{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,15]],"date-time":"2026-06-15T07:54:22Z","timestamp":1781510062055,"version":"3.54.1"},"reference-count":41,"publisher":"Oxford University Press (OUP)","issue":"2","license":[{"start":{"date-parts":[[2025,9,17]],"date-time":"2025-09-17T00:00:00Z","timestamp":1758067200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/pages\/standard-publication-reuse-rights"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32201817"],"award-info":[{"award-number":["32201817"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004731","name":"Zhejiang Provincial Natural Science Foundation","doi-asserted-by":"publisher","award":["LY24C130002"],"award-info":[{"award-number":["LY24C130002"]}],"id":[{"id":"10.13039\/501100004731","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,1,12]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>The 26 kDa \u03b1-globulin, a minor component of rice seed storage proteins (SSPs), is encoded by the single-copy gene \u03b1-globulin 1 (Glb1), which is specifically expressed in the endosperm. However, the specific effects of altered \u03b1-globulin content on SSP composition and grain quality remains poorly understood. In this study, we demonstrated that modulation of \u03b1-globulin levels significantly enhances both total protein content and accumulation of the essential amino acid lysine in Glb1 knockout mutants and overexpression lines. Notably, specific alterations in SSP composition were observed exclusively in Glb1 mutants, with no such changes detected in overexpression lines. Glb1 knockout mutants produced abnormal protein bodies, including small-sized protein body I and center-cracked protein body II. Furthermore, increased proglutelin accumulation in Glb1 mutants triggered endoplasmic reticulum stress, resulting in a higher chalky grain rate. However, the altered \u03b1-globulin content had minimal effects on grain weight and starch properties compared with the wild type. This study elucidates the role of \u03b1-globulin accumulation in SSP reprogramming and provides a potential strategy for enhancing rice endosperm lysine content through the regulation of Glb1 biosynthesis.<\/jats:p>","DOI":"10.1093\/jxb\/eraf410","type":"journal-article","created":{"date-parts":[[2025,9,19]],"date-time":"2025-09-19T22:46:08Z","timestamp":1758321968000},"page":"379-391","source":"Crossref","is-referenced-by-count":4,"title":["The role of \u03b1-globulin accumulation in seed storage protein reprogramming and grain quality in rice"],"prefix":"10.1093","volume":"77","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3380-7287","authenticated-orcid":false,"given":"Feifei","family":"Xu","sequence":"first","affiliation":[{"name":"Institute of Nuclear Agricultural Sciences, College of Agriculture and Biotechnology, Zhejiang University , Zijingang Campus, Hangzhou 310058 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yanni","family":"Zhang","sequence":"additional","affiliation":[{"name":"Institute of Nuclear Agricultural Sciences, College of Agriculture and Biotechnology, Zhejiang University , Zijingang Campus, Hangzhou 310058 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xinyu","family":"Li","sequence":"additional","affiliation":[{"name":"Institute of Nuclear Agricultural Sciences, College of Agriculture and Biotechnology, Zhejiang University , Zijingang Campus, Hangzhou 310058 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jianming","family":"Pan","sequence":"additional","affiliation":[{"name":"Institute of Nuclear Agricultural Sciences, College of Agriculture and Biotechnology, Zhejiang University , Zijingang Campus, Hangzhou 310058 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Meng","family":"Li","sequence":"additional","affiliation":[{"name":"Institute of Nuclear Agricultural Sciences, College of Agriculture and Biotechnology, Zhejiang University , Zijingang Campus, Hangzhou 310058 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jing","family":"Yu","sequence":"additional","affiliation":[{"name":"Institute of Nuclear Agricultural Sciences, College of Agriculture and Biotechnology, Zhejiang University , Zijingang Campus, Hangzhou 310058 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lin","family":"Zhang","sequence":"additional","affiliation":[{"name":"Institute of Nuclear Agricultural Sciences, College of Agriculture and Biotechnology, Zhejiang University , Zijingang Campus, Hangzhou 310058 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yong-Jin","family":"Park","sequence":"additional","affiliation":[{"name":"Department of Plant Resources, College of Industrial Sciences, Kongju National University , Yesan 32439 ,","place":["Republic of Korea"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9606-6615","authenticated-orcid":false,"given":"Jinsong","family":"Bao","sequence":"additional","affiliation":[{"name":"Institute of Nuclear Agricultural Sciences, College of Agriculture and Biotechnology, Zhejiang University , Zijingang Campus, Hangzhou 310058 ,","place":["China"]},{"name":"Hainan Institute, Zhejiang University, Yazhou Bay Science and Technology City , Yazhou District, Sanya 572025 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2025,9,17]]},"reference":[{"key":"2026011209055325900_eraf410-B1","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1186\/s42269-019-0066-1","article-title":"Extraction of high-quality genomic DNA from different plant orders applying a modified CTAB-based method","volume":"43","author":"Aboul-Maaty","year":"2019","journal-title":"Bulletin of the National Research Centre"},{"key":"2026011209055325900_eraf410-B2","doi-asserted-by":"crossref","first-page":"100463","DOI":"10.1016\/j.xplc.2022.100463","article-title":"OPAQUE3, encoding a transmembrane bZIP transcription factor, regulates endosperm storage protein and starch biosynthesis in rice","volume":"3","author":"Cao","year":"2022","journal-title":"Plant Communications"},{"key":"2026011209055325900_eraf410-B3","doi-asserted-by":"crossref","first-page":"111449","DOI":"10.1016\/j.plantsci.2022.111449","article-title":"Efficient breeding of low glutelin content rice germplasm by simultaneous editing multiple glutelin genes via CRISPR\/Cas9","volume":"324","author":"Chen","year":"2022","journal-title":"Plant Science"},{"key":"2026011209055325900_eraf410-B4","doi-asserted-by":"crossref","first-page":"765","DOI":"10.1038\/35037633","article-title":"Messenger RNA targeting of rice seed storage proteins to specific ER subdomains","volume":"407","author":"Choi","year":"2000","journal-title":"Nature"},{"key":"2026011209055325900_eraf410-B5","doi-asserted-by":"crossref","first-page":"4469","DOI":"10.1093\/jxb\/erv212","article-title":"Amylopectin biosynthetic enzymes from developing rice seed form enzymatically active protein complexes","volume":"66","author":"Crofts","year":"2015","journal-title":"Journal of Experimental Botany"},{"key":"2026011209055325900_eraf410-B6","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1093\/jxb\/err262","article-title":"The failure to express a protein disulphide isomerase-like protein results in a floury endosperm and an endoplasmic reticulum stress response in rice","volume":"63","author":"Han","year":"2012","journal-title":"Journal of Experimental Botany"},{"key":"2026011209055325900_eraf410-B7","doi-asserted-by":"crossref","first-page":"946","DOI":"10.1111\/j.1365-313X.2011.04844.x","article-title":"Signal transduction by IRE1-mediated splicing of bZIP50 and other stress sensors in the endoplasmic reticulum stress response of rice","volume":"69","author":"Hayashi","year":"2012","journal-title":"The Plant Journal"},{"key":"2026011209055325900_eraf410-B8","doi-asserted-by":"crossref","first-page":"1999","DOI":"10.1111\/jipb.13176","article-title":"Rice seed storage proteins: biosynthetic pathways and the effects of environmental factors","volume":"63","author":"He","year":"2021","journal-title":"Journal of Integrative Plant Biology"},{"key":"2026011209055325900_eraf410-B9","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1046\/j.1365-313X.1994.6020271.x","article-title":"Efficient transformation of rice (Oryza sativa L.) mediated by Agrobacterium and sequence analysis of the boundaries of the T-DNA","volume":"6","author":"Hiei","year":"1994","journal-title":"The Plant Journal"},{"key":"2026011209055325900_eraf410-B10","doi-asserted-by":"crossref","first-page":"4233","DOI":"10.1093\/jxb\/ern265","article-title":"Characterization of a new rice glutelin gene GluD-1 expressed in the starchy endosperm","volume":"59","author":"Kawakatsu","year":"2008","journal-title":"Journal of Experimental Botany"},{"key":"2026011209055325900_eraf410-B11","doi-asserted-by":"crossref","first-page":"1842","DOI":"10.1104\/pp.110.164343","article-title":"Reducing rice seed storage protein accumulation leads to changes in nutrient quality and storage organelle formation","volume":"154","author":"Kawakatsu","year":"2010","journal-title":"Plant Physiology"},{"key":"2026011209055325900_eraf410-B12","first-page":"109","volume-title":"Rice","author":"Kawakatsu","year":"2019","edition":"4th ed"},{"key":"2026011209055325900_eraf410-B13","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1007\/BF00392095","article-title":"Immunochemical studies on the role of the Golgi complex in protein-body formation in rice seeds","volume":"169","author":"Krishnan","year":"1986","journal-title":"Planta"},{"key":"2026011209055325900_eraf410-B14","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/0168-9452(92)90018-H","article-title":"Characterization and localization of rice (Oryza sativa L.) seed globulins","volume":"81","author":"Krishnan","year":"1992","journal-title":"Plant Science"},{"key":"2026011209055325900_eraf410-B15","doi-asserted-by":"crossref","first-page":"1054","DOI":"10.1126\/science.8235623","article-title":"Rice prolamine protein body biogenesis: a BiP-mediated process","volume":"262","author":"Li","year":"1993","journal-title":"Science"},{"key":"2026011209055325900_eraf410-B16","doi-asserted-by":"crossref","first-page":"1918","DOI":"10.1093\/mp\/sst081","article-title":"OsVPS9A functions cooperatively with OsRAB5A to regulate post-Golgi dense vesicle-mediated storage protein trafficking to the protein storage vacuole in rice endosperm cells","volume":"6","author":"Liu","year":"2013","journal-title":"Molecular Plant"},{"key":"2026011209055325900_eraf410-B17","doi-asserted-by":"crossref","first-page":"101287","DOI":"10.1016\/j.xplc.2025.101287","article-title":"Integrative transcriptogenomic analyses reveal the regulatory network underlying rice eating and cooking quality and identify a role for alpha-globulin in modulating starch and sucrose metabolism","volume":"6","author":"Nawade","year":"2025","journal-title":"Plant Communications"},{"key":"2026011209055325900_eraf410-B18","doi-asserted-by":"crossref","first-page":"14156","DOI":"10.1073\/pnas.0904429106","article-title":"ER membrane-localized oxidoreductase Ero1 is required for disulfide bond formation in the rice endosperm","volume":"106","author":"Onda","year":"2009","journal-title":"Proceedings of the National Academy of Sciences, USA"},{"key":"2026011209055325900_eraf410-B19","doi-asserted-by":"crossref","first-page":"2174","DOI":"10.1093\/plphys\/kiab175","article-title":"Post-Golgi trafficking of rice storage proteins requires the small GTPase Rab7 activation complex MON1\u2013CCZ1","volume":"187","author":"Pan","year":"2021","journal-title":"Plant Physiology"},{"key":"2026011209055325900_eraf410-B20","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1104\/pp.18.00375","article-title":"OsDER1 is an ER-associated protein degradation factor that responds to ER stress","volume":"178","author":"Qian","year":"2018","journal-title":"Plant Physiology"},{"key":"2026011209055325900_eraf410-B21","doi-asserted-by":"crossref","first-page":"410","DOI":"10.1105\/tpc.113.121376","article-title":"GLUTELIN PRECURSOR ACCUMULATION3 encodes a regulator of post-Golgi vesicular traffic essential for vacuolar protein sorting in rice endosperm","volume":"26","author":"Ren","year":"2014","journal-title":"The Plant Cell"},{"key":"2026011209055325900_eraf410-B22","doi-asserted-by":"crossref","first-page":"758","DOI":"10.1105\/tpc.19.00863","article-title":"GPA5 encodes a Rab5a effector required for post-Golgi trafficking of rice storage proteins","volume":"32","author":"Ren","year":"2020","journal-title":"The Plant Cell"},{"key":"2026011209055325900_eraf410-B23","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1007\/978-94-011-4431-5_2","volume-title":"Seed proteins","author":"Shewry","year":"1999"},{"key":"2026011209055325900_eraf410-B24","doi-asserted-by":"crossref","first-page":"947","DOI":"10.1093\/jexbot\/53.370.947","article-title":"Cereal seed storage proteins: structures, properties and role in grain utilization","volume":"53","author":"Shewry","year":"2002","journal-title":"Journal of Experimental Botany"},{"key":"2026011209055325900_eraf410-B25","doi-asserted-by":"crossref","first-page":"401","DOI":"10.1007\/978-94-011-4431-5_17","volume-title":"Seed proteins","author":"Takaiwa","year":"1999"},{"key":"2026011209055325900_eraf410-B26","doi-asserted-by":"crossref","first-page":"568","DOI":"10.1270\/jsbbs.60.568","article-title":"Gene-gene interactions between mutants that accumulate abnormally high amounts of proglutelin in rice seed","volume":"60","author":"Ueda","year":"2010","journal-title":"Breeding Science"},{"key":"2026011209055325900_eraf410-B27","doi-asserted-by":"crossref","first-page":"675","DOI":"10.1111\/j.1365-313X.2010.04453.x","article-title":"Expression of ER quality control-related genes in response to changes in BiP1 levels in developing rice endosperm","volume":"65","author":"Wakasa","year":"2011","journal-title":"Plant Journal"},{"key":"2026011209055325900_eraf410-B28","doi-asserted-by":"crossref","first-page":"812","DOI":"10.1111\/j.1365-313X.2010.04370.x","article-title":"OsRab5a regulates endomembrane organization and storage protein trafficking in rice endosperm cells","volume":"64","author":"Wang","year":"2010","journal-title":"The Plant Journal"},{"key":"2026011209055325900_eraf410-B29","doi-asserted-by":"crossref","first-page":"2850","DOI":"10.1105\/tpc.16.00717","article-title":"GOLGI TRANSPORT 1B regulates protein export from the endoplasmic reticulum in rice endosperm cells","volume":"28","author":"Wang","year":"2016","journal-title":"The Plant Cell"},{"key":"2026011209055325900_eraf410-B30","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1111\/j.1365-313X.2011.04880.x","article-title":"RNA targeting to a specific ER sub-domain is required for efficient transport and packaging of \u03b1-globulins to the protein storage vacuole in developing rice endosperm","volume":"70","author":"Washida","year":"2012","journal-title":"The Plant Journal"},{"key":"2026011209055325900_eraf410-B31","doi-asserted-by":"crossref","first-page":"1397","DOI":"10.1007\/s00122-009-1143-x","article-title":"Amplification of prolamin storage protein genes in different subfamilies of the Poaceae","volume":"119","author":"Xu","year":"2009","journal-title":"Theoretical and Applied Genetics"},{"key":"2026011209055325900_eraf410-B32","doi-asserted-by":"crossref","first-page":"1949","DOI":"10.1038\/s41467-019-09919-y","article-title":"Natural variation of OsGluA2 is involved in grain protein content regulation in rice","volume":"10","author":"Yang","year":"2019","journal-title":"Nature Communications"},{"key":"2026011209055325900_eraf410-B33","doi-asserted-by":"crossref","first-page":"111567","DOI":"10.1016\/j.plantsci.2022.111567","article-title":"Identification of a new allele of soluble starch synthase IIIa involved in the elongation of amylopectin long chains in a chalky rice mutant","volume":"328","author":"Ying","year":"2023","journal-title":"Plant Science"},{"key":"2026011209055325900_eraf410-B34","first-page":"10","article-title":"Extraction and determination of rice protein fractions","volume":"2","author":"Yu","year":"2023","journal-title":"Quality and Safety of Agro-Products"},{"key":"2026011209055325900_eraf410-B35","doi-asserted-by":"crossref","first-page":"6","DOI":"10.1016\/j.rsci.2024.08.005","article-title":"Regulation of yield and eating and cooking quality in rice through seed albumin OsRAL5","volume":"32","author":"Zhao","year":"2025","journal-title":"Rice Science"},{"key":"2026011209055325900_eraf410-B36","doi-asserted-by":"crossref","first-page":"108014","DOI":"10.1016\/j.biotechadv.2022.108014","article-title":"Genetic control of grain appearance quality in rice","volume":"60","author":"Zhao","year":"2022","journal-title":"Biotechnology Advances"},{"key":"2026011209055325900_eraf410-B37","doi-asserted-by":"crossref","first-page":"568","DOI":"10.1111\/pbi.12654","article-title":"Overexpression of the 16-kDa \u03b1-amylase\/trypsin inhibitor RAG2 improves grain yield and quality of rice","volume":"15","author":"Zhou","year":"2017","journal-title":"Plant Biotechnology Journal"},{"key":"2026011209055325900_eraf410-B38","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.carbpol.2018.04.070","article-title":"Physicochemical properties and digestibility of endosperm starches in four indica rice mutants","volume":"195","author":"Zhou","year":"2018","journal-title":"Carbohydrate Polymers"},{"key":"2026011209055325900_eraf410-B39","doi-asserted-by":"crossref","first-page":"1059749","DOI":"10.3389\/fpls.2022.1059749","article-title":"High temperature boosts resistant starch content by altering starch structure and lipid content in rice ssIIIa mutants","volume":"13","author":"Zhou","year":"2022","journal-title":"Frontiers in Plant Science"},{"key":"2026011209055325900_eraf410-B40","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1186\/s12870-019-1911-y","article-title":"OsNHX5-mediated pH homeostasis is required for post-Golgi trafficking of seed storage proteins in rice endosperm cells","volume":"19","author":"Zhu","year":"2019","journal-title":"BMC Plant Biology"},{"key":"2026011209055325900_eraf410-B41","doi-asserted-by":"crossref","first-page":"2192","DOI":"10.1093\/plphys\/kiab099","article-title":"Subunit E isoform 1 of vacuolar H+-ATPase OsVHA enables post-Golgi trafficking of rice seed storage proteins","volume":"187","author":"Zhu","year":"2021","journal-title":"Plant Physiology"}],"container-title":["Journal of Experimental Botany"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/jxb\/advance-article-pdf\/doi\/10.1093\/jxb\/eraf410\/64295532\/eraf410.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/jxb\/article-pdf\/77\/2\/379\/64295532\/eraf410.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/jxb\/article-pdf\/77\/2\/379\/64295532\/eraf410.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,1,12]],"date-time":"2026-01-12T14:06:05Z","timestamp":1768226765000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/jxb\/article\/77\/2\/379\/8256485"}},"subtitle":[],"editor":[{"given":"Scott","family":"Boden","sequence":"additional","affiliation":[{"name":"University of Adelaide ,","place":["Australia"]}],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2025,9,17]]},"references-count":41,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2025,9,17]]},"published-print":{"date-parts":[[2026,1,12]]}},"URL":"https:\/\/doi.org\/10.1093\/jxb\/eraf410","relation":{},"ISSN":["0022-0957","1460-2431"],"issn-type":[{"value":"0022-0957","type":"print"},{"value":"1460-2431","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026,1,12]]},"published":{"date-parts":[[2025,9,17]]}}}