Lecture 3 Regulation of Initiation: Met-tRNA-binding
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1 Institut für Biochemie und Molekulare Medizin Lecture 3 Regulation of Initiation: Met-tRNA-binding Michael Altmann FS 2010
2 Model of initiation eif4g eif4e AAA AAA PABP cap AAA AUG mrna eif4a eif4b ATP eif2-gtp Met-tRNA i eif1 eif5 eif3 eif1-eif2- eif3-eif5- Met-tRNA i TC 40S GTP 80S eif2-eif2b- 60S AAA ATP Elongation eif2b eif5b AAA P
3 Eukaryotic initiation complex TC (ternary complex) 43S-complex 40S + eif2-gtp-initiator trna met + eif3 + eif1 48S-complex 43S + mrna + eif4e/g/h/a/b + PAB (polya)-binding protein 80S-mRNA-initiator trna met complex
4 G-Proteine G-Proteine als molekulare Schalter - G-Proteine sind oft in Signaltransduktionsvorgängen involviert. Es gibt verschiedene Typen von G-Proteinen (heterotrimere Proteine, kleine G-Proteine, etc.). - G-Proteine sind in aktiver Form mit GTP, in inaktiver Form mit beladen. Mit Hilfe eines GTPase-aktivierenden Enzyms (GAP) wird GTP zu und P i hydrolysiert. Zur Wiederaktivierung des G-Proteins wird mit Hilfe eines Guaninnukleotid-austauschenden Proteins (GNRP) gegen GTP ausgetauscht. -> GAPs sind negative Effektoren, GNRPs sind positive Effektoren der G-Protein-Aktivität.
5 Regulation of Met-tRNA i -binding eif2 eif2b 3 subunits: α (becomes phosphorylated), β and χ (binds /GTP) 5 subunits GNRP / GEF (guanine nucleotide exchange factor) of eif2 eif2 kinases GCN2 PKR HRI PERK Unfolded protein response ER stress
6 eif2b Factor: Reactions: Reference: CG Proud Sem. Cell&Dev. Biol. 16: 3-12, 2004
7 exchange reaction * α γ β eif2 α γ β * 100 α γ β * α γ β * minus eif2b α γ β * α γ β eif2b * 50 plus eif2b Filter α γ β * Filter α γ β t (min) *
8 Regulation of eif2b activity (Generation of stable inactive eif2- eif2b complex) (Phosphorylation of α- subunit at ser 51) (Muations leading to degeneration of central nervous system) Reference: GD Pavitt Biochem. Soc. Trans. 33: , 2005
9 Sequesteration of eif2b eif2α-p eif2 kinase eif2α-p + eif2α-p- eif2α-p- + eif2b eif2α-p--eif2b (stable!) -> due to lack of ternary complex (TC) overall translation arrests transiently
10 Reference: CG Proud Sem. Cell&Dev. Biol. 16: 3-12, eif2 kinases
11 Common mechanism of kinase activation Monomer form of kinase is inactive Ligand mediated dimerization of kinase Autophosphorylation of dimerized subunits Binding and phosphorylation of substrate(s) activates cascade Example: Insulin receptor o Upon insulin binding receptor dimerizes o Autophosphorylation of beta-subunits o Binding and phosphorylation of IRS-1 (substrate) activates cascade
12 Dual translational response to amino acid starvation activates Gcn4p 1A 1 1A 1 eif2b (GEF) 1A 1 General translation 1A 1 5 GCN2 eif2(α)~p GCN4 translation Amino acid starvation Amino acid Biosynthetic genes
13 Reinitiation on GCN4 mrna is a sensitive reporter of TC loading Non-Starved WT uorf1 uorf4 GCN4 (Adapted from Leos Valasek)
14 Reinitiation on GCN4 mrna is a sensitive reporter of TC loading Starved WT uorf1 uorf4 GCN4 ON 3-AT resistance
15 Translational regulation of GCN4, a transcription factor Non-starvation conditions Aminoacid starvation Hinnebusch, 2005
16 PKR Interferon Inactive PKR Activators dsrna Inactive PKR Active PKR Phosphorylation of eif2 Inactivators Phosphorylation of transcription factors Translation down Transcription altered Growth down Differentiation Apoptosis Antiviral state
17 PKR and apoptosis Reference: D. Scheuner et al. J.Biol.Chem. 281: , 2006.
18 Reference: M Dey et al. Cell 122: , PKR activation
19 Reference: M Rafie-Kolpin et al. Biochem. 42: , Activation of HRI
20 Reference: SJ Marciniak et al. J. Cell Biol 172: , 2006 Activation of PERK
21 The unfolded protein response Normal situation: Stress situation: Golgi (cleavage) nucleus transcription (stress genes) ATF6 Ire1 ATF6 Ire1 chaperones secreted protein BiP PERK PERK eif2alphaphosphoryl. PP1C inhibition of translation GADD34 stimulation ATF4 ATF6: precursor of transcription factor Ire1: protein kinase and endonuclease (splicing factor) PERK: eif2 kinase BiP: chaperone, folds secreted proteins, keeps ATF6 in ER, inhibits dimerization of Ire1 and PERK BiP is engaged in protein folding: ATF6, Ire1and PERK dimerize ATF6 and BiP stimulate transcription of stress genes PERK inhibits translation Feed back inhibition of the kinase
22 Reference: CM Abbott & CG Proud Trends Biochem. Sci. 29: 25-31, 2004 Translation factors and human diseases
23 Seminar 1 - What are the biol. consequences of mrna transport and localized translation? - Inform yourself about CCA adding enzyme (trna)! - How many rrna genes does a cell need to make 10 6 ribosomes in 6 hours? - What are the most prominent differences in the mechanism of initiation between prokaryotes and eukaryotes? - Why should a cell need so many helicases? - What s about translation in mitochondria? - Are there orthologs of eif s? - How are protein-protein interactions measured? - Why are 10-20% of eif2α-p sufficient to block translation? - Physiological consequences of translational regulation of transcription factors? - Viruses fight against eif2 phosphorylation! How?
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