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. 2019 Mar 25;9:5043. doi: 10.1038/s41598-019-41639-7

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© The Author(s) 2019

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/.

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Hox status imparts unique regenerative response on periosteal cells. (A–H) Representative histologic appearance of the periosteum of frontal, parietal, hyoid bone and tibia with characteristic one-cell-layer thick cambial layer (cl) containing osx-positive bone-lining cells (green cells in immunofluorescence staining)(E–H), and a thicker fibrous layer (fl) in the periphery (n = 5). (I,J) A periosteal scratch injury results in a pure osteogenic response (between dashed line) of the periosteum in the frontal and parietal bone, while (K,L) periostea from the hyoid and tibia responded with a mixed osteochondrogenic response (osx for osteogenesis, sox9 for chondrogenesis)(n = 5). (M,N) Immunofluorescence staining of the periosteal injury confirmed the osteogenic response with Osx-positive cells within the regenerate (between white dashed lines) and an absence of Sox9-expressing cells. (O,P) In contrast, the periosteal response to injury (between white dashed lines) of the hyoid and tibia exhibited both Osx+ and Sox9+ expressing cells, in line with a mixed osteochondrogenic healing response. Abbreviations: c, cortical bone; cl, cambial layer; fl, fibrous layer; p, periosteum.