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ESSENTIALS OF GLYCOBIOLOGY LECTURE 33 GENETIC DISORDERS OF GLYCOSYLATION IN HUMANS Hud Freeze OVERVIEW AND SUMMARY Many Human glycosylation disorders were discovered in the last 10 years, mostly in the N-linked pathway. Disorders are underdiagnosed. Alert physicians. Defects include monosaccharide activation, glycosyl transferases, transporters, biosynthetic glycosidases Golgi trafficking proteins. A few disorders can be treated with monosaccharides Misglycosylation causes some types of muscular dystrophy More glycosylation disorders will be found, phenotypes Will be broad and variable Number of defects GLYCOSYLATION DEFECTS IDENTIFIED N-linked Disorders Muscular Dystrophy 20 20 15 15 10 10 5 5 80- 94 93 95 96 97 98 99 00 01 02-04 Year CONGENITAL DISORDERS OF GLYCOSYLATION TYPE I LLO Dol-PP TYPE II On Protein -N-X-T/S -N-X-T/S CONGENITAL DISORDERS OF GLYCOSYLATION (CDG) TYPE I---Defects in synthesis or transfer of oligosaccharides from Dol-PP carrier to proteins in the lumen of the endoplasmic reticulum TYPE II--Defects in the processing of N-linked oligosaccharides OR in all other types of glycosylation Types Number of Patients Ia-IL-IIa-IIe-Ix or IIx-- >400 (Ia>300) ~20 >50 CDG--often (not always) recognized by glycosylation-dependent alterations in serum transferrin isoelectric focusing pattern Isoelectric focusing of transferrin control CDG patients control 4 2 Sialic acids 0 normal missing chains Sialic acid Galactose altered processing N-acetylglucosamine Mannose 30-40 GENES REQUIRED TO MAKE AND ADD AN ASN-LINKED CHAIN OVERVIEW OF SUGAR METABOLISM IN CELLS Glycolysis Gal Glc ATP Man GlcNAc ATP ATP Pi Gal-1-P Glycogen Glc-6-P Man-6-P Fru-6-P NAD UDP-Glc NAD+ UDP-GlcA GlcN-6-P Glc-1-P Man-l-P GTP AcCoA Dol-P ATP GDP-Man GlcNAc-6-P Dol-P-Glc NADP -CO2 GlcNAc-l-P GalNAc GalNAc-1-P UTP GDP-Fuc Dol-P-Man UDP-GalNAc UDP- GlcNAc ManNAc ATP CMP-Neu5Ac ManNAc-6-P PEP NADP CMP-Neu5Ac Dol-P UTP UDP-Xyl ATP Fuc-1-P -NH3 Glutamine UTP UDP-Glc UDP-Gal Fuc CTP Neu5Ac Neu5Ac-9-P GTP Glycolysis Gal Glc ATP Man GlcNAc ATP ATP Pi Gal-1-P Glycogen Glc-6-P Man-6-P Fru-6-P Fuc-1-P -NH3 Glutamine UTP UDP-Glc UDP-Gal Fuc NAD UDP-Glc NAD+ UDP-GlcA GlcN-6-P Glc-1-P Man-l-P GTP AcCoA Dol-P ATP GTP GDP-Man GlcNAc-6-P Dol-P-Glc NADP -CO2 GlcNAc-l-P GalNAc GalNAc-1-P GDP-Fuc UTP UDP-Xyl ATP Dol-P UTP Dol-P-Man UDP-GalNAc UDP- GlcNAc ManNAc ATP CMP-Neu5Ac Mannose supplements can treat CDG-Ib, Fru-6-P-->Man-6-P defect ManNAc-6-P PEP NADP CMP-Neu5Ac CTP Neu5Ac Neu5Ac-9-P Biosynthesis and Interconve rsion of Monosaccharides. The relative contributions of each under physiological conditions are unknown. - monosaccharides; -control points and - donors; BEFORE MANNOSE DURING MANNOSE 200 160 AT III 120 80 40 0 -15 -13 -11 - 13 -7 -5 -3 -1 1 3 5 1 3 5 1 3 7 9 11 13 15 17 7 anticoagulation (Marcumar6R ) 5 i. v. albumin substitution 4 3 2 1 0 albumin (g/dl) - 15 -9 - 11 -9 -7 -5 -3 -1 7 9 11 13 15 17 10 fecal alpha1 AT (mg/g) 8 6 4 2 0 -15 -13 -11 -9 -7 -5 -3 -1 time in months 5 7 9 11 13 15 17 Therapy for CDG-Ib PMI -6-P Fru-6-P PMM -1-P GDP- Dol-P- CDG defect sugar transporter Man GlcNAc Glycolysis Gal Glc Man Patient is deficient in fucosylation GlcNAc ATP ATP What symptoms would Fru-6-P you expect? Pi Gal-1-P Man-6-P Glc-6-P Glycogen Cause? -NH3 Glutamine UTP UDP-Glc GlcN-6-P Man-l-P Glc-1-P UDP-Gal UDP-Glc What therapy? NAD NAD+ UDP-GlcA GTP AcCoA Dol-P ATP Fuc ATP Fuc-1-P GTP GDP-Man GlcNAc-6-P Dol-P-Glc NADP -CO2 GlcNAc-l-P GalNAc GalNAc-1-P UTP GDP-Fuc UTP UDP-Xyl ATP Dol-P Dol-P-Man UDP-GalNAc UDP- GlcNAc ManNAc ATP Oral fucose supplements used to Treat CDG-I Ic-ManNAc-6-P PEP Defective in Golgi GDP-Fuc Transporter CMP-Neu5Ac NADP CMP-Neu5Ac CTP Neu5Ac Neu5Ac-9-P FUCOSE THERAPY NORMALIZES NEUTROPHIL COUNTS OF ONE CDG-IIc PATIENT Serum Fucose (µM) Normal Range QuickTime™ and a BMP decompressor are needed to see this picture. QuickTime™ and a Photo - JPEG decompressor are needed to see this picture. CLINICAL FEATURES OF CDG PATIENTS Used with permission T. Marquardt. European J. Ped (2003)162 359-379. Dysmorphic features of a baby with CDG-Ia (PMM-deficiency) -6P -6P Ia GDP- Kindly provided by Dr. Thorsten Marquardt -1P DEFECT -6P -6P Ib -1P Ia Dol-P- LLO If Ie GDP- Dol-P- Id Dol-PP Dol-PP Ig Dol-PP Dol-PP Dol-PP Dol-PP Ic Dol-PP Dol-PP On Protein IIb UDP -N-X-T/S -N-X-T/S -N-X-T/S -N-X-T/S -N-X-T/S IIc UDP UDP IIa -N-X-T/S CMP Cytosol GDP- IId -N-X-T/S -N-X-T/S Golgi GDP- -N-X-T/S -N-X-T/S Defect is Shared by N-Linked and O-Linked Pathways -N-X-T/S T/S -N-X-T/S T/S Possible Defect in Sialic Acid Synthesis or Transport? Defects in Multiple Sialyl Transferases? Core1GalT and S T3Gal-I activiti es in control and patient fibroblasts (pmol/mg p rotein/min) Enzyme Control H.D. Core1GalT 3.1 0.6 1.9 0.2 ST3Gal-I 0.97 0.08 0.37 0.05 Nucleotide Sugar Transport is reduced Endogenous acceptor A [3H]CMP-Sia 250 200 150 pmol/mg protein pmol/mg protein Exogenous acceptor (GAP) 100 50 0 250 200 B CMP-[3H]Sia 150 100 50 0 250 C UDP-[3H]Gal 200 D D 300 250 200 150 100 50 0 Control Vmax 306 HD Vmax 92 50 40 30 20 10 0 E 200 F CMP-Sia (uM) Control Vmax 43 HD Vmax 18 UDP-Gal (uM) Control CHO 150 Vmax 182 150 100 100 50 50 0 0 0 3 Time (min) Control Control+TX-100 HD 6 Lec2 0 2 4 6 8 10 12 CMP-Sia (uM) Vmax is reduced, Km remains the same ST-GFP Trafficking is reduced in HD and FD cells T/S ST3Gal-I Recovery FD HD Control Pre-bleach bleach 10min 30min 60 min ldlB=Cog1 ldlC=Cog2 ldlB and ldlC deficient cells are defective in glycosylation Mutation Function Cog7 mutation decreases mRNA and protein Destabilizes COG complex, mislocalizing some COG subunits Decreases trafficking of glycosyltransferases and transporters Decreases enzyme/transporter stability and normal localization Decreases terminal modifications on glycans SUMMARY • • • • • Physicians must know many faces of glycosylation Defective Glycosylation causes disease Only a few of the potential types of CDG are known Simple therapy may help a few CDG patients CDG may be the cause of unexplained cases of developmental delay, coagulopathy, and hypoglycemia • CDG presentations can be very mild and broad WHEN TO CONSIDER CDG? ”EVERY TIME YOU SUSPECT IT…. AND EVERY TIME YOU DON’T” -J. Jaeken Glycolysis Gal Glc ATP Man GlcNAc ATP ATP Pi Gal-1-P Glycogen Glc-6-P Man-6-P Fru-6-P Fuc-1-P -NH3 Glutamine UTP UDP-Glc UDP-Gal Fuc NAD UDP-Glc NAD+ UDP-GlcA GlcN-6-P Glc-1-P Man-l-P GTP AcCoA Dol-P ATP GTP GDP-Man GlcNAc-6-P Dol-P-Glc NADP -CO2 GlcNAc-l-P GalNAc GalNAc-1-P UTP GDP-Fuc UTP UDP-Xyl ATP Dol-P Dol-P-Man UDP-GalNAc UDP- GlcNAc ManNAc ATP CMP-Neu5Ac UDP-GlcNAc epimerase/kinase Defective in two human diseases ManNAc-6-P PEP NADP CMP-Neu5Ac CTP Neu5Ac Neu5Ac-9-P Sialuria and Hereditary Inclusion Body Myopathy-II Sialuria--hypotonia, cerebellar ataxia, and mental retardation Muscular Dystrophies Walker Warburg Syndrome--POMT1 mutations cause 30% MEB-Muscle Eye Brain Disease--POMGnT1(1p32-34) Encodes a b1,2GlcNAc transferase specific for O- Man Fukuyma-type CMD--fukutin (9q31) Putative transferase in golgi (?) Fukutin Related protein--(19q13.3) Putative transferase (?) LARGE- cause of myd mouse, tandem glycosyltransfrases(?) Common Features: Affects a-dystroglycan glycosylation and not b-dystroglycan Hereditary Inclusion Body Myopathy II (9p12-13) UDP-GlcNAc epimerase/kinase used for CMP-Sia O-Mannose: An Emerging Family of Glycans WWS-Mutated transferase O-Mannose: An Emerging Family of Glycans MEB- Mutated transferase MACULAR CORNEAL DYSTROPHY Autosomal recessive Progressive punctate corneal opacity, requires transplant Caused by mutation in GlcNAc-6-Sulfotransferase (CHST6) used for sulfation of keratan sulfate http://www.burnham.org/presentations/hum an_glycosylation_disorders/index.htm