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Among these, the p38 mitogen-activated protein kinases (MAPKs) play a central role in stress and inflammatory signaling and are classically characterized by activation by upstream kinases MKK3 and MKK6. However, accumulating biochemical, structural, and cellular evidence demonstrates that p38\u2014particularly the p38\u03b1 isoform\u2014can also be activated through atypical, MKK3\/6-independent mechanisms driven by direct protein\u2013protein interactions that promote kinase autophosphorylation. In the present review, we synthesize current insights into atypical p38 activation, comparing evidence for signaling mediated by the transforming growth factor \u03b2-activated kinase 1 binding protein 1, zeta-chain-associated protein kinase 70, the Hepatitis C virus core protein, and the Toxoplasma gondii protein GRA24. We examine the molecular and structural determinants of their interactions, the conformational changes that enable cis-autophosphorylation, and the resulting differences in regulation and substrate engagement compared with canonical p38 signaling. We further discuss evidence linking atypical p38 activation to selective pathological outcomes in models of cardiac ischemia, infection, inflammation, and vascular dysfunction. Together, these studies establish atypical p38 signaling as a distinct biochemical regulatory paradigm that expands our understanding of MAPK signaling specificity and context-dependent kinase function.<\/jats:p>","DOI":"10.1042\/bcj20250269","type":"journal-article","created":{"date-parts":[[2026,8,20]],"date-time":"2026-08-20T09:22:57Z","timestamp":1787217777000},"page":"1747-1771","update-policy":"https:\/\/doi.org\/10.1042\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Atypical p38 kinase signaling and pathological impact on human health"],"prefix":"10.1042","volume":"483","author":[{"given":"Fredejah\u00a0T.","family":"Royer","sequence":"first","affiliation":[{"name":"Department of Pharmaceutical and Biomedical Sciences, College of Pharmacy, University of Georgia, Athens 30602, U.S.A."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lillian","family":"Schulz","sequence":"additional","affiliation":[{"name":"Department of Pharmaceutical and Biomedical Sciences, College of Pharmacy, University of Georgia, Athens 30602, U.S.A."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Abby\u00a0E.","family":"Young","sequence":"additional","affiliation":[{"name":"Department of Pharmaceutical and Biomedical Sciences, College of Pharmacy, University of Georgia, Athens 30602, U.S.A."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Johnathan","family":"Burns","sequence":"additional","affiliation":[{"name":"Department of Pharmaceutical and Biomedical Sciences, College of Pharmacy, University of Georgia, Athens 30602, U.S.A."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Emmanuel","family":"Owusu","sequence":"additional","affiliation":[{"name":"Department of Pharmaceutical and Biomedical Sciences, College of Pharmacy, University of Georgia, Athens 30602, U.S.A."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yash","family":"Jadhav","sequence":"additional","affiliation":[{"name":"Department of Pharmaceutical and Biomedical Sciences, College of Pharmacy, University of Georgia, Athens 30602, U.S.A."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kelly","family":"Hanner","sequence":"additional","affiliation":[{"name":"Department of Pharmaceutical and Biomedical Sciences, College of Pharmacy, University of Georgia, Athens 30602, U.S.A."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0481-4920","authenticated-orcid":false,"given":"Neil\u00a0J.","family":"Grimsey","sequence":"additional","affiliation":[{"name":"Department of Pharmaceutical and Biomedical Sciences, College of Pharmacy, University of Georgia, Athens 30602, U.S.A."}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"288","published-online":{"date-parts":[[2026,8,20]]},"reference":[{"key":"2026082005225094700_B1","doi-asserted-by":"publisher","first-page":"4183","DOI":"10.3390\/ijms22084183","article-title":"Atypical p38 signaling, activation, and implications for disease","volume":"22","author":"Burton","year":"2021","journal-title":"Int. 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