RNAポリメラーゼIII
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- an enzyme that catalyzes the formation of new DNA and RNA from an existing strand of DNA or RNA
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出典(authority):フリー百科事典『ウィキペディア（Wikipedia）』「2015/06/12 07:48:51」(JST)
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In eukaryote cells, RNA polymerase III (also called Pol III) transcribes DNA to synthesize ribosomal 5S rRNA, tRNA and other small RNAs. This enzyme complex has a more limited role than the Pol III in prokaryote cells.
The genes transcribed by RNA Pol III fall in the category of "housekeeping" genes whose expression is required in all cell types and most environmental conditions. Therefore the regulation of Pol III transcription is primarily tied to the regulation of cell growth and the cell cycle, thus requiring fewer regulatory proteins than RNA polymerase II. Under stress conditions however, the protein Maf1 represses Pol III activity.[1]
In the process of transcription (by any polymerase) there are three main stages:
- Initiation; requiring construction of the RNA polymerase complex on the gene's promoter.
- Elongation; the synthesis of the RNA transcript.
- Termination; the finishing of RNA transcription and disassembly of the RNA polymerase complex.
Contents
- 1 Initiation
- 1.1 Class I
- 1.2 Class II
- 1.3 Class III
- 2 Termination
- 3 Transcribed RNAs
- 4 See also
- 5 References
Initiation
Initiation: the construction of the polymerase complex on the promoter. Pol III is unusual (compared to Pol II) requiring no control sequences upstream of the gene, instead normally relying on internal control sequences - sequences within the transcribed section of the gene (although upstream sequences are occasionally seen, e.g. U6 snRNA gene has an upstream TATA box as seen in Pol II Promoters).
Class I
Typical stages in 5S rRNA (also termed class I) gene initiation:
- TFIIIA (Transcription Factor for polymerase III A) binds to the intragenic (lying within the transcribed DNA sequence) 5S rRNA control sequence, the C Block (also termed box C).
- TFIIIA Serves as a platform that replaces the A and B Blocks for positioning TFIIIC in an orientation with respect to the start site of transcription that is equivalent to what is observed for tRNA genes.
- Once TFIIIC is bound to the TFIIIA-DNA complex the assembly of TFIIIB proceeds as described for tRNA transcription.
Class II
Typical stages in a tRNA (also termed class II) gene initiation:
- TFIIIC (Transcription Factor for polymerase III C) binds to two intragenic (lying within the transcribed DNA sequence) control sequences, the A and B Blocks (also termed box A and box B).[2]
- TFIIIC acts as an assembly factor that positions TFIIIB to bind to DNA at a site centered approximately 26 base pairs upstream of the start site of transcription. TFIIIB (Transcription Factor for polymerase III B), consists of three subunits: TBP (TATA Binding Protein), the Pol II transcription factor TFIIB-related protein, BRF1 (or Brf2 for transcription of a subset of Pol III-transcribed genes in vertebrates) and BDP1.[3]
- TFIIIB is the transcription factor that assembles Pol III at the start site of transcription. Once TFIIIB is bound to DNA, TFIIIC is no longer required. TFIIIB also plays an essential role in promoter opening.
TFIIIB remains bound to DNA following initiation of transcription by Pol III (unlike bacterial σ factors and most of the basal transcription factors for Pol II transcription). This leads to a high rate of transcriptional reinitiation of Pol III-transcribed genes.
Class III
Typical stages in a U6 snRNA (also termed class III) gene initiation (documented in vertebrates only):
- SNAPc (SNRNA Activating Protein complex) (also termed PBP and PTF) binds to the PSE (Proximal Sequence Element) centered approximately 55 base pairs upstream of the start site of transcription. This assembly is greatly stimulated by the Pol II transcription factors Oct1 and STAF that bind to an enhancer-like DSE (Distal Sequence Element) at least 200 base pairs upstream of the start site of transcription. These factors and promoter elements are shared between Pol II and Pol III transcription of snRNA genes.
- SNAPc acts to assemble TFIIIB at a TATA box centered 26 base pairs upstream of the start site of transcription. It is the presence of a TATA box that specifies that the snRNA gene is transcribed by Pol III rather than Pol II.
- The TFIIIB for U6 snRNA transcription contains a smaller Brf1 paralogue, Brf2.
- TFIIIB is the transcription factor that assembles Pol III at the start site of transcription. Sequence conservation predicts that TFIIIB containing Brf2 also plays a role in promoter opening.
Termination
Polymerase III terminates transcription at small polyTs stretch (5-6). In Eukaryotes, a hairpin loop is not required, as it is in prokaryotes.[citation needed]
Transcribed RNAs
The types of RNAs transcribed from RNA polymerase III includes:
- Transfer RNAs[4]
- 5S ribosomal RNA[4]
- U6 spliceosomal RNA[4]
- RNase P and RNase MRP RNA[4]
- 7SL RNA (the RNA component of the signal recognition particle)[4]
- Vault RNAs[4]
- Y RNA[4]
- SINEs (short interspersed repetitive elements)[4]
- 7SK RNA[4]
- Several microRNAs[4]
- Several small nucleolar RNAs[4]
- Several gene regulatory antisense RNAs[5]
See also
- DNA polymerase III holoenzyme
References
- ^ Vannini, A.; Ringel, R.; Kusser, A. G.; Berninghausen, O.; Kassavetis, G. A.; Cramer, P. (2010). "Molecular Basis of RNA Polymerase III Transcription Repression by Maf1". Cell 143 (1): 59–70. doi:10.1016/j.cell.2010.09.002. PMID 20887893. edit
- ^ Oettel S, Kober I, Seifart KH (October 1998). "The activity binding to the termination region of several pol III genes represents a separate entity and is distinct from a novel component enhancing U6 snRNA transcription.". Nucleic Acids Research 26 (19): 4324–31. doi:10.1093/nar/26.19.4324. PMC 147850. PMID 9742231.
- ^ Ishiguro A, Kassavetis GA, Geiduschek EP (May 2002). "Essential roles of Bdp1, a subunit of RNA polymerase III initiation factor TFIIIB, in transcription and tRNA processing.". Molecular and cellular biology 26 (10): 4324. doi:10.1128/MCB.22.10.3264-3275.2002. PMC 133792. PMID 11971960.
- ^ a b c d e f g h i j k Dieci G, Fiorino G, Castelnuovo M, Teichmann M, Pagano A (December 2007). "The expanding RNA polymerase III transcriptome". Trends Genet. 23 (12): 614–22. doi:10.1016/j.tig.2007.09.001. PMID 17977614.
- ^ Pagano A, Castelnuovo M, Tortelli F, Ferrari R, Dieci G, Cancedda R (February 2007). "New small nuclear RNA gene-like transcriptional units as sources of regulatory transcripts". PLoS Genet. 3 (2): e1. doi:10.1371/journal.pgen.0030001. PMC 1790723. PMID 17274687.
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Transferases: phosphorus-containing groups (EC 2.7)
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2.7.1-2.7.4:
phosphotransferase/kinase
(PO4) |
| 2.7.1: OH acceptor |
- Hexo-
- Gluco-
- Fructo-
- Galacto-
- Phosphofructo-
- 1
- Liver
- Muscle
- Platelet
- 2
- Riboflavin
- Shikimate
- Thymidine
- NAD+
- Glycerol
- Pantothenate
- Mevalonate
- Pyruvate
- Deoxycytidine
- PFP
- Diacylglycerol
- Phosphoinositide 3
- Class I PI 3
- Class II PI 3
- Sphingosine
- Glucose-1,6-bisphosphate synthase
|
|
| 2.7.2: COOH acceptor |
- Phosphoglycerate
- Aspartate kinase
|
|
| 2.7.3: N acceptor |
|
|
| 2.7.4: PO4 acceptor |
- Phosphomevalonate
- Adenylate
- Nucleoside-diphosphate
- Uridylate
- Guanylate
- Thiamine-diphosphate
|
|
|
2.7.6: diphosphotransferase
(P2O7) |
- Ribose-phosphate diphosphokinase
- Thiamine diphosphokinase
|
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2.7.7: nucleotidyltransferase
(PO4-nucleoside) |
| Polymerase |
| DNA polymerase |
- DNA-directed DNA polymerase
- I
- II
- III
- IV
- V
- RNA-directed DNA polymerase
- Reverse transcriptase
- Telomerase
- DNA nucleotidylexotransferase/Terminal deoxynucleotidyl transferase
|
|
| RNA nucleotidyltransferase |
- RNA polymerase/DNA-directed RNA polymerase
- RNA polymerase I
- RNA polymerase II
- RNA polymerase III
- RNA polymerase IV
- Primase
- RNA-dependent RNA polymerase
- PNPase
|
|
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Phosphorolytic
3' to 5' exoribonuclease |
|
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| Nucleotidyltransferase |
- UTP—glucose-1-phosphate uridylyltransferase
- Galactose—1-phosphate uridylyltransferase
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| Guanylyltransferase |
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| Other |
- Recombinase (Integrase)
- Transposase
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|
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| 2.7.8: miscellaneous |
| Phosphatidyltransferases |
- CDP-diacylglycerol—glycerol-3-phosphate 3-phosphatidyltransferase
- CDP-diacylglycerol—serine O-phosphatidyltransferase
- CDP-diacylglycerol—inositol 3-phosphatidyltransferase
- CDP-diacylglycerol—choline O-phosphatidyltransferase
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| Glycosyl-1-phosphotransferase |
- N-acetylglucosamine-1-phosphate transferase
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2.7.10-2.7.13: protein kinase
(PO4; protein acceptor) |
| 2.7.10: protein-tyrosine |
|
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| 2.7.11: protein-serine/threonine |
- see serine/threonine-specific protein kinases
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| 2.7.12: protein-dual-specificity |
- see serine/threonine-specific protein kinases
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| 2.7.13: protein-histidine |
- Protein-histidine pros-kinase
- Protein-histidine tele-kinase
- Histidine kinase
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- Biochemistry overview
- Enzymes overview
- By EC number: 1.1
- 2
- 3
- 4
- 5
- 6
- 7
- 8
- 10
- 11
- 13
- 14
- 15-18
- 2.1
- 3.1
- 4.1
- 5.1
- 6.1-3
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UpToDate Contents
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English Journal
- Separate and concurrent use of 2-deoxy-D-glucose and 3-bromopyruvate in pancreatic cancer cells.
- Xiao H, Li S, Zhang D, Liu T, Yu M, Wang F.SourceDepartment of Colorectal Surgery, China-Japan Union Hospital of Jilin University, Changchun, Jilin 130033, P.R. China.
- Oncology reports.Oncol Rep.2013 Jan;29(1):329-34. doi: 10.3892/or.2012.2085. Epub 2012 Oct 17.
- Unrestrained glycolysis characterizes energy meta-bolism in cancer cells. Thus, antiglycolytic reagents such as 2-deoxy-D-glucose (2-DG) and 3-bromopyruvate (3-BrPA) may be used as anticancer drugs. In the present study, we examined the anticancer effects of 2-DG
- PMID 23076497
Japanese Journal
- アデノウイルスベクターの最近の進展：VA欠失ベクターを中心に
- 近藤 小貴,前川 文,斎藤 泉,鐘ヶ江 裕美
- ウイルス 63(2), 155-164, 2013
- … えた時，残存しているアデノウイルスゲノム領域からPol IIIにより発現している2種類のウイルス随伴RNA (VA RNA)の欠失が必要である．VA RNAはウイルス増殖に必須では無いものの，ウイルス増殖に適した環境を整備する役割を担っており，特に感染後期に大量に発現しているため，VA RNAをトランスに供給する293細胞ではVA欠失AdV作製は困難であ …
- NAID 130004713421
- 新規POLR3A遺伝子変異をみとめた大脳白質形成不全症の1例
- 田村 麻子,丹羽 篤,伊井 裕一郎,佐々木 良元,冨本 秀和,才津 浩智
- 臨床神経学 53(8), 624-629, 2013
- 症例は34歳男性である．兄に類症あり．小学校高学年頃に学力低下で発症し，2次性徴の発現がなく，類宦官体型，軽度小脳失調や強度の近視をみとめたが，歯牙低形成はみられなかった．血液検査上，ゴナドトロピン（LH，FSH）およびテストステロンの低値を，MRIでミエリン低形成，小脳・脳幹萎縮，脳梁低形成を，SPECTで小脳の集積低下をみとめ，遺伝子解析の結果，POLR3A遺伝子の新規c.2350G>A …
- NAID 130004505538
- Monitoring Genetic Diversity of Influenza A(H1N1)pdm09 Virus Circulating during the Post-Pandemic Period in Turkey
- Guldemir Dilek,Kalaycioglu Atila T.,Altas A. Basak [他],Korukluoglu Gulay,Durmaz Riza
- Japanese Journal of Infectious Diseases 66(4), 299-305, 2013
- … A total of 2601 clinical specimens obtained from suspected cases of influenza A(H1N1)pdm09 viral infections were analyzed by real-time reverse transcription polymerase chain reaction. … Viral RNA was detected in 233 (9%) clinical specimens. … On the basis of amino acid substitutions in the HA1 domain, majority of the Turkish isolates were classified in the genetic group v and others in the genetic groups ii, iii, and vi. …
- NAID 130003381692
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