[1] P.J. Ehrlich, L.E. Lanyon, Mechanical strain and bone cell function: a review, Osteoporos. Int. 13 (2002) 688–700.
[2] T. Nakano, K. Kaibara, T. Ishimoto, Y. Tabata, Y. Umakoshi, Biological apatite (BAp) crystallographic orientation and texture as a new index for assessing the microstructure and function of bone regenerated by tissue engineering, Bone 51 (2012) 741–747.
[3] T. Nakano, K. Kaibara, Y. Tabata, N. Nagata, S. Enomoto, E. Marukawa, Y. Umakoshi, Unique alignment and texture of biological apatite crystallites in typical calcified tissues analyzed by microbeam X-ray diffractometer system, Bone 31 (2002) 479–487.
[4] T. Ishimoto, T. Nakano, Y. Umakoshi, M. Yamamoto, Y. Tabata, Degree of biological apatite c-axis orientation rather than bone mineral density controls mechanical function in bone regenerated using recombinant bone morphogenetic protein-2, J. Bone Miner. Res. 28 (2013) 1170–1179.
[5] T. Lang, A. LeBlanc, H. Evans, Y. Lu, H. Genant, A. Yu, Cortical and trabecular bone mineral loss from the spine and hip in long-duration spaceflight, J. Bone Miner. Res. 19 (2004) 1006–1012.
[6] J. Yang, J. Li, X. Cui, W. Li, Y. Xue, P. Shang, H. Zhang, Blocking glucocorticoid signaling in osteoblasts and osteocytes prevents mechanical unloading-induced cortical bone loss, Bone 130 (2020) 115108.
[7] J. Wang, T. Ishimoto, T. Nakano, Unloading-induced degradation of the anisotropic arrangement of collagen/apatite in rat femurs, Calcif. Tissue Int. 100 (2017) 87–94.
[8] A.G. Robling, L.F. Bonewald, The osteocyte: new insights, Annu. Rev. Physiol. 82 (2020) 485–506.
[9] L. Qin, W. Liu, H. Cao, G. Xiao, Molecular mechanosensors in osteocytes, Bone Res 8 (2020) 1–24.
[10] Y. Ishihara, Y. Sugawara, H. Kamioka, N. Kawanabe, H. Kurosaka, K. Naruse, T. Yamashiro, In situ imaging of the autonomous intracellular Ca2+ oscillations of osteoblasts and osteocytes in bone, Bone 50 (2012) 842–852.
[11] V.I. Sikavitsas, J.S. Temenoff, A.G. Mikos, Biomaterials and bone mechanotransduction, Biomaterials 22 (2001) 2581–2593.
[12] R. Ozasa, T. Ishimoto, S. Miyabe, J. Hashimoto, M. Hirao, H. Yoshikawa, T. Nakano, Osteoporosis changes collagen/apatite orientation and Young’s modulus in vertebral cortical bone of rat, Calcif. Tissue Int. 104 (2019) 449–460.
[13] M. Tanaka, A. Matsugaki, T. Ishimoto, T. Nakano, Evaluation of crystallographic orientation of biological apatite at vertebral cortical bone in ovariectomized cynomolgus monkey treated with minodronic acid and alendronate, J. Bone Miner. Metabol. 34 (2016) 234–241.
[14] A. Matsugaki, N. Fujiwara, T. Nakano, Continuous cyclic stretch induces osteoblast alignment and formation of anisotropic collagen fiber matrix, Acta Biomater. 9 (2013) 7227–7235.
[15] A. Matsugaki, G. Aramoto, T. Nakano, The alignment of MC3T3-E1 osteoblasts on steps of slip traces introduced by dislocation motion, Biomaterials 33 (2012) 7327–7335.
[16] A. Matsugaki, Y. Isobe, T. Saku, T. Nakano, Quantitative regulation of bone- mimetic, oriented collagen/apatite matrix structure depends on the degree of osteoblast alignment on oriented collagen substrates, J. Biomed. Mater. Res. A. 103 (2015) 489–499.
[17] K.J. Lewis, D. Frikha-Benayed, J. Louie, S. Stephen, D.C. Spray, M.M. Thi, Z. Seref- Ferlengez, R.J. Majeska, S. Weinbaum, M.B. Schaffler, Osteocyte calcium signals encode strain magnitude and loading frequency in vivo, Proc. Natl. Acad. Sci. Unit. States Am. 114 (2017) 11775–11780.
[18] P.M. Govey, Y.I. Kawasawa, H.J. Donahue, Mapping the osteocytic cell response to fluid flow using RNA-Seq, J. Biomech. 48 (2015) 4327–4332.
[19] A.E. Morrell, G.N. Brown, S.T. Robinson, R.L. Sattler, A.D. Baik, G. Zhen, X. Cao, L. F. Bonewald, W. Jin, L.C. Kam, Mechanically induced Ca 2 oscillations in osteocytes release extracellular vesicles and enhance bone formation, Bone Res 6 (2018) 1–11.
[20] T. Ganesh, L.E. Laughrey, M. Niroobakhsh, N. Lara-Castillo, Multiscale finite element modeling of mechanical strains and fluid flow in osteocyte lacunocanalicular system, Bone (2020) 115328.
[21] T. Adachi, Y. Aonuma, M. Tanaka, M. Hojo, T. Takano-Yamamoto, H. Kamioka, Calcium response in single osteocytes to locally applied mechanical stimulus: differences in cell process and cell body, J. Biomech. 42 (2009) 1989–1995.
[22] M. Oishi, S. Munesue, A. Harashima, M. Nakada, Y. Yamamoto, Y. Hayashi, Aquaporin 1 elicits cell motility and coordinates vascular bed formation by downregulating thrombospondin type-1 domain-containing 7A in glioblastoma, Cancer Med 9 (2020) 3904–3917.
[23] T. Shimasaki, S. Yamamoto, T. Arisawa, Exosome Research and Co-culture study, Biol. Pharm. Bull. 41 (2018) 1311–1321.
[24] Y. Li, S. Yuan, X. Wang, S.K. Tan, J. Mao, Comparison of flow fields in a centrifugal pump among different tracer particles by particle image velocimetry, ASME. J. Fluids Eng. 138 (2016), 061105.
[25] J. Jiang, R. Dingledine, Prostaglandin receptor EP2 in the crosshairs of anti- inflammation, anti-cancer, and neuroprotection, Trends Pharmacol. Sci. 34 (2013) 413–423.
[26] D. Shamir, S. Keila, M. Weinreb, A selective EP4 receptor antagonist abrogates the stimulation of osteoblast recruitment from bone marrow stromal cells by prostaglandin E2 in vivo and in vitro, Bone 34 (2004) 157–162.
[27] A.R. Stern, M.M. Stern1, M.E.V. Dyke, K. Ja¨hn, M. Prideaux, L.F. Bonewald, Isolation and culture of primary osteocytes from the long bones of skeletally mature and aged mice, Biotechniques 52 (2012) 361–373.
[28] A. Matsugaki, D. Yamazaki, T. Nakano, Selective patterning of netrin-1 as a novel guiding cue for anisotropic dendrogenesis in osteocytes, Mater. Sci. Eng. C 108 (2020) 110391.
[29] Y. Kato, J.J. Windle, B.A. Koop, G.R. Mundy, L.F. Bonewald, Establishment of an osteocyte-like cell line, MLO-Y4, J. Bone Miner. Res. 12 (1997) 2014–2023.
[30] L.H. Xu, H. Shao, Y.-H.V. Ma, L. You, OCY454 osteocytes as an in vitro cell model for bone remodeling under mechanical loading, J. Orthop. Res. 37 (2019) 1681–1689.
[31] S.M. Woo, J. Rosser, V. Dusevich, I. Kalajzic, L.F. Bonewald, Cell line IDG-SW3 replicates osteoblast-to-late-osteocyte differentiation in vitro and accelerates bone formation in vivo, J. Bone Miner. Res. 26 (2011) 2634–2646.
[32] G. Nasello, P. Alama´n-Díez, J. Schiavi, M. P´erez, L. McNamara, J.M. García-Aznar, Primary human osteoblasts cultured in a 3D microenvironment create a unique representative model of their differentiation into osteocytes, Front. Bioeng. Biotechnol. 8 (2020) 336.
[33] A. Matsugaki, T. Matuzaka, A. Murakami, P. Wang, T. Nakano, Three-dimensional printing of anisotropic bone mimetic structure with controlled fluid flow stimuli for osteocytes: flow orientation determines the elongation of dendrites, Int. J. Bioprint. 6 (2020) 293.
[34] J. Renaud, M.-G. Martinoli, Development of an insert co-culture system of two cellular types in the absence of cell-cell contact, JoVE J. Vis. Exp. (2016), e54356.
[35] L. Jia, W. Gu, Y. Zhang, Y. Ji, J. Liang, Y. Wen, X. Xu, The crosstalk between HDPSCs and HUCMSCs on proliferation and osteogenic genes expression in coculture system, Int. J. Med. Sci. 14 (2017) 1118–1129.
[36] C.D. Amo, V. Olivares, Matrix architecture plays a pivotal role in 3D osteoblast migration: the effect of interstitial fluid flow, J. Mech. Behav. Biomed. Mater. 83 (2018) 52–62.
[37] R. Ozasa, A. Matsugaki, T. Matsuzaka, T. Ishimoto, H.S. Yun, T. Nakano, Superior alignment of human iPSC-osteoblasts associated with focal adhesion formation stimulated by oriented collagen scaffold, Int. J. Mol. Sci. 22 (2021) 6232.
[38] X. Zhou, J.E. Novotny, L. Wang, Anatomic variations of the lacunar–canalicular system influence solute transport in bone, Bone 45 (2009) 704–710.
[39] C. Price, X. Zhou, W. Li, L. Wang, Real-time measurement of solute transport within the lacunar-canalicular system of mechanically loaded bone: direct evidence for load-induced fluid flow, J. Bone Miner. Res. 26 (2011) 277–285.
[40] A. Matsugaki, G. Aramoto, T. Ninomiya, H. Sawada, S. Hata, T. Nakano, Abnormal arrangement of a collagen/apatite extracellular matrix orthogonal to osteoblast alignment is constructed by a nanoscale periodic surface structure, Biomaterials 37 (2015) 134–143.
[41] S.W. Verbruggen, T.J. Vaughan, L.M. McNamara, Fluid flow in the osteocyte mechanical environment: a fluid–structure interaction approach, Biomech. Model. Mechanobiol. 13 (2014) 85–97.
[42] C.R. Jacobs, C.E. Yellowley, B.R. Davis, Z. Zhou, J.M. Cimbala, H.J. Donahue, Differential effect of steady versus oscillating flow on bone cells, J. Biomech. 31 (1998) 969–976.
[43] J. Li, E. Rose, D. Frances, Y. Sun, L. You, Effect of oscillating fluid flow stimulation on osteocyte mRNA expression, J. Biomech. 45 (2012) 247–251.
[44] M. Hu, G.-W. Tian, D.E. Gibbons, J. Jiao, Y.-X. Qin, Dynamic fluid flow induced mechanobiological modulation of in situ osteocyte calcium oscillations, Arch. Biochem. Biophys. 579 (2015) 55–61.
[45] S. Wang, S. Li, M. Hu, B. Huo, Calcium response in bone cells at different osteogenic stages under unidirectional or oscillatory flow, Biomicrofluidics 13 (2019), 064117.
[46] C. Wittkowske, G.C. Reilly, D. Lacroix, C.M. Perrault, In vitro bone cell models: impact of fluid shear stress on bone formation, Front. Bioeng. Biotechnol. 4 (2016) 87.
[47] I.P. Geoghegan, D.A. Hoey, L.M. McNamara, Estrogen deficiency impairs integrin αvβ3-mediated mechanosensation by osteocytes and alters osteoclastogenic paracrine signalling, Sci. Rep. 9 (2019) 1–15.
[48] M.M. Thi, S.O. Suadicani, M.B. Schaffler, S. Weinbaum, D.C. Spray, Mechanosensory responses of osteocytes to physiological forces occur along processes and not cell body and require αVβ3 integrin, Proc. Natl. Acad. Sci. Unit. States Am. 110 (2013) 21012–21017.
[49] R.M. Delaine-Smith, A. Sittichokechaiwut, G.C. Reilly, Primary cilia respond to fluid shear stress and mediate flow-induced calcium deposition in osteoblasts, Faseb. J. 28 (2014) 430–439.
[50] C. Galli, G. Passeri, G.M. Macaluso, Osteocytes and WNT: the mechanical control of bone formation, J. Dent. Res. 89 (2010) 331–343.
[51] D.A. Hoey, S. Tormey, S. Ramcharan, F.J. O’Brien, C.R. Jacobs, Primary cilia mediated mechanotransduction in human mesenchymal stem cells, Stem Cell. 30 (2012) 2561–2570.
[52] A.G. Robling, et al., Mechanical stimulation of bone in vivo reduces osteocyte expression of Sost/sclerostin, J. Biol. Chem. 283 (2008) 5866–5875.
[53] M.A. Kamel, J.L. Picconi, N. Lara-Castillo, M.L. Johnson, Activation of β-catenin si3naling in MLO-Y4 osteocytic cells versus 2T3 osteoblastic cells by fluid flow shear stress and PGE2: implications for the study of mechanosensation in bone, Bone 47 (2010) 872–881.
[54] M.R. Forwood, Inducible cyclo-oxygenase (COX-2) mediates the induction of bone formation by mechanical loading in vivo, J. Bone Miner. Res. 11 (1996) 1688–1693.
[55] P.P. Cherian, A.J. Siller-Jackson, S. Gu, X. Wang, L.F. Bonewald, E. Sprague, J. X. Jiang, Mechanical strain opens connexin 43 hemichannels in osteocytes: a novel mechanism for the release of prostaglandin, Mol. Biol. Cell 16 (2005) 3100–3106.
[56] Y. Boie, R. Stocco, N. Sawyer, D.M. Slipetz, M.D. Ungrin, F. Neuscha¨fer-Rube, G. P. Püschel, K.M. Metters, M. Abramovitz, Molecular cloning and characterization of the four rat prostaglandin E2 prostanoid receptor subtypes, Eur. J. Pharmacol. 340 (1997) 227–241.
[57] R. Mizuno, K. Kawada, Y. Sakai, Prostaglandin E2/EP signaling in the tumor microenvironment of colorectal cancer, Int. J. Mol. Sci. 20 (2019) 6254.
[58] S. Graham, Z. Gamie, I. Polyzois, A.A. Narvani, K. Tzafetta, E. Tsiridis, M. Heliotis, A. Mantalaris, E. Tsiridis, Prostaglandin EP2 and EP4 receptor agonists in bone formation and bone healing: in vivo and in vitro evidence, Expet Opin. Invest. Drugs 18 (2009) 749–766.
[59] D. Shamir, S. Keila, M. Weinreb, A selective EP4 receptor antagonist abrogates the stimulation of osteoblast recruitment from bone marrow stromal cells by prostaglandin E2 in vivo and in vitro, Bone 34 (2004) 157–162.
[60] K. Müller-Decker, C. Leder, M. Neumann, G. Neufang, F. Marks, G. Fürstenberger, C. Bayerl, J. Schweizer, Expression of cyclooxygenase isozymes during morphogenesis and cycling of pelage hair follicles in mouse skin: precocious onset of the first catagen phase and alopecia upon cyclooxygenase-2 overexpression, J. Invest. Dermatol. 121 (2003) 661–668.
[61] J.J. Egan, G. Gronowicz, G.A. Rodan, Cell density-dependent decrease in cytoskeletal actin and myosin in cultured osteoblastic cells: correlation with cyclic AMP changes, J. Cell. Biochem. 45 (1991) 93–100.
[62] C.H. Tang, R.S. Yang, W.M. Fu, Prostaglandin E2 stimulates fibronectin expression through EP1 receptor, phospholipase C, protein kinase C alpha, and c-Src pathway in primary cultured rat osteoblasts, J. Biol. Chem. 280 (2005) 22907–22916.
[63] A.V. Taubenberger, M.A. Woodruff, H. Bai, D.J. Muller, D.W. Hutmacher, The effect of unlocking RGD-motifs in collagen I on pre-osteoblast adhesion and differentiation, Biomaterials 31 (2010) 2827–2835.
[64] A. Matsugaki, S. Matsumoto, T. Nakano, A novel role of interleukin-6 as a regulatory factor of inflammation-associated deterioration in osteoblast arrangement, Int. J. Mol. Sci. 21 (2020) 6659.