{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,5]],"date-time":"2026-11-05T04:17:38Z","timestamp":1793852258932,"version":"4.6.1"},"reference-count":83,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2021,3,30]],"date-time":"2021-03-30T00:00:00Z","timestamp":1617062400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Cell Dev. Biol."],"abstract":"<jats:p>The current models of osteoclastic bone resorption focus on immobile osteoclasts sitting on the bone surface and drilling a pit into the bone matrix. It recently appeared that many osteoclasts also enlarge their pit by moving across the bone surface while resorbing. Drilling a pit thus represents only the start of a resorption event of much larger amplitude. This prolonged resorption activity significantly contributes to pathological bone destruction, but the mechanism whereby the osteoclast engages in this process does not have an answer within the standard bone resorption models. Herein, we review observations that lead to envision how prolonged resorption is possible through simultaneous resorption and migration. According to the standard pit model, the \u201csealing zone\u201d which surrounds the ruffled border (i.e., the actual resorption apparatus), \u201canchors\u201d the ruffled border against the bone surface to be resorbed. Herein, we highlight that continuation of resorption demands that the sealing zone \u201cglides\u201d inside the cavity. Thereby, the sealing zone emerges as the structure responsible for orienting and displacing the ruffled border, e.g., directing resorption against the cavity wall. Importantly, sealing zone displacement stringently requires thorough collagen removal from the cavity wall - which renders strong cathepsin K collagenolysis indispensable for engagement of osteoclasts in cavity-enlargement. Furthermore, the sealing zone is associated with generation of new ruffled border at the leading edge, thereby allowing the ruffled border to move ahead. The sealing zone and ruffled border displacements are coordinated with the migration of the cell body, shown to be under control of lamellipodia at the leading edge and of the release of resorption products at the rear. We propose that bone resorption demands more attention to osteoclastic models integrating resorption and migration activities into just one cell phenotype.<\/jats:p>","DOI":"10.3389\/fcell.2021.644503","type":"journal-article","created":{"date-parts":[[2021,3,30]],"date-time":"2021-03-30T05:31:47Z","timestamp":1617082307000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":45,"title":["The Mechanism Switching the Osteoclast From Short to Long Duration Bone Resorption"],"prefix":"10.3389","volume":"9","author":[{"given":"Jean-Marie","family":"Delaisse","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kent","family":"S\u00f8e","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Thomas Levin","family":"Andersen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Aleksandra Maria","family":"Rojek","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Niels","family":"Marcussen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2021,3,30]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"603","DOI":"10.1007\/s00418-016-1414-y","article-title":"Early reversal cells in adult human bone remodeling: osteoblastic nature, catabolic functions and interactions with osteoclasts.","volume":"145","author":"Abdelgawad","year":"2016","journal-title":"Histochem. Cell Biol."},{"key":"B2","doi-asserted-by":"publisher","first-page":"181","DOI":"10.1016\/j.bone.2014.07.012","article-title":"Does collagen trigger the recruitment of osteoblasts into vacated bone resorption lacunae during bone remodeling?","volume":"67","author":"Abdelgawad","year":"2014","journal-title":"Bone"},{"key":"B3","doi-asserted-by":"publisher","first-page":"40","DOI":"10.1007\/s002230010047","article-title":"Alendronate disturbs vesicular trafficking in osteoclasts.","volume":"70","author":"Alakangas","year":"2002","journal-title":"Calcif. Tissue Int."},{"key":"B4","doi-asserted-by":"publisher","first-page":"2210","DOI":"10.1083\/jcb.101.6.2210","article-title":"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.","volume":"101","author":"Baron","year":"1985","journal-title":"J. Cell Biol."},{"key":"B5","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1007\/BF02008210","article-title":"The organic-inorganic relationships in bone matrix undergoing osteoclastic resorption.","volume":"16","author":"Bonucci","year":"1974","journal-title":"Calcif. Tissue Res."},{"key":"B6","doi-asserted-by":"publisher","DOI":"10.3390\/ijms21165924","article-title":"Osteoclasts\u2019 ability to generate trenches rather than pits depends on high levels of active cathepsin k and efficient clearance of resorption products.","volume":"21","author":"Borggaard","year":"2020","journal-title":"Int. J. Mol. Sci."},{"key":"B7","doi-asserted-by":"publisher","first-page":"3301","DOI":"10.1091\/mbc.e04-12-1117","article-title":"Dynamin forms a Src kinase-sensitive complex with Cbl and regulates podosomes and osteoclast activity.","volume":"16","author":"Bruzzaniti","year":"2005","journal-title":"Mol. Biol. Cell"},{"key":"B8","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2019.02182","article-title":"Chemokines in physiological and pathological bone remodeling.","volume":"10","author":"Brylka","year":"2019","journal-title":"Front. Immunol."},{"key":"B9","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1242\/jcs.76.1.155","article-title":"Bone cells predispose bone surfaces to resorption by exposure of mineral to osteoclastic contact.","volume":"76","author":"Chambers","year":"1985","journal-title":"J. Cell Sci."},{"key":"B10","doi-asserted-by":"publisher","first-page":"671","DOI":"10.1007\/BF00214590","article-title":"Mammalian collagenase predisposes bone surfaces to osteoclastic resorption.","volume":"241","author":"Chambers","year":"1985","journal-title":"Cell Tissue Res."},{"key":"B11","doi-asserted-by":"publisher","first-page":"424","DOI":"10.1016\/j.semcdb.2008.08.004","article-title":"Vesicular trafficking in osteoclasts.","volume":"19","author":"Coxon","year":"2008","journal-title":"Semin. Cell Dev. Biol."},{"key":"B12","doi-asserted-by":"publisher","DOI":"10.1016\/j.bone.2020.115628","article-title":"Re-thinking the bone remodeling cycle mechanism and the origin of bone loss.","volume":"141","author":"Delaisse","year":"2020","journal-title":"Bone"},{"key":"B13","doi-asserted-by":"publisher","first-page":"77","DOI":"10.1359\/jbmr.2002.17.1.77","article-title":"The bone lining cell: its role in cleaning Howship\u2019s lacunae and initiating bone formation.","volume":"17","author":"Everts","year":"2002","journal-title":"J. Bone Miner. Res."},{"key":"B14","doi-asserted-by":"publisher","first-page":"1219","DOI":"10.1096\/fasebj.13.10.1219","article-title":"Functional heterogeneity of osteoclasts: matrix metalloproteinases participate in osteoclastic resorption of calvarial bone but not in resorption of long bone.","volume":"13","author":"Everts","year":"1999","journal-title":"FASEB J."},{"key":"B15","doi-asserted-by":"publisher","first-page":"11","DOI":"10.4248\/BR201301003","article-title":"Osteoclasts: new insights.","volume":"1","author":"Feng","year":"2013","journal-title":"Bone Res."},{"key":"B16","doi-asserted-by":"publisher","first-page":"32347","DOI":"10.1074\/jbc.273.48.32347","article-title":"The collagenolytic activity of cathepsin K is unique among mammalian proteinases.","volume":"273","author":"Garnero","year":"1999","journal-title":"J. Biol. Chem."},{"key":"B17","doi-asserted-by":"publisher","first-page":"1503","DOI":"10.1242\/jcs.060954","article-title":"Nano-topography sensing by osteoclasts.","volume":"123","author":"Geblinger","year":"2010","journal-title":"J. Cell Sci."},{"key":"B18","doi-asserted-by":"publisher","first-page":"158","DOI":"10.1002\/cbic.200800549","article-title":"Surface-induced regulation of podosome organization and dynamics in cultured osteoclasts.","volume":"10","author":"Geblinger","year":"2009","journal-title":"ChemBioChem"},{"key":"B19","doi-asserted-by":"publisher","first-page":"192","DOI":"10.4161\/cam.27840","article-title":"Podosome organization drives osteoclast-mediated bone resorption.","volume":"8","author":"Georgess","year":"2014","journal-title":"Cell Adhes. Migr."},{"key":"B20","doi-asserted-by":"publisher","DOI":"10.1038\/bonekey.2013.40","article-title":"Novel perspectives on the transcytotic route in osteoclasts.","volume":"2","author":"Hirvonen","year":"2013","journal-title":"Bonekey Rep."},{"key":"B21","doi-asserted-by":"publisher","first-page":"177","DOI":"10.1097\/00003086-197703000-00062","article-title":"The ultrastructure of the osteoclast and its functional implications.","volume":"123","author":"Holtrop","year":"1977","journal-title":"Clin. Orthop. Relat. Res."},{"key":"B22","doi-asserted-by":"publisher","first-page":"1420","DOI":"10.1172\/JCI39104","article-title":"The basics of epithelial-mesenchymal transition.","volume":"119","author":"Kalluri","year":"2009","journal-title":"J. Clin. Invest."},{"key":"B23","doi-asserted-by":"publisher","first-page":"574","DOI":"10.1002\/jbmr.1760","article-title":"Increased presence of capillaries next to remodeling sites in adult human cancellous bone.","volume":"28","author":"Kristensen","year":"2013","journal-title":"J. Bone Miner. Res."},{"key":"B24","doi-asserted-by":"publisher","first-page":"817","DOI":"10.1002\/jbmr.5650060806","article-title":"Kinetics of the osteoclast cytoskeleton during the resorption cycle in vitro.","volume":"6","author":"Lakkakorpi","year":"1991","journal-title":"J. Bone Miner. Res."},{"key":"B25","doi-asserted-by":"publisher","first-page":"21562","DOI":"10.1074\/jbc.M114.559898","article-title":"Chondroitin sulfate promotes activation of cathepsin K.","volume":"289","author":"Lemaire","year":"2014","journal-title":"J. Biol. Chem."},{"key":"B26","doi-asserted-by":"publisher","first-page":"623","DOI":"10.1016\/j.bone.2011.06.014","article-title":"The effects of the cathepsin K inhibitor odanacatib on osteoclastic bone resorption and vesicular trafficking.","volume":"49","author":"Leung","year":"2011","journal-title":"Bone"},{"key":"B27","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1016\/j.tcb.2007.01.002","article-title":"The matrix corroded: podosomes and invadopodia in extracellular matrix degradation.","volume":"17","author":"Linder","year":"2007","journal-title":"Trends Cell Biol."},{"key":"B28","doi-asserted-by":"publisher","first-page":"2079","DOI":"10.1242\/jcs.02390","article-title":"Podosomes at a glance.","volume":"118","author":"Linder","year":"2005","journal-title":"J. Cell Sci."},{"key":"B29","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0000179","article-title":"The architecture of the adhesive apparatus of cultured osteoclasts: from podosome formation to sealing zone assembly.","volume":"2","author":"Luxenburg","year":"2007","journal-title":"PLoS One"},{"key":"B30","doi-asserted-by":"publisher","first-page":"1666","DOI":"10.1242\/jcs.075903","article-title":"Involvement of actin polymerization in podosome dynamics.","volume":"125","author":"Luxenburg","year":"2012","journal-title":"J. Cell Sci."},{"key":"B31","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.54493.sa2","article-title":"Dissecting the phenotypic and functional heterogeneity of mouse inflammatory osteoclasts by the expression of cx3cr1.","volume":"9","author":"Madel","year":"2020","journal-title":"Elife"},{"key":"B32","doi-asserted-by":"publisher","first-page":"637","DOI":"10.1006\/bbrc.1996.0947","article-title":"Mammalian mature osteoclasts as estrogen target cells.","volume":"223","author":"Mano","year":"1996","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"B33","doi-asserted-by":"publisher","DOI":"10.1038\/boneres.2015.32","article-title":"Pit- and trench-forming osteoclasts: a distinction that matters.","volume":"3","author":"Merrild","year":"2015","journal-title":"Bone Res."},{"key":"B34","doi-asserted-by":"publisher","first-page":"104","DOI":"10.1083\/jcb.75.1.104","article-title":"Osteoclast cell-surface changes during the egg-laying cycle in Japanese quail.","volume":"75","author":"Miller","year":"1977","journal-title":"J. Cell Bol."},{"key":"B35","doi-asserted-by":"publisher","DOI":"10.1002\/jbm4.10412","article-title":"Zoledronic acid is not equally potent on osteoclasts generated from different individuals.","volume":"4","author":"M\u00f8ller","year":"","journal-title":"JBMR Plus"},{"key":"B36","doi-asserted-by":"publisher","DOI":"10.3390\/ijms21176368","article-title":"Fusion potential of human osteoclasts in vitro reflects age, menopause, and in vivo bone resorption levels of their donors\u2014a possible involvement of DC-STAMP.","volume":"21","author":"M\u00f8ller","year":"","journal-title":"Int. J. Mol. Sci."},{"key":"B37","doi-asserted-by":"publisher","DOI":"10.1038\/s41413-020-0102-7","article-title":"Aging and menopause reprogram osteoclast precursors for aggressive bone resorption.","volume":"8","author":"M\u00f8ller","year":"","journal-title":"Bone Res."},{"key":"B38","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-018-31159-1","article-title":"Septins are critical regulators of osteoclastic bone resorption.","volume":"8","author":"M\u00f8ller","year":"2018","journal-title":"Sci. Rep."},{"key":"B39","doi-asserted-by":"publisher","first-page":"1641","DOI":"10.1016\/j.yexcr.2008.02.011","article-title":"Recombinant VSV G proteins reveal a novel raft-dependent endocytic pathway in resorbing osteoclasts.","volume":"314","author":"Mulari","year":"2008","journal-title":"Exp. Cell Res."},{"key":"B40","doi-asserted-by":"publisher","first-page":"113","DOI":"10.1034\/j.1600-0854.2003.40206.x","article-title":"Osteoclast ruffled border has distinct subdomains for secretion and degraded matrix uptake.","volume":"4","author":"Mulari","year":"2003","journal-title":"Traffic"},{"key":"B41","doi-asserted-by":"publisher","first-page":"48","DOI":"10.1016\/j.bone.2003.09.001","article-title":"Immunolocalization of matrix metalloproteinase-13 on bone surface under osteoclasts in rat tibia.","volume":"34","author":"Nakamura","year":"2004","journal-title":"Bone"},{"key":"B42","doi-asserted-by":"publisher","first-page":"266","DOI":"10.1126\/science.276.5310.266","article-title":"Trafficking of matrix collagens through bone-resorbing osteoclasts.","volume":"276","author":"Nesbitt","year":"1997","journal-title":"Science"},{"key":"B43","doi-asserted-by":"publisher","first-page":"639","DOI":"10.1042\/BST20180445","article-title":"Membrane trafficking in osteoclasts and implications for osteoporosis.","volume":"47","author":"Ng","year":"2019","journal-title":"Biochem. Soc. Trans."},{"key":"B44","doi-asserted-by":"publisher","first-page":"773","DOI":"10.1111\/jre.12068","article-title":"Fundamental study of osteoclast chemotaxis toward chemoattractants expressed in periodontitis.","volume":"48","author":"Niwa","year":"2013","journal-title":"J. Periodontal. Res."},{"key":"B45","doi-asserted-by":"publisher","first-page":"400","DOI":"10.1007\/s002230001151","article-title":"Morphological and functional features of clasts in low phosphate, vitamin D-deficiency rickets.","volume":"67","author":"Nordahl","year":"2000","journal-title":"Calcif. Tissue Int."},{"key":"B46","doi-asserted-by":"publisher","first-page":"54","DOI":"10.1016\/j.arr.2009.09.005","article-title":"Osteoclast motility: putting the brakes on bone resorption.","volume":"10","author":"Novack","year":"2011","journal-title":"Ageing Res. Rev."},{"key":"B47","doi-asserted-by":"crossref","first-page":"1767","DOI":"10.1242\/jcs.110.15.1767","article-title":"Endocytic pathway from the basal plasma membrane to the ruffled border membrane in bone-resorbing osteoclasts.","volume":"110","author":"Palokangas","year":"1997","journal-title":"J. Cell Sci."},{"key":"B48","doi-asserted-by":"publisher","first-page":"396","DOI":"10.1111\/bph.13383","article-title":"A novel approach to inhibit bone resorption: exosite inhibitors against cathepsin K.","volume":"173","author":"Panwar","year":"2016","journal-title":"Br. J. Pharmacol."},{"key":"B49","doi-asserted-by":"publisher","first-page":"2415","DOI":"10.1002\/jbmr.3227","article-title":"An ectosteric inhibitor of cathepsin K inhibits bone resorption in ovariectomized mice.","volume":"32","author":"Panwar","year":"2017","journal-title":"J. Bone Miner. Res."},{"key":"B50","doi-asserted-by":"publisher","first-page":"5371","DOI":"10.1210\/endo.142.12.8533","article-title":"Estrogen reduces the depth of resorption pits by disturbing the organic bone matrix degradation activity of mature osteoclasts.","volume":"142","author":"Parikka","year":"2001","journal-title":"Endocrinology"},{"key":"B51","first-page":"63","article-title":"Attachment to and phagocytosis of mineral by alveolar bone osteoclasts.","volume":"21","author":"Pierce","year":"1989","journal-title":"J. Submicrosc. Cytol. Pathol."},{"key":"B52","doi-asserted-by":"publisher","first-page":"234","DOI":"10.1038\/nrd3669","article-title":"Inflammatory bone loss: pathogenesis and therapeutic intervention.","volume":"11","author":"Redlich","year":"2012","journal-title":"Nat. Rev. Drug Discov."},{"key":"B53","doi-asserted-by":"publisher","first-page":"477","DOI":"10.1006\/excr.1999.4585","article-title":"Extensive clear zone and defective ruffled border formation in osteoclasts of osteopetrotic (ia\/ia) rats: implications for secretory function.","volume":"251","author":"Reinholt","year":"1999","journal-title":"Exp. Cell Res."},{"key":"B54","doi-asserted-by":"publisher","first-page":"34","DOI":"10.1016\/j.bone.2010.11.008","article-title":"Biochemical and molecular mechanisms of action of bisphosphonates.","volume":"49","author":"Rogers","year":"2011","journal-title":"Bone"},{"key":"B55","doi-asserted-by":"publisher","first-page":"526","DOI":"10.1007\/s00223-013-9786-7","article-title":"Osteoclasts on bone and dentin in vitro: mechanism of trail formation and comparison of resorption behavior.","volume":"93","author":"Rumpler","year":"2013","journal-title":"Calcif. Tissue Int."},{"key":"B56","doi-asserted-by":"publisher","first-page":"13453","DOI":"10.1073\/pnas.95.23.13453","article-title":"Impaired osteoclastic bone resorption leads to osteopetrosis in cathepsin-K-deficient mice.","volume":"95","author":"Saftig","year":"1998","journal-title":"Proc. Natl. Acad. Sci. U S A."},{"key":"B57","doi-asserted-by":"publisher","first-page":"270","DOI":"10.1126\/science.276.5310.270","article-title":"Removal of osteoclast bone resorption products by transcytosis.","volume":"276","author":"Salo","year":"1997","journal-title":"Science"},{"key":"B58","doi-asserted-by":"crossref","first-page":"301","DOI":"10.1242\/jcs.109.2.301","article-title":"Bone-resorbing osteoclasts reveal a dynamic division of basal plasma membrane into two different domains.","volume":"109","author":"Salo","year":"1996","journal-title":"J. Cell Sci."},{"key":"B59","doi-asserted-by":"publisher","first-page":"459","DOI":"10.1016\/j.ejcb.2008.01.001","article-title":"Actin cytoskeletal organisation in osteoclasts: a model to decipher transmigration and matrix degradation.","volume":"87","author":"Saltel","year":"2008","journal-title":"Eur. J. Cell Biol."},{"key":"B60","doi-asserted-by":"publisher","first-page":"5231","DOI":"10.1091\/mbc.e04-06-0522","article-title":"Apatite-mediated actin dynamics in resorbing osteoclasts.","volume":"15","author":"Saltel","year":"2004","journal-title":"Mol. Biol. Cell"},{"key":"B61","doi-asserted-by":"publisher","first-page":"379","DOI":"10.1002\/ar.1092240307","article-title":"Multinucleated cells formed on calcified dentine from mouse bone marrow cells treated with 1\u03b1,25-dihydroxyvitamin D3 have ruffled borders and resorb dentine.","volume":"224","author":"Sasaki","year":"1989","journal-title":"Anat. Rec."},{"key":"B62","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1359\/jbmr.1998.13.1.59","article-title":"The migration of purified osteoclasts through collagen is inhibited by matrix metalloproteinase inhibitors.","volume":"13","author":"Sato","year":"1998","journal-title":"J. Bone Miner. Res."},{"key":"B63","doi-asserted-by":"publisher","first-page":"14932","DOI":"10.1021\/acsami.5b08126","article-title":"Study of osteoclast adhesion to cortical bone surfaces: a correlative microscopy approach for concomitant imaging of cellular dynamics and surface modifications.","volume":"8","author":"Shemesh","year":"2016","journal-title":"ACS Appl. Mater. Interfaces"},{"key":"B64","doi-asserted-by":"publisher","first-page":"2184","DOI":"10.1002\/jbmr.113","article-title":"Glucocorticoids maintain human osteoclasts in the active mode of their resorption cycle.","volume":"25","author":"S\u00f8e","year":"2010","journal-title":"J. Bone Miner. Res."},{"key":"B65","doi-asserted-by":"publisher","first-page":"2026","DOI":"10.1242\/jcs.202036","article-title":"Time-lapse reveals that osteoclasts can move across the bone surface while resorbing.","volume":"130","author":"S\u00f8e","year":"2017","journal-title":"J. Cell Sci."},{"key":"B66","doi-asserted-by":"publisher","first-page":"191","DOI":"10.1016\/j.bone.2013.06.007","article-title":"Steering the osteoclast through the demineralization-collagenolysis balance.","volume":"56","author":"S\u00f8e","year":"2013","journal-title":"Bone"},{"key":"B67","doi-asserted-by":"publisher","first-page":"285","DOI":"10.1016\/S1084952102000587","article-title":"Formation and function of the ruffled border in osteoclasts.","volume":"13","author":"Stenbeck","year":"2002","journal-title":"Semin. Cell Dev. Biol."},{"key":"B68","doi-asserted-by":"crossref","first-page":"1577","DOI":"10.1242\/jcs.113.9.1577","article-title":"A new specialized cell-matrix interaction in actively resorbing osteoclasts.","volume":"113","author":"Stenbeck","year":"2000","journal-title":"J. Cell Sci."},{"key":"B69","doi-asserted-by":"publisher","first-page":"827","DOI":"10.1242\/jcs.00935","article-title":"Endocytic trafficking in actively resorbing osteoclasts.","volume":"117","author":"Stenbeck","year":"2004","journal-title":"J. Cell Sci."},{"key":"B70","doi-asserted-by":"publisher","first-page":"3613","DOI":"10.1210\/en.2005-0717","article-title":"Glucocorticoid regulation of osteoclast differentiation and expression of receptor activator of nuclear factor-\u03baB (NF-\u03baB) ligand, osteoprotegerin, and receptor activator of NF-\u03baB in mouse calvarial bones.","volume":"147","author":"Swanson","year":"2006","journal-title":"Endocrinology"},{"key":"B71","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/s10266-007-0071-y","article-title":"Regulation of osteoclast polarization.","volume":"95","author":"Takahashi","year":"2007","journal-title":"Odontology"},{"key":"B72","doi-asserted-by":"publisher","first-page":"2882","DOI":"10.1091\/mbc.e06-03-0187","article-title":"Cortactin has an essential and specific role in osteoclast actin assembly.","volume":"17","author":"Tehrani","year":"2006","journal-title":"Mol. Biol. Cell"},{"key":"B73","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1242\/jcs.113.3.377","article-title":"The cell biology of osteoclast function.","volume":"113","author":"V\u00e4\u00e4n\u00e4nen","year":"2000","journal-title":"J. Cell Sci."},{"key":"B74","doi-asserted-by":"publisher","DOI":"10.1242\/jcs.236828","article-title":"Probing the mechanical landscape \u2013 new insights into podosome architecture and mechanics.","volume":"132","author":"van den Dries","year":"2019","journal-title":"J. Cell Sci."},{"key":"B75","doi-asserted-by":"publisher","first-page":"919","DOI":"10.1172\/JCI30328","article-title":"Involvement of PLEKHM1 in osteoclastic vesicular transport and osteopetrosis in incisors absent rats and humans.","volume":"117","author":"Van Wesenbeeck","year":"2007","journal-title":"J. Clin. Invest."},{"key":"B76","doi-asserted-by":"publisher","first-page":"240","DOI":"10.1007\/s00223-012-9674-6","article-title":"Glucocorticoid-induced changes in the geometry of osteoclast resorption cavities affect trabecular bone stiffness.","volume":"92","author":"Vanderoost","year":"2013","journal-title":"Calcif. Tissue Int."},{"key":"B77","doi-asserted-by":"publisher","first-page":"1099","DOI":"10.1002\/jbmr.282","article-title":"The Rac1 exchange factor Dock5 is essential for bone resorption by osteoclasts.","volume":"26","author":"Vives","year":"2011","journal-title":"J. Bone Miner. Res."},{"key":"B78","doi-asserted-by":"publisher","first-page":"495","DOI":"10.1111\/j.1600-0854.2011.01292.x","article-title":"At the crossroads of chemistry and cell biology: inhibiting membrane traffic by small molecules.","volume":"13","author":"von Kleist","year":"2012","journal-title":"Traffic"},{"key":"B79","doi-asserted-by":"publisher","first-page":"725","DOI":"10.1002\/jbmr.2068","article-title":"Subcellular elevation of cytosolic free calcium is required for osteoclast migration.","volume":"29","author":"Wheal","year":"2014","journal-title":"J. Bone Miner. Res."},{"key":"B80","doi-asserted-by":"publisher","first-page":"1467","DOI":"10.1007\/s10856-009-3718-0","article-title":"Biomimetic hydroxyapatite coating on glass coverslips for the assay of osteoclast activity in vitro.","volume":"20","author":"Wijenayaka","year":"2009","journal-title":"J. Mater. Sci. Mater. Med."},{"key":"B81","doi-asserted-by":"publisher","first-page":"2584","DOI":"10.1074\/jbc.M805280200","article-title":"Cathepsin K activity-dependent regulation of osteoclast actin ring formation and bone resorption.","volume":"284","author":"Wilson","year":"2009","journal-title":"J. Biol. Chem."},{"key":"B82","doi-asserted-by":"publisher","first-page":"517","DOI":"10.1002\/jbmr.5650030507","article-title":"The distribution of podosomes in osteoclasts cultured on bone laminae: effect of retinol.","volume":"3","author":"Zambonin-Zallone","year":"1988","journal-title":"J. Bone Miner. Res."},{"key":"B83","doi-asserted-by":"publisher","first-page":"39295","DOI":"10.1074\/jbc.M010999200","article-title":"Downregulation of small GTPase Rab7 impairs osteoclast polarization and bone resorption.","volume":"276","author":"Zhao","year":"2001","journal-title":"J. Biol. Chem."}],"container-title":["Frontiers in Cell and Developmental Biology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fcell.2021.644503\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,4,30]],"date-time":"2021-04-30T17:35:18Z","timestamp":1619804118000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fcell.2021.644503\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,3,30]]},"references-count":83,"alternative-id":["10.3389\/fcell.2021.644503"],"URL":"https:\/\/doi.org\/10.3389\/fcell.2021.644503","relation":{},"ISSN":["2296-634X"],"issn-type":[{"value":"2296-634X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,3,30]]},"article-number":"644503"}}