Publications

Conflicts of interest The authors declare no conflict of interest. ACKNOWLEDGMENTS We thank the anonymous reviewers for reviewing this manuscript. REFERENCES Amemiya H, Matsuzaka K, Kokubu E, Ohta S, et al (2008). Cellular responses of rat periodontal ligament cells under hypoxia and re-oxygenation conditions in vitro. J. Periodontal Res. 43: 322-327. http://dx.doi.org/10.1111/j.1600-0765.2007.01032.x Bernet JD, Doles JD, Hall JK, Kelly Tanaka K, et al (2014). p38 MAPK signaling underlies a cell-autonomous loss of stem cell self-renewal in skeletal muscle of aged mice. Nat. Med. 20: 265-271. http://dx.doi.org/10.1038/nm.3465 Corbet EF, et al (2006). Periodontal diseases in Asians. J. Int. Acad. Periodontol. 8: 136-144. Dumitrescu AL, et al (2016). Editorial: Periodontal Disease - A Public Health Problem. Front. Public Health 3: 278. http://dx.doi.org/10.3389/fpubh.2015.00278 Eke PI, Dye BA, Wei L, Slade GD, et al (2015). Update on prevalence of periodontitis in adults in the United States: NHANES 2009 to 2012. J. Periodontol. 86: 611-622. http://dx.doi.org/10.1902/jop.2015.140520 Gay IC, Chen S, MacDougall M, et al (2007). Isolation and characterization of multipotent human periodontal ligament stem cells. Orthod. Craniofac. Res. 10: 149-160. http://dx.doi.org/10.1111/j.1601-6343.2007.00399.x Hackett PH, Roach RC, et al (2001). High-altitude illness. N. Engl. J. Med. 345: 107-114. http://dx.doi.org/10.1056/NEJM200107123450206 Jaiswal RK, Jaiswal N, Bruder SP, Mbalaviele G, et al (2000). Adult human mesenchymal stem cell differentiation to the osteogenic or adipogenic lineage is regulated by mitogen-activated protein kinase. J. Biol. Chem. 275: 9645-9652. http://dx.doi.org/10.1074/jbc.275.13.9645 Jian C, Li C, Ren Y, He Y, et al (2014). Hypoxia augments lipopolysaccharide-induced cytokine expression in periodontal ligament cells. Inflammation 37: 1413-1423. http://dx.doi.org/10.1007/s10753-014-9865-6 Li Q, Yu B, Yang P, et al (2015). Hypoxia-induced HMGB1 in would tissues promotes the osteoblast cell proliferation via activating ERK/JNK signaling. Int. J. Clin. Exp. Med. 8: 15087-15097. Liu Q, Cen L, Zhou H, Yin S, et al (2009). The role of the extracellular signal-related kinase signaling pathway in osteogenic differentiation of human adipose-derived stem cells and in adipogenic transition initiated by dexamethasone. Tissue Eng. Part A 15: 3487-3497. http://dx.doi.org/10.1089/ten.tea.2009.0175 Matsuda N, Morita N, Matsuda K, Watanabe M, et al (1998). Proliferation and differentiation of human osteoblastic cells associated with differential activation of MAP kinases in response to epidermal growth factor, hypoxia, and mechanical stress in vitro. Biochem. Biophys. Res. Commun. 249: 350-354. http://dx.doi.org/10.1006/bbrc.1998.9151 Mattioli-Belmonte M, Teti G, Salvatore V, Focaroli S, et al (2015). Stem cell origin differently affects bone tissue engineering strategies. Front. Physiol. 6: 266. http://dx.doi.org/10.3389/fphys.2015.00266 Park SY, Kim KH, Gwak EH, Rhee SH, et al (2015). Ex vivo bone morphogenetic protein 2 gene delivery using periodontal ligament stem cells for enhanced re-osseointegration in the regenerative treatment of peri-implantitis. J. Biomed. Mater. Res. A 103: 38-47. http://dx.doi.org/10.1002/jbm.a.35145 Qiu X, Zheng M, Song D, Huang L, et al (2016). Notoginsenoside Rb1 inhibits activation of ERK and p38 MAPK pathways induced by hypoxia and hypercapnia. Exp. Ther. Med. 11: 2455-2461. Rodríguez-Carballo E, Gámez B, Ventura F, et al (2016). p38 MAPK Signaling in Osteoblast Differentiation. Front. Cell Dev. Biol. 4: 40. http://dx.doi.org/10.3389/fcell.2016.00040 Seo BM, Miura M, Gronthos S, Bartold PM, et al (2004). Investigation of multipotent postnatal stem cells from human periodontal ligament. Lancet 364: 149-155. http://dx.doi.org/10.1016/S0140-6736(04)16627-0 Somerman MJ, Young MF, Foster RA, Moehring JM, et al (1990). Characteristics of human periodontal ligament cells in vitro. Arch. Oral Biol. 35: 241-247. http://dx.doi.org/10.1016/0003-9969(90)90062-F Sun Y, Liu WZ, Liu T, Feng X, et al (2015). Signaling pathway of MAPK/ERK in cell proliferation, differentiation, migration, senescence and apoptosis. J. Recept. Signal Transduct. Res. 35: 600-604. http://dx.doi.org/10.3109/10799893.2015.1030412 Tang R, Wei F, Wei L, Wang S, et al (2014). Osteogenic differentiated periodontal ligament stem cells maintain their immunomodulatory capacity. J. Tissue Eng. Regen. Med. 8: 226-232. http://dx.doi.org/10.1002/term.1516 Terrizzi AR, Fernandez-Solari J, Lee CM, Bozzini C, et al (2013). Alveolar bone loss associated to periodontal disease in lead intoxicated rats under environmental hypoxia. Arch. Oral Biol. 58: 1407-1414. http://dx.doi.org/10.1016/j.archoralbio.2013.06.010 Trubiani O, Giacoppo S, Ballerini P, Diomede F, et al (2016). Alternative source of stem cells derived from human periodontal ligament: a new treatment for experimental autoimmune encephalomyelitis. Stem Cell Res. Ther. 7: 1. http://dx.doi.org/10.1186/s13287-015-0253-4 Vandana KL, Desai R, Dalvi PJ, et al (2015). Autologous Stem Cell Application in Periodontal Regeneration Technique (SAI-PRT) Using PDLSCs Directly From an Extracted Tooth···An Insight. Int. J. Stem Cells 8: 235-237. http://dx.doi.org/10.15283/ijsc.2015.8.2.235 Wang Z, Wang W, Xu S, Wang S, et al (2016). The role of MAPK signaling pathway in the Her-2-positive meningiomas. Oncol. Rep. 36: 685-695. Wu RX, Bi CS, Yu Y, Zhang LL, et al (2015). Age-related decline in the matrix contents and functional properties of human periodontal ligament stem cell sheets. Acta Biomater. 22: 70-82. http://dx.doi.org/10.1016/j.actbio.2015.04.024 Wu Y, Yang Y, Yang P, Gu Y, et al (2013). The osteogenic differentiation of PDLSCs is mediated through MEK/ERK and p38 MAPK signalling under hypoxia. Arch. Oral Biol. 58: 1357-1368. http://dx.doi.org/10.1016/j.archoralbio.2013.03.011 Xiao X, Li Y, Zhang G, Gao Y, et al (2012). Detection of bacterial diversity in rat’s periodontitis model under imitational altitude hypoxia environment. Arch. Oral Biol. 57: 23-29. http://dx.doi.org/10.1016/j.archoralbio.2011.07.005 Xu CL, Zheng B, Pei JH, Shen SJ, et al (2016). Embelin induces apoptosis of human gastric carcinoma through inhibition of p38 MAPK and NF-κB signaling pathways. Mol. Med. Rep. 14: 307-312. Yang ZH, Zhang XJ, Dang NN, Ma ZF, et al (2009). Apical tooth germ cell-conditioned medium enhances the differentiation of periodontal ligament stem cells into cementum/periodontal ligament-like tissues. J. Periodontal Res. 44: 199-210. http://dx.doi.org/10.1111/j.1600-0765.2008.01106.x Zhang HY, Liu R, Xing YJ, Xu P, et al (2013). Effects of hypoxia on the proliferation, mineralization and ultrastructure of human periodontal ligament fibroblasts in vitro. Exp. Ther. Med. 6: 1553-1559. Zhang QB, Zhang ZQ, Fang SL, Liu YR, et al (2014). Effects of hypoxia on proliferation and osteogenic differentiation of periodontal ligament stem cells: an in vitro and in vivo study. Genet. Mol. Res. 13: 10204-10214. http://dx.doi.org/10.4238/2014.December.4.15

Conflicts of interest The authors declare no conflict of interest. ACKNOWLEDGMENTS We thank the anonymous reviewers for reviewing this manuscript. REFERENCES Amemiya H, Matsuzaka K, Kokubu E, Ohta S, et al (2008). Cellular responses of rat periodontal ligament cells under hypoxia and re-oxygenation conditions in vitro. J. Periodontal Res. 43: 322-327. http://dx.doi.org/10.1111/j.1600-0765.2007.01032.x Bernet JD, Doles JD, Hall JK, Kelly Tanaka K, et al (2014). p38 MAPK signaling underlies a cell-autonomous loss of stem cell self-renewal in skeletal muscle of aged mice. Nat. Med. 20: 265-271. http://dx.doi.org/10.1038/nm.3465 Corbet EF, et al (2006). Periodontal diseases in Asians. J. Int. Acad. Periodontol. 8: 136-144. Dumitrescu AL, et al (2016). Editorial: Periodontal Disease - A Public Health Problem. Front. Public Health 3: 278. http://dx.doi.org/10.3389/fpubh.2015.00278 Eke PI, Dye BA, Wei L, Slade GD, et al (2015). Update on prevalence of periodontitis in adults in the United States: NHANES 2009 to 2012. J. Periodontol. 86: 611-622. http://dx.doi.org/10.1902/jop.2015.140520 Gay IC, Chen S, MacDougall M, et al (2007). Isolation and characterization of multipotent human periodontal ligament stem cells. Orthod. Craniofac. Res. 10: 149-160. http://dx.doi.org/10.1111/j.1601-6343.2007.00399.x Hackett PH, Roach RC, et al (2001). High-altitude illness. N. Engl. J. Med. 345: 107-114. http://dx.doi.org/10.1056/NEJM200107123450206 Jaiswal RK, Jaiswal N, Bruder SP, Mbalaviele G, et al (2000). Adult human mesenchymal stem cell differentiation to the osteogenic or adipogenic lineage is regulated by mitogen-activated protein kinase. J. Biol. Chem. 275: 9645-9652. http://dx.doi.org/10.1074/jbc.275.13.9645 Jian C, Li C, Ren Y, He Y, et al (2014). Hypoxia augments lipopolysaccharide-induced cytokine expression in periodontal ligament cells. Inflammation 37: 1413-1423. http://dx.doi.org/10.1007/s10753-014-9865-6 Li Q, Yu B, Yang P, et al (2015). Hypoxia-induced HMGB1 in would tissues promotes the osteoblast cell proliferation via activating ERK/JNK signaling. Int. J. Clin. Exp. Med. 8: 15087-15097. Liu Q, Cen L, Zhou H, Yin S, et al (2009). The role of the extracellular signal-related kinase signaling pathway in osteogenic differentiation of human adipose-derived stem cells and in adipogenic transition initiated by dexamethasone. Tissue Eng. Part A 15: 3487-3497. http://dx.doi.org/10.1089/ten.tea.2009.0175 Matsuda N, Morita N, Matsuda K, Watanabe M, et al (1998). Proliferation and differentiation of human osteoblastic cells associated with differential activation of MAP kinases in response to epidermal growth factor, hypoxia, and mechanical stress in vitro. Biochem. Biophys. Res. Commun. 249: 350-354. http://dx.doi.org/10.1006/bbrc.1998.9151 Mattioli-Belmonte M, Teti G, Salvatore V, Focaroli S, et al (2015). Stem cell origin differently affects bone tissue engineering strategies. Front. Physiol. 6: 266. http://dx.doi.org/10.3389/fphys.2015.00266 Park SY, Kim KH, Gwak EH, Rhee SH, et al (2015). Ex vivo bone morphogenetic protein 2 gene delivery using periodontal ligament stem cells for enhanced re-osseointegration in the regenerative treatment of peri-implantitis. J. Biomed. Mater. Res. A 103: 38-47. http://dx.doi.org/10.1002/jbm.a.35145 Qiu X, Zheng M, Song D, Huang L, et al (2016). Notoginsenoside Rb1 inhibits activation of ERK and p38 MAPK pathways induced by hypoxia and hypercapnia. Exp. Ther. Med. 11: 2455-2461. Rodríguez-Carballo E, Gámez B, Ventura F, et al (2016). p38 MAPK Signaling in Osteoblast Differentiation. Front. Cell Dev. Biol. 4: 40. http://dx.doi.org/10.3389/fcell.2016.00040 Seo BM, Miura M, Gronthos S, Bartold PM, et al (2004). Investigation of multipotent postnatal stem cells from human periodontal ligament. Lancet 364: 149-155. http://dx.doi.org/10.1016/S0140-6736(04)16627-0 Somerman MJ, Young MF, Foster RA, Moehring JM, et al (1990). Characteristics of human periodontal ligament cells in vitro. Arch. Oral Biol. 35: 241-247. http://dx.doi.org/10.1016/0003-9969(90)90062-F Sun Y, Liu WZ, Liu T, Feng X, et al (2015). Signaling pathway of MAPK/ERK in cell proliferation, differentiation, migration, senescence and apoptosis. J. Recept. Signal Transduct. Res. 35: 600-604. http://dx.doi.org/10.3109/10799893.2015.1030412 Tang R, Wei F, Wei L, Wang S, et al (2014). Osteogenic differentiated periodontal ligament stem cells maintain their immunomodulatory capacity. J. Tissue Eng. Regen. Med. 8: 226-232. http://dx.doi.org/10.1002/term.1516 Terrizzi AR, Fernandez-Solari J, Lee CM, Bozzini C, et al (2013). Alveolar bone loss associated to periodontal disease in lead intoxicated rats under environmental hypoxia. Arch. Oral Biol. 58: 1407-1414. http://dx.doi.org/10.1016/j.archoralbio.2013.06.010 Trubiani O, Giacoppo S, Ballerini P, Diomede F, et al (2016). Alternative source of stem cells derived from human periodontal ligament: a new treatment for experimental autoimmune encephalomyelitis. Stem Cell Res. Ther. 7: 1. http://dx.doi.org/10.1186/s13287-015-0253-4 Vandana KL, Desai R, Dalvi PJ, et al (2015). Autologous Stem Cell Application in Periodontal Regeneration Technique (SAI-PRT) Using PDLSCs Directly From an Extracted Tooth···An Insight. Int. J. Stem Cells 8: 235-237. http://dx.doi.org/10.15283/ijsc.2015.8.2.235 Wang Z, Wang W, Xu S, Wang S, et al (2016). The role of MAPK signaling pathway in the Her-2-positive meningiomas. Oncol. Rep. 36: 685-695. Wu RX, Bi CS, Yu Y, Zhang LL, et al (2015). Age-related decline in the matrix contents and functional properties of human periodontal ligament stem cell sheets. Acta Biomater. 22: 70-82. http://dx.doi.org/10.1016/j.actbio.2015.04.024 Wu Y, Yang Y, Yang P, Gu Y, et al (2013). The osteogenic differentiation of PDLSCs is mediated through MEK/ERK and p38 MAPK signalling under hypoxia. Arch. Oral Biol. 58: 1357-1368. http://dx.doi.org/10.1016/j.archoralbio.2013.03.011 Xiao X, Li Y, Zhang G, Gao Y, et al (2012). Detection of bacterial diversity in rat’s periodontitis model under imitational altitude hypoxia environment. Arch. Oral Biol. 57: 23-29. http://dx.doi.org/10.1016/j.archoralbio.2011.07.005 Xu CL, Zheng B, Pei JH, Shen SJ, et al (2016). Embelin induces apoptosis of human gastric carcinoma through inhibition of p38 MAPK and NF-κB signaling pathways. Mol. Med. Rep. 14: 307-312. Yang ZH, Zhang XJ, Dang NN, Ma ZF, et al (2009). Apical tooth germ cell-conditioned medium enhances the differentiation of periodontal ligament stem cells into cementum/periodontal ligament-like tissues. J. Periodontal Res. 44: 199-210. http://dx.doi.org/10.1111/j.1600-0765.2008.01106.x Zhang HY, Liu R, Xing YJ, Xu P, et al (2013). Effects of hypoxia on the proliferation, mineralization and ultrastructure of human periodontal ligament fibroblasts in vitro. Exp. Ther. Med. 6: 1553-1559. Zhang QB, Zhang ZQ, Fang SL, Liu YR, et al (2014). Effects of hypoxia on proliferation and osteogenic differentiation of periodontal ligament stem cells: an in vitro and in vivo study. Genet. Mol. Res. 13: 10204-10214. http://dx.doi.org/10.4238/2014.December.4.15

Bessho H, Honda S, Kondo N and Negi A (2011). The association of age-related maculopathy susceptibility 2 polymorphisms with phenotype in typical neovascular age-related macular degeneration and polypoidal choroidal vasculopathy. Mol. Vis. 17: 977-982. PMid:21541271 PMCid:3084225   Chang YC, Chang TJ, Jiang YD, Kuo SS, et al. (2007). Association study of the genetic polymorphisms of the transcription factor 7-like 2 (TCF7L2) gene and type 2 diabetes in the Chinese population. Diabetes 56: 2631-2637. http://dx.doi.org/10.2337/db07-0421 PMid:17579206   Ciardella AP, Donsoff IM, Huang SJ, Costa DL, et al. (2004). Polypoidal choroidal vasculopathy. Surv. Ophthalmol. 49: 25-37. http://dx.doi.org/10.1016/j.survophthal.2003.10.007 PMid:14711438   DeAngelis MM, Ji F, Kim IK, Adams S, et al. (2007). Cigarette smoking, CFH, APOE, ELOVL4, and risk of neovascular age-related macular degeneration. Arch. Ophthalmol. 125: 49-54. http://dx.doi.org/10.1001/archopht.125.1.49 PMid:17210851   Dewan A, Liu M, Hartman S, Zhang SS, et al. (2006). HTRA1 promoter polymorphism in wet age-related macular degeneration. Science 314: 989-992. http://dx.doi.org/10.1126/science.1133807 PMid:17053108   Fritsche LG, Loenhardt T, Janssen A, Fisher SA, et al. (2008). Age-related macular degeneration is associated with an unstable ARMS2 (LOC387715) mRNA. Nat. Genet. 40: 892-896. http://dx.doi.org/10.1038/ng.170 PMid:18511946   Fuse N, Mengkegale M, Miyazawa A, Abe T, et al. (2011). Polymorphisms in ARMS2 (LOC387715) and LOXL1 genes in the Japanese with age-related macular degeneration. Am. J. Ophthalmol. 151: 550-556. http://dx.doi.org/10.1016/j.ajo.2010.08.048 PMid:21236409   Gotoh N, Nakanishi H, Hayashi H, Yamada R, et al. (2009). ARMS2 (LOC387715) variants in Japanese patients with exudative age-related macular degeneration and polypoidal choroidal vasculopathy. Am. J. Ophthalmol. 147: 1037- 41, 1041.   Gotoh N, Yamashiro K, Nakanishi H, Saito M, et al. (2010). Haplotype analysis of the ARMS2/HTRA1 region in Japanese patients with typical neovascular age-related macular degeneration or polypoidal choroidal vasculopathy. Jpn. J. Ophthalmol. 54: 609-614. http://dx.doi.org/10.1007/s10384-010-0865-2 PMid:21191724   Hayashi H, Yamashiro K, Gotoh N, Nakanishi H, et al. (2010). CFH and ARMS2 variations in age-related macular degeneration, polypoidal choroidal vasculopathy, and retinal angiomatous proliferation. Invest. Ophthalmol. Vis. Sci. 51: 5914-5919. http://dx.doi.org/10.1167/iovs.10-5554 PMid:20574013   Imamura Y, Engelbert M, Iida T, Freund KB, et al. (2010). Polypoidal choroidal vasculopathy: a review. Surv. Ophthalmol. 55: 501-515. http://dx.doi.org/10.1016/j.survophthal.2010.03.004 PMid:20850857   Jakobsdottir J, Conley YP, Weeks DE, Mah TS, et al. (2005). Susceptibility genes for age-related maculopathy on chromosome 10q26. Am. J. Hum. Genet. 77: 389-407. http://dx.doi.org/10.1086/444437 PMid:16080115 PMCid:1226205   Kondo N, Honda S, Ishibashi K, Tsukahara Y, et al. (2007). LOC387715/HTRA1 variants in polypoidal choroidal vasculopathy and age-related macular degeneration in a Japanese population. Am. J. Ophthalmol. 144: 608-612. http://dx.doi.org/10.1016/j.ajo.2007.06.003 PMid:17692272   Laude A, Cackett PD, Vithana EN, Yeo IY, et al. (2010). Polypoidal choroidal vasculopathy and neovascular age-related macular degeneration: same or different disease? Prog. Retin. Eye Res. 29: 19-29. http://dx.doi.org/10.1016/j.preteyeres.2009.10.001 PMid:19854291   Lee KY, Vithana EN, Mathur R, Yong VH, et al. (2008). Association analysis of CFH, C2, BF, and HTRA1 gene polymorphisms in Chinese patients with polypoidal choroidal vasculopathy. Invest. Ophthalmol. Vis. Sci. 49: 2613- 2619. http://dx.doi.org/10.1167/iovs.07-0860 PMid:18515590   Leske MC, Wu SY, Hennis A, Nemesure B, et al. (2006). Nine-year incidence of age-related macular degeneration in the Barbados Eye Studies. Ophthalmology 113: 29-35. http://dx.doi.org/10.1016/j.ophtha.2005.08.012 PMid:16290049   Lima LH, Schubert C, Ferrara DC, Merriam JE, et al. (2010). Three major loci involved in age-related macular degeneration are also associated with polypoidal choroidal vasculopathy. Ophthalmology 117: 1567-1570. http://dx.doi.org/10.1016/j.ophtha.2009.12.018 PMid:20378180 PMCid:2901561   Machida S, Takahashi T, Gotoh N, Yoshimura N, et al. (2010). Monozygotic twins with polypoidal choroidal vasuculopathy. Clin. Ophthalmol. 4: 793-800. http://dx.doi.org/10.2147/OPTH.S11003 PMid:20689796 PMCid:2915866   Maruko I, Iida T, Saito M, Nagayama D, et al. (2007). Clinical characteristics of exudative age-related macular degeneration in Japanese patients. Am. J. Ophthalmol. 144: 15-22. http://dx.doi.org/10.1016/j.ajo.2007.03.047 PMid:17509509   Nakanishi H, Yamashiro K, Yamada R, Gotoh N, et al. (2010). Joint effect of cigarette smoking and CFH and LOC387715/ HTRA1 polymorphisms on polypoidal choroidal vasculopathy. Invest. Ophthalmol. Vis. Sci. 51: 6183-6187. http://dx.doi.org/10.1167/iovs.09-4948 PMid:20688737   Rivera A, Fisher SA, Fritsche LG, Keilhauer CN, et al. (2005). Hypothetical LOC387715 is a second major susceptibility gene for age-related macular degeneration, contributing independently of complement factor H to disease risk. Hum. Mol. Genet. 14: 3227-3236. http://dx.doi.org/10.1093/hmg/ddi353 PMid:16174643   Sakurada Y, Kubota T, Mabuchi F, Imasawa M, et al. (2008). Association of LOC387715 A69S with vitreous hemorrhage in polypoidal choroidal vasculopathy. Am. J. Ophthalmol. 145: 1058-1062. http://dx.doi.org/10.1016/j.ajo.2008.02.007 PMid:18400199   Sakurada Y, Kubota T, Imasawa M, Mabuchi F, et al. (2010). Association of LOC387715 A69S genotype with visual prognosis after photodynamic therapy for polypoidal choroidal vasculopathy. 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