{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,28]],"date-time":"2026-09-28T01:56:10Z","timestamp":1790560570421,"version":"4.1.0"},"reference-count":42,"publisher":"CSIRO Publishing","issue":"10","license":[{"start":{"date-parts":[[2026,9,5]],"date-time":"2026-09-05T00:00:00Z","timestamp":1788566400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/doi.org\/10.1071\/journalslicense"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,10,23]]},"abstract":"<jats:p>Salt stress is a major environmental factor limiting crop growth and productivity. We found a long-chain acyl-CoA synthetase (LACS) gene ZmLACS9 that was associated with salt tolerance traits in maize (Zea mays). Mutation of ZmLACS9 impaired plant growth, while overexpression of ZmLACS9 improved plant growth vigour under salt stress conditions. Metabolome and transcriptome analyses indicated that ZmLACS9 may regulate the expression of lipid trafficking-related genes, thereby influencing the concentrations of galactolipids and phospholipid in the photosynthetic membrane under salt stress. Under salt stress conditions, ultrastructural observation and chlorophyll fluorescence parameters showed that the mutation of ZmLACS9 led to significant impairment of the number of thylakoid layer structures and PSII activity, while overexpression of ZmLACS9 markedly improved the number of chloroplast thylakoid grana lamella and PSII activity. Furthermore, yeast one-hybrid and LUC transient expression assays found that ZmWRKY17 could bind to the promoter of ZmLACS9. Collectively, our study provides candidate genes for breeding maize varieties with higher stress resistance.<\/jats:p>","DOI":"10.1071\/fp26135","type":"journal-article","created":{"date-parts":[[2026,9,28]],"date-time":"2026-09-28T01:21:35Z","timestamp":1790558495000},"source":"Crossref","is-referenced-by-count":0,"title":["Lipidomic and transcriptomic analyses reveal that the gene ZmLACS9 enhances salt stress tolerance by regulating chloroplast lipid metabolism and maintaining photosynthetic performance in maize"],"prefix":"10.1071","volume":"53","author":[{"given":"Cheng","family":"Chi","sequence":"first","affiliation":[{"name":"ASchool of Biological Science and Technology, University of Jinan, Jinan 250022, China."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Libo","family":"Wang","sequence":"additional","affiliation":[{"name":"BSichuan Light-Industry Research & Design Institute Co., Ltd., Chengdu 610081, China."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hongtao","family":"Li","sequence":"additional","affiliation":[{"name":"CLianyungang Academy of Agricultural Sciences, Lianyungang 222000, China."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qinyou","family":"Xu","sequence":"additional","affiliation":[{"name":"ASchool of Biological Science and Technology, University of Jinan, Jinan 250022, China."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"MingZi","family":"Shi","sequence":"additional","affiliation":[{"name":"ASchool of Biological Science and Technology, University of Jinan, Jinan 250022, China."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Menghan","family":"Sun","sequence":"additional","affiliation":[{"name":"ASchool of Biological Science and Technology, University of Jinan, Jinan 250022, China."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Haojie","family":"Pi","sequence":"additional","affiliation":[{"name":"ASchool of Biological Science and Technology, University of Jinan, Jinan 250022, China."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5398-2552","authenticated-orcid":true,"given":"Hui","family":"Li","sequence":"additional","affiliation":[{"name":"ASchool of Biological Science and Technology, University of Jinan, Jinan 250022, China."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhaohua","family":"Ding","sequence":"additional","affiliation":[{"name":"DMaize Research Institute, Shandong Academy of Agricultural Sciences, Jinan 250100, China."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4706-586X","authenticated-orcid":true,"given":"Jiantang","family":"Zhu","sequence":"additional","affiliation":[{"name":"ASchool of Biological Science and Technology, University of Jinan, Jinan 250022, China."}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"67","published-online":{"date-parts":[[2026,9,28]]},"reference":[{"key":"2026092721213302100_R1","doi-asserted-by":"publisher","first-page":"64","DOI":"10.1016\/j.plaphy.2020.08.042","article-title":"Salinity induced physiological and biochemical changes in plants: an omic approach towards salt stress tolerance.","volume":"156","author":"Arif","year":"2020","journal-title":"Plant Physiology and Biochemistry"},{"key":"2026092721213302100_R2","doi-asserted-by":"publisher","first-page":"32","DOI":"10.1016\/j.jplph.2016.03.018","article-title":"Changes in chloroplast lipid contents and chloroplast ultrastructure in Sulla carnosa and Sulla coronaria leaves under salt stress.","volume":"198","author":"Bejaoui","year":"2016","journal-title":"Journal of Plant Physiology"},{"key":"2026092721213302100_R3","doi-asserted-by":"publisher","first-page":"71","DOI":"10.1146\/annurev.cellbio.042308.113414","article-title":"Mechanisms of lipid transport involved in organelle biogenesis in plant cells.","volume":"25","author":"Benning","year":"2009","journal-title":"Annual Review of Cell and Developmental Biology"},{"key":"2026092721213302100_R4"},{"key":"2026092721213302100_R5","doi-asserted-by":"publisher","first-page":"470","DOI":"10.1016\/j.bbabio.2013.09.007","article-title":"Glycerolipids in photosynthesis: composition, synthesis and trafficking.","volume":"1837","author":"Boudi\u00e8re","year":"2014","journal-title":"Biochimica et Biophysica Acta (BBA) - Bioenergetics"},{"key":"2026092721213302100_R6","doi-asserted-by":"publisher","first-page":"2633","DOI":"10.1093\/bioinformatics\/btm308","article-title":"TASSEL: software for association mapping of complex traits in diverse samples.","volume":"23","author":"Bradbury","year":"2007","journal-title":"Bioinformatics"},{"key":"2026092721213302100_R7","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-14027-y","article-title":"Natural variation of an EF-hand Ca2+-binding-protein coding gene confers saline-alkaline tolerance in maize.","volume":"11","author":"Cao","year":"2020","journal-title":"Nature Communications"},{"key":"2026092721213302100_R8","doi-asserted-by":"publisher","first-page":"735","DOI":"10.1046\/j.1365-313x.1998.00343.x","article-title":"Floral dip: a simplified method for Agrobacterium-mediated transformation of Arabidopsis thaliana.","volume":"16","author":"Clough","year":"1998","journal-title":"The Plant Journal"},{"key":"2026092721213302100_R9","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1046\/j.1365-313X.2002.01405.x","article-title":"Two long-chain acyl-CoA synthetases from Arabidopsis thaliana involved in peroxisomal fatty acid \u03b2-oxidation.","volume":"32","author":"Fulda","year":"2002","journal-title":"The Plant Journal"},{"key":"2026092721213302100_R10","doi-asserted-by":"publisher","first-page":"489","DOI":"10.1016\/S1360-1385(00)01767-2","article-title":"Self-regulation of the lipid content of membranes by non-bilayer lipids: a hypothesis.","volume":"5","author":"Garab","year":"2000","journal-title":"Trends in Plant Science"},{"key":"2026092721213302100_R11","doi-asserted-by":"publisher","first-page":"726","DOI":"10.1111\/tpj.12241","article-title":"The role of lipid metabolism in the acquisition of desiccation tolerance in Craterostigma plantagineum: a comparative approach.","volume":"75","author":"Gasulla","year":"2013","journal-title":"The Plant Journal"},{"key":"2026092721213302100_R12","doi-asserted-by":"publisher","first-page":"1029","DOI":"10.1111\/pce.12666","article-title":"Lipid signalling in plant responses to abiotic stress.","volume":"39","author":"Hou","year":"2016","journal-title":"Plant, Cell & Environment"},{"key":"2026092721213302100_R13","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2016.01464","article-title":"Genome-wide analysis and expression profiling of the SUC and SWEET gene families of sucrose transporters in oilseed rape (Brassica napus L.).","volume":"7","author":"Jian","year":"2016","journal-title":"Frontiers in Plant Science"},{"key":"2026092721213302100_R14"},{"key":"2026092721213302100_R15","doi-asserted-by":"publisher","DOI":"10.3390\/ijms21062223","article-title":"Functional analysis of rice long-chain acyl-CoA synthetase 9 (OsLACS9) in the chloroplast envelope membrane.","volume":"21","author":"Kitajima-Koga","year":"2020","journal-title":"International Journal of Molecular Sciences"},{"key":"2026092721213302100_R16","doi-asserted-by":"publisher","first-page":"565","DOI":"10.1007\/s10265-016-0827-y","article-title":"Role of membrane glycerolipids in photosynthesis, thylakoid biogenesis and chloroplast development.","volume":"129","author":"Kobayashi","year":"2016","journal-title":"Journal of Plant Research"},{"key":"2026092721213302100_R17","doi-asserted-by":"publisher","first-page":"10270","DOI":"10.1073\/pnas.0702061104","article-title":"EXECUTER1- and EXECUTER2-dependent transfer of stress-related signals from the plastid to the nucleus of Arabidopsis thaliana.","volume":"104","author":"Lee","year":"2007","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"key":"2026092721213302100_R18","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1038\/ng.2484","article-title":"Genome-wide association study dissects the genetic architecture of oil biosynthesis in maize kernels.","volume":"45","author":"Li","year":"2013","journal-title":"Nature Genetics"},{"key":"2026092721213302100_R19","doi-asserted-by":"publisher","first-page":"1404","DOI":"10.1104\/pp.19.00894","article-title":"A sequence-indexed Mutator insertional library for maize functional genomics study.","volume":"181","author":"Liang","year":"2019","journal-title":"Plant Physiology"},{"key":"2026092721213302100_R20","doi-asserted-by":"publisher","DOI":"10.1007\/s11738-017-2501-y","article-title":"Antioxidants and unsaturated fatty acids are involved in salt tolerance in Peanut.","volume":"39","author":"Liu","year":"2017","journal-title":"Acta Physiologiae Plantarum"},{"key":"2026092721213302100_R21","doi-asserted-by":"publisher","first-page":"335","DOI":"10.1007\/s40626-018-0129-y","article-title":"Exogenous application of gibberellic acid participates in up-regulation of lipid biosynthesis under salt stress in rice.","volume":"30","author":"Liu","year":"2018","journal-title":"Theoretical and Experimental Plant Physiology"},{"key":"2026092721213302100_R22","doi-asserted-by":"publisher","first-page":"1937","DOI":"10.1111\/pbi.13607","article-title":"Molecular dissection of maize seedling salt tolerance using a genome-wide association analysis method.","volume":"19","author":"Luo","year":"2021","journal-title":"Plant Biotechnology Journal"},{"issue":"3","key":"2026092721213302100_R23","doi-asserted-by":"publisher","first-page":"473","DOI":"10.1034\/j.1399-3054.1994.920316.x","article-title":"Plasma membrane lipid alterations induced by NaCl in winter wheat roots.","volume":"92","author":"Magdy","year":"1994","journal-title":"Physiologia Plantarum"},{"key":"2026092721213302100_R24","doi-asserted-by":"publisher","first-page":"416","DOI":"10.1007\/s12229-015-9156-4","article-title":"Role of the plasma membrane in saline conditions: lipids and proteins.","volume":"81","author":"Mansour","year":"2015","journal-title":"The Botanical Review"},{"key":"2026092721213302100_R25","doi-asserted-by":"publisher","first-page":"226","DOI":"10.1126\/science.1191803","article-title":"Freezing tolerance in plants requires lipid remodeling at the outer chloroplast membrane.","volume":"330","author":"Moellering","year":"2010","journal-title":"Science"},{"key":"2026092721213302100_R26","doi-asserted-by":"publisher","first-page":"651","DOI":"10.1146\/annurev.arplant.59.032607.092911","article-title":"Mechanisms of salinity tolerance.","volume":"59","author":"Munns","year":"2008","journal-title":"Annual Review of Plant Biology"},{"key":"2026092721213302100_R27","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1111\/ppl.12404","article-title":"Salinity induces membrane structure and lipid changes in maize mesophyll and bundle sheath chloroplasts.","volume":"157","author":"Omoto","year":"2016","journal-title":"Physiologia Plantarum"},{"key":"2026092721213302100_R28","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1016\/j.plaphy.2011.10.003","article-title":"Arabidopsis LONG-CHAIN ACYL-COA SYNTHETASE 1 (LACS1), LACS2, and LACS3 facilitate fatty acid uptake in yeast.","volume":"51","author":"Pulsifer","year":"2012","journal-title":"Plant Physiology and Biochemistry"},{"key":"2026092721213302100_R29"},{"key":"2026092721213302100_R30","doi-asserted-by":"publisher","first-page":"1903","DOI":"10.1104\/pp.15.01935","article-title":"Nonphotochemical chlorophyll fluorescence quenching: mechanism and effectiveness in protecting plants from photodamage.","volume":"170","author":"Ruban","year":"2016","journal-title":"Plant Physiology"},{"key":"2026092721213302100_R31","doi-asserted-by":"publisher","first-page":"6926","DOI":"10.1073\/pnas.88.16.6926","article-title":"Differential effects of plant sterols on water permeability and on acyl chain ordering of soybean phosphatidylcholine bilayers.","volume":"88","author":"Schuler","year":"1991","journal-title":"Proceedings of the National Academy of Sciences"},{"key":"2026092721213302100_R32","first-page":"1","article-title":"Insights into the physiological and biochemical responses of plants to salt stress.","volume":"39","author":"Shahid","year":"2020","journal-title":"Critical Reviews in Plant Sciences"},{"key":"2026092721213302100_R33","doi-asserted-by":"publisher","first-page":"1710","DOI":"10.1104\/pp.003269","article-title":"Arabidopsis contains nine long-chain acyl-coenzyme a synthetase genes that participate in fatty acid and glycerolipid metabolism.","volume":"129","author":"Shockey","year":"2002","journal-title":"Plant Physiology"},{"key":"2026092721213302100_R34","doi-asserted-by":"publisher","first-page":"1473","DOI":"10.1073\/pnas.97.4.1473","article-title":"Self-assembly of large, ordered lamellae from non-bilayer lipids and integral membrane proteins in vitro.","volume":"97","author":"Simidjiev","year":"2000","journal-title":"Proceedings of the National Academy of Sciences"},{"key":"2026092721213302100_R35","doi-asserted-by":"publisher","first-page":"623","DOI":"10.1007\/s11099-010-0080-x","article-title":"Increase in unsaturated fatty acids in membrane lipids of Suaeda salsa L. enhances protection of photosystem II under high salinity.","volume":"48","author":"Sui","year":"2010","journal-title":"Photosynthetica"},{"key":"2026092721213302100_R36","doi-asserted-by":"publisher","first-page":"861","DOI":"10.1080\/07388551.2019.1616669","article-title":"Avenues of the membrane transport system in adaptation of plants to abiotic stresses.","volume":"39","author":"Vishwakarma","year":"2019","journal-title":"Critical Reviews in Biotechnology"},{"key":"2026092721213302100_R37","doi-asserted-by":"publisher","DOI":"10.1016\/j.stress.2023.100271","article-title":"Genome-wide analysis of the maize LACS gene family and functional characterization of the ZmLACS9 responses to heat stress.","volume":"10","author":"Wang","year":"2023","journal-title":"Plant Stress"},{"key":"2026092721213302100_R38"},{"key":"2026092721213302100_R39","doi-asserted-by":"publisher","first-page":"37","DOI":"10.1111\/tpj.15273","article-title":"Mechanisms and functions of membrane lipid remodeling in plants.","volume":"107","author":"Yu","year":"2021","journal-title":"The Plant Journal"},{"key":"2026092721213302100_R40","doi-asserted-by":"publisher","first-page":"1450","DOI":"10.1093\/treephys\/tpaa079","article-title":"An apple long-chain acyl-CoA synthetase 2 gene enhances plant resistance to abiotic stress by regulating the accumulation of cuticular wax.","volume":"40","author":"Zhang","year":"2020","journal-title":"Tree Physiology"},{"key":"2026092721213302100_R41","doi-asserted-by":"publisher","first-page":"2018","DOI":"10.1093\/pcp\/pcaa116","article-title":"Choline-mediated lipid reprogramming as a dominant salt tolerance mechanism in grass species lacking glycine betaine.","volume":"61","author":"Zhang","year":"2020","journal-title":"Plant & Cell Physiology"},{"key":"2026092721213302100_R42","doi-asserted-by":"publisher","DOI":"10.1186\/s13068-022-02254-3","article-title":"Overexpression the BnLACS9 could increase the chlorophyll and oil content in Brassica napus.","volume":"16","author":"Zhu","year":"2023","journal-title":"Biotechnology for Biofuels and Bioproducts"}],"container-title":["Functional Plant Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/connectsci.au\/fp\/article-pdf\/doi\/10.1071\/FP26135\/1971936\/fp26135.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/connectsci.au\/fp\/article-pdf\/doi\/10.1071\/FP26135\/1971936\/fp26135.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,28]],"date-time":"2026-09-28T01:21:37Z","timestamp":1790558497000},"score":1,"resource":{"primary":{"URL":"https:\/\/connectsci.au\/fp\/article\/53\/10\/FP26135\/275988\/Lipidomic-and-transcriptomic-analyses-reveal-that"}},"subtitle":[],"editor":[{"given":"Suleyman","family":"Allakhverdiev","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2026,9,28]]},"references-count":42,"journal-issue":{"issue":"10","published-print":{"date-parts":[[2026,10,23]]}},"URL":"https:\/\/doi.org\/10.1071\/fp26135","relation":{},"ISSN":["1445-4408","1445-4416"],"issn-type":[{"value":"1445-4408","type":"print"},{"value":"1445-4416","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,28]]},"article-number":"FP26135"}}