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RNA polymerase-2
Function:
- RNA polymerase 2 transcribes genes into mRNA at ~ 50 bases/sec
- synthesis of mRNA, snRNAs U1-U5 & U7
- at the start of transcription, a single stranded DNA template strand of the promoter is positioned within the central active site cleft of RNA polymerase 2
- a bridging helix emanates from POLR2A & crosses the cleft near the catalytic site & is thought to promote translocation of RNA polymerase 2 by acting as a ratchet that moves the RNA-DNA hybrid through the active site by switching from straight to bent conformations at each step of nucleotide addition
- during transcription elongation, RNA polymerase 2 moves on the template as the transcript elongates
- elongation is influenced by the phosphorylation status of the C-terminal domain (CTD) of POLR2A which serves as a platform for assembly of factors that regulate transcription initiation, elongation, termination & mRNA processing
- binding of ribonucleoside triphosphate to the RNA polymerase 2 transcribing complex probably involves a two- step mechanism
- initial binding seems to occur at the entry (E) site & involves a Mg+2 temporarily coordinated by 3 conserved Asp of the two largest RNA polymerase 2 subunits (POLR2A & POLR2B)
- the ribonucleoside triphosphate is transferred by a rotation to the nucelotide addition (A) site for pairing with the template DNA
- the catalytic A site involves 3 conserved Asp of POLR2A permanently coordinating a 2nd Mg+2
- RNA polymerase 2 interacts with accessory proteins that modulate transcription including:
a) elongation factor ELL2
b) transcription elongation factors necessary for efficient RNA polymerase 2 transcription elongation past template- encoded arresting sites
- holoenzyme contains RNA polymerase 2 elongator complex
Structure:
- 12 subunits
- yeast RNA polymerase 2 complex consists of 12 subunits all of which have been cloned
- in mammalian systems many subunit genes have not yet been cloned
- 2 largest subunits comprise at least part of the catalytic site, but the functions of other subunits are not yet clear
- consist of core element with a central large cleft, a clamp element that moves to open & close the cleft & jaws that are thought to grab the incoming DNA template
- the RNA polymerase 2 core element consists of POLR2A & POLR2B (2 largest subunits)
Interactions
molecular events
Related
RNA polymerase 2 elongator complex
RNA polymerase II elongation factor ELL2 (ELL2)
RNA polymerase II elongation factor ELL; eleven-nineteen lysine-rich leukemia protein (ELL, C19orf17)
transcription
General
phosphoprotein
regulatory molecule
RNA polymerase
Properties
SUBUNITS: RNA polymerase-2 L'
MOTIF: Zn finger C2H2-type
SITE: 71-87
EFFECTOR-BOUND: Zn+2
MOTIF: Zn+2-binding site
SITE: 71-71
Zn+2-binding site
SITE: 74-74
Zn+2-binding site
SITE: 81-81
Zn+2-binding site
SITE: 84-84
Zn+2-binding site
SITE: 111-111
Zn+2-binding site
SITE: 114-114
Tyr phosphorylation site {Y145}
Zn+2-binding site
SITE: 154-154
Zn+2-binding site
SITE: 184-184
Mg+2-binding site
SITE: 495-495
Mg+2-binding site
SITE: 497-497
Mg+2-binding site
SITE: 499-499
Bridging helix {833-845}
repeat {1593-1960}
MOTIF: consensus repeat {1593-1960}
SEQUENCE-IS: Y-[ST]-P-[STQ]-[ST]-P-[SRTEVKGN]
Ser phosphorylation site {S1878}
Tyr phosphorylation site {Y1923}
Ser phosphorylation site {S1927}
Thr phosphorylation site {T1933}
Ser phosphorylation site {S1934}
RNA polymerase-2 L
MOTIF: Mg+2-binding site
SITE: 792-792
Zn finger C4-type
SITE: 1119-1140
EFFECTOR-BOUND: Zn+2
MOTIF: Zn+2-binding site
SITE: 1119-1119
Zn+2-binding site
SITE: 1122-1122
Zn+2-binding site
SITE: 1137-1137
Zn+2-binding site
SITE: 1140-1140
RNA polymerase-2 p33
MOTIF: cysteine-rich region {88-97}
RNA polymerases 1, 2, & 3 subunit RPABC1
RNA polymerase-2 p19
RNA polymerase-2 p16
RNA polymerase 2 subunit RPB9
MOTIF: Zn finger C4-type
SITE: 17-42
EFFECTOR-BOUND: Zn+2
MOTIF: Zn+2-binding site
SITE: 17-17
Zn+2-binding site
SITE: 20-20
Zn+2-binding site
SITE: 39-39
Zn+2-binding site
SITE: 42-42
Ser phosphorylation site {S73}
Zinc finger
NAME: Zinc finger
SITE: 82-124
EFFECTOR-BOUND: Zn+2
MOTIF: Zn+2-binding site
SITE: 86-86
Zn+2-binding site
SITE: 89-89
Zn+2-binding site
SITE: 114-114
Zn+2-binding site
SITE: 119-119
RNA polymerases 1, 2, & 3 subunit RPABC2
MOTIF: Ser phosphorylation site {S2}
RNA polymerase-2 p13.3
DNA-directed RNA polymerases 1, 2, & 3 subunit RPABC5
MOTIF: Zn+2-binding site
SITE: 7-7
Zn+2-binding site
SITE: 10-10
Zn+2-binding site
SITE: 44-44
Zn+2-binding site
SITE: 45-45
RNA polymerase 1, 2, 3 17.1 kD subunit (RPABC3, RPB17, RPB8, hRPB8, POLR2H)
RNA polymerase I, 2, 2 7.0 kD subunit
MOTIF: Zn finger C4-type
SITE: 19-39
EFFECTOR-BOUND: Zn+2
RNA polymerase 2-associated protein 1
COMPARTMENT: cell nucleus
MOTIF: Ser phosphorylation site {S603}
Thr phosphorylation site {T1116}
References
- UI89295557
- Buratowski S.
The basics of basal transcription by RNA polymerase II.
Cell. 1994 Apr 8;77(1):1-3. Review.
PMID: 8156586
Component-of
Tat-SF complex
Components
DNA-directed RNA polymerases 1, 2, & 3 subunit RPABC5 (RNA polymerase 2 p7.6 kD subunit, RPB10, RPB7.6, RNA polymerase 2 subunit L, POLR2L, RPABC5)
RNA polymerase 1, 2, 3 17.1 kD subunit (RPABC3, RPB17, RPB8, hRPB8, POLR2H)
RNA polymerase 2 13.3 kD subunit; RPB11; RPB11a; RNA polymerase 2 subunit J (POLR2J, POLR2J1)
RNA polymerase 2 140 kD subunit (RPB2, RNA polymerase 2 subunit B, RNA polymerase 2 subunit 2, POLR2B)
RNA polymerase 2 33 kD subunit (RPB33, RPB31, RNA polymerase 2 subunit C, POLR2C)
RNA polymerase 2 largest subunit; RPB1; RNA polymerase 2 subunit A; RNA polymerase 2 subunit B1 (POLR2A)
RNA polymerase 2 p16 kD subunit (RPB4, RNA polymerase 2 subunit D, POLR2D)
RNA polymerase 2 p19 kD subunit (RPB7, RNA polymerase 2 subunit G, POLR2G)
RNA polymerase 2 subunit RPB9 (RNA polymerase II p14.5 kD subunit, RPB14.5, RNA polymerase 2 subunit I, POLR2I, RPB9)
RNA polymerase 2-associated protein 1 (RPAP1, KIAA1403)
RNA polymerase I, 2, 2 7.0 kD subunit
RNA polymerase II elongation factor ELL3 (ELL3)
RNA polymerase II subunit A C-terminal domain phosphatase; TFIIF-associating CTD phosphatase (CTDP1, FCP1)
RNA polymerases 1, 2, & 3 subunit RPABC1 (RNA polymerases 1, 2, & 3 subunit ABC1, RNA polymerase 2 23 kD subunit, RNA polymerase 2 subunit E, RNA polymerase 2 subunit 5, POLR2E, RPABC1, RPB25, RBP5, XAP4)
RNA polymerases 1, 2, & 3 subunit RPABC2 (RPB6, RNA polymerase 2 subunit F, POLR2F, POLRF, ABC2, RPABC14.4, RPB14.4, RPC15)
TBP-associated factor 2N; RNA-binding protein 56; RBP56; TAF15 RNA polymerase 2; TATA box-binding protein-associated factor 68-kd (TAF2N)