Biology of Bone Tissue: Structure, Function, and Factors That Influence Bone Cells

BioMed Research International

Volumn 2015, Issue , 2015, Pages

  Florencio Silva, Rinaldo ^a   Sasso, Gisela Rodrigues Da Silva ^a   Sasso Cerri, Estela ^b   Simões, Manuel Jesus ^a   Cerri, Paulo Sérgio ^b  

^a FEDERAL UNIVERSITY OF SÃO PAULO   (Brazil)

^b SÃO PAULO STATE UNIVERSITY UNESP   (Brazil)

Author keywords

[No Author keywords available]

Indexed keywords

BONE CELL; BONE MATRIX; BONE REMODELING; BONE TISSUE; CELL FUNCTION; CELL STRUCTURE; HOMEOSTASIS; HUMAN; MALE; OSTEOBLAST; OSTEOCLAST; OSTEOCYTE; OSTEOLYSIS; REVIEW; ANIMAL; BIOLOGICAL MODEL; BONE; BONE MINERALIZATION; CELL COMMUNICATION; CYTOLOGY; PHYSIOLOGY; STRUCTURE ACTIVITY RELATION;

ANIMALS; BONE AND BONES; BONE REMODELING; CALCIFICATION, PHYSIOLOGIC; CELL COMMUNICATION; HUMANS; MODELS, BIOLOGICAL; OSTEOBLASTS; OSTEOCLASTS; STRUCTURE-ACTIVITY RELATIONSHIP;

EID: 84938149709     PISSN: 23146133     EISSN: 23146141     Source Type: Journal    
DOI: 10.1155/2015/421746     Document Type: Review

References (211)

1. J. A. Buckwalter, M. J. Glimcher, R. R. Cooper, and R. Recker, "Bone biology. I: structure, blood supply, cells, matrix, and mineralization, " Instructional Course Lectures, vol. 45, pp. 371-386, 1996.
2. P. A. Downey and M. I. Siegel, "Bone biology and the clinical implications for osteoporosis, " Physical Therapy, vol. 86, no. 1, pp. 77-91, 2006.
3. A. G. Robling, A. B. Castillo, and C. H. Turner, "Biomechanical and molecular regulation of bone remodeling, " Annual Review of Biomedical Engineering, vol. 8, pp. 455-498, 2006.
4. H. K. Datta, W. F. Ng, J. A. Walker, S. P. Tuck, and S. S. Varanasi, "The cell biology of bone metabolism, " Journal of Clinical Pathology, vol. 61, no. 5, pp. 577-587, 2008.
5. B. Clarke, "Normal bone anatomy and physiology, " Clinical Journal of the American Society of Nephrology, vol. 3, no. 3, pp. 131-139, 2008.
6. G. Karsenty, H. M. Kronenberg, and C. Settembre, "Genetic control of bone formation, " Annual Review of Cell and Developmental Biology, vol. 25, pp. 629-648, 2009.
7. S. L. Teitelbaum, "Osteoclasts: what do they do and how do they do it?" The American Journal of Pathology, vol. 170, no. 2, pp. 427-435, 2007.
8. L. F. Bonewald, "The amazing osteocyte, " Journal of Bone and Mineral Research, vol. 26, no. 2, pp. 229-238, 2011.
9. V. Everts, J. M. Delaissié, W. Korper et al., "The bone lining cell: its role in cleaning Howship's lacunae and initiating bone formation, " Journal of Bone and Mineral Research, vol. 17, no. 1, pp. 77-90, 2002.
10. N. A. Sims and J. H. Gooi, "Bone remodeling: Multiple cellular interactions required for coupling of bone formation and resorption, " Seminars in Cell and Developmental Biology, vol. 19, no. 5, pp. 444-451, 2008.
11. K. Matsuo and N. Irie, "Osteoclast-osteoblast communication, " Archives of Biochemistry and Biophysics, vol. 473, no. 2, pp. 201-209, 2008.
12. H. M. Frost, "Tetracycline-based histological analysis of bone remodeling, " Calcified Tissue Research, vol. 3, no. 1, pp. 211-237, 1969.
13. E. M. Hauge, D. Qvesel, E. F. Eriksen, L. Mosekilde, and F. Melsen, "Cancellous bone remodeling occurs in specialized compartments lined by cells expressing osteoblastic markers, " Journal of Bone and Mineral Research, vol. 16, no. 9, pp. 1575-1582, 2001.
14. T. L. Andersen, T. E. Sondergaard, K. E. Skorzynska et al., "A physical mechanism for coupling bone resorption and formation in adult human bone, " American Journal of Pathology, vol. 174, no. 1, pp. 239-247, 2009.
15. S. L. Dallas, M. Prideaux, and L. F. Bonewald, "The osteocyte: an endocrine cell ... and more, " Endocrine Reviews, vol. 34, no. 5, pp. 658-690, 2013.
16. S. Khosla, M. J. Oursler, and D. G. Monroe, "Estrogen and the skeleton, " Trends in Endocrinology and Metabolism, vol. 23, no. 11, pp. 576-581, 2012.
17. C. Sobacchi, A. Schulz, F. P. Coxon, A. Villa, and M. H. Helfrich, "Osteopetrosis: genetics, treatment and new insights into osteoclast function, " Nature Reviews Endocrinology, vol. 9, no. 9, pp. 522-536, 2013.
18. L. G. Raisz and G. A. Rodan, "Embryology and cellular biology of bone, " in Metabolic Bone Disease and Clinically Related Disorders, L. V. Avioli and S.M. Krane, Eds., pp. 1-22, Academic Press, New York, NY, USA, 3rd edition, 1998.
19. T. C. A. Phan, J. Xu, and M. H. Zheng, "Interaction between osteoblast and osteoclast: impact in bone disease, " Histology and Histopathology, vol. 19, no. 4, pp. 1325-1344, 2004.
20. J. C. Crockett, D. J. Mellis, D. I. Scott, and M. H. Helfrich, "New knowledge on critical osteoclast formation and activation pathways from study of rare genetic diseases of osteoclasts: focus on the RANK/RANKL axis, " Osteoporosis International, vol. 22, no. 1, pp. 1-20, 2011.
21. S. Fukumoto and T. J. Martin, "Bone as an endocrine organ, " Trends in Endocrinology andMetabolism, vol. 20, no. 5, pp. 230-236, 2009.
22. M. Capulli, R. Paone, and N. Rucci, "Osteoblast and osteocyte: games without frontiers, " Archives of Biochemistry and Biophysics, vol. 561, pp. 3-12, 2014.
23. S. C. Marks Jr. and S. N. Popoff, "Bone cell biology: the regulation of development, structure, and function in the skeleton, " American Journal of Anatomy, vol. 183, no. 1, pp. 1-44, 1988.
24. P. D. Damoulis and P. V. Hauschka, "Nitric oxide acts in conjunction with proinflammatory cytokines to promote cell death in osteoblasts, " Journal of Bone andMineral Research, vol. 12, no. 3, pp. 412-422, 1997.
25. A. E. Grigoriadis, J. N. M. Heersche, and J. E. Aubin, "Differentiation of muscle, fat, cartilage, and bone from progenitor cells present in a bone-derived clonal cell population: effect of dexamethasone, " Journal of Cell Biology, vol. 106, no.6, pp. 2139-2151, 1988.
26. P. Ducy, R. Zhang, V. Geoffroy, A. L. Ridall, and G. Karsenty, "Osf2/Cbfa1: a transcriptional activator of osteoblast differentiation, " Cell, vol. 89, no. 5, pp. 747-754, 1997.
27. T. Komori, H. Yagi, S. Nomura et al., "Targeted disruption of Cbfa1 results in a complete lack of bone formation owing to maturational arrest of osteoblasts, " Cell, vol. 89, no. 5, pp. 755-764, 1997.
28. M. Fakhry, E. Hamade, B. Badran, R. Buchet, and D. Magne, "Molecular mechanisms of mesenchymal stem cell differentiation towards osteoblasts, "World Journal of Stem Cells, vol. 5, no. 4, pp. 136-148, 2013.
29. K. Nakashima, X. Zhou, G. Kunkel et al., "Thenovel zinc fingercontaining transcription factorOsterix is required for osteoblast differentiation and bone formation, " Cell, vol. 108, no. 1, pp. 17-29, 2002.
30. D. A. Glass II, P. Bialek, J. D. Ahn et al., "Canonical Wnt signaling in differentiated osteoblasts controls osteoclast differentiation, " Developmental Cell, vol. 8, no. 5, pp. 751-764, 2005.
31. H. Hu, M. J. Hilton, X. Tu, K. Yu, D. M. Ornitz, and F. Long, "Sequential roles of Hedgehog and Wnt signaling in osteoblast development, " Development, vol. 132, no. 1, pp. 49-60, 2005.
32. K. Kapinas, C. Kessler, T. Ricks, G. Gronowicz, and A. M. Delany, "miR-29modulatesWnt signaling in human osteoblasts through a positive feedback loop, " The Journal of Biological Chemistry, vol. 285, no. 33, pp. 25221-25231, 2010.
33. Y. Zhang, R.-L. Xie, C. M. Croce et al., "A program of microRNAs controls osteogenic lineage progression by targeting transcription factor Runx2, " Proceedings of the National Academy of Sciences of the United States of America, vol. 108, no. 24, pp. 9863-9868, 2011.
34. A. Y. Montero, Y. Okada, M. Tomita et al., "Disruption of the fibroblast growth factor-2 gene results in decreased bone mass and bone formation, " The Journal of Clinical Investigation, vol. 105, no. 8, pp. 1085-1093, 2000.
35. A. M. Buo and J. P. Stains, "Gap junctional regulation of signal transduction in bone cells, " FEBS Letters, vol. 588, no. 8, pp. 1315-1321, 2014.
36. Z. O. Hamidouche, O. Fromigué, U. Nuber et al., "Autocrine fibroblast growth factor 18 mediates dexamethasone-induced osteogenic differentiation of murine mesenchymal stem cells, " Journal of Cellular Physiology, vol. 224, no. 2, pp. 509-515, 2010.
37. M. Q. Hassan, Y. Maeda, H. Taipaleenmaki et al., "miR-218 directs a Wnt signaling circuit to promote differentiation of osteoblasts and osteomimicry of metastatic cancer cells, " The Journal of Biological Chemistry, vol. 287, no. 50, pp. 42084-42092, 2012.
38. M. Tomé, P. López-Romero, C. Albo et al., "miR-335 orchestrates cell proliferation, migration and differentiation in human mesenchymal stem cells, " Cell Death and Differentiation, vol. 18, no. 6, pp. 985-995, 2011.
39. A. M. Flenniken, L. R. Osborne, N. Anderson et al., "A Gja1 missense mutation in a mouse model of oculodentodigital dysplasia, " Development, vol. 132, no. 19, pp. 4375-4386, 2005.
40. H. C. Anderson, "Matrix vesicles and calcification, " Current Rheumatology Reports, vol. 5, no. 3, pp. 222-226, 2003.
41. Y. Yoshiko, G. A. Candeliere, N. Maeda, and J. E. Aubin, "Osteoblast autonomous Pi regulation via Pit1 plays a role in bone mineralization, "Molecular and Cellular Biology, vol. 27, no. 12, pp. 4465-4474, 2007.
42. V. E. Arana-Chavez, A. M. V. Soares, and E. Katchburian, "Junctions between early developing osteoblasts of rat calvaria as revealed by freeze-fracture and ultrathin section electron microscopy, " Archives of Histology and Cytology, vol. 58, no. 3, pp. 285-292, 1995.
43. M. J. Glimcher, "The nature of the mineral phase in bone, " in Metabolic Bone Disease, M. J. Glimcher, Ed., pp. 23-50, Academic Press, San Diego, Calif, USA, 1998.
44. G. Boivin and P. J. Meunier, "The degree of mineralization of bone tissue measured by computerized quantitative contact microradiography, " Calcified Tissue International, vol. 70, no. 6, pp. 503-511, 2002.
45. G. Boivin, Y. Bala, A. Doublier et al., "Therole of mineralization and organic matrix in the microhardness of bone tissue from controls and osteoporotic patients, " Bone, vol. 43, no. 3, pp. 532-538, 2008.
46. S. C. Manolagas, "Birth and death of bone cells: basic regulatory mechanisms and implications for the pathogenesis and treatment of osteoporosis, " Endocrine Reviews, vol. 21, no. 2, pp. 115-137, 2000.
47. A. M. Parfitt, "Bone-forming cells in clinical conditions, " in Bone, Vol 1:The Osteoblast and Osteocyte, B. K. Hall, Ed., pp. 351-429, Telford Press, CRC Press, Boca Raton, Fla, USA, 1990.
48. R. L. Jilka, R. S. Weinstein, T. Bellido, A. M. Parfitt, and S. C. Manolagas, "Osteoblast programmed cell death (apoptosis): modulation by growth factors and cytokines, " Journal of Bone and Mineral Research, vol. 13, no. 5, pp. 793-802, 1998.
49. P. S. Cerri, "Osteoblasts engulf apoptotic bodies during alveolar bone formation in the rat maxilla, " Anatomical Record A, vol. 286, no. 1, pp. 833-840, 2005.
50. S. C. Miller, L. de Saint-Georges, B. M. Bowman, and W. S. S. Jee, "Bone lining cells: structure and function, " Scanning Microscopy, vol. 3, no. 3, pp. 953-961, 1989.
51. E. M. Aarden, E. H. Burger, and P. J. Nijweide, "Function of osteocytes in bone, " Journal of Cellular Biochemistry, vol. 55, no. 3, pp. 287-299, 1994.
52. H. J. Donahue, K. J. McLeod, C. T. Rubin et al., "Cell-to-cell communication in osteoblastic networks: cell line-dependent hormonal regulation of gap junction function, " Journal of Bone and Mineral Research, vol. 10, no. 6, pp. 881-889, 1995.
53. J. R. Mosley, "Osteoporosis and bone functional adaptation: mechanobiological regulation of bone architecture in growing and adult bone, a review, " Journal of Rehabilitation Research and Development, vol. 37, no. 2, pp. 189-199, 2000.
54. T. A. Franz-Odendaal, B.K. Hall, and P. E. Witten, "Buried alive: how osteoblasts become osteocytes, " Developmental Dynamics, vol. 235, no. 1, pp. 176-190, 2006.
55. G. Y. Rochefort, S. Pallu, and C. L. Benhamou, "Osteocyte: the unrecognized side of bone tissue, " Osteoporosis International, vol. 21, no. 9, pp. 1457-1469, 2010.
56. C. Palumbo, S. Palazzini, D. Zaffe, and G. Marotti, "Osteocyte differentiation in the tibia of newborn rabbit: an ultrastructural study of the formation of cytoplasmic processes, " Acta Anatomica, vol. 137, no. 4, pp. 350-358, 1990.
57. J. D. Currey, "The many adaptations of bone, " Journal of Biomechanics, vol. 36, no. 10, pp. 1487-1495, 2003.
58. M. B. Schaffler, W.-Y. Cheung, R. Majeska, and O. Kennedy, "Osteocytes: master orchestrators of bone, " Calcified Tissue International, vol. 94, no. 1, pp. 5-24, 2014.
59. K. Zhang, C. Barragan-Adjemian, L. Ye et al., "E11/gp38 selective expression in osteocytes: regulation by mechanical strain and role in dendrite elongation, "Molecular and Cellular Biology, vol. 26, no. 12, pp. 4539-4552, 2006.
60. A. Wetterwald, W. Hofstetter, M. G. Cecchini et al., "Characterization and cloning of the E11 antigen, a marker expressed by rat osteoblasts and osteocytes, " Bone, vol. 18, no. 2, pp. 125-132, 1996.
61. E. Schulze, M. Witt, M. Kasper, C. W. G. M. Löwik, and R. H. W. Funk, "Immunohistochemical investigations on the differentiation marker protein E11 in rat calvaria, calvaria cell culture and the osteoblastic cell line ROS 17/2.8, "Histochemistry and Cell Biology, vol. 111, no. 1, pp. 61-69, 1999.
62. Y. Mikuni-Takagaki, Y. Kakai, M. Satoyoshi et al., "Matrix mineralization and the differentiation of osteocyte-like cells in culture, " Journal of Bone andMineral Research, vol. 10, no. 2, pp. 231-242, 1995.
63. K. E. S. Poole, R. L. van Bezooijen, N. Loveridge et al., "Sclerostin is a delayed secreted product of osteocytes that inhibits bone formation, " The FASEB Journal, vol. 19, no. 13, pp. 1842-1844, 2005.
64. S. Ubaidus, M. Li, S. Sultana et al., "FGF23 is mainly synthesized by osteocytes in the regularly distributed osteocytic lacunar canalicular system established after physiological bone remodeling, " Journal of ElectronMicroscopy, vol. 58, no. 6, pp. 381-392, 2009.
65. S. C. Manolagas, "Choreography from the tomb: an emerging role of dying osteocytes in the purposeful, and perhaps not so purposeful, targeting of bone remodeling, " BoneKEy-Osteovision, vol. 3, no. 1, pp. 5-14, 2006.
66. R. Civitelli, F. Lecanda, N. R. Jørgensen, and T. H. Steinberg, "Intercellular junctions and cell-cell communication in bone, " in Principles of Bone Biology, J. P. Bilezikan, L. Raisz, and G. A. Rodan, Eds., pp. 287-302, Academic Press, San Diego, Calif, USA, 2002.
67. L. C. Johnson, "The kinetics of skeletal remodeling, " Birth Defects Original Article Series, vol. 2, no. 1, pp. 66-142, 1966.
68. M. G. Mullender, D. D. Van Der Meer, R. Huiskes, and P. Lips, "Osteocyte density changes in aging and osteoporosis, " Bone, vol. 18, no. 2, pp. 109-113, 1996.
69. L. F. Bonewald, "Osteocytes as dynamic multifunctional cells, " Annals of the New York Academy of Sciences, vol. 1116, pp. 281-290, 2007.
70. B. S. Noble, H. Stevens, N. Loveridge, and J. Reeve, "Identification of apoptotic changes in osteocytes in normal and pathological human bone, " Bone, vol. 20, no. 3, pp. 273-282, 1997.
71. J. I. Aguirre, L. I. Plotkin, S. A. Stewart et al., "Osteocyte apoptosis is induced by weightlessness in mice and precedes osteoclast recruitment and bone loss, " Journal of Bone and Mineral Research, vol. 21, no. 4, pp. 605-615, 2006.
72. L. I. Plotkin, "Apoptotic osteocytes and the control of targeted bone resorption, " Current Osteoporosis Reports, vol. 12, no. 1, pp. 121-126, 2014.
73. T. Bellido, "Osteocyte-driven bone remodeling, " Calcified Tissue International, vol. 94, no. 1, pp. 25-34, 2014.
74. F. Boabaid, P. S. Cerri, and E. Katchburian, "Apoptotic bone cells may be engulfed by osteoclasts during alveolar bone resorption in young rats, " Tissue and Cell, vol. 33, no. 4, pp. 318-325, 2001.
75. P. S. Cerri, F. Boabaid, and E. Katchburian, "Combined TUNEL and TRAP methods suggest that apoptotic bone cells are inside vacuoles of alveolar bone osteoclasts in young rats, " Journal of Periodontal Research, vol. 38, no. 2, pp. 223-226, 2003.
76. A. P. S. Faloni, E. Sasso-Cerri, E. Katchburian, and P. S. Cerri, "Decrease in the number and apoptosis of alveolar bone osteoclasts in estrogen-treated rats, " Journal of Periodontal Research, vol. 42, no. 3, pp. 193-201, 2007.
77. M. L. K. Tate, "'Whither flows the fluid in bone?' An osteocyte's perspective, " Journal of Biomechanics, vol. 36, no. 10, pp. 1409-1424, 2003.
78. Z. Xiao, S. Zhang, J. Mahlios et al., "Cilia-like structures and polycystin-1 in osteoblasts/osteocytes and associated abnormalities in skeletogenesis and Runx2 expression, " The Journal of Biological Chemistry, vol. 281, no. 41, pp. 30884-30895, 2006.
79. A. Santos, A. D. Bakker, B. Zandieh-Doulabi, J. M. A. de Blieck- Hogervorst, and J. Klein-Nulend, "Early activation of the β- catenin pathway in osteocytes is mediated by nitric oxide, phosphatidyl inositol-3 kinase/Akt, and focal adhesion kinase, " Biochemical and Biophysical Research Communications, vol. 391, no. 1, pp. 364-369, 2010.
80. E. H. Burger and J. Klein-Nulend, "Mechanotransduction in bone-role of the lacuno-canalicular network, " The FASEB Journal, vol. 13, no. 8, pp. S101-S112, 1999.
81. B. F. Boyce, D. E. Hughes, K. R. Wright, L. Xing, and A. Dai, "Recent advances in bone biology provide insight into the pathogenesis of bone diseases, " Laboratory Investigation, vol. 79, no. 2, pp. 83-94, 1999.
82. M. P. Yavropoulou and J. G. Yovos, "Osteoclastogenesis- current knowledge and future perspectives, " Journal of Musculoskeletal Neuronal Interactions, vol. 8, no. 3, pp. 204-216, 2008.
83. H. Takayanagi, "Osteoimmunology: shared mechanisms and crosstalk between the immune and bone systems, " Nature Reviews Immunology, vol. 7, no. 4, pp. 292-304, 2007.
84. K. Kim, S. H. Lee, J. H. Kim, Y. Choi, and N. Kim, "NFATc1 induces osteoclast fusion via up-regulation of osteoclast fusion and increased bone formation, " Nature Medicine, vol. 12, no. 12, pp. 1403-1409, 2006.
85. H. Yoshida, S.-I. Hayashi, T. Kunisada et al., "Themurinemutation osteopetrosis is in the coding region of the macrophage colony stimulating factor gene, " Nature, vol. 345, no. 6274, pp. 442-444, 1990.
86. J. Sodek and M. D. McKee, "Molecular and cellular biology of alveolar bone, " Periodontology 2000, vol. 24, no. 1, pp. 99-126, 2000.
87. B. F. Boyce and L. Xing, "Functions of RANKL/RANK/OPG in bone modeling and remodeling, " Archives of Biochemistry and Biophysics, vol. 473, no. 2, pp. 139-146, 2008.
88. R. Longhini, P. A. de Oliveira, A. P. de Souza Faloni, E. Sasso-Cerri, and P. S. Cerri, "Increased apoptosis in osteoclasts and decreased RANKL immunoexpression in periodontium of cimetidine-treated rats, " Journal of Anatomy, vol. 222, no. 2, pp. 239-247, 2013.
89. R. Longhini, P. A. de Oliveira, E. Sasso-Cerri, and P. S. Cerri, "Cimetidine reduces alveolar bone loss in induced periodontitis in rat molars, " Journal of Periodontology, vol. 85, no. 8, pp. 1115-1125, 2014.
90. M. Matsumoto, M. Kogawa, S. Wada et al., "Essential role of p38 mitogen-activated protein kinase in cathepsin K gene expression during osteoclastogenesis through association of NFATc1 and PU.1, " The Journal of Biological Chemistry, vol. 279, no. 44, pp. 45969-45979, 2004.
91. T. Miyamoto, "The dendritic cell-specific transmembrane protein DC-STAMP is essential for osteoclast fusion and osteoclast bone-resorbing activity, " Modern Rheumatology, vol. 16, no. 6, pp. 341-342, 2006.
92. Y. Kobayashi, N. Udagawa, and N. Takahashi, "Action of RANKL and OPG for osteoclastogenesis, " Critical Reviews in Eukaryotic Gene Expression, vol. 19, no. 1, pp. 61-72, 2009.
93. A. P. De Souza Faloni, T. Schoenmaker, A. Azari et al., "Jaw and long bone marrows have a different osteoclastogenic potential, " Calcified Tissue International, vol. 88, no. 1, pp. 63-74, 2011.
94. M. Mulari, J. Vääräniemi, and H. K. Väänänen, "Intracellular membrane trafficking in bone resorbing osteoclasts, " Microscopy Research and Technique, vol. 61, no. 6, pp. 496-503, 2003.
95. V. E. Arana-Chavez and V. Bradaschia-Correa, "Clastic cells: mineralized tissue resorption in health and disease, " The International Journal of Biochemistry&Cell Biology, vol. 41, no. 3, pp. 446-450, 2009.
96. P. T. Lakkakorpi, M. A. Horton, M. H. Helfrich, E.-K. Karhukorpi, and H. K. Vaananen, "Vitronectin receptor has a role in bone resorption but does not mediate tight sealing zone attachment of osteoclasts to the bone surface, " Journal of Cell Biology, vol. 115, no. 4, pp. 1179-1186, 1991.
97. F. Saltel, O. Destaing, F. Bard, D. Eichert, and P. Jurdic, "Apatitemediated actin dynamics in resorbing osteoclasts, " Molecular Biology of the Cell, vol. 15, no. 12, pp. 5231-5241, 2004.
98. C. Luxenburg, D. Geblinger, E. Klein et al., "The architecture of the adhesive apparatus of cultured osteoclasts: from podosome formation to sealing zone assembly, " PLoS ONE, vol. 2, no. 1, article e179, 2007.
99. A. Chabadel, I. Bañon-Rodríguez, D. Cluet et al., "CD44 and β3 integrin organize two functionally distinct actin-based domains in osteoclasts, " Molecular Biology of the Cell, vol. 18, no. 12, pp. 4899-4910, 2007.
100. U. Kornak, D. Kasper, M. R. Bösl et al., "Loss of the CIC-7 chloride channel leads to osteopetrosis in mice and man, " Cell, vol. 104, no. 2, pp. 205-215, 2001.
101. A. R. Graves, P. K. Curran, C. L. Smith, and J. A. Mindell, "The Cl-/H+ antiporter ClC-7 is the primary chloride permeation pathway in lysosomes, " Nature, vol. 453, no. 7196, pp. 788-792, 2008.
102. T. Yamaza, T. Goto, T. Kamiya, Y. Kobayashi, H. Sakai, and T. Tanaka, "Study of immunoelectron microscopic localization of cathepsin K in osteoclasts and other bone cells in the mouse femur, " Bone, vol. 23, no. 6, pp. 499-509, 1998.
103. J. Ljusberg, Y. Wang, P. Lång et al., "Proteolytic excision of a repressive loop domain in tartrate-resistant acid phosphatase by cathepsin K in osteoclasts, " The Journal of Biological Chemistry, vol. 280, no. 31, pp. 28370-28381, 2005.
104. A. P. de Souza Faloni, E. Sasso-Cerri, F. R. G. Rocha, E. Katchburian, and P. S. Cerri, "Structural and functional changes in the alveolar bone osteoclasts of estrogen-treated rats, " Journal of Anatomy, vol. 220, no. 1, pp. 77-85, 2012.
105. X. Feng and J. M. McDonald, "Disorders of bone remodeling, " Annual Review of Pathology: Mechanisms of Disease, vol. 6, pp. 121-145, 2011.
106. E. Seeman and P. D. Delmas, "Bone quality-the material and structural basis of bone strength and fragility, " The New England Journal of Medicine, vol. 354, no. 21, pp. 2250-2261, 2006.
107. S. Kimura, A. Nagai, T. Onitsuka et al., "Induction of experimental periodontitis in mice with Porphyromonas gingivalisadhered ligatures, " Journal of Periodontology, vol. 71, no. 7, pp. 1167-1173, 2000.
108. H. Hasturk, A. Kantarci, N. Ebrahimi et al., "Topical H2 antagonist prevents periodontitis in a rabbit model, " Infection and Immunity, vol. 74, no. 4, pp. 2402-2414, 2006.
109. J. Katz, Q.-B. Yang, P. Zhang et al., "Hydrolysis of epithelial junctional proteins by Porphyromonas gingivalis gingipains, " Infection and Immunity, vol. 70, no. 5, pp. 2512-2518, 2002.
110. C. H. Li and S. Amar, "Morphometric, histomorphometric, and microcomputed tomographic analysis of periodontal inflammatory lesions in a murine model, " Journal of Periodontology, vol. 78, no. 6, pp. 1120-1128, 2007.
111. N. Jain, G. K. Jain, S. Javed et al., "Recent approaches for the treatment of periodontitis, " Drug Discovery Today, vol. 13, no. 21-22, pp. 932-943, 2008.
112. J. F. Charles and A. O. Aliprantis, "Osteoclasts: more than 'bone eaters', " Trends inMolecularMedicine, vol. 20, no. 8, pp.449-459, 2014.
113. A. L. Boskey, L. Spevak, E. Paschalis, S. B. Doty, and M. D. McKee, "Osteopontin deficiency increases mineral content and mineral crystallinity in mouse bone, " Calcified Tissue International, vol. 71, no. 2, pp. 145-154, 2002.
114. A. Aszódi, J. F. Bateman, E. Gustafsson, R. Boot-Handford, and R. Fässler, "Mammalian skeletogenesis and extracellular matrix: what can we learn from knockout mice?" Cell Structure and Function, vol. 25, no. 2, pp. 73-84, 2000.
115. J. Christoffersen and W. J. Landis, "A contribution with review to the description of mineralization of bone and other calcified tissues in vivo, " Anatomical Record, vol. 230, no. 4, pp. 435-450, 1991.
116. K. Yagami, J.-Y. Suh, M. Enomoto-Iwamoto et al., "Matrix GLA protein is a developmental regulator of chondrocyte mineralization and, when constitutively expressed, blocks endochondral and intramembranous ossification in the limb, " The Journal of Cell Biology, vol. 147, no. 5, pp. 1097-1108, 1999.
117. J. Green, S. Schotland, D. J. Stauber, C. R. Kleeman, and T. L. Clemens, "Cell-matrix interaction in bone: type I collagen modulates signal transduction in osteoblast-like cells, " The American Journal of Physiology-Cell Physiology, vol. 268, no. 5, pp. C1090-C1103, 1995.
118. E. Sornay-Rendu, S. Boutroy, F. Munoz, and P. D. Delmas, "Alterations of cortical and trabecular architecture are associated with fractures in postmenopausal women, partially independent of decreasedBMDmeasured byDXA: theOF-ELY study, " Journal of Bone and Mineral Research, vol. 22, no. 3, pp. 425-433, 2007.
119. D. Vashishth, "Collagen glycation and its role in fracture properties of bone, " Journal ofMusculoskeletal Neuronal Interactions, vol. 5, article 316, 2005.
120. S. Viguet-Carrin, P. Garnero, and P. D. Delmas, "The role of collagen in bone strength, "Osteoporosis International, vol. 17, no. 3, pp. 319-336, 2006.
121. M. Saito, K. Fujii, S. Soshi, and T. Tanaka, "Reductions in degree of mineralization and enzymatic collagen cross-links and increases in glycation-induced pentosidine in the femoral neck cortex in cases of femoral neck fracture, " Osteoporosis International, vol. 17, no. 7, pp. 986-995, 2006.
122. S. Y. Tang, U. Zeenath, and D. Vashishth, "Effects of nonenzymatic glycation on cancellous bone fragility, " Bone, vol. 40, no. 4, pp. 1144-1151, 2007.
123. M. Owen and M. R. Shetlar, "Uptake of 3H-glucosamine by osteoclasts, " Nature, vol. 220, no. 5174, pp. 1335-1336, 1968.
124. C. C. Johnston, D. M. Smith, and A. R. Severson, "Bone resorption and matrix hexosamine labeling, " in Calcium, Parathyroid Hormone and the Calcitonins, R. V. Talmage and P. L. Munson, Eds., pp. 327-337, Excerpta Medica Foundation, Amsterdam, The Netherlands, 1972.
125. A. R. Severson, P. F. Rothberg, R. M. Pratt, and J. F. Goggins, "Effect of parathyroid hormone on the incorporation of 3H- glucosamine into hyaluronic acid in bone organ culture, " Endocrinology, vol. 92, no. 4, pp. 1282-1285, 1973.
126. R. A. Luben, J. F. Goggins, and L. G. Raisz, "Stimulation by parathyroid hormone of bone hyaluronate synthesis in organ culture, " Endocrinology, vol. 94, no. 3, pp. 737-745, 1974.
127. C. W. Prince, "Roles of hyaluronan in bone resorption, " BMC Musculoskeletal Disorders, vol. 5, article 12, 2004.
128. R. Zohar, "Signals between cells and matrix mediate bone regeneration, " in Bone Regeneration, P. H. Tal, Ed., InTech, 2012, http://www.intechopen.com/books/bone-regeneration/signalsbetween- cells-and-matrix-mediate-bone-regeneration.
129. D. Zimmerman, F. Jin, P. Leboy, S. Hardy, and C. Damsky, "Impaired bone formation in transgenic mice resulting from altered integrin function in osteoblasts, "Developmental Biology, vol. 220, no. 1, pp. 2-15, 2000.
130. C. H. Damsky, "Extracellular matrix-integrin interactions in osteoblast function and tissue remodeling, " Bone, vol. 25, no. 1, pp. 95-96, 1999.
131. P. J. Marie, "Role of N-cadherin in bone formation, " Journal of Cellular Physiology, vol. 190, no. 3, pp. 297-305, 2002.
132. M. H. Helfrich, G. Stenbeck, M. A. Nesbitt et al., "Integrins and adhesion molecules, " in Principles of Bone Biology, J. P. Bilezikan, L. G. Raisz, and T. J. Martin, Eds., pp. 385-424, Academic Press, Elsevier, San Diego, Calif, USA, 2008.
133. M. H. Helfrich, S. A. Nesbitt, E. L. Dorey, and M. A. Horton, "Rat osteoclasts adhere to a wide range of RGD(Arg-Gly-Asp) peptide-containing proteins, including the bone sialoproteins and fibronectin, via a β3 integrin, " Journal of Bone and Mineral Research, vol. 7, no. 3, pp. 335-343, 1992.
134. M.H. Helfrich, S. A. Nesbitt, P. T. Lakkakorpi et al., "β1 integrins and osteoclast function: involvement in collagen recognition and bone resorption, " Bone, vol. 19, no. 4, pp. 317-328, 1996.
135. G. Mbalaviele, S. S. Chan, and R. Civitelli, "Cell-cell adhesion and signaling through cadherins: connecting bone cells in their microenvironment, " Journal of Bone and Mineral Research, vol. 21, no. 12, pp. 1821-1827, 2006.
136. Y. Wang, L.M. McNamara, M.B. Schaffler, and S. Weinbaum, "A model for the role of integrins in flow induced mechanotransduction in osteocytes, " Proceedings of the National Academy of Sciences of the United States of America, vol. 104, no. 40, pp. 15941-15946, 2007.
137. L. M. McNamara, R. J. Majeska, S. Weinbaum, V. Friedrich, and M. B. Schaffler, "Attachment of osteocyte cell processes to the bone matrix, " Anatomical Record, vol. 292, no. 3, pp. 355-363, 2009.
138. J. B. Litzenberger, J.-B. Kim, P. Tummala, and C. R. Jacobs, "β1 Integrins mediate mechanosensitive signaling pathways in Osteocytes, " Calcified Tissue International, vol. 86, no. 4, pp. 325-332, 2010.
139. L.-D. You, S. Weinbaum, S. C. Cowin, and M. B. Schaffler, "Ultrastructure of the osteocyte process and its pericellular matrix, " Anatomical Record Part A: Discoveries in Molecular, Cellular, and Evolutionary Biology, vol. 278, no. 2, pp. 505-513, 2004.
140. K. J. Noonan, J. W. Stevens, R. Tammi, M. Tammi, J. A. Hernandez, and R. J. Midura, "Spatial distribution of CD44 and hyaluronan in the proximal tibia of the growing rat, " Journal of Orthopaedic Research, vol. 14, no. 4, pp. 573-581, 1996.
141. W. R. Thompson, S. Modla, B. J. Grindel et al., "Perlecan/Hspg2 deficiency alters the pericellular space of the lacunocanalicular system surrounding osteocytic processes in cortical bone, " Journal of Bone and Mineral Research, vol. 26, no. 3, pp. 618-629, 2011.
142. H. Kamioka, Y. Kameo, Y. Imai et al., "Microscale fluid flow analysis in a human osteocyte canaliculus using a realistic highresolution image-based three-dimensional model, " Integrative Biology, vol. 4, no. 10, pp. 1198-1206, 2012.
143. A. Miyauchi, M. Gotoh, H. Kamioka et al., "αvβ3 Integrin ligands enhance volume-sensitive calcium influx in mechanically stretched osteocytes, " Journal of Bone and Mineral Metabolism, vol. 24, no. 6, pp. 498-504, 2006.
144. K. Piekarski and M. Munro, "Transport mechanism operating between blood supply and osteocytes in long bones, " Nature, vol. 269, no. 5623, pp. 80-82, 1977.
145. S. Weinbaum, S. C. Cowin, and Y. Zeng, "A model for the excitation of osteocytes by mechanical loading-induced bone fluid shear stresses, " Journal of Biomechanics, vol. 27, no. 3, pp. 339-360, 1994.
146. T. Negishi-Koga and H. Takayanagi, "Bone cell communication factors and Semaphorins, " BoneKEy Reports, vol. 1, article 183, 2012.
147. L. M. Calvi, N. A. Sims, J. L. Hunzelman et al., "Activated parathyroid hormone/parathyroid hormone-related protein receptor in osteoblastic cells differentially affects cortical and trabecular bone, " The Journal of Clinical Investigation, vol. 107, no. 3, pp. 277-286, 2001.
148. D. Miao and A. Scutt, "Recruitment, augmentation and apoptosis of rat osteoclasts in 1, 25-(OH)2D3 response to shortterm treatment with 1, 25-dihydroxyvitamin D3 in vivo, " BMC Musculoskeletal Disorders, vol. 7, pp. 3-16, 2002.
149. M. Zaidi, A. M. Inzerillo, B. S. Moonga, P. J. R. Bevis, and C. L.-H. Huang, "Forty years of calcitonin-where are we now? A tribute to the work of Iain Macintyre, FRS, " Bone, vol. 30, no. 5, pp. 655-663, 2002.
150. D. Vanderschueren, L. Vandenput, S. Boonen, M. K. Lindberg, R. Bouillon, and C. Ohlsson, "Androgens and bone, " Endocrine Reviews, vol. 25, no. 3, pp. 389-425, 2004.
151. A. Tomkinson, J. Reeve, R.W. Shaw, and B. S. Noble, "The death of osteocytes via apoptosis accompanies estrogen withdrawal in human bone, " Endocrine Society Journals and Publications, vol. 82, pp. 3128-3135, 1997.
152. S. Kousteni, J.-R. Chen, T. Bellido et al., "Reversal of bone loss in mice by nongenotropic signaling of sex steroids, " Science, vol. 298, no. 5594, pp. 843-846, 2002.
153. K. B. Emerton, B. Hu, A. A. Woo et al., "Osteocyte apoptosis and control of bone resorption following ovariectomy in mice, " Bone, vol. 46, no. 3, pp. 577-583, 2010.
154. D. E. Hughes, A. Dai, J. C. Tiffee, H. H. Li, G. R. Munoy, and B. F. Boyce, "Estrogen promotes apoptosis of murine osteoclasts mediated by TGF-β, " Nature Medicine, vol. 2, no. 10, pp. 1132-1136, 1996.
155. L. C. Hofbauer and A. E. Heufelder, "Role of receptor activator of nuclear factor-κB ligand and osteoprotegerin in bone cell biology, " Journal ofMolecular Medicine, vol. 79, no. 5-6, pp. 243-253, 2001.
156. S. Kawamoto, S. Ejiri, E. Nagaoka, and H. Ozawa, "Effects of oestrogen deficiency on osteoclastogenesis in the rat periodontium, " Archives of Oral Biology, vol. 47, no. 1, pp. 67-73, 2002.
157. G. Eghbali-Fatourechi, S. Khosla, A. Sanyal, W. J. Boyle, D. L. Lacey, and B. L. Riggs, "Role of RANK ligand in mediating increased bone resorption in early postmenopausal women, " The Journal of Clinical Investigation, vol. 111, no. 8, pp. 1221-1230, 2003.
158. K. Väänänen, "Mechanism of osteoclast mediated bone resorption- rationale for the design of new therapeutics, " Advanced Drug Delivery Reviews, vol. 57, no. 7, pp. 959-971, 2005.
159. L. J. Robinson, B. B. Yaroslavskiy, R. D. Griswold et al., "Estrogen inhibits RANKL-stimulated osteoclastic differentiation of human monocytes through estrogen and RANKL-regulated interaction of estrogen receptor-α with BCAR1 and Traf6, " Experimental Cell Research, vol. 315, no. 7, pp. 1287-1301, 2009.
160. R. Pacifici, "Estrogen, cytokines, and pathogenesis of postmenopausal osteoporosis, " Journal of Bone and Mineral Research, vol. 11, no. 8, pp. 1043-1051, 1996.
161. S. Cenci, M. N. Weitzmann, C. Roggia et al., "Estrogen deficiency induces bone loss by enhancing T-cell production of TNF-α, "The Journal of Clinical Investigation, vol. 106, no. 10, pp. 1229-1237, 2000.
162. M. J. Oursler, P. Osdoby, J. Pyfferoen, B. L. Riggs, and T.C. Spelsberg, "Avian osteoclasts as estrogen target cells, " Proceedings of the National Academy of Sciences of the United States of America, vol. 88, no. 15, pp. 6613-6617, 1991.
163. M. Cruzoé-Souza, E. Sasso-Cerri, and P. S. Cerri, "Immunohistochemical detection of estrogen receptor β in alveolar bone cells of estradiol-treated female rats: possible direct action of estrogen on osteoclast life span, " Journal of Anatomy, vol. 215, no. 6, pp. 673-681, 2009.
164. M. Martin-Millan, M. Almeida, E. Ambrogini et al., "The estrogen receptor-α in osteoclasts mediates the protective effects of estrogens on cancellous but not cortical bone, " Molecular Endocrinology, vol. 24, no. 2, pp. 323-334, 2010.
165. H. M. Frost, Intermediary Organization of the Skeleton, CRC Press, Boca Raton, Fla, USA, 1986.
166. F. Elefteriou, "Regulation of bone remodeling by the central and peripheral nervous system, " Archives of Biochemistry and Biophysics, vol. 473, no. 2, pp. 231-236, 2008.
167. T. Nakashima, M. Hayashi, T. Fukunaga et al., "Evidence for osteocyte regulation of bone homeostasis through RANKL expression, " Nature Medicine, vol. 17, no. 10, pp. 1231-1234, 2011.
168. J. Xiong, M. Onal, R. L. Jilka, R. S. Weinstein, S. C. Manolagas, and C. A. O'Brien, "Matrix-embedded cells control osteoclast formation, " Nature Medicine, vol. 17, no. 10, pp. 1235-1241, 2011.
169. G. A. Howard, B. L. Bottemiller, R. T. Turner, J. I. Rader, and D. J. Baylink, "Parathyroid hormone stimulates bone formation and resorption in organ culture: evidence for a coupling mechanism, " Proceedings of the National Academy of Sciences of the United States of America, vol. 78, no. 5, pp. 3204-3208, 1981.
170. T. A. Linkhart, S. Mohan, and D. J. Baylink, "Growth factors for bone growth and repair: IGF, TGF beta and BMP, " Bone, vol. 19, no. 1, 1996.
171. T. Segovia-Silvestre, A. V. Neutzsky-Wulff, M. G. Sorensen et al., "Advances in osteoclast biology resulting from the study of osteopetrotic mutations, " Human Genetics, vol. 124, no. 6, pp. 561-577, 2009.
172. T. Negishi-Koga, M. Shinohara, N. Komatsu et al., "Suppression of bone formation by osteoclastic expression of semaphorin 4D, " Nature Medicine, vol. 17, no. 11, pp. 1473-1480, 2011.
173. K. Suzuki, A. Kumanogoh, and H. Kikutani, "Semaphorins and their receptors in immune cell interactions, " Nature Immunology, vol. 9, no. 1, pp. 17-23, 2008.
174. G. Delorme, F. Saltel, E. Bonnelye, P. Jurdic, and I. Machuca- Gayet, "Expression and function of semaphorin 7A in bone cells, " Biology of the Cell, vol. 97, no. 7, pp. 589-597, 2005.
175. A. L. M. Sutton, X. Zhang, D. R. Dowd, Y. P. Kharode, B. S. Komm, and P. N. MacDonald, "Semaphorin 3B is a 1, 25- dihydroxyvitamin D3-induced gene in osteoblasts that promotes osteoclastogenesis and induces osteopenia in mice, " Molecular Endocrinology, vol. 22, no. 6, pp. 1370-1381, 2008.
176. A. Hughes, J. Kleine-Albers, M. H. Helfrich, S. H. Ralston, and M. J. Rogers, "A Class III semaphorin (Sema3e) inhibits mouse osteoblast migration and decreases osteoclast formation in vitro, " Calcified Tissue International, vol. 90, no. 2, pp. 151-162, 2012.
177. M. Hayashi, T. Nakashima, M. Taniguchi, T. Kodama, A. Kumanogoh, and H. Takayanagi, "Osteoprotection by semaphorin 3A, " Nature, vol. 485, no. 7396, pp. 69-74, 2012.
178. C. Zhao, N. Irie, Y. Takada et al., "Bidirectional ephrinB2-EphB4 signaling controls bone homeostasis, " Cell Metabolism, vol. 4, no. 2, pp. 111-121, 2006.
179. E. H. Allan, K. D. Häusler, T. Wei et al., "EphrinB2 regulation by PTH and PTHrP revealed by molecular profiling in differentiating osteoblasts, " Journal of Bone andMineral Research, vol. 23, no. 8, pp. 1170-1181, 2008.
180. L. Pederson, M. Ruan, J. J. Westendorf, S. Khosla, and M. J. Oursler, "Regulation of bone formation by osteoclasts involves Wnt/BMP signaling and the chemokine sphingosine- 1-phosphate, " Proceedings of the National Academy of Sciences of the United States of America, vol. 105, no. 52, pp. 20764-20769, 2008.
181. J. Klein-Nulend, C. M. Semeins, N. E. Ajubi, P. J. Nijweide, and E. H. Burger, "Pulsating fluid flow increases nitric oxide (NO) synthesis by osteocytes but not periosteal fibroblasts- correlation with prostaglandin upregulation, " Biochemical and Biophysical Research Communications, vol. 217, no. 2, pp. 640-648, 1995.
182. P. P. Cherian, A. J. Siller-Jackson, S. Gu et al., "Mechanical strain opens connexin 43 hemichannels in osteocytes: a novel mechanism for the release of prostaglandin, " Molecular Biology of the Cell, vol. 16, no. 7, pp. 3100-3106, 2005.
183. D. C. Genetos, C. J. Kephart, Y. Zhang, C. E. Yellowley, and H. J. Donahue, "Oscillating fluid flow activation of gap junction hemichannels induces ATP release from MLO-Y4 osteocytes, " Journal of Cellular Physiology, vol. 212, no. 1, pp. 207-214, 2007.
184. X. L. Lu, B. Huo, M. Park, and X. E. Guo, "Calcium response in osteocytic networks under steady and oscillatory fluid flow, " Bone, vol. 51, no. 3, pp. 466-473, 2012.
185. A. G. Robling, T. Bellido, and C. H. Turner, "Mechanical stimulation in vivo reduces osteocyte expression of sclerostin, " Journal ofMusculoskeletal Neuronal Interactions, vol. 6, no. 4, p. 354, 2006.
186. A. G. Robling, P. J. Niziolek, L. A. Baldridge et al., "Mechanical stimulation of bone in vivo reduces osteocyte expression of Sost/sclerostin, " Journal of Biological Chemistry, vol. 283, no. 9, pp. 5866-5875, 2008.
187. A. Gaudio, P. Pennisi, C. Bratengeier et al., "Increased sclerostin serum levels associated with bone formation and resorption markers in patients with immobilization-induced bone loss, " The Journal of Clinical Endocrinology &Metabolism, vol. 95, no. 5, pp. 2248-2253, 2010.
188. T. Moriishi, R. Fukuyama, M. Ito et al., "Osteocyte network; a negative regulatory system for bone mass augmented by the induction of Rankl in osteoblasts and sost in osteocytes at unloading, " PLoS ONE, vol. 7, no. 6, Article ID e40143, 2012.
189. L. F. Bonewald and M. L. Johnson, "Osteocytes, mechanosensing andWnt signaling, " Bone, vol. 42, no. 4, pp. 606-615, 2008.
190. O. Verborgt, G. J. Gibson, and M. B. Schaffler, "Loss of osteocyte integrity in association with microdamage and bone remodeling after fatigue in vivo, " Journal of Bone and Mineral Research, vol. 15, no. 1, pp. 60-67, 2000.
191. L. Cardoso, B. C. Herman, O. Verborgt, D. Laudier, R. J. Majeska, and M. B. Schaffler, "Osteocyte apoptosis controls activation of intracortical resorption in response to bone fatigue, " Journal of Bone and Mineral Research, vol. 24, no. 4, pp. 597-605, 2009.
192. N. N. Taniwaki and E. Katchburian, "Ultrastructural and lanthanum tracer examination of vapidly resorbing rat alveolar bone suggests that osteoclasts internalize dying bone cells, " Cell and Tissue Research, vol. 293, no. 1, pp. 173-176, 1998.
193. G. Gu, M. Mulari, Z. Peng, T.A. Hentunen, and H.K.Väänänen, "Death of osteocytes turns off the inhibition of osteoclasts and triggers local bone resorption, " Biochemical and Biophysical Research Communications, vol. 335, no. 4, pp. 1095-1101, 2005.
194. O. D. Kennedy, B. C. Herman, D. M. Laudier, R. J. Majeska, H. B. Sun, and M. B. Schaffler, "Activation of resorption in fatigue-loaded bone involves both apoptosis and active proosteoclastogenic signaling by distinct osteocyte populations, " Bone, vol. 50, no. 5, pp. 1115-1122, 2012.
195. A. C. Wu, N. A. Morrison, W. L. Kelly, and M. R. Forwood, "MCP-1 expression is specifically regulated during activation of skeletal repair and remodeling, " Calcified Tissue International, vol. 92, no. 6, pp. 566-575, 2013.
196. N. Bivi, K. W. Condon, M. R. Allen et al., "Cell autonomous requirement of connexin 43 for osteocyte survival: consequences for endocortical resorption and periosteal bone formation, " Journal of Bone and Mineral Research, vol. 27, no. 2, pp. 374-389, 2012.
197. L. I. Plotkin, "Connexin 43 hemichannels and intracellular signaling in bone cells, " Frontiers in Physiology, vol. 5, article 131, 2014.
198. J. Yang, R. Shah, A. G. Robling et al., "HMGB1 is a bone-active cytokine, " Journal of Cellular Physiology, vol. 214, no. 3, pp. 730-739, 2008.
199. J. R. Klune, R. Dhupar, J. Cardinal, T. R. Billiar, and A. Tsung, "HMGB1: endogenous danger signaling, " Molecular Medicine, vol. 14, no. 7-8, pp. 476-484, 2008.
200. Z. Zhou, J.-Y. Han, C.-X. Xi et al., "HMGB1 regulates RANKLinduced osteoclastogenesis in a manner dependent on RAGE, " Journal of Bone and Mineral Research, vol. 23, no. 7, pp. 1084-1096, 2008.
201. S. E. Harris, M. MacDougall, D. Horn et al., "Meox2Cremediated disruption of CSF-1 leads to osteopetrosis and osteocyte defects, " Bone, vol. 50, no. 1, pp. 42-53, 2012.
202. A. R. Guntur and C. J. Rosen, "Bone as an endocrine organ, " Endocrine Practice, vol. 18, no. 5, pp. 758-762, 2012.
203. G. Karsenty and M. Ferron, "The contribution of bone towholeorganism physiology, " Nature, vol. 481, no. 7381, pp. 314-320, 2012.
204. M. Ferron, E. Hinoi, G. Karsenty, and P. Ducy, "Osteocalcin differentially regulates β cell and adipocyte gene expression and affects the development of metabolic diseases in wild-type mice, " Proceedings of the National Academy of Sciences of the United States of America, vol. 105, no. 13, pp. 5266-5270, 2008.
205. C. B. Confavreux, "Bone: from a reservoir of minerals to a regulator of energy metabolism, " Kidney International, vol. 79, no. 121, pp. S14-S19, 2011.
206. F. Oury, G. Sumara, O. Sumara et al., "Endocrine regulation of male fertility by the skeleton, " Cell, vol. 144, no. 5, pp. 796-809, 2011.
207. F. Oury, L. Khrimian, C. A. Denny et al., "Maternal and offspring pools of osteocalcin influence brain development and functions, " Cell, vol. 155, no. 1, pp. 228-241, 2013.
208. L. D. Quarles, "Endocrine functions of bone in mineral metabolism regulation, " The Journal of Clinical Investigation, vol. 118, no. 12, pp. 3820-3828, 2008.
209. A. Martin, S. Liu, V. David et al., "Bone proteins PHEX and DMP1 regulate fibroblastic growth factor Fgf23 expression in osteocytes through a common pathway involving FGF receptor (FGFR) signaling, " The FASEB Journal, vol. 25, no. 8, pp. 2551-2562, 2011.
210. M. Sato, N. Asada, Y. Kawano et al., "Osteocytes regulate primary lymphoid organs and fat metabolism, " Cell Metabolism, vol. 18, no. 5, pp. 749-758, 2013.
211. M. C. Walsh, N. Kim, Y. Kadono et al., "Osteoimmunology: interplay between the immune system and bone metabolism, " Annual Review of Immunology, vol. 24, pp. 33-63, 2006.