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Candidate genes, identified through whole-genome resequencing, are consistent with possible roles in chloroplast proteostasis.<\/jats:bold>\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Abstract<\/jats:title>\n                    <jats:p>\n                      Variegation mutants provide valuable insights into chloroplast biogenesis. We characterized a newly identified variegated barley mutant, in which the phenotype is controlled by two epistatic recessive genes. The causal loci,\n                      <jats:italic>var4<\/jats:italic>\n                      and\n                      <jats:italic>var5<\/jats:italic>\n                      , were mapped on chromosomes 2H and 3H. We used genotyping-by-sequencing, linkage mapping, and PACE genotyping, followed by whole-genome resequencing to identify candidate genes. Our two top candidate genes are an\n                      <jats:italic>NBR1-like selective autophagy receptor<\/jats:italic>\n                      gene and a\n                      <jats:italic>DNAJ-domain-containing<\/jats:italic>\n                      gene, respectively. Based on their homology-based functional annotation, both candidates could be implicated in chloroplast proteostasis, regulating protein import, folding, and\/or degradation. While functional confirmation is still required, mild, independent defects in proteostasis from each mutation may act synergistically to surpass a functional threshold, impairing chloroplast development in early leaves while allowing partial recovery in later stages. These findings reveal a novel digenic basis for variegation in barley and identify candidate genes and pathways linked to chloroplast proteostasis.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1007\/s00122-026-05370-y","type":"journal-article","created":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T04:43:42Z","timestamp":1792039422000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Leaf variegation in a barley EMS mutant is controlled by two epistatic mutations"],"prefix":"10.1007","volume":"139","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-9802-1787","authenticated-orcid":false,"given":"H\u00e9l\u00e8ne","family":"Pidon","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shobhashree Rajulareddy","family":"Nagireddy","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5213-4030","authenticated-orcid":false,"given":"Michael","family":"Melzer","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7338-0946","authenticated-orcid":false,"given":"Axel","family":"Himmelbach","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3011-8731","authenticated-orcid":false,"given":"Nils","family":"Stein","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,10,14]]},"reference":[{"issue":"2","key":"5370_CR1","doi-asserted-by":"publisher","first-page":"174","DOI":"10.1038\/nbt.2095","volume":"30","author":"A Abe","year":"2012","unstructured":"Abe A, Kosugi S, Yoshida K, Natsume S, Takagi H, Kanzaki H, Matsumura H, Yoshida K, Mitsuoka C, Tamiru M, Innan H, Cano L, Kamoun S, Terauchi R (2012) Genome sequencing reveals agronomically important loci in rice using MutMap. 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