{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,18]],"date-time":"2026-03-18T23:24:11Z","timestamp":1773876251490,"version":"3.50.1"},"reference-count":70,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,3,18]],"date-time":"2026-03-18T00:00:00Z","timestamp":1773792000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Vet. Sci."],"abstract":"<jats:p>\n                    Cashmere, a fiber of high economic value, is produced by secondary hair follicles (SHFs), whose growth depends on the proliferation and apoptosis of SHF-derived dermal papilla cells (SHF-DPCs). All-trans retinoic acid (ATRA), a metabolite of vitamin A, has shown inconsistent effects on hair follicle biology, with reports of both inhibition and stimulation. To clarify the role of ATRA in follicular development, we investigated its effects on the proliferation and apoptosis of\n                    <jats:italic>in vitro<\/jats:italic>\n                    \u2013cultured SHF-DPCs isolated from\n                    <jats:italic>Capra hircus<\/jats:italic>\n                    (Albas cashmere goats). SHF-DPCs were obtained from scapular skin, and the optimal ATRA concentration (10\n                    <jats:sup>\u22124<\/jats:sup>\n                    \u202fM) and treatment duration (24\u202fh) were identified using Cell Counting Kit-8, 5-ethynyl-2\u2032-deoxyuridine, and terminal deoxynucleotidyl transferase dUTP nick end labeling assays. Cells were then assigned to an ATRA-treated group (10\n                    <jats:sup>\u22124<\/jats:sup>\n                    \u202fM) or a control group. Proliferation, apoptosis, and cell cycle progression were evaluated, followed by transcriptome sequencing. Transcriptomic analysis indicated enrichment of the transforming growth factor-\n                    <jats:italic>\u03b2<\/jats:italic>\n                    (TGF-\n                    <jats:italic>\u03b2<\/jats:italic>\n                    ) signaling pathway. Therefore, mRNA and protein levels of TGF-\n                    <jats:italic>\u03b2<\/jats:italic>\n                    2\u2014a follicle growth\u2013related regulator\u2014and Smad2\/3 phosphorylation were examined. The TGF-\n                    <jats:italic>\u03b2<\/jats:italic>\n                    type I\/II receptor inhibitor LY2109761 was used to further validate the involvement of this pathway. ATRA inhibited SHF-DPC proliferation by inducing G1-phase cell cycle arrest and promoted apoptosis by upregulating Bax expression while downregulating Bcl-2 protein expression. Transcriptomic analysis showed that ATRA upregulated\n                    <jats:italic>TGF\u03b22<\/jats:italic>\n                    expression in SHF-DPCs. Gene Ontology analysis revealed enrichment of genes related to proliferation and apoptosis, such as \u201ccell migration\u201d and \u201ccell cycle.\u201d Kyoto Encyclopedia of Genes and Genomes pathway analysis identified the TGF-\n                    <jats:italic>\u03b2<\/jats:italic>\n                    signaling pathway as a major regulatory target. Treatment with LY2109761 partially reversed ATRA-induced reductions in proliferation, increases in apoptosis, and elevations in Smad2\/3 phosphorylation. Overall, ATRA inhibits proliferation and promotes apoptosis in SHF-DPCs, likely through the TGF-\n                    <jats:italic>\u03b2<\/jats:italic>\n                    2\/Smad2\/3 pathway. These findings provide novel insights into ATRA-mediated regulation of hair growth and offer a theoretical basis for future research on cashmere goat SHF development.\n                  <\/jats:p>","DOI":"10.3389\/fvets.2026.1765302","type":"journal-article","created":{"date-parts":[[2026,3,18]],"date-time":"2026-03-18T15:50:24Z","timestamp":1773849024000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["All-trans retinoic acid modulates proliferation and apoptosis of secondary hair follicle\u2013dermal papilla cells in cashmere goats via the TGF-\u03b22\/Smad2\/3 pathway"],"prefix":"10.3389","volume":"13","author":[{"given":"Yadong","family":"Gao","sequence":"first","affiliation":[{"name":"Key Laboratory of Reproductive Regulation and Breeding of Grassland Livestock, School of Life Sciences, Inner Mongolia University","place":["Hohhot, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Guizhen","family":"Gao","sequence":"additional","affiliation":[{"name":"Key Laboratory of Reproductive Regulation and Breeding of Grassland Livestock, School of Life Sciences, Inner Mongolia University","place":["Hohhot, China"]},{"name":"Peking University Cancer Hospital (Inner Mongolia Campus), Affiliated Cancer Hospital of Inner Mongolia Medical University","place":["Hohhot, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Xiao","family":"Zhang","sequence":"additional","affiliation":[{"name":"Key Laboratory of Reproductive Regulation and Breeding of Grassland Livestock, School of Life Sciences, Inner Mongolia University","place":["Hohhot, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Wei","family":"Lian","sequence":"additional","affiliation":[{"name":"Key Laboratory of Reproductive Regulation and Breeding of Grassland Livestock, School of Life Sciences, Inner Mongolia University","place":["Hohhot, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Xueyong","family":"Wu","sequence":"additional","affiliation":[{"name":"Key Laboratory of Reproductive Regulation and Breeding of Grassland Livestock, School of Life Sciences, Inner Mongolia University","place":["Hohhot, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Kun","family":"Cui","sequence":"additional","affiliation":[{"name":"Key Laboratory of Reproductive Regulation and Breeding of Grassland Livestock, School of Life Sciences, Inner Mongolia University","place":["Hohhot, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Lei","family":"Zhu","sequence":"additional","affiliation":[{"name":"Key Laboratory of Reproductive Regulation and Breeding of Grassland Livestock, School of Life Sciences, Inner Mongolia University","place":["Hohhot, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Guoqing","family":"Jiang","sequence":"additional","affiliation":[{"name":"Key Laboratory of Reproductive Regulation and Breeding of Grassland Livestock, School of Life Sciences, Inner Mongolia University","place":["Hohhot, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ziyang","family":"Xu","sequence":"additional","affiliation":[{"name":"Key Laboratory of Reproductive Regulation and Breeding of Grassland Livestock, School of Life Sciences, Inner Mongolia University","place":["Hohhot, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Jiawei","family":"Wang","sequence":"additional","affiliation":[{"name":"Key Laboratory of Reproductive Regulation and Breeding of Grassland Livestock, School of Life Sciences, Inner Mongolia University","place":["Hohhot, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Mingli","family":"Peng","sequence":"additional","affiliation":[{"name":"Key Laboratory of Reproductive Regulation and Breeding of Grassland Livestock, School of Life Sciences, Inner Mongolia University","place":["Hohhot, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Rui","family":"Ding","sequence":"additional","affiliation":[{"name":"Key Laboratory of Reproductive Regulation and Breeding of Grassland Livestock, School of Life Sciences, Inner Mongolia University","place":["Hohhot, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Dongjun","family":"Liu","sequence":"additional","affiliation":[{"name":"Key Laboratory of Reproductive Regulation and Breeding of Grassland Livestock, School of Life Sciences, Inner Mongolia University","place":["Hohhot, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ming","family":"Cang","sequence":"additional","affiliation":[{"name":"Key Laboratory of Reproductive Regulation and Breeding of Grassland Livestock, School of Life Sciences, Inner Mongolia University","place":["Hohhot, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Fei","family":"Hao","sequence":"additional","affiliation":[{"name":"Key Laboratory of Reproductive Regulation and Breeding of Grassland Livestock, School of Life Sciences, Inner Mongolia University","place":["Hohhot, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1965","published-online":{"date-parts":[[2026,3,18]]},"reference":[{"key":"ref1","doi-asserted-by":"publisher","first-page":"216","DOI":"10.1046\/j.0022-202x.2001.01670.x","article-title":"Molecular mechanisms regulating hair follicle development","volume":"118","author":"Millar","year":"2002","journal-title":"J Invest 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