Mus musculus Myelin
              Associated Glycoprotein (MAG)
            
                Lara O'Callaghan '23 and Maggie Williard '23 
        
         
        
         Contents:
        
        
         
         I. Introduction
        
        
        
          
          
          
        
         The myelin sheath is essential for the nervous system beause it
          insulates the axon and improves signal transmission by increasing the
          speed and efficiency of the electrical impulse. Myelin-associated
          glycoproteins (MAG) are specialized proteins that aid in the
          stabilization, construction, regulation, and maintenance of the myelin
          sheath. MAG acts as a bridge between the axon cell and the myelin
          sheath to maintain the periaxonal diameter, known as myelin-axon
          spacing (Fig. 1). MAG is a membrane glycoprotein and a member of the
          Ig superfamily (immunoglobulin protein family). If MAG is compromised
          via misfolding or autoimmune attacks, neurodegenerative disorders such
          as multiple sclerosis can develop. 
        MAG proteins are dimers that interact with glycolipids to adhere to
          the myelin sheath and bind to the axon to provide structure.
          Additionally, MAG inhibits axon regeneration and controls myelin
          formation. MAG consists of five Ig domains in a homodimeric
          arrangement with membrane-proximal domains Ig5 and Ig4. MAG binds to
          specialized lipids that reside on the axon membrane. MAG also has
          trans-membrane non-structured tails that reach beyond the myelin to
          interact with other proteins. Short linkers connect the MAG domains,
          creating a rigid structure which allows for bidirectional interactions
          between the myelin and the axon. MAG is post-translationally modified.
          The protein structure of MAG was solved in Mus musculus, the common
          house mouse. 
         
          
        
          Figure 1. Structure of
              Myelin-Associate Glycoprotein. (UMC Utrecht, 2016)
          
           II. General Structure
           
        Mag is a
          
          ; this means it is a protein composed of two chemically identical
          polypeptide chains (Monomer 1 and Monomer
            2) that mirror each other. Each
          
          is an asymetric unit composed of five domains labled Ig1,
          Ig2, Ig3,
          Ig4 and Ig5.
          . MAG is primarily composed of
          
          but it also contains several
          
          alpha helical structures. The Ig4 and Ig5 domains link the two
          monomers together to form the homodimer; their binding forms two
          equivalent hemi-interfaces that are largely hydrophobic and
          hydrophilic. The Ig1 and Ig2 domains create the biggest interface
          where Ig2 loops at the N-terminal side to interact with Ig1's A2-B
          side. However, Ig3, Ig4, and Ig5 are only formed in a head-to-tail
          manner at loops in the
          
          head and
          
          tail sides of the Ig domains.
 
        
         III. Specific Structures and Binding Interactions
        
         Each Ig domain is joined by
          
          , creating a rigid structure that enables MAG to connect the myelin
          and the axon. This allows MAG to moderate communication between the
          membranes. The Ig1 domain's N-terminal has a V-type Ig fold; this is
          consistent with other Siglec proteins. Meanwhile, the Ig3 and Ig4
          domains are C2-type. The Ig2 and Ig5
          domains have a C1-type Ig fold
          .
          MAG has an inter-domain disulfide bond between Ig1 and Ig2. MAG is
          post-translationally modified at many sites, containing seven
          
          . More specifically, five of these disulfide
            bridges are cannonical for the Ig domains. An inter-domain
          disulfide is formed between Ig1 and Ig2 by cysteines 37 and 165.
          Another intra-domain disulfide in Ig5 is formed by cysteines 421 and
          430. MAG contains a plethora of
          
          whose binding can cause conformational changes, affecting the overall
          structure. Ig1 plays an essential role in the recognition of ligands.
          Mag also contains 6 strong covalent bonds. Additionally, MAG goes
          under an N-linked glycosylation at the dimerization interface, which
          acts as a post-translational modification. MAG has eight
          
          sites. As the body develops, MAG glycosylation changes, and
          myelination deficiencies are correlated with abnormal glycosylation. 
        
         IV. MAG Mutagenesis
        
        Mutations in MAG can result in a plethora of aberrant functions.
          Mutations can often cause MAG misfolding or anti-MAG autoimmunity.
          This can have negative effects such as demyelination and
          neurodegenerative disorders. Myelin deficiency in the CNS and PNS is
          the result of the mutation of a gene that links signal transduction to
          RNA metabolism. The mutation of
          
          , which makes contact with sialic acid, on MAG reduces the potency of
          the inhibitory activity of MAG.
        
        
         VI. References
        Goodsell DS. 2020 Jul 1. Myelin-associated
              Glycoprotein. RCSB PDB. doi:10.2210/rcsb_pdb/mom_2020_7.
        
        Lossos A, Elazar N, Lerer I, Schueler-Furman
              O, Fellig Y, Glick B, Zimmerman B-E, Azulay H, Dotan S, Goldberg
              S, et al. 2015. Myelin-associated glycoprotein gene mutation
              causes Pelizaeus-Merzbacher disease-like disorder. Brain.
              138(9):2521-2536. 
        Pronker MF, Lemstra S, Snijder J, Heck AJR,
              Thies-Weesie DME, Pasterkamp RJ, Janssen BJC. 2016. Structural
              basis of myelin-associated glycoprotein adhesion and signalling. Nature
                Communications. 7(1):13584. doi:10.1038/ncomms13564.
        
        Quarles RH. 2007. Myelin-associated
              glycoprotein (MAG): past, present and beyond. Journal of
                Neurochemistry. 100(6):1431-1448.
              doi:10.1111/j.1471-4159.2006.04319.x. 
        UMC Utrecht. Structure of protein connecting
              neuron and myelin clarified (December 6, 2016). 
         Vinson M, Strijbos PJLM, Rowles A, Facci L,
              Moore SE, Simmons DL, Walsh FS. 2001. Myelin-associated
              Glycoprotein Interacts with Ganglioside GT1b: A MECHANISM FOR
              NEURITE OUTGROWTH INHIBITION*. Journal of Biological
                Chemistry. 276(23):20280-20285. doi:10.1074/jbc.M100345200.
        
         Yang LJ, Zeller CB, Shaper NL, Kiso M,
              Hasegawa A, Shapiro RE, Schnaar RL. 1996. Gangliosides are
              neuronal ligands for myelin-associated glycoprotein.Proc Natl
                Acad Sci USA. 93 (2):814. doi:10.1073/pnas.93.2.814.
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