• 1
    Coxon FP, Taylor A. Vesicular trafficking in osteoclasts. Semin Cell Dev Biol 2008;19:424433.
  • 2
    Teitelbaum SL. Bone resorption by osteoclasts. Science 2000;289: 15041508.
  • 3
    Vaananen HK, Zhao H, Mulari M, Halleen JM. The cell biology of osteoclast function. J Cell Sci 2000;113:377381.
  • 4
    Bossard MJ, Tomaszek TA, Thompson SK, Amegadzie BY, Hanning CR, Jones C, Kurdyla JT, McNulty DE, Drake FH, Gowen M, Levy MA. Proteolytic activity of human osteoclast cathepsin K. Expression, purification, activation, and substrate identification. J Biol Chem 1996;271:1251712524.
  • 5
    Bromme D, Okamoto K, Wang BB, Biroc S. Human cathepsin O2, a matrix protein-degrading cysteine protease expressed in osteoclasts. Functional expression of human cathepsin O2 in Spodoptera frugiperda and characterization of the enzyme. J Biol Chem 1996;271:21262132.
  • 6
    Angel NZ, Walsh N, Forwood MR, Ostrowski MC, Cassady AI, Hume DA. Transgenic mice overexpressing tartrate-resistant acid phosphatase exhibit an increased rate of bone turnover. J Bone Miner Res 2000;15:103110.
  • 7
    Halleen JM, Raisanen S, Salo JJ, Reddy SV, Roodman GD, Hentunen TA, Lehenkari PP, Kaija H, Vihko P, Vaananen HK. Intracellular fragmentation of bone resorption products by reactive oxygen species generated by osteoclastic tartrate-resistant acid phosphatase. J Biol Chem 1999;274:2290722910.
  • 8
    Sun P, Sleat DE, Lecocq M, Hayman AR, Jadot M, Lobel P. Acid phosphatase 5 is responsible for removing the mannose 6-phosphate recognition marker from lysosomal proteins. Proc Natl Acad Sci U S A 2008;105:1659016595.
  • 9
    Baron R, Neff L, Brown W, Courtoy PJ, Louvard D, Farquhar MG. Polarized secretion of lysosomal enzymes: co-distribution of cation-independent mannose-6-phosphate receptors and lysosomal enzymes along the osteoclast exocytic pathway. J Cell Biol 1988;106:18631872.
  • 10
    Baron R, Neff L, Louvard D, Courtoy PJ. Cell-mediated extracellular acidification and bone resorption: evidence for a low pH in resorbing lacunae and localization of a 100-kD lysosomal membrane protein at the osteoclast ruffled border. J Cell Biol 1985;101:22102222.
  • 11
    Czupalla C, Mansukoski H, Riedl T, Thiel D, Krause E, Hoflack B. Proteomic analysis of lysosomal acid hydrolases secreted by osteoclasts: implications for lytic enzyme transport and bone metabolism. Mol Cell Proteomics 2006;5:134143.
  • 12
    Goto T, Yamaza T, Tanaka T. Cathepsins in the osteoclast. J Electron Microsc (Tokyo) 2003;52:551558.
  • 13
    Littlewood-Evans A, Kokubo T, Ishibashi O, Inaoka T, Wlodarski B, Gallagher JA, Bilbe G. Localization of cathepsin K in human osteoclasts by in situ hybridization and immunohistochemistry. Bone 1997;20:8186.
  • 14
    Mulari M, Vaaraniemi J, Vaananen HK. Intracellular membrane trafficking in bone resorbing osteoclasts. Microsc Res Tech 2003;61:496503.
  • 15
    Toyomura T, Murata Y, Yamamoto A, Oka T, Sun-Wada GH, Wada Y, Futai M. From lysosomes to the plasma membrane: localization of vacuolar-type H+ -ATPase with the a3 isoform during osteoclast differentiation. J Biol Chem 2003;278:2202322030.
  • 16
    Yamaza T, Goto T, Kamiya T, Kobayashi Y, Sakai H, Tanaka T. Study of immunoelectron microscopic localization of cathepsin K in osteoclasts and other bone cells in the mouse femur. Bone 1998;23:499509.
  • 17
    Gelfman CM, Vogel P, Issa TM, Turner CA, Lee WS, Kornfeld S, Rice DS. Mice lacking alpha/beta subunits of GlcNAc-1- phosphotransferase exhibit growth retardation, retinal degeneration, and secretory cell lesions. Invest Ophthalmol Vis Sci 2007;48:52215228.
  • 18
    Lee WS, Payne BJ, Gelfman CM, Vogel P, Kornfeld S. Murine UDP-GlcNAc:lysosomal enzyme N-acetylglucosamine-1-phospho transferase lacking the gamma-subunit retains substantial activity toward acid hydrolases. J Biol Chem 2007;282:2719827203.
  • 19
    Holt OJ, Gallo F, Griffiths GM. Regulating secretory lysosomes. J Biochem 2006;140:712.
  • 20
    Sachse M, Ramm G, Strous G, Klumperman J. Endosomes: multipurpose designs for integrating housekeeping and specialized tasks. Histochem Cell Biol 2002;117:91104.
  • 21
    Blott EJ, Griffiths GM. Secretory lysosomes. Nat Rev Mol Cell Biol 2002;3:122131.
  • 22
    Peters PJ, Borst J, Oorschot V, Fukuda M, Krahenbuhl O, Tschopp J, Slot JW, Geuze HJ. Cytotoxic T lymphocyte granules are secretory lysosomes, containing both perforin and granzymes. J Exp Med 1991;173:10991109.
  • 23
    Klumperman J, Hille A, Veenendaal T, Oorschot V, Stoorvogel W, von Figura K, Geuze HJ. Differences in the endosomal distributions of the two mannose 6-phosphate receptors. J Cell Biol 1993;121:9971010.
  • 24
    van Meel E, Klumperman J. Imaging and imagination: understanding the endo-lysosomal system. Histochem Cell Biol 2008;129:253266.
  • 25
    Ljusberg J, Wang Y, Lang P, Norgard M, Dodds R, Hultenby K, Ek-Rylander B, Andersson G. Proteolytic excision of a repressive loop domain in tartrate-resistant acid phosphatase by cathepsin K in osteoclasts. J Biol Chem 2005;280:2837028381.
  • 26
    Goto T, Tsukuba T, Ayasaka N, Yamamoto K, Tanaka T. Immunocytochemical localization of cathepsin D in the rat osteoclast. Histochemistry 1992;97:1318.
  • 27
    Reinholt FP, Hultenby K, Heinegard D, Marks SC Jr, Norgard M, Anderson G. Extensive clear zone and defective ruffled border formation in osteoclasts of osteopetrotic (ia/ia) rats: implications for secretory function. Exp Cell Res 1999;251:477491.
  • 28
    Goto T, Kiyoshima T, Moroi R, Tsukuba T, Nishimura Y, Himeno M, Yamamoto K, Tanaka T. Localization of cathepsins B, D, and L in the rat osteoclast by immuno-light and -electron microscopy. Histochemistry 1994;101:3340.
  • 29
    Dodds RA, James IE, Rieman D, Ahern R, Hwang SM, Connor JR, Thompson SD, Veber DF, Drake FH, Holmes S, Lark MW, Gowen M. Human osteoclast cathepsin K is processed intracellularly prior to attachment and bone resorption. J Bone Miner Res 2001;16:478486.
  • 30
    Rieman DJ, McClung HA, Dodds RA, Hwang SM, Holmes MW, James IE, Drake FH, Gowen M. Biosynthesis and processing of cathepsin K in cultured human osteoclasts. Bone 2001;28: 282289.
  • 31
    Zhao H, Ito Y, Chappel J, Andrews NW, Teitelbaum SL, Ross FP. Synaptotagmin VII regulates bone remodeling by modulating osteoclast and osteoblast secretion. Dev Cell 2008;14:914925.
  • 32
    Zhao H, Laitala-Leinonen T, Parikka V, Vaananen HK. Downregulation of small GTPase Rab7 impairs osteoclast polarization and bone resorption. J Biol Chem 2001;276:3929539302.
  • 33
    Boonen M, van Meel E, Oorschot V, Klumperman J, Kornfeld S. Vacuolization of mucolipidosis type II mouse exocrine gland cells represents accumulation of autolysosomes. Mol Biol Cell 2011;22: 11351147.
  • 34
    Nesbitt SA, Horton MA. Trafficking of matrix collagens through bone-resorbing osteoclasts. Science 1997;276:266269.
  • 35
    Salo J, Lehenkari P, Mulari M, Metsikko K, Vaananen HK. Removal of osteoclast bone resorption products by transcytosis. Science 1997;276:270273.
  • 36
    Lacey DL, Timms E, Tan HL, Kelley MJ, Dunstan CR, Burgess T, Elliott R, Colombero A, Elliott G, Scully S, Hsu H, Sullivan J, Hawkins N, Davy E, Capparelli C et al. Osteoprotegerin ligand is a cytokine that regulates osteoclast differentiation and activation. Cell 1998;93:165176.
  • 37
    Griffiths GM, Isaaz S. Granzymes A and B are targeted to the lytic granules of lymphocytes by the mannose-6-phosphate receptor. J Cell Biol 1993;120:885896.
  • 38
    Dittmer F, Ulbrich EJ, Hafner A, Schmahl W, Meister T, Pohlmann R, von Figura K. Alternative mechanisms for trafficking of lysosomal enzymes in mannose 6-phosphate receptor-deficient mice are cell type-specific. J Cell Sci 1999;112:15911597.
  • 39
    Glickman JN, Kornfeld S. Mannose 6-phosphate-independent targeting of lysosomal enzymes in I-cell disease B lymphoblasts. J Cell Biol 1993;123:99108.
  • 40
    Rijnboutt S, Aerts HM, Geuze HJ, Tager JM, Strous GJ. Mannose 6-phosphate-independent membrane association of cathepsin D, glucocerebrosidase, and sphingolipid-activating protein in HepG2 cells. J Biol Chem 1991;266:48624868.
  • 41
    Rijnboutt S, Kal AJ, Geuze HJ, Aerts H, Strous GJ. Mannose 6-phosphate-independent targeting of cathepsin D to lysosomes in HepG2 cells. J Biol Chem 1991;266:2358623592.
  • 42
    Canuel M, Korkidakis A, Konnyu K, Morales CR. Sortilin mediates the lysosomal targeting of cathepsins D and H. Biochem Biophys Res Commun 2008;373:292297.
  • 43
    Lefrancois S, Canuel M, Zeng J, Morales CR. Inactivation of sortilin (a novel lysosomal sorting receptor) by dominant negative competition and RNA interference. Biol Proced Online 2005;7:1725.
  • 44
    Vogel P, Payne BJ, Read R, Lee WS, Gelfman CM, Kornfeld S. Comparative pathology of murine mucolipidosis types II and IIIC. Vet Pathol 2009;46:313324.
  • 45
    Hochman JA, Treem WR, Dougherty F, Bentley RC. Mucolipidosis II (I-cell disease) presenting as neonatal cholestasis. J Inherit Metab Dis 2001;24:603604.
  • 46
    Kabra M, Gulati S, Kaur M, Sharma J, Singh A, Chopra V, Menon PS, Kalra V. I-cell disease (mucolipidosis II). Indian J Pediatr 2000;67: 683687.
  • 47
    Patriquin HB, Kaplan P, Kind HP, Giedion A. Neonatal mucolipidosis II (I-cell disease): clinical and radiologic features in three cases. AJR Am J Roentgenol 1977;129:3743.
  • 48
    Pazzaglia UE, Beluffi G, Campbell JB, Bianchi E, Colavita N, Diard F, Gugliantini P, Hirche U, Kozlowski K, Marchi A, Nayanar V, Pagani G. Mucolipidosis II: correlation between radiological features and histopathology of the bones. Pediatr Radiol 1989;19:406413.
  • 49
    Glickman JN, Morton PA, Slot JW, Kornfeld S, Geuze HJ. The biogenesis of the MHC class II compartment in human I-cell disease B lymphoblasts. J Cell Biol 1996;132:769785.
  • 50
    Filgueira L. Fluorescence-based staining for tartrate-resistant acidic phosphatase (TRAP) in osteoclasts combined with other fluorescent dyes and protocols. J Histochem Cytochem 2004;52:411414.
  • 51
    Kamiya T, Kobayashi Y, Kanaoka K, Nakashima T, Kato Y, Mizuno A, Sakai H. Fluorescence microscopic demonstration of cathepsin K activity as the major lysosomal cysteine proteinase in osteoclasts. J Biochem 1998;123:752759.
  • 52
    Valenzano KJ, Kallay LM, Lobel P. An assay to detect glycoproteins that contain mannose 6-phosphate. Anal Biochem 1993;209:156162.
  • 53
    Takeshita S, Kaji K, Kudo A. Identification and characterization of the new osteoclast progenitor with macrophage phenotypes being able to differentiate into mature osteoclasts. J Bone Miner Res 2000;15:14771488.
  • 54
    Slot JW, Geuze HJ. Cryosectioning and immunolabeling. Nat Protoc 2007;2:24802491.
  • 55
    de Wit H, Lichtenstein Y, Geuze HJ, Kelly RB, van der Sluijs P, Klumperman J. Synaptic vesicles form by budding from tubular extensions of sorting endosomes in PC12 cells. Mol Biol Cell 1999;10: 41634176.
  • 56
    Boonen M, Vogel P, Platt KA, Dahms N, Kornfeld S. Mice lacking mannose 6-phosphate uncovering enzyme activity have a milder phenotype than mice deficient for N-acetylglucosamine-1-phosphotransferase activity. Mol Biol Cell 2009;20:43814389.
  • 57
    Varki A, Kornfeld S. The spectrum of anionic oligosaccharides released by endo-beta-N-acetylglucosaminidase H from glycoproteins. Structural studies and interactions with the phosphomannosyl receptor. J Biol Chem 1983;258:28082818.