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E2 ubiquitin-conjugating enzymes share a conserved UBC fold in which surface loops fine-tune catalysis and partner interactions, yet the roles of individual loops remain incompletely defined. Here, we identify loop 3\u2014a component of the \u201cbackside\u201d \u03b22\u2013\u03b23 hairpin\u2014as a conserved structural and allosteric element in Rad6-family E2s. Structural and bioinformatic analyses of yeast Rad6 and its human homologs (UBE2A\/UBE2B) reveal that loop 3 forms an overlapping triple \u03b2-turns, with variable first turn and a highly conserved second\/third turn that links catalytic regulation to E3 ligase engagement. Systematic mutagenesis of the yeast Rad6 backside \u03b2-turn (residues 42\u201351) shows that this element is required in vivo for Bre1-dependent histone H2B Lys123 monoubiquitination, Rad18-dependent PCNA monoubiquitination, and Ubr1\/Ubr2-dependent polyubiquitination and degradation of Sml1 and N-end rule substrates, and related biological processes. Charge-reversal mutations at backside \u03b2-turn Glu49 and Asp50 disrupt E3 binding, whereas cancer-relevant substitutions in kink-inducing prolines (Pro43\/Pro47) impair mono- and polyubiquitination without abolishing E3 interactions. Certain backside \u03b2-turn mutations, including cancer-relevant variants, compromise steady-state levels following DNA damage, revealing them as conditional null or loss-of-function alleles. Nuclear magnetic resonance spectroscopy demonstrates that Pro43\/Pro47 mutations induce long-range structural perturbations from backside \u03b2-turn into the front-face catalytic pocket, correlating with reduced in vitro ubiquitination activity. Deletion or alanine replacement of the \u03b2-turn destabilizes yeast Rad6 and human UBE2A\/UBE2B. Together, these findings establish the loop 3\/backside \u03b2-turn as a critical structural element of Rad6-family enzymes.<\/jats:p>","DOI":"10.1042\/bcj20260108","type":"journal-article","created":{"date-parts":[[2026,5,7]],"date-time":"2026-05-07T10:44:03Z","timestamp":1778150643000},"page":"1007-1027","update-policy":"https:\/\/doi.org\/10.1042\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["The backside \u03b2-turn is a key structural element of Rad6-family E2 ubiquitin-conjugating enzymes"],"prefix":"10.1042","volume":"483","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-5152-3715","authenticated-orcid":false,"given":"Prakash\u00a0K.","family":"Shukla","sequence":"first","affiliation":[{"name":"1Huntsman Cancer Institute, University of Utah School of Medicine, Salt Lake City, UT 84112, U.S.A."},{"name":"2Center for Bioseparation Technology, Vellore Institute of Technology, Vellore, Tamil Nadu 632014, India"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Andrew\u00a0M.","family":"Leng","sequence":"additional","affiliation":[{"name":"1Huntsman Cancer Institute, University of Utah School of Medicine, Salt Lake City, UT 84112, U.S.A."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Hui-Hsuan","family":"Chen","sequence":"additional","affiliation":[{"name":"1Huntsman Cancer Institute, University of Utah School of Medicine, Salt Lake City, UT 84112, U.S.A."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6900-0074","authenticated-orcid":false,"given":"Rajarshi","family":"Ganguly","sequence":"additional","affiliation":[{"name":"1Huntsman Cancer Institute, University of Utah School of Medicine, Salt Lake City, UT 84112, U.S.A."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8920-7750","authenticated-orcid":false,"given":"Nicholas","family":"Newell","sequence":"additional","affiliation":[{"name":"3Independent Scientist, Reading, MA 01867, U.S.A."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9956-8354","authenticated-orcid":false,"given":"Mahesh\u00a0B.","family":"Chandrasekharan","sequence":"additional","affiliation":[{"name":"1Huntsman Cancer Institute, University of Utah School of Medicine, Salt Lake City, UT 84112, U.S.A."},{"name":"4Department of Radiation Oncology, University of Utah School of Medicine, Salt Lake City, UT 84112, U.S.A."},{"name":"5Division of Hematology and Hematological Malignancies, Department of Internal Medicine, University of Utah School of Medicine, Salt Lake City, UT 84112, U.S.A."}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"288","published-online":{"date-parts":[[2026,5,21]]},"reference":[{"key":"2026052105113757500_B1","doi-asserted-by":"publisher","first-page":"864","DOI":"10.1073\/pnas.74.3.864","article-title":"Isopeptide linkage between nonhistone and histone 2A polypeptides of chromosomal conjugate-protein A24","volume":"74","author":"Goldknopf","year":"1977","journal-title":"Proc. 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