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This is mediated by the steroidogenic acute regulatory protein (STARD1), containing a mitochondrial import sequence followed by a cholesterol\u2010binding START domain. Although mutations in this protein have been linked to lipoid congenital adrenal hyperplasia (LCAH), the mechanism of steroidogenesis regulation by STARD1 remains debatable. It has been hypothesized to involve a molten\u2010globule structural transition and interaction with 14\u20103\u20103 proteins. In this study, we aimed to address the structural basis for the 14\u20103\u20103\u2010STARD1 interaction. We show that, while the isolated START domain does not interact with 14\u20103\u20103, this interaction is enabled by STARD1 phosphorylation at Ser57, close to the mitochondrial peptide cleavage site. Biochemical analysis of the STARD1 affinity toward 14\u20103\u20103 and crystal structures of 14\u20103\u20103 complexes with Ser57 and Ser195 phosphopeptides suggest distinct roles of site\u2010specific phosphorylations in recruiting 14\u20103\u20103, to modulate STARD1 activity, processing and import to the mitochondria. Phosphorylation at Ser195 creates a unique conditional site that could only bind to 14\u20103\u20103 upon partial unfolding of the START domain. Overall, our findings on the interaction between 14\u20103\u20103 and STARD1 may have potential clinical implications for patients with LCAH.<\/jats:p>","DOI":"10.1111\/febs.15474","type":"journal-article","created":{"date-parts":[[2020,7,6]],"date-time":"2020-07-06T22:59:35Z","timestamp":1594076375000},"page":"3944-3966","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":33,"title":["Molecular basis for the recognition of steroidogenic acute regulatory protein by the 14\u20103\u20103 protein family"],"prefix":"10.1111","volume":"287","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-3933-4529","authenticated-orcid":false,"given":"Kristina V.","family":"Tugaeva","sequence":"first","affiliation":[{"name":"Federal Research Center of Biotechnology of the Russian Academy of Sciences A.N. Bach Institute of Biochemistry Moscow Russia"},{"name":"Department of Biochemistry School of Biology M.V. 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