Transcrip)on Regula)on And Gene Expression in Eukaryotes Cycle G2 (lecture 13709) FS 2014 P Ma?hias & RG Clerc
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1 Transcrip)on Regula)on And Gene Expression in Eukaryotes Cycle G2 (lecture 13709) FS 2014 P Ma?hias & RG Clerc P. Ma?hias, March 5th, 2014 The Basics of Transcrip-on (2) General Transcrip-on Factors: TBP/TFIID TBP Associated Factors: TAFs (Elonga-on of Transcrip-on) h?p://
2 GTFs: recognize the core promoter Factors needed for expression of nearly all Class II promoters Biochemical Fractionation (R. Roeder, Rockefeller University) led to identification of 6 fractions that contain proteins that permit correct transcription initiation by RNA pol II These fractions were labeled TF II: A, B, D, E, F, H. (Fraction C was an artefact.) Of these TFIID, TFIIA and TFIIB bind specific proximal elements (others such as TFIIH bind DNA but are not thought to have strict DNA sequence preference) TFIID nucleates the assembly of the pre-initation complex (PIC)
3 Eukaryotic Pol II promoter: upstream and core NF-kB Fos Oct-1 CBFA-1 CREB etc Transcription activators TATA INR RNAPII! TFIIH! TFIIF! TFIIE! TFIIA! TFIIB! TFIID! GENE upstream (regulatory, enhancer) core Basal machinery RNA PolII 12 TFIIA 3 TFIIB 1 TFIID (TBP) ca. 14 (1) TFIIE 2 TFIIF 2 TFIIH ca. 9
4 The core promoters in more detail BRE TATA box Initiator DPE G/CG/CG/CCGCC TATAAA PyPyANT/APyPy GA/TCGTG TFIIB TFIID TFIID TFIID TFIIA TBP TAF II 250 TAF II 150 TAF II 60 TAF II 40 TATA-containing TATA-less
5 TFIID TFIID is a mul-protein complex: TBP + TAFs (~10-14 different subunits) The subunit responsible for recognizing the TATA box is called TATA- Binding Protein ( TBP ) TBP can specifically bind to TATA- boxes on its own In vitro TBP is sufficient for basal transcrip-on, not for ac-vated transcrip-on
6 Akhtar and Veenstra Cell & Bioscience 2011, 1:23 h?p://
7 TBP: a GTF essential for all polymerases
8 TAFIIs of TFIID: mostly conserved proteins ranging from 30 à 250 kda
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10 TBP-related factors
11 Akhtar and Veenstra Cell & Bioscience 2011, 1:23 h?p://
12 Akhtar and Veenstra Cell & Bioscience 2011, 1:23 h?p://
13 TBP-related factors
14 TBP-related factors specific complexes
15 Promoters with TBP and TRFs
16 Goodrich & Tjian, NRG Aug 2010
17 Albright and Tjian (2000). TAFs revisited: more data reveal new twists and confirm old ideas. Gene 242, 1-13.
18 Beyond TFIID: Multiple TAF complexes Albright and Tjian (2000). TAFs revisited: more data reveal new twists and confirm old ideas. Gene 242, 1-13.
19 Beyond TFIID: Multiple TAF complexes
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21 Summary IIB! IIA! TAFs! TBP! TFIID! }! Recognize core promoter! IIF! Targets Pol II! to promoter! Modulates helicase! IIE! Helicase! IIH! helicase! CTD protein kinase! protein kinase! IIE! IIB! IIA! Pol IIa! TBP! Inr! IIH! CTD of large subunit of Pol II! IIF!
22 Cell and stage- specific dynamics in basal factors D Alessio et al., Molecular Cell 36, December 24, 2009
23 Goodrich & Tjian, NRG Aug 2010
24 Goodrich & Tjian, NRG Aug 2010
25 Gene bookmarking during mitosis Goodrich & Tjian, NRG Aug 2010
26 TATA vs TATA- less promoters 6000 PICs localized in yeast TATA- less promoters: have TATA- like elements S Buratowski, Nature Vol 483, 2012 (News & Views) See Rhee & Pugh, Nature Vol 483, 2012
27 A short story ca years ago: What are the in vivo targets of transcrip-on ac-vators??? East coast vs West coast, Yeast vs Metazoans, In vivo vs in vitro,...
28 TBP vs TFIID: TAFs as universal coactivators?? Sp1 + TFIID TBP GTFs + In vitro txn assay
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32 Which part of the promoter is TAF responsive?
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35 Transcription Elongation In vivo rates of transcription = nucl/min. Baseline in vitro rates = nucleotides/min. frequent pausing and arrests even with chromatin-free DNA Large number of proteins and protein complexes that regulate elongation of transcription Suppress pausing Promote pausing Overcome transcription arrest Enable elongation through chromatin
36 Elongation factors for RNA Pol II Suppress pausing and/or arrest TFIIF P-TEFb (CTD kinase) S II (TF II S) CSB (Cockayne syndrome binding pro.) ELL 1/2, Elongins certain transcription activators (e.g. heat shock factor) Promotes pausing/arrest DSIF (DRB sensitivity inducing factor) NELF (binds early during RNA elongation) Enable elongation through chromatin FACT RSF Elongator (HAT associated with Pol II)
37 Elongation NMCB, August 2006
38 Elongation
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