gene that the by to a synthesis was at the the between this and the tRNA are R. V. A. S. J. and R. J. Mol. in Scholar). revealed that the of tRNA in the was than in the and this represented termination efficiency The of tRNA to from with 5 more termination efficiency 3 A that 3 that the difference in in the and was We also examined the of that are related to B the in 3 A and with an complementary to an of three as well as the same for revealed that to the of by in these and that it to a similar to than the from This that the of is to a in but is more with a in the specific of the as a The presence of a termination at the terminator was from 3 more dTs but clear in as in 3 D. This evidence that a small of termination occurs at a terminator but a We that efficient termination occurs at 5 in S. pombe and that is a reporter of pol III termination in termination is efficient and to extracts pombe that were active for tRNA transcription we that be by the of the extracts that were active for tRNA transcription. The that the transcripts were by pol III as synthesis the presence of transcript on the of the tract transcript was by an of pol III and α-amanitin at an intermediate efficiency was at of and and was found to be at and similar to in pol III the To compare termination on the same template and for sequence transcription complexes were on gene and S. pombe extracts as the of TFs and pol III. We examined genes that in the the tract of which is by the in In this transcripts that are at are to the pol III at 4 with a but of transcription to 4 pombe pol III at 5 with a but of the transcript 4 cerevisiae pol III at 5 but with lower efficiency than S. pombe at 5 and The in the of the are the for termination at the that the species in the in 4 4 as transcripts that are on termination at similar to was in Although we have the of these species 4 to the transcript and the on 3 This that this extract is for as are S. M. Willis I. Soll D. 1990; PubMed Scopus Google Scholar). The of in the the of the human on in Ref. H. Goodier J.L. J. D.R. Maraia R.J. Mol. Cell. Biol. 1998; 18: PubMed Scopus Google Scholar). of the and was and termination efficiency was to the This revealed that termination by S. cerevisiae pol III was lower at 6 than was S. pombe at 5 4 pombe pols III with efficiency at 6 this of efficiency was by S. cerevisiae pol III it We that human pol III is the most to termination at a minimal dT(n) tract cerevisiae was the and S. pombe was Another of the termination pattern was apparent in these Although human and S. pombe pol III 4 and 5 for efficient termination, exhibit a distinct as revealed by a in transcripts at the A and The is as for S. cerevisiae pol as the of transcripts more in 4 of the of that appears in 5 is in as appears in the was more for S. pombe This is with the idea that termination by S. cerevisiae pol III a dT(n) tract may be a more of a than occurs in the pol III complexes E.P. Tocchini-Valentini G.P. Annu. Rev. Biochem. 1988; 57: 873-914Crossref PubMed Scopus (446) Google Scholar). A gene a terminator was to the sensitivity to α-amanitin of the three pols III pol III was the most to S. pombe was and S. cerevisiae was revealed that S. pombe pol III was by α-amanitin 5 as for the S. pombe RNA gene D. 1999; 8: Google that the here may reveal an accurate for the human S. pombe and human pols III are to α-amanitin and S. cerevisiae is by this toxin that these P. Proc. Natl. Acad. Sci. U. S. A. PubMed Scopus Google Scholar). We have developed a tRNA suppressor gene function in S. pombe is on accurate and efficient termination by pol and we it to termination in and in We described for the first extract from S. pombe that is active for tRNA transcription, the results and that pol III termination occurs efficiently at 5 in S. pombe. on the same demonstrated that S. cerevisiae pol III requires a tract than human S. pombe pols III for termination and supported the idea that this enzyme terminates more than S. pombe pols III in a tract E.P. Tocchini-Valentini G.P. Annu. Rev. Biochem. 1988; 57: 873-914Crossref PubMed Scopus (446) Google Scholar). the of this in extract was a that described transcription of RNA in a extract of S. D. 1999; 8: Google Scholar). The that RNA synthesis was to α-amanitin, and that the enzyme responsible was pol III tRNA and 5 S rRNA in the same tRNA transcription in was it is the S. pombe extract be active for tRNA transcription. In this it is to that an upstream of the S. pombe RNA gene, was transcribed efficiently in extract but in the S. pombe extract D. 1999; 8: Google Scholar). This suggests that the internal promoter was recognized by factors in extract and that a comparable was in the S. pombe results that is a in tRNA transcription from S. pombe A that we to from with is that sensitivity to this clear evidence of the distinct of the pols III examined in eukaryotic RNA that to α-amanitin have been to a conserved of the subunits of the RNA from and in Ref. 28Thuillier V. Brun I. Sentenac A. Werner M. EMBO J. 1996; 15: 618-629Crossref PubMed Scopus (37) Google Scholar). A of this was by in the RNA polymerase that termination X. C.W. J. Biol. Chem. 1995; Full Text Full Text PDF PubMed Scopus Google Scholar, R. C. G. R. Genes Dev. 1994; 8: PubMed Scopus Google Scholar, K. D. E. V. R. S.A. A. J. Biol. Chem. 1995; Full Text Full Text PDF PubMed Scopus Google Scholar, D.J. Proc. Natl. Acad. Sci. U. S. A. 1992; 89: PubMed Scopus Google Scholar). is that the known of α-amanitin is of RNA by RNA pol an that is to multisubunit RNA Cell. 1994; Full Text PDF PubMed Scopus Google Scholar, D.R. H. J. Biol. Chem. 1995; Full Text Full Text PDF PubMed Scopus Google Scholar). to a between termination signal recognition and α-amanitin here is that RNA is for pol III termination S. Riva M. Schultz P. Sentenac A. Carles C. Genes Dev. 1998; 12: 3857-3871Crossref PubMed Scopus (150) Google Scholar). it α-amanitin inhibit RNA and/or termination by pol III. We that was in from a that in S. cerevisiae. In with that results that 5 signal termination by S. cerevisiae pol III in Hall B.D. EMBO J. PubMed Scopus Google Scholar). in termination was with than 6 in that more dTs than pombe a comparable suppressor can be a to gene and in this of the transcription may be associated with to factors D.J. Proc. Natl. Acad. Sci. U. S. A. 1992; 89: PubMed Scopus Google Scholar and references therein). The La, which the terminus of pol III transcripts, may be a of as human been shown to regulate pol III recycling and associated activities in a manner in an in H. Sakulich A.L. Goodier J.L. Zhang X. Qin J. Maraia R.J. Cell. 1997; 88: 707-715Abstract Full Text Full Text PDF PubMed Scopus (94) Google Scholar, 19Goodier J.L. Maraia R.J. J. Biol. Chem. 1998; 273: 26110-26116Abstract Full Text Full Text PDF PubMed Scopus (25) Google Scholar, 21Maraia R.J. Kenan D.J. Keene J.D. Mol. Cell. Biol. 1994; 14: 2147-2158Crossref PubMed Scopus (135) Google Scholar). The TFIIIC subunits some of which bind at near the may also a in termination, as a of human and associated factors have been to affect termination and recycling by pol III Z. Roeder R.G. Mol. Cell. 1998; 1: 749-757Abstract Full Text Full Text PDF PubMed Scopus (67) Google Scholar). of also that pol III termination is a (17Dieci G. Sentenac A. Cell. 1996; 84: 245-252Abstract Full Text Full Text PDF PubMed Scopus (147) Google Scholar, 26Shaaban S.A. Bobkova E.V. Chudzik D.M. Hall B.D. Mol. Cell. Biol. 1996; 16: 6468-6476Crossref PubMed Scopus (26) Google Scholar, 27Chedin S. Riva M. Schultz P. Sentenac A. Carles C. Genes Dev. 1998; 12: 3857-3871Crossref PubMed Scopus (150) Google Scholar, 28Thuillier V. Brun I. Sentenac A. Werner M. EMBO J. 1996; 15: 618-629Crossref PubMed Scopus (37) Google E.V. Hall B.D. J. Biol. Chem. 1997; Full Text Full Text PDF PubMed Scopus (26) Google Scholar). The described here can be in with systems to further these of pol III termination in and We I. Willis and of H. for and D. Y. and an for
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