AGS3 and AGS4 in G-protein Signaling. Joe B. Blumer, Ph.D. Cell and Molecular Pharmacology Medical University of South Carolina
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1 AGS3 and AGS4 in G-protein Signaling Joe B. Blumer, Ph.D. Cell and Molecular Pharmacology Medical University of South Carolina
2 stimulus extracellular Plasma membrane intracellular G-protein G-protein G-protein Signal regulator (Accessory protein) effectors Expanded functional roles for G-proteins
3 Activators of G-protein Signaling GROUP I (GEF) (activate G i but not G s, G 16 in yeast functional assay) AGS1 - (RASD1, DexRas1)- ras-related protein GROUP II (GDI) (activate G i but not G s, G 16 in yeast functional assay) AGS3 - (Gpsm1) four GPR motifs AGS4 - (Gpsm3, G18.1b) - three GPR motifs AGS5 - (Gpsm2, LGN, mpins) - four GPR motifs AGS6 - (RGS12) - one GPR motif GROUP III Bind G AGS2 - (tctex-1) - light chain of cytoplasmic dynein AGS7 - (Trip13) AGS8 - KIAA1866 AGS9 - (Rpn10) - proteosome component AGS10 - (Go ) Group II AGS3 AGS4 AGS5 TPR GPR
4 GDI (GPR/GoLoco motif) GPR GDP + GDP GTP GAP/ RGS P i GTP + GDP GPCR* Effectors Effectors
5 AGS3 is widely distributed in the brain and is developmentally regulated day AGS3 brainstem cerebellum cortex hippocampus midbrain OFB--olfactory bulb CB--cerebellum HP--hippocampus MED-- medulla VTA--ventral tegmental area SN--substantia nigra NA--nucleus accumbens ST--striatum PFC--prefrontal cortex CC--cerebral cortex THAL--thalamus HYPO--hypothalamus AMY--amygdala AGS3 50 cortical neuronal cultures AGS days in culture AGS3 Blumer et al., J Biol Chem :
6 Exploring the in vivo role of AGS proteins: Generation of a conditional AGS3 (Gpsm1) null mouse A Gpsm1 Targeting Vector WT C A ATG c 565bp neo r exon: exon: 2 3 d B H B kda rat brain mouse brain +/+ +/- -/- AGS3 Gpsm1 null exon: c 2 310bp d * Gi 3 bp /+ +/- -/- WT Gpsm1 null AGS5 (LGN) Blumer et al., Endocrinology (8):
7 WT Gpsm1 -/- Cerebellum Cerebral cortex Nissl stained sagittal brain sections 8 week male littermates
8 Run time (sec) Motor coordination and learning appear unaffected in AGS3 (Gpsm1) null mice Accelerating Rotarod Performance Test Day 1 Day 2 Day Rotation Speed (rpm) WT, n = 12 Gpsm1 -/-, n = 12
9 Spatial learning and memory appear unaffected in AGS3 (Gpsm1) null mice Radial arm water maze
10 Redcutions in AGS3 expression in PFC or NAc blocks reinstatement of drug-seeking behavior in rats
11 Cocaine-induced locomotor sensitization in WT vs AGS3 (Gpsm1) null mice
12 Summary: Behavioral studies of AGS3 (Gpsm1) null mice AGS3 knockout mice are viable and fertile No obvious alterations in brain cellularity or morphology Initial behavioral testing revealed no obvious differences in motor coordination or in spatial learning and working memory Locomotor sensitization to cocaine is similar to wildtype mice Phase II phenotyping: Targeted disruption of AGS3 by tissue-specific or inducible Cre expression
13 GDI (GPR/GoLoco motif) How is this regulated?? GPR GDP + GDP GTP GAP/ RGS P i GTP + GDP GPCR* Effectors Effectors
14 GTP S Binding (% of control) AGS3 (650 aa) TPR TPR TPR TPR TPR TPR TPR Gi GDP Gi GDP Gi GDP Gi GDP GPR GPR GPR GPR Gi GDP Gi GDP Gi GDP AGS4 (160 aa) GPR GPR GPR Input GST AGS4 AGS3- GPR Gi 1/2 Gi 3 AGS4 pcdna AGS4 Gi 3 Gi 3 G s G q G Input IP: Gi 3 Beads GST AGS4 AGS3-GPR log [AGS], M Cao et al., J Biol Chem. 2004; 279(26):
15 Bioluminescence Resonance Energy Transfer (BRET): A tool to measure AGS-G i interaction dynamics AGS4-RLuc Rluc + Gi 1 YFP Pfleger & Eidne, Biochem. J. (2005) 385, Gales et al., Nature Structural & Molecular Biology - 13, (2006)
16 AGS4 G i BRET signals are specific, saturable and inhibited by excess G AGS4-RLuc AGS4-Q/A-RLuc Rluc Rluc + Gi 1 YFP
17 Agonist-induced GPCR activation decreases AGS4 G i interaction + AGS4-Rluc Rluc G i1 YFP + 2A-AR AGS4-Luc G i1-yfp G i ng 2A -AR ng AGS4-Luc ng G i1-yfp
18 Agonist-induced decreases AGS4 Gi interaction are blocked by antagonist and pertussis toxin + AGS4-Rluc Rluc G i1 YFP + 2A-AR
19 G i redistributes AGS4 to the plasma membrane AGS4-GFP AGS4-GFP + G i3-wt AGS4-GFP + G i3-q204l AGS4-Q/A-GFP AGS4-Q/A-GFP + G i3-wt
20 Agonist-induced translocation of AGS4 from membrane to cytosol + AGS4-Rluc Rluc G i1 YFP + 2A-AR A B RLU x AGS4-Rluc G i1-yfp A-AR UK AGS4-Luc membrane * cytosol vehicle UK * RFU x AGS4-Rluc G i1-yfp membrane cytosol vehicle UK G i1-yfp A-AR UK14304
21 Summary: GPCR regulation of AGS4 G i BRET AGS4 G i BRET signals are decreased by G i- linked GPCR activation This effect is blocked by antagonist and pertussis toxin Receptor activation may result in either dissociation or conformational rearrangement of a pre-formed AGS4 G i complex Agonist-induced translocation of AGS4 from membrane to cytosol suggests AGS4 G i dissociation Is the receptor directly interfacing with the AGS4 G i complex?
22 AGS4 G i complexes are proximal to GPCRs and agonist-sensitive AGS4-Rluc Rluc + 2A-AR-Venus Venus
23 AGS4 interacts with G i-linked GPCRs in a G i-dependent manner
24 Summary and Perspective G i G G i??? AGS4 - The interaction of AGS4 with G i as measured by BRET is robust, saturable and specific - Receptor activation results in a decrease in AGS4 G i BRET that is blocked by antagonist and PTX - AGS4 2A -AR BRET signals are G i-dependent and reduced by receptor activation; this effect is blocked by antagonist and PTX - The data suggest that the interaction of AGS4 with G i enhances the presence of AGS4 at the cell surface whereupon receptor activation then dissociates AGS4 from G i and the cell cortex and into the cytosol - The possibility that proteins like AGS4 with multiple GPR motifs may seed G complexes that interface with receptor provides an interesting alternative mechanism for signal processing through a GPCR
25 Acknowledgements Michel Bouvier, Ph.D. Billy Breton University of Montreal Greg Tall, Ph.D. University of Rochester Lakshmi A. Devi, Ph.D. Mount Sinai Medical Center Thom Saunders, Ph.D. University of Michigan Ranney Mize, Ph.D. Luis Marrero LSU Health Sciences Center Blumer Lab: Ellen Maher Melissa O Connor Ph.D. MUSC: Stephen M. Lanier, Ph.D. Sadik Oner, Ph.D. Ali Vural Heather Bainbridge, M.S. Alejandra Pacchioni, Ph.D. Peter Kalivas, Ph.D. L. Judson Chandler, Ph.D. Lawrence Middaugh, Ph.D. Heather Boger, Ph.D. Claudia Umphlet Supported by NIH R01GM (Blumer), NS24821 & DA (Stephen M. Lanier, Ph.D.)
26
27 Membrane targeting AGS4 is not sufficient to observe BRET with G i1-yfp G i1 YFP myr-ags4-rluc + + Rluc 2A-AR G i1-n149i YFP
28 Membrane targeting AGS4 is not sufficient to observe BRET with G i1-yfp + myr-ags4-rluc Rluc G i1 YFP + 2A-AR
29 AGS4-Rluc shows reduced BRET with RGSinsensitive G i1-yfp AGS4-Rluc Rluc + G i1-wt G i1-g183s YFP YFP
30 AGS4 G i BRET signals are reduced in G i-yfp mutants but are unaltered by PTX G i1-wt YFP AGS4-Rluc Rluc + G i1-n149i YFP G i1-q204l YFP
31 AGS4 2A -AR BRET requires wild-type G i AGS4-Rluc Rluc + 2A-AR-Venus Venus G i3 G s
32 AGS4 interacts with G i-linked GPCRs in a G i-dependent manner + G i3 + G i3 + G i3 + G i3 receptor: agonist: 2-AR-Venus UK14,304 CXCR4-Venus CXCL12 MOR-YFP DAMGO 2-AR-Venus Isoproterenol
33 Agonist Regulation of the AGS4 G i complex: Dose Response & Timecourse + AGS4-Rluc Rluc G i1 YFP + 2A-AR
34 AGS4 G i complexes are proximal to GPCRs and agonist-sensitive AGS4-Rluc Rluc + 2A-AR-Venus Venus
35 Agonist-regulated AGS4 GPCR BRET signals are PTX sensitive AGS4-Rluc Rluc + 2A-AR-Venus Venus
36 Classical paradigm of G protein signaling R Unexpected Roles for G protein Signaling Accessory Proteins G G E Expanded functional roles for G-protein signaling
37 Phase I phenotyping: Behavioral Studies Brain Histology/Morphology Accelerating Rotarod Perfomance Radial Arm Water Maze Cocaine-induced Locomotor Sensitization
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