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Amino Acid Residues Implicated in the Interaction of Melanocortin Ligands and Their Receptors: a Study of MC2R Selectivity

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Amino Acid Residues Implicated in the Interaction of Melanocortin Ligands and Their Receptors: a Study of MC2R Selectivity University of Denver Digital Commons @ DU Electronic Theses and Dissertations Graduate Studies 1-1-2009 Amino Acid Residues Implicated in the Interaction of Melanocortin Ligands and Their Receptors: A Study of MC2R Selectivity Kristopher D. Veo University of Denver Follow this and additional works at: https://digitalcommons.du.edu/etd Part of the Biology Commons, and the Molecular Biology Commons Recommended Citation Veo, Kristopher D., "Amino Acid Residues Implicated in the Interaction of Melanocortin Ligands and Their Receptors: A Study of MC2R Selectivity" (2009). Electronic Theses and Dissertations. 945. https://digitalcommons.du.edu/etd/945 This Thesis is brought to you for free and open access by the Graduate Studies at Digital Commons @ DU. It has been accepted for inclusion in Electronic Theses and Dissertations by an authorized administrator of Digital Commons @ DU. For more information, please contact [email protected],[email protected]. Amino Acid Residues Implicated in the Interaction of Melanocortin Ligands and their Receptors: A Study of MC2R Selectivity __________ A Thesis Presented to The Faculty of Natural Sciences and Mathematics University of Denver __________ In Partial Fulfillment of the Requirements for the Degree Master’s of Science __________ by Kristopher D. Veo August 2009 Advisor: Dr. Robert M. Dores Author: Kristopher D. Veo Title: Amino acid residues implicated in the interaction of Melanocortin ligands and their receptors: A study of MC2R selectivity Advisor: Dr. Robert M. Dores Degree Date: August 2009 ABSTRACT Melanocortin receptor ligand selectivity has been a question not easily answered. The inability to functionally express melanocortin 2 receptor (MC2R) has inhibited the study of why MC2R is only stimulated by ACTH, a melanocortin hormone. With the recent discovery of the MC2R accessory protein (MRAP), creating a heterologous system is now feasible. Using a general cell line like CHO-K1 cells, which do not express endogenous MCRs, we were able to create a heterologous expression system and test the selectivity of MC2R using analog variants of ACTH(1-24). Our results indicate an amino acid requirement in the C-terminal portion of ACTH(1-24) for activation, which supports the 2-step method of activation hypothesized for MC2R. This site, the tetra basic cleavage site, when altered does not stimulate cAMP production and does not compete with ACTH(1-24) for binding. We also demonstrate the potential for a non-mammalian MC2R system in cloning full length Silurana tropicalis MC2R and completed localization studies with this system with MRAP using CHO-K1 cells. ii Table of Contents Introduction.......................................................................................................................1 Overview...............................................................................................................1 Melanocortin Receptor Family .............................................................................1 POMC ...................................................................................................................2 Cloning the First MCRs........................................................................................5 MCR Locations and Functions .............................................................................6 MCR Gene Evolution ...........................................................................................6 MCR Classification.............................................................................................11 MRAP .................................................................................................................15 MC2R Selectivity Issues.....................................................................................15 MC2R Activation Hypothesis.............................................................................16 Goals ...................................................................................................................20 Materials and Methods....................................................................................................22 Cloning MC2R....................................................................................................22 Human MC2R.....................................................................................................24 MC2R and MRAP Co-Localization Studies.......................................................30 Mammalian Cell Nucleofection..........................................................................33 cAMP EIA Assay................................................................................................33 Results ............................................................................................................................35 Studies using an endogenous MCR system: GT1-7 cell line and MC4R...........35 Initial MC2R endogenous system and heterologous expression system ............42 Human and Frog MC2R Localization with MRAP in CHO cells ......................45 Activation Assay using CHO cells and hMC2R+MRAP ...................................46 CHO/hMC2R+MRAP Ligand Competition Assays...........................................52 Discussion.......................................................................................................................58 GT1-7 ACTH(1-24) & NDP-MSH DRCs ..........................................................59 GT1-7 Experiments.............................................................................................60 INS-1 & Cloudman M3 Experiments .................................................................62 CHO Heterologous System Localization Experiments.......................................63 CHO Heterologous System DRCs......................................................................64 CHO Heterologous System Single Analog Activation Experiments..................64 CHO Heterologous System Competition Assays ...............................................66 Future Directions ................................................................................................67 Bibliography ...................................................................................................................71 iii INTRODUCTION Overview: Ligand selectivity exhibited by receptors in the melanocortin receptor family is a question that has been hard to answer until recently. Without the discovery of the MC2R accessory protein (MRAP), obtaining functional MC2R in a heterologous system was nearly impossible. The focus of this thesis is three-fold: first, using endogenous expression systems to analyze a representative from each MCR class that will allow for functional studies in native surroundings. The second goal was creation a heterologous system for functional expression system of MC2R that can be easily cultured. The final goal was to look at the interaction and the ligand sequence requirement for activation of MCRs will be investigated using analog variants of ACTH hormone. Melanocortin Receptor Family: Melanocortin receptors (MCRs) are a family of G-protein coupled receptors (GPCR) activated by melanocortin peptides (Cone, 2006). The G protein-coupled receptor superfamily is divided into five different classes (Class A to Class E/F; Fredriksson et al., 2003). Class A is the rhodopsin or rhodopsin-like family, which includes the melanocortin receptors. Other families of GPCRs are adhesion, secretin, glutamate, and frizzled/TAS2. A unifying feature of the GPCRs is that each receptor has seven membrane spanning domains that are connected by internal and external polypeptide loop domains (Gantz and Fong, 2003). While the 3-dimensional structure of 1 melanocortin receptors has not been resolved, the structures of other GPCRs have been determined, such as the bovine rhodopsin and the human β-adrenergic receptor (Cone, 2006). GPCRs typically activate second messenger systems in order to relay messages the receptors receive. Melanocortin receptors are no different in that they relay messages using the adenylate cyclase/cAMP messenger pathway to propagate a biological response (Gantz and Fong, 2003). Another interesting feature about melanocortin receptor stimulation is the association of melanocortin signaling to intracellular systems that 2+ increase intracellular Ca via activation of IP3 (Konda et al., 1994), and induce either janus kinase/signal transducer and activator of transcription (Buggy, 1998), the influx of extracellular Ca2+ (Kojima et al., 1985), as well as activation of MAP kinase (Englaro et al., 1995), and other PKCs (Kapas et al., 1995). The activation of MCRs is derived from proopiomelanocortin, or POMC, which is the common precursor for different stimulatory ligands that activate melanocortin receptors. POMC: The melanocortin peptides come from a large preprohormone complex called proopiomelanocortin (or POMC) that is cleaved into many hormone products that may or may not undergo further processing (Pritchard et al., 2002; Fig. 1). Processing of the POMC gene product is tissue specific in that main cleavage products are made via processing by prohormone convertase 1/3 (PC1/3) in the anterior lobe of the pituitary and a distinct set of end-products can be generated from this precursor in the intermediate pituitary and hypothalamus through the action of prohormone convertases 2 {PC2} 2 Figure 1: Processing of the preprohormone POMC POMC is processed into several different melanocortin ligands depending on the tissue. Primary
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