55:
1187:
311:
322:
388:, the "ruffled border", that opposes the surface of the bone tissue. This extensively folded or ruffled border facilitates bone removal by dramatically increasing the cell surface for secretion and uptake of the resorption compartment contents and is a morphologic characteristic of an osteoclast that is actively resorbing bone.
40:
287:
subosteoclastic compartment. These enzymes digest and degrade collagen and other organic components of decalcified bone matrix. The degradation products are phagocytosed by osteoclasts at the ruffled border. Because of their phagocytic properties, osteoclasts are considered to be a component of the
731:
In the 1980s and 90s the physiology of typical osteoclasts was studied in detail. With the isolation of the ruffled border, ion transport across it was studied directly in biochemical detail. Energy-dependent acid transport was verified and the postulated proton pump purified. With the successful
337:
to osteoclasts, very large cells that may reach 100 μm in diameter occur. These may have dozens of nuclei, and typically express major osteoclast proteins but have significant differences from cells in living bone because of the not-natural substrate. The size of the multinucleated assembled
732:
culture of osteoclasts, it became apparent that they are organized to support the massive transport of protons for acidification of the resorption compartment and solubilization of the bone mineral. This includes ruffled border Cl permeability to control membrane potential and basolateral Cl/HCO
286:
and thus create an acidic microenvironment, which increases solubility of bone mineral, resulting in the release and re-entry of bone minerals into the cytoplasm of osteoclasts to be delivered to nearby capillaries. After the removal of minerals, collagenase and gelatinase are secreted into the
269:. The actin filaments enable the cell membrane surrounding the sealing zone to be anchored firmly to the bony wall of Howship's lacunae. In this way, a closed subosteoclastic compartment is created between the ruffled border and the bone that is undergoing resorption. The osteoclasts secrete
739:
The effectiveness of its ion secretion depends upon the osteoclast forming an effective seal around the resorption compartment. The positioning of this "sealing zone" appears to be mediated by integrins expressed on the osteoclast surface. With the sealing zone in place, the multinucleated
744:
of the mineral and degraded collagen from the ruffled border to the free membrane of the cell, and its release into the extracellular compartment. This activity completes the bone resorption, and both the mineral components and collagen fragments are released to the general circulation.
608:
disease, characterised by a lack of functional cathepsin K expression. Knockout studies of cathepsin K in mice lead to an osteopetrotic phenotype, which, is partially compensated by increased expression of proteases other that cathepsin K and enhanced osteoclastogenesis.
281:
and hydrolytic enzymes into this compartment. Resorption of bone matrix by the osteoclasts involves two steps: (1) dissolution of inorganic components (minerals), and (2) digestion of organic component of the bone matrix. The osteoclasts pump hydrogen ions into
612:
Cathepsin K has an optimal enzymatic activity in acidic conditions. It is synthesized as a proenzyme with a molecular weight of 37kDa, and upon activation by autocatalytic cleavage, is transformed into the mature, active form with a molecular weight of ~27kDa.
616:
Upon polarization of the osteoclast over the site of resorption, cathepsin K is secreted from the ruffled border into the resorptive pit. Cathepsin K transmigrates across the ruffled border by intercellular vesicles and is then released by the functional
695:(MMPs) comprise a family of more than 20 zinc-dependent endopeptidases. The role of matrix metalloproteinases (MMPs) in osteoclast biology is ill-defined, but in other tissue they have been linked with tumor promoting activities, such as activation of
291:(MPS). The activity of osteoclasts is controlled by hormones and cytokines. Calcitonin, a hormone of thyroid gland, suppresses the osteoclastic activity. The osteoclasts do not have receptors for parathyroid hormone (PTH). However, PTH stimulates the
461:. NF-κβ activation is stimulated almost immediately after RANKL-RANK interaction occurs and is not upregulated. NFATc1 stimulation, however, begins ~24–48 hours after binding occurs and its expression has been shown to be RANKL dependent.
401:
are of the same lineage, and osteoblasts fuse together to form osteoclasts. After years of controversy it is now clear that these cells develop from the self fusion of macrophages. It was in the beginning of 1980 that the
456:
and defects of tooth eruption, along with an absence or deficiency of osteoclasts. RANKL activates NF-κβ (nuclear factor-κβ) and NFATc1 (nuclear factor of activated t cells, cytoplasmic, calcineurin-dependent 1) through
241:. The resorption bays are created by the erosive action of osteoclasts on the underlying bone. The border of the lower part of an osteoclast exhibits finger-like processes due to the presence of deep infoldings of the
468:(OPG), which is produced by osteoblasts and binds to RANKL thereby preventing interaction with RANK. While osteoclasts are derived from the hematopoietic lineage, osteoblasts are derived from mesenchymal stem cells.
396:
Since their discovery in 1873 there has been considerable debate about their origin. Three theories were dominant: from 1949 to 1970 the connective tissue origin was popular, which stated that osteoclasts and
722:
MMPs expressed by the osteoclast include MMP-9, -10, -12, and -14. apart from MMP-9, little is known about their relevance to the osteoclast, however, high levels of MMP-14 are found at the sealing zone.
740:
osteoclast reorganizes itself. Developing the highly invaginated ruffled membrane apposing the resorption compartment allows massive secretory activity. In addition, it permits the vesicular
554:, water and other substances. Dysfunction of the carbonic anhydrase has been documented to cause some forms of osteopetrosis. Hydrogen ions are pumped against a high concentration gradient by
1935:
719:
is believed to be involved in bone resorption and in osteoclast differentiation, as knockout mice revealed decreased osteoclast numbers, osteopetrosis, and decreased bone resorption.
871:
876:
1045:
Basle MF, Mazaud P, Malkani K, Chretien MF, Moreau MF, Rebel A (1988). "Isolation of osteoclasts from
Pagetic bone tissue: morphometry and cytochemistry on isolated cells".
1560:
Blair HC, Teitelbaum SL, Tan HL, Koziol CM, Schlesinger PH (June 1991). "Passive chloride permeability charge coupled to H(+)-ATPase of avian osteoclast ruffled membrane".
480:. Osteoclasts lie in small cavities called Howship's lacunae, formed from the digestion of the underlying bone. The sealing zone is the attachment of the osteoclast's
237:
Osteoclasts are found on those surfaces of bone that are undergoing resorption. On such surfaces, the osteoclasts are seen to be located in shallow depressions called
824:: broken). To avoid confusion, the cell was originally termed osotoclast. When the surgical instrument went out of use, the cell became known by its present name.
147:
2524:
1928:
249:. The ruffled border lies in contact with the bone surface within a resorption bay. The periphery of the ruffled border is surrounded by a ring-like zone of
2442:
2055:
777:. Osteoporosis occurs when there is an imbalance between the bone resorption activities of osteoclasts and the bone formation activities of osteoblasts.
574:
and matrix metalloprotease (MMP) groups, are released to digest the organic components of the matrix. These enzymes are released into the compartment by
338:
osteoclast allows it to focus the ion transport, protein secretory and vesicular transport capabilities of many macrophages on a localized area of bone.
333:
and human osteoclasts on bone typically have four nuclei and are 150–200 μm in diameter. When osteoclast-inducing cytokines are used to convert
1921:
1149:
844:
986:
1002:
Piper K, Boyde A, Jones SJ (1992). "The relationship between the number of nuclei of an osteoclast and its resorptive capability in vitro".
430:
411:
1876:
353:. Osteoclasts are characterized by a cytoplasm with a homogeneous, "foamy" appearance. This appearance is due to a high concentration of
626:
600:
involved in the degradation of type I collagen and other noncollagenous proteins. Mutations in the cathepsin K gene are associated with
374:
314:
1738:
1667:
Salo J, Lehenkari P, Mulari M, Metsikkö K, Väänänen HK (April 1997). "Removal of osteoclast bone resorption products by transcytosis".
2517:
154:
1427:
Blair HC, Teitelbaum SL, Ghiselli R, Gluck S (August 1989). "Osteoclastic bone resorption by a polarized vacuolar proton pump".
54:
441:
activation of tyrosine kinase Src. Both of these molecules are necessary for osteoclastogenesis and are widely involved in the
47:
of an osteoclast displaying typical distinguishing characteristics: a large cell with multiple nuclei and a "foamy" cytosol.
1290:
Nijweide PJ, Burger EH, Feyen JH (October 1986). "Cells of bone: proliferation, differentiation, and hormonal regulation".
295:
to secrete the cytokine called osteoclast-stimulating factor, which is a potent stimulator of the osteoclastic activity.
142:
850:
Osteoclasts play a major role in orthodontic tooth movement and pathologic migration of periodontally compromised teeth.
705:
is associated with the bone microenvironment. It is expressed by osteoclasts, and is known to be required for osteoclast
2630:
2325:
2133:
403:
288:
2510:
1904:
833:
442:
2635:
1511:
Teti A, Blair HC, Teitelbaum SL, Kahn AJ, Koziol C, Konsek J, Zambonin-Zallone A, Schlesinger PH (January 1989).
692:
976:
2907:
2387:
1889:
130:
118:
709:
and is a powerful gelatinase. Transgenic mice lacking MMP-9 develop defects in bone development, intraosseous
298:
An odontoclast (/odon·to·clast/; o-don´to-klast) is an osteoclast associated with absorption of the roots of
2489:
2408:
2277:
622:
596:
that is mainly expressed in osteoclasts, and is secreted into the resorptive pit. Cathepsin K is the major
1702:
Nesbitt SA, Horton MA (April 1997). "Trafficking of matrix collagens through bone-resorbing osteoclasts".
859:
449:
176:
780:
Osteoclast activity is also mediated by the interaction of two molecules produced by osteoblasts, namely
2558:
423:
1636:
Väänänen HK, Zhao H, Mulari M, Halleen JM (February 2000). "The cell biology of osteoclast function".
683:
cathepsin L knockout mice compared to wild-type and another report finding no skeletal abnormalities.
135:
2902:
2897:
2784:
2484:
1436:
1254:
590:
488:. Attachment to the bone matrix is facilitated by integrin receptors, such as αvβ3, via the specific
484:
to the underlying bone. Sealing zones are bounded by belts of specialized adhesion structures called
1597:"Characterization of the osteoclast ruffled border chloride channel and its role in bone resorption"
2806:
2272:
2247:
758:
438:
2816:
2418:
1179:
1027:
661:
593:
497:
350:
1775:"RANK ligand and osteoprotegerin: paracrine regulators of bone metabolism and vascular function"
664:
is generally unknown within bone, and they are expressed at much lower levels than cathepsin K.
346:
In bone, osteoclasts are found in pits in the bone surface which are called resorption bays, or
1873:
2591:
2586:
2282:
1853:
1835:
1796:
1719:
1684:
1649:
1618:
1577:
1542:
1493:
1452:
1409:
1358:
1307:
1270:
1227:
1171:
1111:
1062:
1019:
982:
954:
907:
816:
An osteoclast can also be an instrument used to fracture and reset bones (the origin is Greek
657:
567:
489:
354:
2831:
2821:
2734:
2359:
2262:
2077:
1843:
1827:
1786:
1746:
1711:
1676:
1641:
1608:
1569:
1532:
1524:
1483:
1444:
1399:
1389:
1348:
1338:
1299:
1262:
1217:
1161:
1101:
1093:
1054:
1011:
944:
531:
501:
366:
2779:
2738:
2308:
2097:
2029:
1880:
1378:"Impact of Air Pollutants on Oxidative Stress in Common Autophagy-Mediated Aging Diseases"
805:
801:
781:
672:
601:
465:
434:
406:
was recognized as precursor of osteoclasts. Osteoclast formation requires the presence of
370:
299:
226:
202:
1470:
Mattsson JP, Schlesinger PH, Keeling DJ, Teitelbaum SL, Stone DK, Xie XS (October 1994).
1848:
1815:
1440:
1258:
381:. Osteoclast rough endoplasmic reticulum is sparse, and the Golgi complex is extensive.
2867:
2596:
2374:
2354:
2303:
2147:
2092:
2072:
2067:
1989:
1953:
1885:
1831:
1791:
1774:
1404:
1377:
1353:
1326:
1131:
1106:
1081:
837:
706:
696:
509:
1537:
1512:
1488:
1471:
530:
into the resorptive cavity, acidifying and aiding dissolution of the mineralized bone
452:), and is essential in osteoclastogenesis. RANKL knockout mice exhibit a phenotype of
310:
2891:
2872:
2553:
2454:
2364:
2009:
1573:
1472:"Isolation and reconstitution of a vacuolar-type proton pump of osteoclast membranes"
1166:
1058:
949:
932:
770:
668:
605:
481:
453:
415:
385:
330:
258:
242:
1183:
2677:
2625:
2617:
2571:
2479:
2413:
2382:
2242:
2107:
1999:
1962:
1945:
1031:
774:
741:
710:
680:
653:
563:
523:
347:
270:
1513:"Cytoplasmic pH regulation and chloride/bicarbonate exchange in avian osteoclasts"
1303:
1266:
1913:
1715:
1680:
1082:"Giant osteoclasts after long-term bisphosphonate therapy: diagnostic challenges"
123:
2836:
2576:
2156:
2039:
766:
649:
645:
641:
637:
633:
586:
555:
516:
493:
419:
378:
292:
278:
274:
221:
216:. The osteoclast disassembles and digests the composite of hydrated protein and
198:
94:
1325:
Michou, Laëtitia; Numan, Mohamed; Amiable, Nathalie; Brown, Jacques P. (2015).
2857:
2799:
2758:
2743:
2651:
2563:
2267:
2237:
2229:
2191:
2174:
2164:
2102:
1994:
1895:
1751:
762:
676:
477:
398:
334:
210:
70:
44:
1839:
1613:
1596:
933:"Osteoclasts and odontoclasts: signaling pathways to development and disease"
2862:
2852:
2826:
2794:
2748:
2729:
2724:
2656:
2545:
2537:
2474:
2320:
2252:
2203:
2169:
2034:
2014:
2004:
1448:
1394:
1097:
571:
254:
250:
194:
1857:
1800:
1653:
1413:
1362:
1274:
1231:
1115:
958:
17:
1723:
1688:
1645:
1622:
1581:
1546:
1497:
1456:
1311:
1222:
1205:
1066:
1023:
476:
Once activated, osteoclasts move to areas of microfracture in the bone by
369:. This permits characterization of osteoclasts by their staining for high
2811:
2447:
2341:
2220:
2208:
2184:
2179:
1967:
1900:
1816:"Topography-mediated immunomodulation in osseointegration; Ally or Enemy"
1175:
754:
597:
575:
559:
485:
362:
358:
213:
1343:
736:
exchange to maintain cytosolic pH in physiologically acceptable ranges.
384:
At a site of active bone resorption, the osteoclast forms a specialized
321:
2753:
2687:
2349:
2196:
1015:
535:
231:
217:
39:
1528:
1150:"The ultrastructure of the osteoclast and its functional implications"
160:
2774:
2682:
2502:
2437:
2313:
1977:
716:
788:. These molecules also regulate differentiation of the osteoclast.
422:, thus requiring direct contact between these cells and osteoclast
2392:
785:
578:. Of these hydrolytic enzymes, cathepsin K is of most importance.
407:
320:
309:
106:
1595:
Schlesinger PH, Blair HC, Teitelbaum SL, Edwards JC (July 1997).
1382:
International
Journal of Environmental Research and Public Health
2708:
2703:
2608:
2533:
702:
632:
Several other cathepsins are expressed in osteoclasts including
458:
206:
83:
2506:
1917:
621:. Within these intercellular vesicles, cathepsin K, along with
1814:
Shirazi, Sajjad; Ravindran, Sriram; Cooper, Lyndon F. (2022).
496:. The osteoclast releases hydrogen ions through the action of
414:. These membrane-bound proteins are produced by neighbouring
201:. This function is critical in the maintenance, repair, and
1245:
S. L. Teitelbaum (2000), "Bone
Resorption by Osteoclasts",
1204:
Väänänen HK, Zhao H, Mulari M, Halleen JM (February 2000).
1327:"Paget's disease of bone: An osteoimmunological disorder?"
429:
M-CSF acts through its receptor on the osteoclast, c-fms (
1376:
Numan, Mohamed; Brown, Jacques; Michou, Laëtitia (2015).
894:
Medical
Histology by Laiq Hussain Siddiqui (6th Edition)
832:
Giant osteoclasts can occur in some diseases, including
699:
and are required for tumor metastasis and angiogenesis.
230:. This process also helps regulate the level of blood
902:
900:
872:
List of human cell types derived from the germ layers
562:. This enzyme has been targeted in the prevention of
410:(receptor activator of nuclear factor κβ ligand) and
1773:
Schoppet M, Preissner KT, Hofbauer LC (April 2002).
325:
Illustrated cross-section of an activated osteoclast
2845:
2767:
2717:
2696:
2670:
2644:
2616:
2607:
2544:
2467:
2427:
2401:
2373:
2340:
2293:
2228:
2219:
2155:
2146:
2126:
2054:
1976:
1961:
1952:
1739:"Fun facts about bones: More than just scaffolding"
877:
List of distinct cell types in the adult human body
141:
129:
117:
105:
100:
89:
79:
69:
64:
32:
1779:Arteriosclerosis, Thrombosis, and Vascular Biology
1130:
847:, necessitating extraction of the affected teeth.
629:, further degrades the bone extracellular matrix.
843:In cats, abnormal odontoclast activity can cause
448:RANKL is a member of the tumour necrosis family (
1285:
1283:
2518:
1929:
1909:on the role of osteoclasts in bone remodeling
492:Arg-Gly-Asp in bone matrix proteins, such as
220:at a molecular level by secreting acid and a
8:
412:M-CSF (Macrophage colony-stimulating factor)
981:. Jaypee Brothers Publishers. p. 155.
464:Osteoclast differentiation is inhibited by
2613:
2525:
2511:
2503:
2225:
2152:
2060:
1982:
1973:
1958:
1936:
1922:
1914:
1154:Clinical Orthopaedics and Related Research
671:have been mixed, with a report of reduced
1888:at the U.S. National Library of Medicine
1847:
1790:
1750:
1612:
1536:
1487:
1403:
1393:
1352:
1342:
1221:
1206:"The cell biology of osteoclast function"
1165:
1105:
948:
59:Illustration showing a single osteoclast
2443:Megakaryocyte–erythroid progenitor cell
887:
845:feline odontoclastic resorptive lesions
773:, is one of the factors in the disease
445:of monocyte/macrophage derived cells.
158:
29:
1517:The Journal of Clinical Investigation
753:Osteoclasts are regulated by several
7:
1331:Drug Design, Development and Therapy
970:
968:
926:
924:
431:colony-stimulating factor 1 receptor
1601:The Journal of Biological Chemistry
1476:The Journal of Biological Chemistry
910:. Farlex, The Free Dictionary. 2007
375:tartrate resistant acid phosphatase
317:positive osteoclast in cell culture
315:Tartrate resistant acid phosphatase
239:resorption bays (Howship's lacunae)
1832:10.1016/j.biomaterials.2022.121903
1792:10.1161/01.ATV.0000012303.37971.DA
1562:The American Journal of Physiology
1193:from the original on Mar 25, 2024.
931:Wang Z, McCauley LK (March 2011).
761:(PTH) from the parathyroid gland,
25:
1907:Random42 Scientific Communication
1737:Dance, Amber (23 February 2022).
800:is an osteoclast associated with
27:Cell that breaks down bone tissue
1574:10.1152/ajpcell.1991.260.6.C1315
1167:10.1097/00003086-197703000-00062
975:Chatterjee K (1 December 2006).
950:10.1111/j.1601-0825.2010.01718.x
582:Cathepsin K and other cathepsins
155:Anatomical terms of microanatomy
53:
38:
858:Osteoclasts were discovered by
1:
1489:10.1016/S0021-9258(17)31485-0
1304:10.1152/physrev.1986.66.4.855
1267:10.1126/science.289.5484.1504
1080:Jain N, Weinstein RS (2009).
765:from the thyroid gland, and
193: 'broken') is a type of
2134:Extramedullary hematopoiesis
1716:10.1126/science.276.5310.266
1681:10.1126/science.276.5310.270
1148:Holtrop ME, King GJ (1977).
1059:10.1016/8756-3282(88)90020-8
978:Essentials of Oral Histology
289:mononuclear phagocyte system
1640:. 113 ( Pt 3) (3): 377–81.
769:(IL-6). This last hormone,
767:growth factor interleukin 6
284:subosteoclastic compartment
2924:
1137:(39th ed.). Elsevier.
404:monocyte phagocytic system
2300:Antigen-presenting cells
2088:
2063:
2025:
1985:
1752:10.1146/knowable-022222-1
1129:Standring S, ed. (2005).
693:matrix metalloproteinases
687:Matrix metalloproteinases
656:. The function of these
570:, such as members of the
361:. These vacuoles include
329:An osteoclast is a large
186: 'bone' and
153:
52:
37:
2388:Nucleated red blood cell
1944:Myeloid blood cells and
1890:Medical Subject Headings
1614:10.1074/jbc.272.30.18636
558:, specifically a unique
253:which is devoid of cell
245:; this border is called
2490:Hematopoietic stem cell
2409:Leukocyte extravasation
2278:Foreign-body giant cell
1638:Journal of Cell Science
1449:10.1126/science.2528207
1395:10.3390/ijerph120202289
1210:Journal of Cell Science
1098:10.1038/nrrheum.2009.87
834:Paget's disease of bone
802:absorption of the roots
713:, and fracture repair.
667:Studies on cathepsin L
623:reactive oxygen species
566:. In addition, several
1896:The Life of Osteoclast
1004:Anatomy and Embryology
437:-receptor, leading to
326:
318:
261:. This zone is called
75:osteoclast progenitors
2559:fibrocartilage callus
1646:10.1242/jcs.113.3.377
1292:Physiological Reviews
1223:10.1242/jcs.113.3.377
828:Clinical significance
812:Alternate use of term
727:Osteoclast physiology
324:
313:
224:, a process known as
2485:Hematopoietic system
2273:Langhans giant cells
1568:(6 Pt 1): C1315-24.
589:is a collagenolytic
2248:Alveolar macrophage
1441:1989Sci...245..855B
1344:10.2147/DDDT.S88845
1259:2000Sci...289.1504T
1253:(5484): 1504–1508,
759:parathyroid hormone
439:secondary messenger
433:), a transmembrane
331:multinucleated cell
2419:Intrinsic immunity
2283:Touton giant cells
1879:2013-05-25 at the
1016:10.1007/BF00185977
662:aspartic proteases
568:hydrolytic enzymes
498:carbonic anhydrase
327:
319:
136:H2.00.03.7.00005
2885:
2884:
2881:
2880:
2790:connective tissue
2785:Volkmann's canals
2500:
2499:
2463:
2462:
2336:
2335:
2263:Epithelioid cells
2142:
2141:
2122:
2121:
2118:
2117:
2050:
2049:
1743:Knowable Magazine
1529:10.1172/jci113863
1086:Nat Rev Rheumatol
988:978-81-8061-865-9
594:cysteine protease
197:that breaks down
169:
168:
164:
16:(Redirected from
2915:
2832:medullary cavity
2807:Sharpey's fibres
2780:Haversian canals
2735:epiphyseal plate
2725:subchondral bone
2614:
2527:
2520:
2513:
2504:
2434:Precursor cells
2360:Promegakaryocyte
2226:
2153:
2078:Promegakaryocyte
2061:
1983:
1974:
1959:
1938:
1931:
1924:
1915:
1862:
1861:
1851:
1811:
1805:
1804:
1794:
1770:
1764:
1763:
1761:
1759:
1754:
1734:
1728:
1727:
1699:
1693:
1692:
1664:
1658:
1657:
1633:
1627:
1626:
1616:
1607:(30): 18636–43.
1592:
1586:
1585:
1557:
1551:
1550:
1540:
1508:
1502:
1501:
1491:
1482:(40): 24979–82.
1467:
1461:
1460:
1424:
1418:
1417:
1407:
1397:
1388:(2): 2289–2305.
1373:
1367:
1366:
1356:
1346:
1322:
1316:
1315:
1287:
1278:
1277:
1242:
1236:
1235:
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1201:
1195:
1194:
1192:
1169:
1145:
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1120:
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1077:
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1036:
1035:
999:
993:
992:
972:
963:
962:
952:
928:
919:
918:
916:
915:
904:
895:
892:
619:secretory domain
490:amino acid motif
367:acid phosphatase
161:edit on Wikidata
57:
45:Light micrograph
42:
30:
21:
2923:
2922:
2918:
2917:
2916:
2914:
2913:
2912:
2908:Skeletal system
2888:
2887:
2886:
2877:
2841:
2763:
2739:epiphyseal line
2713:
2692:
2666:
2640:
2631:intramembranous
2603:
2540:
2531:
2501:
2496:
2459:
2423:
2402:Immune response
2397:
2375:Red blood cells
2369:
2332:
2309:Langerhans cell
2304:Dendritic cells
2289:
2215:
2138:
2114:
2098:Proerythroblast
2084:
2046:
2030:Monocytopoiesis
2021:
1965:
1948:
1942:
1881:Wayback Machine
1870:
1865:
1813:
1812:
1808:
1772:
1771:
1767:
1757:
1755:
1736:
1735:
1731:
1710:(5310): 266–9.
1701:
1700:
1696:
1675:(5310): 270–3.
1666:
1665:
1661:
1635:
1634:
1630:
1594:
1593:
1589:
1559:
1558:
1554:
1510:
1509:
1505:
1469:
1468:
1464:
1435:(4920): 855–7.
1426:
1425:
1421:
1375:
1374:
1370:
1324:
1323:
1319:
1289:
1288:
1281:
1244:
1243:
1239:
1203:
1202:
1198:
1190:
1160:(123): 177–96.
1147:
1146:
1142:
1128:
1127:
1123:
1079:
1078:
1074:
1044:
1043:
1039:
1001:
1000:
996:
989:
974:
973:
966:
930:
929:
922:
913:
911:
906:
905:
898:
893:
889:
885:
868:
856:
830:
814:
806:deciduous teeth
794:
782:osteoprotegerin
751:
735:
729:
689:
673:trabecular bone
604:, a hereditary
602:pycnodysostosis
584:
560:vacuolar-ATPase
553:
549:
545:
541:
520:
513:
505:
482:plasma membrane
474:
466:osteoprotegerin
443:differentiation
435:tyrosine kinase
394:
344:
308:
300:deciduous teeth
259:actin filaments
257:but is rich in
227:bone resorption
165:
60:
48:
28:
23:
22:
15:
12:
11:
5:
2921:
2919:
2911:
2910:
2905:
2900:
2890:
2889:
2883:
2882:
2879:
2878:
2876:
2875:
2870:
2865:
2860:
2855:
2849:
2847:
2843:
2842:
2840:
2839:
2834:
2829:
2824:
2819:
2814:
2809:
2804:
2803:
2802:
2797:
2787:
2782:
2777:
2771:
2769:
2765:
2764:
2762:
2761:
2756:
2751:
2746:
2741:
2732:
2727:
2721:
2719:
2715:
2714:
2712:
2711:
2706:
2700:
2698:
2694:
2693:
2691:
2690:
2685:
2680:
2674:
2672:
2668:
2667:
2665:
2664:
2659:
2654:
2648:
2646:
2642:
2641:
2639:
2638:
2633:
2628:
2622:
2620:
2611:
2605:
2604:
2602:
2601:
2600:
2599:
2594:
2589:
2581:
2580:
2579:
2574:
2566:
2561:
2556:
2550:
2548:
2542:
2541:
2532:
2530:
2529:
2522:
2515:
2507:
2498:
2497:
2495:
2494:
2493:
2492:
2482:
2477:
2471:
2469:
2465:
2464:
2461:
2460:
2458:
2457:
2452:
2451:
2450:
2445:
2440:
2431:
2429:
2425:
2424:
2422:
2421:
2416:
2411:
2405:
2403:
2399:
2398:
2396:
2395:
2390:
2385:
2379:
2377:
2371:
2370:
2368:
2367:
2362:
2357:
2355:Megakaryoblast
2352:
2346:
2344:
2338:
2337:
2334:
2333:
2331:
2330:
2329:
2328:
2318:
2317:
2316:
2311:
2306:
2297:
2295:
2291:
2290:
2288:
2287:
2286:
2285:
2280:
2275:
2265:
2260:
2255:
2250:
2245:
2240:
2234:
2232:
2223:
2217:
2216:
2214:
2213:
2212:
2211:
2201:
2200:
2199:
2189:
2188:
2187:
2177:
2172:
2167:
2161:
2159:
2150:
2148:Myeloid tissue
2144:
2143:
2140:
2139:
2137:
2136:
2130:
2128:
2124:
2123:
2120:
2119:
2116:
2115:
2113:
2112:
2111:
2110:
2105:
2100:
2093:Erythropoiesis
2089:
2086:
2085:
2083:
2082:
2081:
2080:
2075:
2073:Megakaryoblast
2068:Thrombopoiesis
2064:
2058:
2052:
2051:
2048:
2047:
2045:
2044:
2043:
2042:
2037:
2026:
2023:
2022:
2020:
2019:
2018:
2017:
2012:
2007:
2002:
1997:
1990:Granulopoiesis
1986:
1980:
1971:
1956:
1950:
1949:
1943:
1941:
1940:
1933:
1926:
1918:
1912:
1911:
1898:
1893:
1883:
1869:
1868:External links
1866:
1864:
1863:
1806:
1765:
1729:
1694:
1659:
1628:
1587:
1552:
1503:
1462:
1419:
1368:
1317:
1279:
1237:
1196:
1140:
1133:Gray's Anatomy
1121:
1072:
1037:
994:
987:
964:
920:
896:
886:
884:
881:
880:
879:
874:
867:
864:
855:
852:
838:bisphosphonate
829:
826:
813:
810:
793:
790:
750:
747:
733:
728:
725:
697:growth factors
688:
685:
583:
580:
551:
547:
543:
539:
528:ruffled border
526:) through the
518:
511:
503:
473:
470:
393:
390:
343:
340:
307:
304:
247:ruffled border
167:
166:
157:
151:
150:
145:
139:
138:
133:
127:
126:
121:
115:
114:
109:
103:
102:
98:
97:
91:
87:
86:
81:
77:
76:
73:
67:
66:
62:
61:
58:
50:
49:
43:
35:
34:
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
2920:
2909:
2906:
2904:
2901:
2899:
2896:
2895:
2893:
2874:
2871:
2869:
2866:
2864:
2861:
2859:
2856:
2854:
2851:
2850:
2848:
2844:
2838:
2835:
2833:
2830:
2828:
2825:
2823:
2820:
2818:
2815:
2813:
2810:
2808:
2805:
2801:
2798:
2796:
2793:
2792:
2791:
2788:
2786:
2783:
2781:
2778:
2776:
2773:
2772:
2770:
2766:
2760:
2757:
2755:
2752:
2750:
2747:
2745:
2742:
2740:
2736:
2733:
2731:
2728:
2726:
2723:
2722:
2720:
2716:
2710:
2707:
2705:
2702:
2701:
2699:
2695:
2689:
2686:
2684:
2681:
2679:
2676:
2675:
2673:
2669:
2663:
2660:
2658:
2655:
2653:
2650:
2649:
2647:
2643:
2637:
2634:
2632:
2629:
2627:
2624:
2623:
2621:
2619:
2615:
2612:
2610:
2606:
2598:
2595:
2593:
2590:
2588:
2585:
2584:
2582:
2578:
2575:
2573:
2570:
2569:
2567:
2565:
2562:
2560:
2557:
2555:
2554:perichondrium
2552:
2551:
2549:
2547:
2543:
2539:
2535:
2528:
2523:
2521:
2516:
2514:
2509:
2508:
2505:
2491:
2488:
2487:
2486:
2483:
2481:
2478:
2476:
2473:
2472:
2470:
2466:
2456:
2455:Myelomonocyte
2453:
2449:
2446:
2444:
2441:
2439:
2436:
2435:
2433:
2432:
2430:
2426:
2420:
2417:
2415:
2412:
2410:
2407:
2406:
2404:
2400:
2394:
2391:
2389:
2386:
2384:
2381:
2380:
2378:
2376:
2372:
2366:
2365:Megakaryocyte
2363:
2361:
2358:
2356:
2353:
2351:
2348:
2347:
2345:
2343:
2339:
2327:
2324:
2323:
2322:
2319:
2315:
2312:
2310:
2307:
2305:
2302:
2301:
2299:
2298:
2296:
2292:
2284:
2281:
2279:
2276:
2274:
2271:
2270:
2269:
2266:
2264:
2261:
2259:
2256:
2254:
2251:
2249:
2246:
2244:
2243:Kupffer cells
2241:
2239:
2236:
2235:
2233:
2231:
2227:
2224:
2222:
2218:
2210:
2207:
2206:
2205:
2202:
2198:
2195:
2194:
2193:
2190:
2186:
2183:
2182:
2181:
2178:
2176:
2173:
2171:
2168:
2166:
2163:
2162:
2160:
2158:
2154:
2151:
2149:
2145:
2135:
2132:
2131:
2129:
2125:
2109:
2106:
2104:
2101:
2099:
2096:
2095:
2094:
2091:
2090:
2087:
2079:
2076:
2074:
2071:
2070:
2069:
2066:
2065:
2062:
2059:
2057:
2053:
2041:
2038:
2036:
2033:
2032:
2031:
2028:
2027:
2024:
2016:
2013:
2011:
2010:Metamyelocyte
2008:
2006:
2003:
2001:
1998:
1996:
1993:
1992:
1991:
1988:
1987:
1984:
1981:
1979:
1975:
1972:
1969:
1964:
1960:
1957:
1955:
1954:Hematopoiesis
1951:
1947:
1939:
1934:
1932:
1927:
1925:
1920:
1919:
1916:
1910:
1908:
1905:Animation by
1902:
1899:
1897:
1894:
1891:
1887:
1884:
1882:
1878:
1875:
1872:
1871:
1867:
1859:
1855:
1850:
1845:
1841:
1837:
1833:
1829:
1825:
1821:
1817:
1810:
1807:
1802:
1798:
1793:
1788:
1785:(4): 549–53.
1784:
1780:
1776:
1769:
1766:
1753:
1748:
1744:
1740:
1733:
1730:
1725:
1721:
1717:
1713:
1709:
1705:
1698:
1695:
1690:
1686:
1682:
1678:
1674:
1670:
1663:
1660:
1655:
1651:
1647:
1643:
1639:
1632:
1629:
1624:
1620:
1615:
1610:
1606:
1602:
1598:
1591:
1588:
1583:
1579:
1575:
1571:
1567:
1563:
1556:
1553:
1548:
1544:
1539:
1534:
1530:
1526:
1523:(1): 227–33.
1522:
1518:
1514:
1507:
1504:
1499:
1495:
1490:
1485:
1481:
1477:
1473:
1466:
1463:
1458:
1454:
1450:
1446:
1442:
1438:
1434:
1430:
1423:
1420:
1415:
1411:
1406:
1401:
1396:
1391:
1387:
1383:
1379:
1372:
1369:
1364:
1360:
1355:
1350:
1345:
1340:
1336:
1332:
1328:
1321:
1318:
1313:
1309:
1305:
1301:
1298:(4): 855–86.
1297:
1293:
1286:
1284:
1280:
1276:
1272:
1268:
1264:
1260:
1256:
1252:
1248:
1241:
1238:
1233:
1229:
1224:
1219:
1216:(3): 377–81.
1215:
1211:
1207:
1200:
1197:
1189:
1185:
1181:
1177:
1173:
1168:
1163:
1159:
1155:
1151:
1144:
1141:
1135:
1134:
1125:
1122:
1117:
1113:
1108:
1103:
1099:
1095:
1091:
1087:
1083:
1076:
1073:
1068:
1064:
1060:
1056:
1052:
1048:
1041:
1038:
1033:
1029:
1025:
1021:
1017:
1013:
1009:
1005:
998:
995:
990:
984:
980:
979:
971:
969:
965:
960:
956:
951:
946:
943:(2): 129–42.
942:
938:
937:Oral Diseases
934:
927:
925:
921:
909:
908:"Odontoclast"
903:
901:
897:
891:
888:
882:
878:
875:
873:
870:
869:
865:
863:
861:
853:
851:
848:
846:
841:
839:
835:
827:
825:
823:
819:
811:
809:
807:
803:
799:
791:
789:
787:
783:
778:
776:
772:
768:
764:
760:
756:
748:
746:
743:
737:
726:
724:
720:
718:
714:
712:
708:
704:
700:
698:
694:
686:
684:
682:
678:
674:
670:
669:knockout mice
665:
663:
659:
655:
651:
647:
643:
639:
635:
630:
628:
625:generated by
624:
620:
614:
610:
607:
606:osteopetrotic
603:
599:
595:
592:
588:
581:
579:
577:
573:
569:
565:
561:
557:
537:
533:
529:
525:
521:
514:
507:
499:
495:
491:
487:
483:
479:
471:
469:
467:
462:
460:
455:
454:osteopetrosis
451:
446:
444:
440:
436:
432:
427:
425:
421:
417:
416:stromal cells
413:
409:
405:
400:
391:
389:
387:
386:cell membrane
382:
380:
376:
372:
368:
364:
360:
356:
352:
349:
341:
339:
336:
332:
323:
316:
312:
305:
303:
301:
296:
294:
290:
285:
280:
276:
272:
271:hydrogen ions
268:
264:
260:
256:
252:
248:
244:
243:cell membrane
240:
235:
233:
229:
228:
223:
219:
215:
212:
208:
204:
200:
196:
192:
188:
185:
181:
178:
177:Ancient Greek
174:
162:
156:
152:
149:
146:
144:
140:
137:
134:
132:
128:
125:
122:
120:
116:
113:
110:
108:
104:
99:
96:
93:Breakdown of
92:
88:
85:
82:
78:
74:
72:
68:
63:
56:
51:
46:
41:
36:
31:
19:
2789:
2678:bone mineral
2661:
2636:endochondral
2626:primary bone
2618:Ossification
2572:chondroblast
2414:Phagocytosis
2383:Reticulocyte
2257:
2157:Granulocytes
2108:Reticulocyte
2000:Promyelocyte
1963:Myelopoiesis
1906:
1823:
1820:Biomaterials
1819:
1809:
1782:
1778:
1768:
1756:. Retrieved
1742:
1732:
1707:
1703:
1697:
1672:
1668:
1662:
1637:
1631:
1604:
1600:
1590:
1565:
1561:
1555:
1520:
1516:
1506:
1479:
1475:
1465:
1432:
1428:
1422:
1385:
1381:
1371:
1337:: 4695–707.
1334:
1330:
1320:
1295:
1291:
1250:
1246:
1240:
1213:
1209:
1199:
1157:
1153:
1143:
1132:
1124:
1092:(6): 341–6.
1089:
1085:
1075:
1050:
1046:
1040:
1010:(4): 291–9.
1007:
1003:
997:
977:
940:
936:
912:. Retrieved
890:
857:
849:
842:
831:
821:
817:
815:
797:
795:
779:
775:osteoporosis
757:, including
752:
742:transcytosis
738:
730:
721:
715:
711:angiogenesis
701:
690:
681:heterozygous
666:
634:cathepsins B
631:
618:
615:
611:
585:
564:osteoporosis
556:proton pumps
527:
475:
463:
447:
428:
395:
383:
365:filled with
345:
328:
297:
283:
267:sealing zone
266:
262:
246:
238:
236:
225:
190:
187:
183:
180:
172:
170:
112:osteoclastus
111:
2903:Human cells
2898:Macrophages
2837:bone marrow
2577:chondrocyte
2268:giant cells
2258:Osteoclasts
2238:Histiocytes
2230:Macrophages
2040:Promonocyte
1886:Osteoclasts
1874:MedicineNet
820:: bone and
798:odontoclast
792:Odontoclast
786:RANK ligand
591:papain-like
587:Cathepsin K
494:osteopontin
420:osteoblasts
399:osteoblasts
392:Development
379:cathepsin K
377:(TRAP) and
335:macrophages
293:osteoblasts
279:cathepsin K
275:collagenase
222:collagenase
199:bone tissue
101:Identifiers
95:bone tissue
18:Odontoclast
2892:Categories
2827:trabeculae
2822:canaliculi
2800:periosteum
2759:epicondyle
2744:metaphysis
2704:cancellous
2662:osteoclast
2652:osteoblast
2564:metaphysis
2192:Eosinophil
2175:Neutrophil
2165:Myeloblast
2103:Normoblast
1995:Myeloblast
1826:: 121903.
1053:(1): 1–6.
914:2013-11-06
883:References
840:toxicity.
763:calcitonin
749:Regulation
677:homozygous
478:chemotaxis
424:precursors
371:expression
263:clear zone
255:organelles
203:remodeling
175:(from
173:osteoclast
33:Osteoclast
2868:irregular
2795:endosteum
2768:Structure
2749:diaphysis
2730:epiphysis
2657:osteocyte
2546:Cartilage
2538:cartilage
2475:Phagocyte
2342:Platelets
2321:Monoblast
2253:Microglia
2221:Monocytes
2204:Mast cell
2170:Band cell
2035:Monoblast
2015:Band cell
2005:Myelocyte
1840:1878-5905
862:in 1873.
707:migration
576:lysosomes
572:cathepsin
486:podosomes
363:lysosomes
348:Howship's
306:Structure
251:cytoplasm
211:vertebral
195:bone cell
191:(clastos)
71:Precursor
2873:sesamoid
2812:enthesis
2709:cortical
2448:CFU-GEMM
2209:CFU-Mast
2185:CFU-Baso
2180:Basophil
1968:CFU-GEMM
1901:Random42
1877:Archived
1858:36410109
1849:10148651
1801:11950689
1654:10639325
1414:25690002
1363:26316708
1275:10968780
1232:10639325
1188:Archived
1184:30010574
1116:19491914
959:20659257
866:See also
860:Kölliker
755:hormones
658:cysteine
598:protease
472:Function
359:vacuoles
355:vesicles
342:Location
214:skeleton
184:(osteon)
90:Function
80:Location
2817:lacunae
2754:condyle
2718:Regions
2688:osteoid
2597:fibrous
2592:elastic
2587:hyaline
2350:CFU-Meg
2197:CFU-Eos
2127:General
1758:8 March
1724:9092478
1704:Science
1689:9092479
1669:Science
1623:9228032
1582:1829326
1547:2910910
1498:7929181
1457:2528207
1437:Bibcode
1429:Science
1405:4344726
1354:4544727
1312:3532144
1255:Bibcode
1247:Science
1107:2860596
1067:2837260
1032:7570118
1024:1416078
854:History
822:klastos
351:lacunae
232:calcium
218:mineral
209:of the
189:κλαστός
124:D010010
65:Details
2846:Shapes
2775:osteon
2683:ossein
2671:Matrix
2583:Types
2568:Cells
2480:Plasma
2438:CFU-GM
2314:CFU-DL
1978:CFU-GM
1946:plasma
1892:(MeSH)
1856:
1846:
1838:
1799:
1722:
1687:
1652:
1621:
1580:
1545:
1538:303666
1535:
1496:
1455:
1412:
1402:
1361:
1351:
1310:
1273:
1230:
1182:
1176:856515
1174:
1114:
1104:
1065:
1030:
1022:
985:
957:
818:osteon
717:MMP-13
652:, and
532:matrix
182:ὀστέον
2858:short
2697:Types
2645:Cells
2468:Other
2428:Other
2393:CFU-E
2294:Other
1191:(PDF)
1180:S2CID
1028:S2CID
534:into
408:RANKL
207:bones
179:
159:[
148:66781
107:Latin
2863:flat
2853:long
2609:Bone
2536:and
2534:Bone
1854:PMID
1836:ISSN
1797:PMID
1760:2022
1720:PMID
1685:PMID
1650:PMID
1619:PMID
1578:PMID
1543:PMID
1494:PMID
1453:PMID
1410:PMID
1359:PMID
1308:PMID
1271:PMID
1228:PMID
1172:PMID
1112:PMID
1063:PMID
1047:Bone
1020:PMID
983:ISBN
955:PMID
836:and
784:and
771:IL-6
703:MMP9
691:The
679:and
660:and
627:TRAP
459:RANK
418:and
357:and
119:MeSH
84:Bone
2326:MPS
2056:MEP
1844:PMC
1828:doi
1824:291
1787:doi
1747:doi
1712:doi
1708:276
1677:doi
1673:276
1642:doi
1609:doi
1605:272
1570:doi
1566:260
1533:PMC
1525:doi
1484:doi
1480:269
1445:doi
1433:245
1400:PMC
1390:doi
1349:PMC
1339:doi
1300:doi
1263:doi
1251:289
1218:doi
1214:113
1162:doi
1158:123
1102:PMC
1094:doi
1055:doi
1012:doi
1008:186
945:doi
804:of
796:An
675:in
546:, H
538:, H
517:HCO
450:TNF
373:of
265:or
205:of
171:An
143:FMA
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