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Background: a bit about… Important roles in: •Glycoprotein biosynthesis, quality control & catabolism •Multiple forms of α-mannosidases in mammalian cells, differing in specificity, function and cellular location Classified into 2 groups conserved through eukaryotic evolution Class I Class II - Narrow specificity for α-1, 2mannosidic linkages - Broad specificity: hydrolyse α(1→ 2), α(1→3) and α(1→6) mannosidic linkages -Further specificity within family for isomeric species produced Man a1-2 Mana1-2 Mana1-6 Mana1-3 Mana1-6 Manß1-4GlcNAcß1-4GlcNAcß1-Asn Man a 1-2 Mana1-2 Mana1-3 CELL ERAD Cytosol persistently misfolded M8B / M8C Monosaccharide constituents Lysosome misfolded ER Protein and Oligosaccharide Processing in the Cytosol of ERAD + G1-3M8N2 + M8N2 Chitobiase Proteasome M8N1 Chitobiase Cytosolicαmannosidase Fast M5N1 G1-3M8N1 Cytosolicαmannosidase lysosome G1-3M5N1 Cyt α-man ? G1-3M4N1 Plasma membrane ? G1-3M4N1 Cyt α-man? Slow M4N1 Known α-mannosidase inhibitors OH H N HO N HO HO OH OH HO Deoxymannojorimycin (DMJ) HO O HO H N O N H Kifunensin (Kif) Weak general inhibitor of mammalian α-mannosidases OH HO OH HO Swainsonine (SW) OH 1, 4 deoxy-1,4-imino Dmannitol (DIM) Inhibits lysosomal mannosidase and Golgi mannosidase ER mannosidase I and Man9-mannosidase Higher conc.: Will start to inhibit other α-mannosidases HO H H Potent and specific inhibitor of Golgi mannosidase I Low conc.: Golgi mannosidase I and Man9-mannosidase H N Novel 7-membered imino-sugars: inhibitory effect on cellular α-mannosidases (Kukushkin & Butters; unpublished data) N HO N OH HO 148 OH N HO OH HO OH 149 N OH HO 266 OH OH HO OH 265 Project Aims The aims of my project were: To define α-mannosidase targets of novel inhibitors in cells and compare these to known αmannosidase inhibitors This will be accomplished by: 1.Evaluation of cytosolic and lysosomal α-mannosidase inhibition by measuring free oligosaccharides (FOS) in HL60 and MDBK cell lines. 2.Increasing our knowledge of mannosidases within their cellular pathways in cell lines above. 3.Using glycoprotein precipitation, isolation and PNGase glycan release methods to observe effects of inhibitors on glycan species of total cellular glycoprotein. 4.Identification of lysosomal FOS species following cellular fractionation. Two cell lines will be used that have differing biosynthetic and catabolic pathways: HL60, a human promyelocytic leukaemia cell line and MDBK, a bovine kidney cell line. 5.Estimation of ER and Golgi α-mannosidase inhibition by using Fluorescence-activated cell sorting (FACS) to analysing glycoprotein derived oligosaccharides on the cell surfacee following treatment with α-mannosidase inhibitors. HPLC analysis of 2-AA fluorescently labelled free oligosaccharides (FOS) from HL60 cells treated with known α-mannosidase inhibitors Fluorescence (arbitrary units) Untreated HL60 M5N M7N M4N DMJ treated (1mM) DIM treated (100μM) M9N G1M5 M9N/N2 M5N G1M5 M4N M7N/N2 M7N/N2 M8N/N2 M8N G1M9N? M9N/N2 M7N/N2 M8N/N2 HPLC analysis of 2-AA fluorescently labelled free oligosaccharides (FOS) from MDBK cells treated with known α-mannosidase inhibitors 3000 M5N Fluorescence (arbitrary units) Untreated MDBK cells M4N G1M5N M3N2 M4N2 M9N/N2 6000 DIM treated (100μM) M3N2 M5N M4N2 M5N2 M6N2 M7N/N2 M8N2 M9N/N2 HPLC analysis of 2-AA fluorescently labelled free oligosaccharides (FOS) from MDBK cells treated with Kif at increasing concentrations (1μM- 100μM) - Untreated M5N G1M5 M4N2 M9N Fluorescence (arbitrary units) - + 1μM Kif M5N M4N2 G1M5 M6N2 M8N2 M7N2 M9N + 10μM Kif M8N2 M7N2 M9N/N2 M8N2 + 100μM Kif M7N2 M9N/N2 22 23 24 25 26 27 28 29 30 31 32 Minutes 33 34 35 36 37 38 39 40 41 HPLC analysis of 2-AA fluorescently labelled free oligosaccharides (FOS) from HL60 cells treated with 148 and 149 M5N G1M5 Fluorescence (arbitrary units) M4N SATIN M8N M9N - Fluorescence M8N M9N M6N2 G1M5 M6N G1M9 M7N/ N2 M5N M4N M5N M4N 22 24 26 28 M8N G1M5 M6N2 M7N/ M6N N2 30 32 Minutes 34 M9N G1M9 36 38 40 42 HPLC analysis of 2-AA fluorescently labelled free oligosaccharides (FOS) from MDBK cells treated with 148 and 149 M5N 1200 Fluorescence (arbitrary units) G1M5 M4N2 M3N2 - M3N2 1800 M4N2 M5N M8N2 M6N2 M7N2 M9N/N2 M5N2 6000 M4N2 M5N M7N2 M6N2 M8N2 M9N/N2 M5N2 18 20 22 24 26 28 30 Minutes 32 34 36 38 40 42 HPLC analysis of 2-AA fluorescently labelled free oligosaccharides (FOS) from HL60 cells treated with 265 and 266 1600 Untreated M5N Fluorescence (arbitrary units) M9N M3N 4000 M4N M6N + 100μM 265 M3N M5N M4N M6N 2000 + 100μM 266 M5N M4N M3N 12 M9N 14 16 18 20 22 24 26 Minutes M9N 28 30 32 34 36 38 40 42 HPLC analysis of 2-AA fluorescently labelled free oligosaccharides (FOS) from MDBK cells treated with 265 and 266 1300 Fluorescence (arbitrary units) M3N2 SATIN - Fluorescence M3N2 6000 M3N2 6000 0 12.00 14.00 16.00 18.00 20.00 22.00 24.00 26.00 28.00 Minutes 30.00 32.00 34.00 36.00 38.00 40.00 42.00 Flow cytometry data representing % change in ConA binding in Kif and 148 treated HL60 cells over 24hrs and 72 hrs Effect on total cellular glycoprotein upon inhibitor treatment Untreated HL60 600 1000 % increase in ConA binding after 24hrs % increase in ConA binding after 72hrs + 100μM Kif 68.1% 136% + 100μM 148 13.9% 34.2% + DMJ (1mM) SATIN - Fluorescence fluorescence 1200 + 148 (100μM) + 265 (100μM) 1000 1000 + 266 (100μM) SATIN - Fluorescence 2500 + Kif (100μM) 22 24 26 M9N2 28 30 32 34 36 Elution time (minutes) 38 40 42 44 Cellular fractionation of untreated HL60 and MDBK cells HL60 Cells Lysosomal Activity MDBK Cells Lysosomal Activity 1.4 1.2 0.5 Fraction 9 lysosomal activity/ ug protein Lysosomal Activity/ ug protein 0.6 0.4 0.3 0.2 0.1 Fraction 8 1 0.8 0.6 0.4 0.2 0 0 0 2 4 6 8 10 12 14 0 -0.1 2 4 6 8 10 12 14 -0.2 Cell Fraction Number Fluorescence (arbitrary units) Fraction number 16 18 20 22 24 26 28 30 32 34 36 38 40 42 12 14 Elution time (minutes) 16 18 20 22 24 26 28 30 32 34 36 38 40 42 In Summary Known α-mannosidase inhibitors: - DMJ: low potency, low specificity α-mannosidase inhibitor. Does not inhibit cytosolic mannosidase at high concentration (1mM) but most likely inhibiting ER/Golgi and partially inhibiting lysosomal α-mannosidases. - Kif: Known strong inhibitor of ER Mannosidase I and Golgi mannosidases. No inhibition of ER Man II due to accumulation of M8C. - Very high potency, inhibitory effects observed from 1μM conc. No inhibition of cytosolic/lysosomal αmannosidases. Selective for ER/Golgi mannosidases and the biosynthetic pathway for glycoprotein maturation. - DIM: Selective for lysosomal α-mannosidase and lysosomal α1,6 mannosidase. Partial inhibition of catabolism of high mannose structures, strong inhibition of breakdown of core N-linked glycan isomer M3aN2. Novel α-mannosidase inhibitors - 148 & 149: Very similar inhibitory patterns in the cell. Inhibits cytosolic α-mannosidase, and lysosomal mannosidase in a similar way to DIM. Subtle inhibition of conversion of oligomannose to complex glycan maturation to the cell surface. - 265: No inhibition of cytosolic α-mannosidase. Targets lysosomal α-mannosidase and lysosomal α1, 6 mannosidase with particularly potent inhibition for the catabolism of core M3aN2. - 266: No inhibition of cytosolic α-mannosidase. Inhibits lysosomal α-mannosidase with particularly potent inhibition for the catabolism of core M3aN2. Partial inhibition of catabolism of high oligomannose glycans by lysosomal αmannosidase . Low levels of inhibition for α1,6 mannosidase. Future work • Cellular fractionation with added inhibitor followed by FOS isolation and characterisation to confirm and clarify target compartments of novel inhibitors within the cell and to specify enzymes inhibited. • Carry out Flow Cytometry experiment with 266 and 265 to analyse effect on glycoprotein maturation at the cell surface • To confirm the cytosolic inhibition of 148 and 149. Proposed experiment: Addition of 148/149 following treatment of cells with glucosidase inhibitor such as NB-DMJ to create more triglucosylated species. If 148/149 inhibit in the cytosol, will generate more glucosylated highmannose structures from FOS analysis. • May want to develop a more selective or & more potent inhibitor of the cytosolic α-mannosidase: drug screening with a range of 148,149-like structures. Acknowledgements Dr Terry Butters Dr David Neville Dr Dominic Alonzi Stephanie Boomkamp Professor Raymond Dwek