{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T13:33:02Z","timestamp":1791379982235,"version":"4.3.3"},"reference-count":37,"publisher":"SAGE Publications","license":[{"start":{"date-parts":[[2026,6,22]],"date-time":"2026-06-22T00:00:00Z","timestamp":1782086400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"},{"start":{"date-parts":[[2026,6,22]],"date-time":"2026-06-22T00:00:00Z","timestamp":1782086400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/journals.sagepub.com\/page\/policies\/text-and-data-mining-license"}],"funder":[{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["23K16081"],"award-info":[{"award-number":["23K16081"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["25K20340"],"award-info":[{"award-number":["25K20340"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["journals.sagepub.com"],"crossmark-restriction":true},"short-container-title":["J Dent Res"],"abstract":"<jats:p>Tooth extraction is a fundamental dental intervention for managing oral infections; however, the resulting tooth loss is associated with systemic health risks, including cognitive decline and increased mortality. Alveolar bone resorption following tooth extraction significantly alters jaw morphology, complicating subsequent functional rehabilitation. Despite its clinical significance, the cellular and molecular mechanisms governing bone regeneration within extraction sockets remain poorly understood. Here, we identified group 2 innate lymphoid cells (ILC2s) as pivotal regulators of alveolar bone regeneration following tooth extraction. Using single-cell RNA sequencing and flow cytometry, we demonstrated that ILC2s accumulate and persist within extraction sockets beyond the acute inflammatory phase. Genetic lymphocyte ablation revealed that loss of all ILCs severely impairs bone formation, highlighting their essential role in socket regeneration. Following tooth extraction, ILC2s exhibited increased interleukin (IL)\u201313 production and promoted bone formation via IL-13\u2013mediated activation of bone marrow\u2013derived mesenchymal stromal cells. Single-cell RNA sequencing and histological analyses suggested that gingival fibroblasts are a major source of IL-33 in extraction sockets during mucosal healing. IL-33 activated ILC2s, increasing IL-13 production and promoting osteogenesis, whereas IL-33 neutralization suppressed ILC2 activation and impaired socket bone regeneration. Collectively, these findings suggest that the gingival fibroblast\u2013IL-33\u2013ILC2\u2013IL-13 axis may contribute to the regulation of alveolar bone regeneration following tooth extraction. This study provides a biological basis for developing novel therapeutic strategies to optimize postextraction bone preservation.<\/jats:p>","DOI":"10.1177\/00220345261449746","type":"journal-article","created":{"date-parts":[[2026,6,22]],"date-time":"2026-06-22T17:21:17Z","timestamp":1782148877000},"update-policy":"https:\/\/doi.org\/10.1177\/sage-journals-update-policy","source":"Crossref","is-referenced-by-count":1,"title":["Gingival Fibroblast-Driven Osteoimmunology via the IL-33\u2013ILC2\u2013IL-13 Axis"],"prefix":"10.1177","author":[{"given":"Y.","family":"Sato","sequence":"first","affiliation":[{"name":"Division of Molecular &amp; Regenerative Prosthodontics, Tohoku University Graduate School of Dentistry, Sendai, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0002-1560-1741","authenticated-orcid":false,"given":"T.","family":"Kondo","sequence":"additional","affiliation":[{"name":"Division of Molecular &amp; Regenerative Prosthodontics, Tohoku University Graduate School of Dentistry, Sendai, Japan"},{"name":"Department of Next-Generation Dental Material Engineering, Tohoku University Graduate School of Dentistry, Sendai, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"K.","family":"Otake","sequence":"additional","affiliation":[{"name":"Department of Next-Generation Dental Material Engineering, Tohoku University Graduate School of Dentistry, Sendai, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"S.","family":"Ambo","sequence":"additional","affiliation":[{"name":"Division of Molecular &amp; Regenerative Prosthodontics, Tohoku University Graduate School of Dentistry, Sendai, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2759-055X","authenticated-orcid":false,"given":"A.","family":"Ashry","sequence":"additional","affiliation":[{"name":"Division of Molecular &amp; Regenerative Prosthodontics, Tohoku University Graduate School of Dentistry, Sendai, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"K.","family":"Engkatanachai","sequence":"additional","affiliation":[{"name":"Division of Molecular &amp; Regenerative Prosthodontics, Tohoku University Graduate School of Dentistry, Sendai, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0200-8132","authenticated-orcid":false,"given":"H.","family":"Egusa","sequence":"additional","affiliation":[{"name":"Division of Molecular &amp; Regenerative Prosthodontics, Tohoku University Graduate School of Dentistry, Sendai, Japan"},{"name":"Department of Next-Generation Dental Material Engineering, Tohoku University Graduate School of Dentistry, Sendai, Japan"},{"name":"Center of Excellence for Dental Stem Cell Biology, Faculty of Dentistry, Chulalongkorn University, Bangkok, Thailand"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"179","published-online":{"date-parts":[[2026,6,22]]},"reference":[{"key":"e_1_3_3_2_1","doi-asserted-by":"crossref","unstructured":"Akashi Y et al. 2023. Cyclic pressure-induced cytokines from gingival fibroblasts stimulate osteoclast activity: clinical implications for alveolar bone loss in denture wearers. J Prosthodont Res. 67(1):77\u201386. https:\/\/doi.org\/10.2186\/jpr.jpr_d_21_00238","DOI":"10.2186\/jpr.JPR_D_21_00238"},{"key":"e_1_3_3_3_1","doi-asserted-by":"crossref","unstructured":"Alam A et al. 2022. Fungal mycobiome drives IL-33 secretion and type 2 immunity in pancreatic cancer. Cancer Cell. 40(2):153\u2013167.e11. https:\/\/doi.org\/10.1016\/j.ccell.2022.01.003","DOI":"10.1016\/j.ccell.2022.01.003"},{"key":"e_1_3_3_4_1","doi-asserted-by":"crossref","unstructured":"Amisaki M et al. 2025. IL-33-activated ILC2s induce tertiary lymphoid structures in pancreatic cancer. Nature. 638(8052):1076\u20131084. https:\/\/doi.org\/10.1038\/s41586-024-08426-5","DOI":"10.1038\/s41586-024-08426-5"},{"key":"e_1_3_3_5_1","doi-asserted-by":"crossref","unstructured":"Ando Y et al. 2024. The neutrophil-osteogenic cell axis promotes bone destruction in periodontitis. Int J Oral Sci. 16(1):18. https:\/\/doi.org\/10.1038\/s41368-023-00275-8","DOI":"10.1038\/s41368-023-00275-8"},{"key":"e_1_3_3_6_1","doi-asserted-by":"crossref","unstructured":"Barlow JL et al. 2013. IL-33 is more potent than IL-25 in provoking IL-13-producing nuocytes (type 2 innate lymphoid cells) and airway contraction. J Allergy Clin Immunol. 132(4):933\u2013941. https:\/\/doi.org\/10.1016\/j.jaci.2013.05.012","DOI":"10.1016\/j.jaci.2013.05.012"},{"key":"e_1_3_3_7_1","doi-asserted-by":"crossref","unstructured":"Brestoff JR et al. 2015. Group 2 innate lymphoid cells promote beiging of white adipose tissue and limit obesity. Nature. 519(7542):242\u2013246. https:\/\/doi.org\/10.1038\/nature14115","DOI":"10.1038\/nature14115"},{"key":"e_1_3_3_8_1","doi-asserted-by":"crossref","unstructured":"Ding X et al. 2016. IL-33-driven ILC2\/eosinophil axis in fat is induced by sympathetic tone and suppressed by obesity. J Endocrinol. 231(1):35\u201348. https:\/\/doi.org\/10.1530\/joe-16-0229","DOI":"10.1530\/JOE-16-0229"},{"key":"e_1_3_3_9_1","doi-asserted-by":"crossref","unstructured":"Griesenauer B Paczesny S. 2017. The ST2\/IL-33 axis in immune cells during inflammatory diseases. Front Immunol. 8:475. https:\/\/doi.org\/10.3389\/fimmu.2017.00475","DOI":"10.3389\/fimmu.2017.00475"},{"key":"e_1_3_3_10_1","doi-asserted-by":"crossref","unstructured":"Kamal A Kamal M Hasnat A Mohsin A Samsudin AR. 2025. Healing of tooth extraction wound socket do we have current understanding? J Oral Maxillofac Surg Med Pathol. 37(1):8\u201322. https:\/\/doi.org\/10.1016\/j.Ajoms.2024.07.009","DOI":"10.1016\/j.ajoms.2024.07.009"},{"key":"e_1_3_3_11_1","doi-asserted-by":"crossref","unstructured":"Koka S Gupta A. 2018. Association between missing tooth count and mortality: a systematic review. J Prosthodont Res. 62(2):134\u2013151. https:\/\/doi.org\/10.1016\/j.jpor.2017.08.003","DOI":"10.1016\/j.jpor.2017.08.003"},{"key":"e_1_3_3_12_1","doi-asserted-by":"crossref","unstructured":"Kondo T Bhattarai S Hatakeyama M Egusa H. 2026. CAD\/CAM-fabricated zirconia removable dental prosthesis with telescopic crowns: a case report. Int J Prosthodont. 39(1):89\u201393. https:\/\/doi.org\/10.11607\/ijp.9262","DOI":"10.11607\/ijp.9262"},{"key":"e_1_3_3_13_1","doi-asserted-by":"crossref","unstructured":"Kondo T Gleason A Okawa H Hokugo A Nishimura I. 2023. Mouse gingival single-cell transcriptomic atlas identified a novel fibroblast subpopulation activated to guide oral barrier immunity in periodontitis. ELife. 12:RP88183. https:\/\/doi.org\/10.7554\/elife.88183","DOI":"10.7554\/eLife.88183.3"},{"key":"e_1_3_3_14_1","doi-asserted-by":"crossref","unstructured":"Kondo T Kanayama K Egusa H Nishimura I. 2023. Current perspectives of residual ridge resorption: pathological activation of oral barrier osteoclasts. J Prosthodont Res. 67(1):12\u201322. https:\/\/doi.org\/10.2186\/jpr.jpr_d_21_00333","DOI":"10.2186\/jpr.JPR_D_21_00333"},{"key":"e_1_3_3_15_1","doi-asserted-by":"crossref","unstructured":"Kondo T et al. 2024a. Zinc- and fluoride-releasing bioactive glass as a novel bone substitute. J Dent Res. 103(5):526\u2013535. https:\/\/doi.org\/10.1177\/00220345241231772","DOI":"10.1177\/00220345241231772"},{"key":"e_1_3_3_16_1","doi-asserted-by":"crossref","unstructured":"Kondo T Yamada M Egusa H. 2024b. Innate immune regulation in dental implant osseointegration. J Prosthodont Res. 68(4):511\u2013521. https:\/\/doi.org\/10.2186\/jpr.jpr_d_23_00198","DOI":"10.2186\/jpr.JPR_D_23_00198"},{"key":"e_1_3_3_17_1","doi-asserted-by":"crossref","unstructured":"Lerner UH. 2006. Inflammation-induced bone remodeling in periodontal disease and the influence of post-menopausal osteoporosis. J Dent Res. 85(7):596\u2013607. https:\/\/doi.org\/10.1177\/154405910608500704","DOI":"10.1177\/154405910608500704"},{"key":"e_1_3_3_18_1","doi-asserted-by":"crossref","unstructured":"Lin W et al. 2024. Early infiltrating NKT lymphocytes attenuate bone regeneration through secretion of CXCL2. Sci Adv. 10(20):eadl6343. https:\/\/doi.org\/10.1126\/sciadv.adl6343","DOI":"10.1126\/sciadv.adl6343"},{"key":"e_1_3_3_19_1","doi-asserted-by":"crossref","unstructured":"Liu A et al. 2024. The IL -33\/ST2 axis is protective against acute inflammation during the course of periodontitis. Nat Commun. 15(1):2707. https:\/\/doi.org\/10.1038\/s41467-024-46746-2","DOI":"10.1038\/s41467-024-46746-2"},{"key":"e_1_3_3_20_1","doi-asserted-by":"crossref","unstructured":"Ma X et al. 2025. Association between denture restoration for tooth loss and cognitive impairment: a systematic review and meta-analysis. J Prosthodont Res. 69(3):313\u2013320. https:\/\/doi.org\/10.2186\/jpr.jpr_d_24_00060","DOI":"10.2186\/jpr.JPR_D_24_00060"},{"key":"e_1_3_3_21_1","doi-asserted-by":"crossref","unstructured":"Momiuchi Y et al. 2021. Group 2 innate lymphoid cells in bone marrow regulate osteoclastogenesis in a reciprocal manner via RANKL GM-CSF and IL-13. Int Immunol. 33(11):573\u2013585. https:\/\/doi.org\/10.1093\/intimm\/dxab062","DOI":"10.1093\/intimm\/dxab062"},{"key":"e_1_3_3_22_1","doi-asserted-by":"crossref","unstructured":"Monticelli LA et al. 2011. Innate lymphoid cells promote lung-tissue homeostasis after infection with influenza virus. Nat Immunol. 12(11):1045\u20131054. https:\/\/doi.org\/10.1031\/ni.2131","DOI":"10.1038\/ni.2131"},{"key":"e_1_3_3_23_1","doi-asserted-by":"crossref","unstructured":"Moro K et al. 2010. Innate production of T(H)2 cytokines by adipose tissue-associated c-Kit(+)Sca-1(+) lymphoid cells. Nature. 463(7280):540\u2013544. https:\/\/doi.org\/10.1038\/nature08636","DOI":"10.1038\/nature08636"},{"key":"e_1_3_3_24_1","doi-asserted-by":"crossref","unstructured":"Neill DR et al. 2010. Nuocytes represent a new innate effector leukocyte that mediates type-2 immunity. Nature. 464(7293):1367\u20131370. https:\/\/doi.org\/10.1038\/nature08900","DOI":"10.1038\/nature08900"},{"key":"e_1_3_3_25_1","doi-asserted-by":"crossref","unstructured":"Ohtake T Kondo T Bhattarai S Harata T Egusa H. 2025. A technique for denture fabrication using 3D-printed duplicated removable partial dentures. Int J Prosthodont. [published online ahead of print]. https:\/\/doi.org\/10.11607\/ijp.9423","DOI":"10.11607\/ijp.9423"},{"key":"e_1_3_3_26_1","doi-asserted-by":"crossref","unstructured":"Okawa H et al. 2022. Mechanism of bisphosphonate-related osteonecrosis of the jaw (BRONJ) revealed by targeted removal of legacy bisphosphonate from jawbone using competing inert hydroxymethylene diphosphonate. ELife. 11:e76207. https:\/\/doi.org\/10.7554\/elife.76207","DOI":"10.7554\/eLife.76207"},{"key":"e_1_3_3_27_1","doi-asserted-by":"crossref","unstructured":"Omata Y et al. 2018. Group 2 innate lymphoid cells attenuate inflammatory arthritis and protect from bone destruction in mice. Cell Rep. 24(1):169\u2013180. https:\/\/doi.org\/10.1016\/j.celrep.2018.06.005","DOI":"10.1016\/j.celrep.2018.06.005"},{"key":"e_1_3_3_28_1","doi-asserted-by":"crossref","unstructured":"Omata Y et al. 2020. Type 2 innate lymphoid cells inhibit the differentiation of osteoclasts and protect from ovariectomy-induced bone loss. Bone. 136:115335. https:\/\/doi.org\/10.1016\/j.bone.2020.115335","DOI":"10.1016\/j.bone.2020.115335"},{"key":"e_1_3_3_29_1","doi-asserted-by":"crossref","unstructured":"Omata Y et al. 2022. Inflammatory arthritis and bone metabolism regulated by type 2 innate and adaptive immunity. Int J Mol Sci. 23(3):1104. https:\/\/doi.org\/10.3390\/ijms23031104","DOI":"10.3390\/ijms23031104"},{"key":"e_1_3_3_30_1","doi-asserted-by":"crossref","unstructured":"Otake T et al. 2026. Zinc- and fluoride-containing bioactive glass enhances angiogenesis-mediated bone regeneration via M2d macrophage activation. Sci Rep. 16(1):11351. https:\/\/doi.org\/10.1038\/s41598-026-44931-5","DOI":"10.1038\/s41598-026-44931-5"},{"key":"e_1_3_3_31_1","doi-asserted-by":"crossref","unstructured":"Shah MS et al. 2025. Unveiling the dental cellular landscape: a comprehensive review of single-cell RNA sequencing in human dental tissues. Jpn Dent Sci Rev. 61:301\u2013316. https:\/\/doi.org\/10.1016\/j.jdsr.2025.11.004","DOI":"10.1016\/j.jdsr.2025.11.004"},{"key":"e_1_3_3_32_1","doi-asserted-by":"crossref","unstructured":"Tsukasaki M et al. 2018. Host defense against oral microbiota by bone-damaging T cells. Nat Commun. 9(1):701. https:\/\/doi.org\/10.1038\/s41467-018-03147-6","DOI":"10.1038\/s41467-018-03147-6"},{"key":"e_1_3_3_33_1","doi-asserted-by":"crossref","unstructured":"Vivier E et al. 2018. Innate lymphoid cells: 10 years on. Cell. 174(5):1054\u20131066. https:\/\/doi.org\/10.1016\/j.cell.2018.07.017","DOI":"10.1016\/j.cell.2018.07.017"},{"key":"e_1_3_3_34_1","doi-asserted-by":"crossref","unstructured":"Walker JA Barlow JL McKenzie AN. 2013. Innate lymphoid cells\u2014how did we miss them? Nat Rev Immunol. 13(2):75\u201387. https:\/\/doi.org\/10.1038\/nri3349","DOI":"10.1038\/nri3349"},{"key":"e_1_3_3_35_1","doi-asserted-by":"crossref","unstructured":"Wang M Gao M Yi Z. 2023. Biological effects of IL-33\/ST2 axis on oral diseases: autoimmune diseases and periodontal diseases. Int Immunopharmacol. 122:110524. https:\/\/doi.org\/10.1016\/j.intimp.2023.110524","DOI":"10.1016\/j.intimp.2023.110524"},{"key":"e_1_3_3_36_1","doi-asserted-by":"crossref","unstructured":"Wang X et al. 2025. Macrophage-derived amphiregulin promoted the osteogenic differentiation of chondrocytes through EGFR\/Yap axis and TGF-\u03b2 activation. Bone. 190:117275. https:\/\/doi.org\/10.1016\/j.bone.2024.117275","DOI":"10.1016\/j.bone.2024.117275"},{"key":"e_1_3_3_37_1","doi-asserted-by":"crossref","unstructured":"Yin G Zhao C Pei W. 2022. Crosstalk between macrophages and innate lymphoid cells (ILCs) in diseases. Int Immunopharmacol. 110:108937. https:\/\/doi.org\/10.1016\/j.intimp.2022.108937","DOI":"10.1016\/j.intimp.2022.108937"},{"key":"e_1_3_3_38_1","doi-asserted-by":"crossref","unstructured":"Zhang Y et al. 2023. Group 2 innate lymphoid cells protect mice from abdominal aortic aneurysm formation via IL5 and eosinophils. Adv Sci (Weinh). 10(7):e2206958. https:\/\/doi.org\/10.1002\/advs.202206958","DOI":"10.1002\/advs.202206958"}],"container-title":["Journal of Dental Research"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/journals.sagepub.com\/doi\/pdf\/10.1177\/00220345261449746","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/journals.sagepub.com\/doi\/full-xml\/10.1177\/00220345261449746","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/journals.sagepub.com\/doi\/pdf\/10.1177\/00220345261449746","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,26]],"date-time":"2026-06-26T13:17:10Z","timestamp":1782479830000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.sagepub.com\/doi\/10.1177\/00220345261449746"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,6,22]]},"references-count":37,"alternative-id":["10.1177\/00220345261449746"],"URL":"https:\/\/doi.org\/10.1177\/00220345261449746","relation":{},"ISSN":["0022-0345","1544-0591"],"issn-type":[{"value":"0022-0345","type":"print"},{"value":"1544-0591","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,6,22]]},"article-number":"00220345261449746"}}