1.
J. A. Siddiqui, N. C. Partridge, Physiological Bone Remodeling: Systemic Regulation and
Growth Factor Involvement. Physiology (Bethesda, Md.) 31, 233-245 (2016).
2.
E. Hsu, R. Pacifici, From Osteoimmunology to Osteomicrobiology: How the Microbiota
and the Immune System Regulate Bone. Calcif Tissue Int 102, 512-521 (2018).
3.
K. Okamoto, H. Takayanagi, Osteoimmunology. Cold Spring Harbor perspectives in
medicine 9, (2019).
4.
T. J. Guzik, R. Korbut, T. Adamek-Guzik, Nitric oxide and superoxide in inflammation
and immune regulation. J Physiol Pharmacol : an official journal of the Polish
Physiological Society 54, 469-487 (2003).
5.
H. Hikiji, T. Takato, T. Shimizu, S. Ishii, The roles of prostanoids, leukotrienes, and
platelet-activating factor in bone metabolism and disease. Prog Lipid Res 47, 107-126
(2008).
6.
K. Ghimire, H. M. Altmann, A. C. Straub, J. S. Isenberg, Nitric oxide: what's new to
NO? Am J Physiol Cell Physiol 312, C254-c262 (2017).
7.
S. Moncada, R. M. Palmer, E. A. Higgs, Nitric oxide: physiology, pathophysiology, and
pharmacology. Pharmacological Reviews 43, 109 (1991).
8.
T. Okuno, T. Yokomizo, T. Hori, M. Miyano, T. Shimizu, Leukotriene B4 receptor and
the function of its helix 8. J Biol Chem 280, 32049-32052 (2005).
9.
T. R. Arnett, I. R. Orriss, Metabolic properties of the osteoclast. Bone 115, 25-30
(2018).
10.
M. Hukkanen et al., Cytokine-stimulated expression of inducible nitric oxide synthase by
mouse, rat, and human osteoblast-like cells and its functional role in osteoblast metabolic
activity. Endocrinology 136, 5445-5453 (1995).
11.
C. W. Löwik, P. H. Nibbering, M. van de Ruit, S. E. Papapoulos, Inducible production of
nitric oxide in osteoblast-like cells and in fetal mouse bone explants is associated with
suppression of osteoclastic bone resorption. J Clin Invest 93, 1465-1472 (1994).
12.
I. MacIntyre et al., Osteoclastic inhibition: an action of nitric oxide not mediated by
cyclic GMP. Proc Natl Acad Sci U S A 88, 2936-2940 (1991).
13.
N. Takahashi et al., Osteoblastic cells are involved in osteoclast formation.
Endocrinology 123, 2600-2602 (1988).
14.
J. K. Liao, W. S. Shin, W. Y. Lee, S. L. Clark, Oxidized low-density lipoprotein decreases
the expression of endothelial nitric oxide synthase. J Biol Chem 270, 319-324 (1995).
15.
C. R. Lyons, G. J. Orloff, J. M. Cunningham, Molecular cloning and functional
expression of an inducible nitric oxide synthase from a murine macrophage cell line. J
130
Biol Chem 267, 6370-6374 (1992).
16.
A. J. Celeste et al., Isolation of the human gene for bone gla protein utilizing mouse and
rat cDNA clones. The EMBO journal 5, 1885-1890 (1986).
17.
L. C. Green et al., Analysis of nitrate, nitrite, and [15N]nitrate in biological fluids.
Analytical biochemistry 126, 131-138 (1982).
18.
M. Noda et al., cDNA cloning of alkaline phosphatase from rat osteosarcoma (ROS
17/2.8) cells. J Bone Miner Res 2, 161-164 (1987).
19.
M. Gowen, D. D. Wood, E. J. Ihrie, M. K. McGuire, R. G. Russell, An interleukin 1 like
factor stimulates bone resorption in vitro. Nature 306, 378-380 (1983).
20.
D. R. Bertolini, G. E. Nedwin, T. S. Bringman, D. D. Smith, G. R. Mundy, Stimulation
of bone resorption and inhibition of bone formation in vitro by human tumour necrosis
factors. Nature 319, 516-518 (1986).
21.
C. M. Maragos et al., Complexes of .NO with nucleophiles as agents for the controlled
biological release of nitric oxide. Vasorelaxant effects. J Med Chem 34, 3242-3247
(1991).
22.
W. S. Shin et al., Autocrine and paracrine effects of endothelium-derived relaxing factor
on intracellular Ca2+ of endothelial cells and vascular smooth muscle cells. Identification
by two-dimensional image analysis in coculture. J Biol Chem 267, 20377-20382 (1992).
23.
Y. Wang et al., Contribution of sustained Ca2+ elevation for nitric oxide production in
endothelial cells and subsequent modulation of Ca2+ transient in vascular smooth
muscle cells in coculture. J Biol Chem 271, 5647-5655 (1996).
24.
J. A. Riancho et al., Expression and functional role of nitric oxide synthase in osteoblastlike cells. J Bone Miner Res 10, 439-446 (1995).
25.
E. Bornefalk, S. Ljunghall, A. G. Johansson, K. Nilsson, O. Ljunggren, Interleukin-1 beta
induces cyclic AMP formation in isolated human osteoblasts: a signalling mechanism
that is not related to enhanced prostaglandin formation. Bone and mineral 27, 97-107
(1994).
26.
A. J. de Brum-Fernandes et al., Expression of prostaglandin endoperoxide synthase-1
and prostaglandin endoperoxide synthase-2 in human osteoblasts. Biochem Biophys Res
Commun 198, 955-960 (1994).
27.
J. S. Beckman, T. W. Beckman, J. Chen, P. A. Marshall, B. A. Freeman, Apparent
hydroxyl radical production by peroxynitrite: implications for endothelial injury from
nitric oxide and superoxide. Proc Natl Acad Sci U S A 87, 1620-1624 (1990).
28.
A. Inoue, Y. Hiruma, S. Hirose, A. Yamaguchi, H. Hagiwara, Reciprocal regulation by
cyclic nucleotides of the differentiation of rat osteoblast-like cells and mineralization of
nodules. Biochem Biophys Res Commun 215, 1104-1110 (1995).
131
29.
H. Hagiwara et al., cGMP produced in response to ANP and CNP regulates proliferation
and differentiation of osteoblastic cells. Am J Physiol 270, C1311-1318 (1996).
30.
J. Lian et al., Structure of the rat osteocalcin gene and regulation of vitamin Ddependent expression. Proc Natl Acad Sci U S A 86, 1143-1147 (1989).
31.
G. S. Stein, J. B. Lian, Molecular Mechanisms Mediating Developmental and Hormone
Regulated Expression of Genes in Osteoblasts. (Academic Press, New York, NY, ed.
1st, 1993).
32.
H. Kawaguchi, C. C. Pilbeam, J. R. Harrison, L. G. Raisz, The role of prostaglandins in
the regulation of bone metabolism. Clin Orthop Relat Res 36-46 (1995).
33.
H. Ozawa et al., Effect of a continuously applied compressive pressure on mouse
osteoblast-like cells (MC3T3-E1) in vitro. J Cell Physiol 142, 177-185 (1990).
34.
B. Brüne, E. G. Lapetina, Activation of a cytosolic ADP-ribosyltransferase by nitric
oxide-generating agents. J Biol Chem 264, 8455-8458 (1989).
35.
U. C. Garg, A. Hassid, Nitric oxide decreases cytosolic free calcium in Balb/c 3T3
fibroblasts by a cyclic GMP-independent mechanism. J Biol Chem 266, 9-12 (1991).
36.
D. C. Hooper et al., Uric acid, a natural scavenger of peroxynitrite, in experimental
allergic encephalomyelitis and multiple sclerosis. Proc Natl Acad Sci U S A 95, 675-680
(1998).
37.
H. Ischiropoulos, L. Zhu, J. S. Beckman, Peroxynitrite formation from macrophagederived nitric oxide. Arch Biochem Biophys 298, 446-451 (1992).
38.
H. Hikiji et al., Direct action of nitric oxide on osteoblastic differentiation. FEBS letters
410, 238-242 (1997).
39.
M. Weinreb, D. Shinar, G. A. Rodan, Different pattern of alkaline phosphatase,
osteopontin, and osteocalcin expression in developing rat bone visualized by in situ
hybridization. J Bone Miner Res 5, 831-842 (1990).
40.
C. S. Lader, A. M. Flanagan, Prostaglandin E2, interleukin 1alpha, and tumor necrosis
factor-alpha increase human osteoclast formation and bone resorption in vitro.
Endocrinology 139, 3157-3164 (1998).
41.
C. Szabó, B. Zingarelli, A. L. Salzman, Role of poly-ADP ribosyltransferase activation in
the vascular contractile and energetic failure elicited by exogenous and endogenous
nitric oxide and peroxynitrite. Circ Res 78, 1051-1063 (1996).
42.
H. Kawaguchi et al., In vivo gene transfection of human endothelial cell nitric oxide
synthase in cardiomyocytes causes apoptosis-like cell death. Identification using Sendai
virus-coated liposomes. Circulation 95, 2441-2447 (1997).
43.
A. G. Estévez et al., Examining apoptosis in cultured cells after exposure to nitric oxide
and peroxynitrite. Methods in enzymology 301, 393-402 (1999).
132
44.
Y. W. Xie, M. S. Wolin, Role of nitric oxide and its interaction with superoxide in the
suppression of cardiac muscle mitochondrial respiration. Involvement in response to
hypoxia/reoxygenation. Circulation 94, 2580-2586 (1996).
45.
J. S. Beckmann et al., Extensive nitration of protein tyrosines in human atherosclerosis
detected by immunohistochemistry. Biol Chem Hoppe Seyler 375, 81-88 (1994).
46.
D. M. Evans, S. H. Ralston, Nitric oxide and bone. J Bone Miner Res 11, 300-305
(1996).
47.
H. MC, L. JA, Local regulators of bone: IL-1, TNF, lymphotoxin, interferon 7, IL-8, IL-
10, IL-4, the LIF/IL-6 family, and additional cytokines., Principles of Bone Biology
(Academic Press, San Diego, CA, 1996).
48.
D. B. Evans, M. Thavarajah, J. A. Kanis, Involvement of prostaglandin E2 in the
inhibition of osteocalcin synthesis by human osteoblast-like cells in response to cytokines
and systemic hormones. Biochem Biophys Res Commun 167, 194-202 (1990).
49.
R. S. Taichman, P. V. Hauschka, Effects of interleukin-1 beta and tumor necrosis factoralpha on osteoblastic expression of osteocalcin and mineralized extracellular matrix in
vitro. Inflammation 16, 587-601 (1992).
50.
R. S. Lewis, S. Tamir, S. R. Tannenbaum, W. M. Deen, Kinetic analysis of the fate of
nitric oxide synthesized by macrophages in vitro. J Biol Chem 270, 29350-29355 (1995).
51.
H. Ischiropoulos et al., Peroxynitrite-mediated tyrosine nitration catalyzed by superoxide
dismutase. Arch Biochem Biophys 298, 431-437 (1992).
52.
C. R. White et al., Superoxide and peroxynitrite in atherosclerosis. Proc Natl Acad Sci U
S A 91, 1044-1048 (1994).
53.
J. J. Poderoso et al., Nitric oxide regulates oxygen uptake and hydrogen peroxide release
by the isolated beating rat heart. Am J Physiol 274, C112-119 (1998).
54.
P. D. Damoulis, P. V. Hauschka, Nitric oxide acts in conjunction with proinflammatory
cytokines to promote cell death in osteoblasts. J Bone Miner Res 12, 412-422 (1997).
55.
Y. Hou, J. Wang, J. Ramirez, P. G. Wang, in Methods in Enzymology. (Academic Press,
San Diego, CA, 1999), vol. 301, pp. 242-249.
56.
S. W. Fox, T. J. Chambers, J. W. Chow, Nitric oxide is an early mediator of the increase
in bone formation by mechanical stimulation. Am J Physiol 270, E955-960 (1996).
57.
C. Melchiorri et al., Enhanced and coordinated in vivo expression of inflammatory
cytokines and nitric oxide synthase by chondrocytes from patients with osteoarthritis.
Arthritis Rheum 41, 2165-2174 (1998).
58.
A. W. Ford-Hutchinson, M. A. Bray, M. V. Doig, M. E. Shipley, M. J. Smith,
Leukotriene B, a potent chemokinetic and aggregating substance released from
polymorphonuclear leukocytes. Nature 286, 264-265 (1980).
133
59.
B. Samuelsson, S. E. Dahlén, J. A. Lindgren, C. A. Rouzer, C. N. Serhan, Leukotrienes
and lipoxins: structures, biosynthesis, and biological effects. Science (New York, N.Y.)
237, 1171-1176 (1987).
60.
T. Yokomizo, T. Izumi, K. Chang, Y. Takuwa, T. Shimizu, A G-protein-coupled receptor
for leukotriene B4 that mediates chemotaxis. Nature 387, 620-624 (1997).
61.
T. Yokomizo, K. Kato, K. Terawaki, T. Izumi, T. Shimizu, A second leukotriene B(4)
receptor, BLT2. A new therapeutic target in inflammation and immunological disorders.
The J Exp Med 192, 421-432 (2000).
62.
T. Yokomizo, T. Izumi, T. Shimizu, Leukotriene B4: metabolism and signal
transduction. Arch Biochem Biophys 385, 231-241 (2001).
63.
T. Yokomizo, K. Kato, H. Hagiya, T. Izumi, T. Shimizu, Hydroxyeicosanoids bind to and
activate the low affinity leukotriene B4 receptor, BLT2. J Biol Chem 276, 12454-12459
(2001).
64.
T. Okuno et al., 12(S)-Hydroxyheptadeca-5Z, 8E, 10E-trienoic acid is a natural ligand
for leukotriene B4 receptor 2. Journal Exp Med 205, 759-766 (2008).
65.
L. Iversen, K. Kragballe, V. A. Ziboh, Significance of leukotriene-A4 hydrolase in the
pathogenesis of psoriasis. Skin pharmacol : the official journal of the Skin Pharmacology
Society 10, 169-177 (1997).
66.
Z. Csoma et al., Increased leukotrienes in exhaled breath condensate in childhood
asthma. Am J Respir Crit Care Med 166, 1345-1349 (2002).
67.
A. T. Cole et al., Mucosal factors inducing neutrophil movement in ulcerative colitis: the
role of interleukin 8 and leukotriene B4. Gut 39, 248-254 (1996).
68.
B. J. Zimmerman, D. J. Guillory, M. B. Grisham, T. S. Gaginella, D. N. Granger, Role of
leukotriene B4 in granulocyte infiltration into the postischemic feline intestine.
Gastroenterol 99, 1358-1363 (1990).
69.
L. B. Klickstein, C. Shapleigh, E. J. Goetzl, Lipoxygenation of arachidonic acid as a
source of polymorphonuclear leukocyte chemotactic factors in synovial fluid and tissue in
rheumatoid arthritis and spondyloarthritis. J Clin Invest 66, 1166-1170 (1980).
70.
E. M. Davidson, S. A. Rae, M. J. Smith, Leukotriene B4, a mediator of inflammation
present in synovial fluid in rheumatoid arthritis. Ann Rheum Dis 42, 677-679 (1983).
71.
N. Ahmadzadeh, M. Shingu, M. Nobunaga, T. Tawara, Relationship between
leukotriene B4 and immunological parameters in rheumatoid synovial fluids.
Inflammation 15, 497-503 (1991).
72.
R. J. Griffiths et al., Leukotriene B4 plays a critical role in the progression of collageninduced arthritis. Proc Natl Acad Sci U S A 92, 517-521 (1995).
73.
H. Yasuda et al., A novel molecular mechanism modulating osteoclast differentiation and
134
function. Bone 25, 109-113 (1999).
74.
T. Suda, Y. Ueno, K. Fujii, T. Shinki, Vitamin D and bone. J Cell Biochem 88, 259-266
(2003).
75.
S. Meghji, J. R. Sandy, A. M. Scutt, W. Harvey, M. Harris, Stimulation of bone
resorption by lipoxygenase metabolites of arachidonic acid. Prostaglandins 36, 139-149
(1988).
76.
C. Garcia et al., Leukotriene B4 stimulates osteoclastic bone resorption both in vitro and
in vivo. J Bone Miner Res 11, 1619-1627 (1996).
77.
J. B. Smith, M. K. Haynes, Rheumatoid arthritis--a molecular understanding. Ann Intern
Med 136, 908-922 (2002).
78.
N. D. Kim, R. C. Chou, E. Seung, A. M. Tager, A. D. Luster, A unique requirement for
the leukotriene B4 receptor BLT1 for neutrophil recruitment in inflammatory arthritis. J
Exp Med 203, 829-835 (2006).
79.
W. H. Shao, A. Del Prete, C. B. Bock, B. Haribabu, Targeted disruption of leukotriene
B4 receptors BLT1 and BLT2: a critical role for BLT1 in collagen-induced arthritis in
mice. J Immunol 176, 6254-6261 (2006).
80.
M. Hegen et al., Cytosolic phospholipase A2alpha-deficient mice are resistant to
collagen-induced arthritis. J Exp Med 197, 1297-1302 (2003).
81.
M. Chen et al., Neutrophil-derived leukotriene B4 is required for inflammatory arthritis.
J Exp Med 203, 837-842 (2006).
82.
H. J. Showell, R. Breslow, M. J. Conklyn, G. P. Hingorani, K. Koch, Characterization of
the pharmacological profile of the potent LTB4 antagonist CP-105,696 on murine LTB4
receptors in vitro. Br J Pharmacol 117, 1127-1132 (1996).
83.
K. Terawaki et al., Absence of leukotriene B4 receptor 1 confers resistance to airway
hyperresponsiveness and Th2-type immune responses. J Immunol 175, 4217-4225
(2005).
84.
R. J. Sells Galvin, C. L. Gatlin, J. W. Horn, T. R. Fuson, TGF-beta enhances osteoclast
differentiation in hematopoietic cell cultures stimulated with RANKL and M-CSF.
Biochem Biophys Res Commun 265, 233-239 (1999).
85.
Y. Okada et al., Prostaglandin G/H synthase-2 is required for maximal formation of
osteoclast-like cells in culture. J Clin Invest 105, 823-832 (2000).
86.
Y. Iizuka et al., Characterization of a mouse second leukotriene B4 receptor, mBLT2:
BLT2-dependent ERK activation and cell migration of primary mouse keratinocytes. J
Biol Chem 280, 24816-24823 (2005).
87.
A. R. Villanueva, L. Ilnicki, H. M. Frost, R. Arnstein, Measurement of the bone
formation rate in a case of familial hypophosphatemic vitamin D-resistant rickets. J Lab
135
Clin Med 67, 973-982 (1966).
88.
H. M. Frost, Preparation of thin undecalcified bone sections by rapid manual method.
Stain technol 33, 273-277 (1958).
89.
D. Rickard, A. Harris, R. Turner, S. Khosla, T. C. Spelsberg, in Principals of Bone
Biology, J. P. Bilesikian, L. G. Raisz, G. A. Rodan, Eds. (Academic Press, San Diego, CA,
2002), vol. 2, pp. 655-676.
90.
H. Takayanagi et al., T-cell-mediated regulation of osteoclastogenesis by signalling
cross-talk between RANKL and IFN-gamma. Nature 408, 600-605 (2000).
91.
L. Li, A. Khansari, L. Shapira, D. T. Graves, S. Amar, Contribution of interleukin-11 and
prostaglandin(s) in lipopolysaccharide-induced bone resorption in vivo. Infect Immun
70, 3915-3922 (2002).
92.
C. Miyaura et al., An essential role of cytosolic phospholipase A2alpha in prostaglandin
E2-mediated bone resorption associated with inflammation. J Exp Med 197, 1303-1310
(2003).
93.
N. Tapon, A. Hall, Rho, Rac and Cdc42 GTPases regulate the organization of the actin
cytoskeleton. Curr Opin Cell Biol 9, 86-92 (1997).
94.
V. Kölsch, P. G. Charest, R. A. Firtel, The regulation of cell motility and chemotaxis by
phospholipid signaling. J Cell Sci 121, 551-559 (2008).
95.
K. Väänanen, H. Zhao, in Principals of Bone Biology, J. P. Bilesikian, L. G. Raisz, G. A.
Rodan, Eds. (Academic Press, San Diego, CA, 2002), vol. 1, chap. 127-140.
96.
H. K. Väänänen, H. Zhao, in Principals of Bone Biology, J. P. Bilesikian, L. G. Raisz, T.
J. Martin, Eds. (Academic Press, San Diego, 2008), vol. 1, pp. 193-210.
97.
L. T. Duong, A. Sanjay, W. Home, R. Baron, G. A. Rodan, in Principals of Bone Biology,
J. P. Bilesikian, L. G. Raisz, G. A. Rodan, Eds. (Academic Press, San Diego, CA, 2002),
vol. 1, pp. 141-150.
98.
R. Pacifici et al., Spontaneous release of interleukin 1 from human blood monocytes
reflects bone formation in idiopathic osteoporosis. Proc Natl Acad Sci U S A 84, 46164620 (1987).
99.
R. Pacifici et al., Effect of surgical menopause and estrogen replacement on cytokine
release from human blood mononuclear cells. Proc Natl Acad Sci U S A 88, 5134-5138
(1991).
100.
R. T. Turner, B. L. Riggs, T. C. Spelsberg, Skeletal effects of estrogen. Endocr Rev 15,
275-300 (1994).
101.
J. Ciampolini, K. G. Harding, Pathophysiology of chronic bacterial osteomyelitis. Why
do antibiotics fail so often? Postgrad Med J 76, 479-483 (2000).
102.
L. Duplomb, M. Dagouassat, P. Jourdon, D. Heymann, Concise review: embryonic stem
136
cells: a new tool to study osteoblast and osteoclast differentiation. Stem cells (Dayton,
Ohio) 25, 544-552 (2007).
103.
H. Hikiji, S. Ishii, H. Shindou, T. Takato, T. Shimizu, Absence of platelet-activating
factor receptor protects mice from osteoporosis following ovariectomy. J Clin Invest 114,
85-93 (2004).
104.
N. Uozumi et al., Role of cytosolic phospholipase A2 in allergic response and parturition.
Nature 390, 618-622 (1997).
105.
G. Camussi, C. Tetta, F. Bussolino, C. Baglioni, Tumor necrosis factor stimulates human
neutrophils to release leukotriene B4 and platelet-activating factor. Induction of
phospholipase A2 and acetyl-CoA:1-alkyl-sn-glycero-3-phosphocholine O2acetyltransferase activity and inhibition by antiproteinase. Europ J Biochem 182, 661666 (1989).
106.
P. Conti et al., The combination of interleukin 1 plus tumor necrosis factor causes
greater generation of LTB4, thromboxanes and aggregation on human macrophages
than these compounds alone. Prog Cinic Biol Res 301, 541-545 (1989).
107.
M. E. Surette, N. Dallaire, N. Jean, S. Picard, P. Borgeat, Mechanisms of the priming
effect of lipopolysaccharides on the biosynthesis of leukotriene B4 in chemotactic
peptide-stimulated human neutrophils. Faseb J 12, 1521-1531 (1998).
108.
J. A. Rankin, I. Sylvester, S. Smith, T. Yoshimura, E. J. Leonard, Macrophages cultured
in vitro release leukotriene B4 and neutrophil attractant/activation protein (interleukin
8) sequentially in response to stimulation with lipopolysaccharide and zymosan. J Clin
Invest 86, 1556-1564 (1990).
109.
A. Fukuda et al., Regulation of osteoclast apoptosis and motility by small GTPase
binding protein Rac1. J Bone Miner Res 20, 2245-2253 (2005).
110.
N. Ishida et al., CCR1 acts downstream of NFAT2 in osteoclastogenesis and enhances
cell migration. J Bone Miner Res 21, 48-57 (2006).
111.
Y. Wang et al., Identifying the relative contributions of Rac1 and Rac2 to
osteoclastogenesis. J Bone Miner Res 23, 260-270 (2008).
112.
M. Yagi et al., DC-STAMP is essential for cell-cell fusion in osteoclasts and foreign body
giant cells. J Expb Med 202, 345-351 (2005).
113.
A. Sanjay et al., Cbl associates with Pyk2 and Src to regulate Src kinase activity,
alpha(v)beta(3) integrin-mediated signaling, cell adhesion, and osteoclast motility. J Cell
Biol 152, 181-195 (2001).
114.
S. L. Teitelbaum, Bone resorption by osteoclasts. Science (New York, N.Y.) 289, 15041508 (2000).
115.
C. N. Serhan et al., Reduced inflammation and tissue damage in transgenic rabbits
137
overexpressing 15-lipoxygenase and endogenous anti-inflammatory lipid mediators. J
Immunol 171, 6856-6865 (2003).
116.
H. Hasturk et al., Resolvin E1 regulates inflammation at the cellular and tissue level and
restores tissue homeostasis in vivo. J Immunol 179, 7021-7029 (2007).
117.
B. S. Herrera et al., An endogenous regulator of inflammation, resolvin E1, modulates
osteoclast differentiation and bone resorption. Br J Pharmacol 155, 1214-1223 (2008).
118.
C. MacLean et al., Systematic review: comparative effectiveness of treatments to prevent
fractures in men and women with low bone density or osteoporosis. Ann Intern Med
148, 197-213 (2008).
119.
K. Henriksen, J. Bollerslev, V. Everts, M. A. Karsdal, Osteoclast activity and subtypes as
a function of physiology and pathology--implications for future treatments of
osteoporosis. Endocr Rev 32, 31-63 (2011).
120.
R. Baron, L. Neff, D. Louvard, P. J. Courtoy, Cell-mediated extracellular acidification
and bone resorption: evidence for a low pH in resorbing lacunae and localization of a
100-kD lysosomal membrane protein at the osteoclast ruffled border. J Cell Biol 101,
2210-2222 (1985).
121.
J. A. Wemmie, M. P. Price, M. J. Welsh, Acid-sensing ion channels: advances, questions
and therapeutic opportunities. Trends Neurosci 29, 578-586 (2006).
122.
K. Seuwen, M. G. Ludwig, R. M. Wolf, Receptors for protons or lipid messengers or
both? J Recept Signal Transduct Res 26, 599-610 (2006).
123.
M. G. Ludwig et al., Proton-sensing G-protein-coupled receptors. Nature 425, 93-98
(2003).
124.
N. Murakami, T. Yokomizo, T. Okuno, T. Shimizu, G2A is a proton-sensing G-proteincoupled receptor antagonized by lysophosphatidylcholine. J Biol Chem 279, 4248442491 (2004).
125.
S. Ishii, Y. Kihara, T. Shimizu, Identification of T cell death-associated gene 8 (TDAG8)
as a novel acid sensing G-protein-coupled receptor. J Biol Chem 280, 9083-9087 (2005).
126.
J. Q. Wang et al., TDAG8 is a proton-sensing and psychosine-sensitive G-proteincoupled receptor. J Biol Chem 279, 45626-45633 (2004).
127.
D. S. Im, C. E. Heise, T. Nguyen, B. F. O'Dowd, K. R. Lynch, Identification of a
molecular target of psychosine and its role in globoid cell formation. J Cell Biol 153, 429434 (2001).
128.
C. Mogi et al., Involvement of proton-sensing TDAG8 in extracellular acidificationinduced inhibition of proinflammatory cytokine production in peritoneal macrophages. J
Immunol 182, 3243-3251 (2009).
129.
Y. Ihara et al., The G protein-coupled receptor T-cell death-associated gene 8 (TDAG8)
138
facilitates tumor development by serving as an extracellular pH sensor. Proc Natl Acad
Sci U S A 107, 17309-17314 (2010).
130.
R. Turner, D. Rickard, U. T. Iwaniec, T. C. Spelsberg, in Principals of Bone Biology, J.
P. Bilesikian, L. G. Raisz, T. J. Martin, Eds. (Academic Press, San Diego, 2008), vol. 1,
pp. 855-886.
131.
K. Horie et al., Characterization of Sleeping Beauty transposition and its application to
genetic screening in mice. Mol Cell Biol 23, 9189-9207 (2003).
132.
C. G. Radu et al., Normal immune development and glucocorticoid-induced thymocyte
apoptosis in mice deficient for the T-cell death-associated gene 8 receptor. Mol Cell Biol
26, 668-677 (2006).
133.
N. Tosa et al., Critical function of T cell death-associated gene 8 in glucocorticoidinduced thymocyte apoptosis. Int Immunol 15, 741-749 (2003).
134.
W. C. Horne, L. T. Duong, A. Sanjay, R. Baron, in Principals of Bone Biology, J. P.
Bilesikian, L. G. Raisz, T. J. Martin, Eds. (Academic Press, San Diego, 2008), vol. 1, pp.
221-236.
135.
H. Niwa, K. Yamamura, J. Miyazaki, Efficient selection for high-expression transfectants
with a novel eukaryotic vector. Gene 108, 193-199 (1991).
136.
A. Bruzzaniti et al., Dynamin forms a Src kinase-sensitive complex with Cbl and
regulates podosomes and osteoclast activity. Mol Biol Cell 16, 3301-3313 (2005).
137.
A. J. Ridley, A. Hall, The small GTP-binding protein rho regulates the assembly of focal
adhesions and actin stress fibers in response to growth factors. Cell 70, 389-399 (1992).
138.
M. A. Chellaiah et al., Rho-A is critical for osteoclast podosome organization, motility,
and bone resorption. J Biol Chem 275, 11993-12002 (2000).
139.
M. Uehata et al., Calcium sensitization of smooth muscle mediated by a Rho-associated
protein kinase in hypertension. Nature 389, 990-994 (1997).
140.
M. K. Osako et al., Estrogen inhibits vascular calcification via vascular RANKL system:
common mechanism of osteoporosis and vascular calcification. Circ Res 107, 466-475
(2010).
141.
H. Feng et al., Myocyte enhancer factor 2 and microphthalmia-associated transcription
factor cooperate with NFATc1 to transactivate the V-ATPase d2 promoter during
RANKL-induced osteoclastogenesis. J Biol Chem 284, 14667-14676 (2009).
142.
D. Q. Yang et al., V-ATPase subunit ATP6AP1 (Ac45) regulates osteoclast
differentiation, extracellular acidification, lysosomal trafficking, and protease exocytosis
in osteoclast-mediated bone resorption. J Bone Miner Res 27, 1695-1707 (2012).
143.
K. Iwai et al., RGS18 acts as a negative regulator of osteoclastogenesis by modulating the
acid-sensing OGR1/NFAT signaling pathway. J Bone Miner Res 22, 1612-1620 (2007).
139
144.
A. Pereverzev et al., Extracellular acidification enhances osteoclast survival through an
NFAT-independent, protein kinase C-dependent pathway. Bone 42, 150-161 (2008).
145.
M. Yang et al., Expression of and role for ovarian cancer G-protein-coupled receptor 1
(OGR1) during osteoclastogenesis. J Biol Chem 281, 23598-23605 (2006).
146.
M. Chellaiah et al., Gelsolin deficiency blocks podosome assembly and produces
increased bone mass and strength. J Cell Biol 148, 665-678 (2000).
147.
S. Linder, P. Kopp, Podosomes at a glance. J Cell Sci 118, 2079-2082 (2005).
148.
B. Zhao, TNF and Bone Remodeling. Current osteoporosis reports 15, 126-134 (2017).
140
...