{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,13]],"date-time":"2026-06-13T18:24:03Z","timestamp":1781375043731,"version":"3.54.1"},"reference-count":36,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,5,12]],"date-time":"2026-05-12T00:00:00Z","timestamp":1778544000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"},{"start":{"date-parts":[[2026,5,12]],"date-time":"2026-05-12T00:00:00Z","timestamp":1778544000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Mol Biol Rep"],"published-print":{"date-parts":[[2026,12]]},"DOI":"10.1007\/s11033-026-11949-x","type":"journal-article","created":{"date-parts":[[2026,5,12]],"date-time":"2026-05-12T08:29:42Z","timestamp":1778574582000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Genome wide identification, evolution and expression analysis of NHX gene family in cultivated barley (Hordeum vulgare. L) grown under salinity"],"prefix":"10.1007","volume":"53","author":[{"given":"Warda","family":"Saoudi","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chedly","family":"Abdelly","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hatem","family":"Boubakri","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,5,12]]},"reference":[{"key":"11949_CR1","doi-asserted-by":"publisher","DOI":"10.1016\/j.plgene.2019.100178","volume":"18","author":"R Yarra","year":"2019","unstructured":"Yarra R (2019) The wheat NHX gene family: potential role in improving salinity stress tolerance of plants. Plant Gene 18:100178","journal-title":"Plant Gene"},{"key":"11949_CR2","doi-asserted-by":"publisher","DOI":"10.1016\/j.stress.2023.100266","volume":"10","author":"K Parveen","year":"2023","unstructured":"Parveen K, Saddique MAB, Rehman SU, Ali Z, Aziz I, Shamsi IH, Muneer MA (2023) Identification and characterization of salt stress-responsive NHX gene family in chickpea. Plant Stress 10:100266","journal-title":"Plant Stress"},{"key":"11949_CR3","doi-asserted-by":"publisher","DOI":"10.1016\/j.envexpbot.2024.105690","volume":"220","author":"M Qu","year":"2024","unstructured":"Qu M, Havsh\u00f8i NW, Huang X, Shabala L, Yu M, Fuglsang AT, Shabala S (2024) Understanding the mechanistic basis of ameliorative effects of boron on salinity in barley (Hordeum vulgare). Environ Exp Bot 220:105690","journal-title":"Environ Exp Bot"},{"key":"11949_CR4","doi-asserted-by":"publisher","first-page":"1701","DOI":"10.3390\/plants13121701","volume":"13","author":"S Liu","year":"2024","unstructured":"Liu S, An Z, Li Y, Yang R, Lai Z (2024) Genome-wide identification of the Cation\/Proton Antiporter (CPA) gene family and functional analysis of AtrNHX8 under salt stress. Plants Basel 13:1701","journal-title":"Plants Basel"},{"key":"11949_CR5","doi-asserted-by":"publisher","DOI":"10.3389\/fgene.2024.1396375","volume":"15","author":"J Yihong","year":"2024","unstructured":"Yihong J, Zhen L, Chang L, Ziying S, Ning Z, Meiqing S, Yuhui L, Lei W (2024) Genome wide identification and drought stress-induced expression analysis of the NHX gene family in potato. Front Genet 15:1396375","journal-title":"Front Genet"},{"key":"11949_CR6","doi-asserted-by":"publisher","DOI":"10.3389\/fgene.2020.00964","volume":"11","author":"X Fu","year":"2020","unstructured":"Fu X, Lu Z, Wei H, Zhang J, Yang X, Wu A, Ma L, Kang M, Lu J, Wang H, Yu S (2020) Genome-wide identification and expression analysis of the NHX (Sodium\/Hydrogen Antiporter) gene family in cotton. Front Genet 11:964","journal-title":"Front Genet"},{"issue":"7","key":"11949_CR7","doi-asserted-by":"publisher","DOI":"10.3390\/genes11070803","volume":"11","author":"U Akram","year":"2020","unstructured":"Akram U, Song Y, Liang C, Abid MA, Askari M, Myat AA, Abbas M, Malik W, Ali Z, Guo S, Zhang R, Meng Z (2020) Genome-wide characterization and expression analysis of NHX gene family under salinity stress in Gossypium barbadense and its comparison with Gossypium hirsutum. Genes Basel 11(7):803","journal-title":"Genes Basel"},{"key":"11949_CR8","doi-asserted-by":"publisher","first-page":"356","DOI":"10.1016\/j.ygeno.2019.02.015","volume":"112","author":"H Sharma","year":"2020","unstructured":"Sharma H, Taneja M, Upadhyay SK (2020) Identification, characterization and expression profiling of cation-proton antiporter superfamily in Triticum aestivum L. and functional analysis of TaNHX4-B. Genomics 112:356\u2013370","journal-title":"Genomics"},{"key":"11949_CR9","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.96.4.1480","author":"A Roberto","year":"1999","unstructured":"Roberto A, Gaxiola RR, Amir S, Paula G, Seth LA (1999) The Arabidopsis thaliana proton transporters, AtNhx1 and Avp1, can function in cation detoxification in yeast. Proc Natl Acad Sci USA. https:\/\/doi.org\/10.1073\/pnas.96.4.1480","journal-title":"Proc Natl Acad Sci USA"},{"key":"11949_CR10","doi-asserted-by":"publisher","first-page":"3482","DOI":"10.1105\/tpc.111.089581","volume":"23","author":"E Bassil","year":"2011","unstructured":"Bassil E, Tajima H, Liang YC, Ohto MA, Ushijima K, Nakano R, Esumi T, Coku A, Belmonte M, Blumwald E (2011) The Arabidopsis Na+\/H+ antiporters NHX1 and NHX2 control vacuolar pH and K+ homeostasis to regulate growth, flower development, and reproduction. Plant Cell 23:3482\u20133497","journal-title":"Plant Cell"},{"key":"11949_CR11","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/s13205-020-02555-0","volume":"11","author":"S Joshi","year":"2021","unstructured":"Joshi S, Kaur K, Khare T, Srivastava AK, Suprasanna P, Kumar V (2021) Genome-wide identification, characterization and transcriptional profiling of NHX-type (Na+\/H+) antiporters under salinity stress in soybean. 3 Biotech 11:1\u201317","journal-title":"3 Biotech"},{"key":"11949_CR12","doi-asserted-by":"crossref","unstructured":"Wu GQ, Wang JL, Li SJ (2019) Genome-wide identification of Na+\/H+ antiporter (NHX) genes in sugar beet (Beta vulgaris L.) and their regulated expression under salt stress. Genes 10(5):401","DOI":"10.3390\/genes10050401"},{"key":"11949_CR13","doi-asserted-by":"publisher","first-page":"109","DOI":"10.1016\/j.jplph.2017.07.015","volume":"218","author":"N Li","year":"2017","unstructured":"Li N, Wang X, Ma B, Du C, Zheng L, Wang Y (2017) Expression of a Na+\/H+ antiporter RtNHX1 from recretohalophyte Reaumuria trigyna improved salt tolerance of transgenic Arabidopsis thaliana. J Plant Physiol 218:109\u2013120","journal-title":"J Plant Physiol"},{"issue":"3","key":"11949_CR14","doi-asserted-by":"publisher","first-page":"964","DOI":"10.1111\/pbi.12451","volume":"14","author":"AK Bao","year":"2016","unstructured":"Bao AK, Du BQ, Touil L, Kang P, Wang QL, Wang SM (2016) Co-expression of tonoplast Cation\/H+ antiporter and H+\u2010pyrophosphatase from xerophyte Zygophyllum xanthoxylum improves alfalfa plant growth under salinity, drought and field conditions. Plant Biotechnol J 14(3):964\u2013975","journal-title":"Plant Biotechnol J"},{"key":"11949_CR15","doi-asserted-by":"publisher","first-page":"771920","DOI":"10.3389\/fpls.2022.771920","volume":"13","author":"V Kaur","year":"2022","unstructured":"Kaur V, Aravind J, Manju, Narendra P, Rana JC, Anjula P, Ashok K (2022) Phenotypic characterization, genetic diversity assessment in 6,778 accessions of Barley (Hordeum vulgare L. ssp. vulgare) germplasm conserved in national gene bank of India and development of a core set. Front Plant Sci 13:771920","journal-title":"Front Plant Sci"},{"key":"11949_CR16","doi-asserted-by":"publisher","DOI":"10.1016\/j.envexpbot.2024.105852","volume":"225","author":"W Saoudi","year":"2024","unstructured":"Saoudi W, Boubakri H, Taamalli W, Debez A, Abdelly C (2024) Differential reduction in cadmium accumulation in barley under cadmium-salt co-exposure involves specific HMA genes and antioxidative responses. Environ Exp Bot 225:105852","journal-title":"Environ Exp Bot"},{"issue":"5","key":"11949_CR17","doi-asserted-by":"publisher","DOI":"10.1111\/jac.12756","volume":"210","author":"S Singh","year":"2024","unstructured":"Singh S, Yadav SK, Arya SS, Jacob SR, Gautam RK, Singh GP, Kaur V (2024) Unravelling morpho-physiological mechanism and candidate genes associated with salinity tolerance in superior haplotypes of barley (Hordeum vulgare L.). J Agron Crop Sci 210(5):e12756","journal-title":"J Agron Crop Sci"},{"key":"11949_CR18","unstructured":"Hewitt EJ (1966) Sand and water culture methods used in the study of plant nutrition. Tech Commun Bur Hort. 22:431\u2013446"},{"key":"11949_CR19","doi-asserted-by":"publisher","DOI":"10.1016\/j.gene.2022.146591","volume":"833","author":"H Boubakri","year":"2022","unstructured":"Boubakri H, Najjar E, Jihnaoui N, Chihaoui S, Barhoumi F, Jebara M (2022) Genome-wide identification, characterization and expression analysis of glutaredoxin gene family (Grxs) in Phaseolus vulgaris. Gene 833:146591","journal-title":"Gene"},{"issue":"4","key":"11949_CR20","doi-asserted-by":"publisher","first-page":"402","DOI":"10.1006\/meth.2001.1262","volume":"25","author":"KJ Livak","year":"2001","unstructured":"Livak KJ, Schmittgen TD (2001) Analysis of relative gene expression data using real-time quantitative PCR and the 2(-Delta Delta C(T)) method. Methods 25(4):402\u2013408","journal-title":"Methods"},{"issue":"12","key":"11949_CR21","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0028150","volume":"6","author":"Y Wang","year":"2011","unstructured":"Wang Y, Wang X, Tang H, Tan X, Ficklin SP, Feltus FA, Paterson AH (2011) Modes of gene duplication contribute differently to genetic novelty and redundancy, but show parallels across divergent angiosperms. PloS one 6(12):e28150","journal-title":"PloS one"},{"key":"11949_CR22","doi-asserted-by":"publisher","first-page":"236","DOI":"10.3390\/genes9050236","volume":"9","author":"H Kumari","year":"2018","unstructured":"Kumari H, Kumar S, Ramesh K, Palakolanu SR, Mark N, Prakash A, Shah T, Henderson A, Srivastava R, Rajasheker G (2018) Genome-wide identification and analysis of Arabidopsis sodium proton antiporter (NHX) and human sodium proton exchanger (NHE) homologs in Sorghum bicolor. Genes 9:236","journal-title":"Genes"},{"key":"11949_CR23","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-017-02894-8","volume":"7","author":"FX Tian","year":"2017","unstructured":"Tian FX, Chang E, Li Y, Sun P, Hu JJ, Zhang J (2017) Expression and integrated network analyses revealed functional divergence of NHX-type Na+\/H+ exchanger genes in poplar. Sci Rep 7:2607","journal-title":"Sci Rep"},{"key":"11949_CR24","doi-asserted-by":"publisher","first-page":"73","DOI":"10.1016\/j.plantsci.2005.02.030","volume":"169","author":"YY Wu","year":"2005","unstructured":"Wu YY, Chen QJ, Chen M, Chen J, Wang XC (2005) Salt-tolerant transgenic perennial ryegrass (Lolium perenne L.) obtained by Agrobacterium tumefaciens- mediated transformation of the vacuolar Na+\/H+ antiporter gene. Plant Sci 169:73","journal-title":"Plant Sci"},{"key":"11949_CR25","doi-asserted-by":"publisher","first-page":"356","DOI":"10.1007\/s12011-010-8662-9","volume":"139","author":"G Wu","year":"2011","unstructured":"Wu G, Wang Q, Bao A, Wang S (2011) Amiloride reduces sodium transport and accumulation in the succulent xerophyte Zygophyllum xanthoxylum under salt conditions. Biol Trace Elem Res 139:356\u2013367","journal-title":"Biol Trace Elem Res"},{"key":"11949_CR26","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2018.00809","volume":"9","author":"Z Xu","year":"2018","unstructured":"Xu Z, Raza Q, Xu L, He X, Huang Y, Yi J (2018) GmWRKY49, a salt-responsive nuclear protein, improved root length and governed better salinity tolerance in transgenic Arabidopsis. Front Plant Sci 9:809","journal-title":"Front Plant Sci"},{"key":"11949_CR27","doi-asserted-by":"publisher","first-page":"11444","DOI":"10.1073\/pnas.191389398","volume":"98","author":"RA Gaxiola","year":"2001","unstructured":"Gaxiola RA, Li JL, Undurraga S, Dang LM, Fink GR (2001) Drought- and salt-tolerant plants result from overexpression of the AVP1 H+-pump. Proc Natl Acad Sci U S A 98:11444\u201311449","journal-title":"Proc Natl Acad Sci U S A"},{"key":"11949_CR28","doi-asserted-by":"publisher","first-page":"341","DOI":"10.1016\/j.plaphy.2020.05.012","volume":"154","author":"MR Romero-Aranda","year":"2020","unstructured":"Romero-Aranda MR, Fern\u00e1ndez PG, L\u00f3pez-Tienda JR, L\u00f3pez-Diaz MR, Espinosa J, Granum E, Traverso J\u00c1, Pineda B, Garcia-Sogo B, Moreno V (2020) Na+ transporter HKT1;2 reduces flower Na+ content and considerably mitigates the decline in tomato fruit yields under saline conditions. Plant Physiol Biochem 154:341\u2013352","journal-title":"Plant Physiol Biochem"},{"key":"11949_CR29","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2016.01296","volume":"7","author":"KA Wilkins","year":"2016","unstructured":"Wilkins KA, Matthus E, Swarbreck SM, Davies JM (2016) Calcium-mediated abiotic stress signaling in roots. Front Plant Sci 7:1296","journal-title":"Front Plant Sci"},{"key":"11949_CR30","doi-asserted-by":"publisher","first-page":"232","DOI":"10.1016\/j.plantsci.2008.10.009","volume":"176","author":"AK Bao","year":"2008","unstructured":"Bao AK, Wang SM, Wu GQ, Xi JJ, Wang CM (2008) Overexpression of the Arabidopsis H+ -PPase enhanced resistance to salt and drought stress in transgenic alfalfa (Medicago sativa L). Plant Sci 176:232\u2013240","journal-title":"Plant Sci"},{"key":"11949_CR31","doi-asserted-by":"crossref","unstructured":"Mani SMN, Shi SJ, Mao JJ, Dong LH, Su YL, Wang Q, Liu H (2015) The calcium sensor CBL-CIPK is involved in plant\u2019s response to abiotic stresses. Int J Genom 2015:1\u201310","DOI":"10.1155\/2015\/493191"},{"key":"11949_CR32","doi-asserted-by":"publisher","first-page":"80","DOI":"10.3389\/fpls.2019.00080","volume":"10","author":"SV Isayenkov","year":"2019","unstructured":"Isayenkov SV, Maathuis FJ (2019) Plant salinity stress: many unanswered questions remain. Front Plant Sci 10:80","journal-title":"Front Plant Sci"},{"issue":"5","key":"11949_CR33","doi-asserted-by":"publisher","first-page":"529","DOI":"10.1046\/j.1365-313X.2002.01309.x","volume":"30","author":"S Yokoi","year":"2002","unstructured":"Yokoi S, Quintero FJ, Cubero B, Ruiz MT, Bressan RA, Hasegawa PM, Pardo JM (2002) Differential expression and function of Arabidopsis thaliana NHX Na+\/H+ antiporters in the salt stress response. Plant J 30(5):529\u2013539","journal-title":"Plant J"},{"key":"11949_CR34","doi-asserted-by":"publisher","first-page":"17","DOI":"10.1016\/j.plaphy.2013.12.013","volume":"76","author":"W Lu","year":"2014","unstructured":"Lu W, Guo C, Li X, Duan W, Ma C, Zhao M, Gu J, Du X, Liu Z, Xiao K (2014) Overexpression of TaNHX3, a vacuolar Na+\/H+ antiporter gene in wheat, enhances salt stress tolerance in tobacco by improving related physiological processes. Plant Physiol Biochem 76:17\u201328","journal-title":"Plant Physiol Biochem"},{"key":"11949_CR35","doi-asserted-by":"publisher","first-page":"809988","DOI":"10.3389\/fgene.2021.809988","volume":"12","author":"Z Jabeen","year":"2022","unstructured":"Jabeen Z, Irshad F, Hussain N, Han Y, ZhangG (2022) NHX-type Na+\/H+ antiporter gene expression under different salt levels and allelic diversity of HvNHX in wild and cultivated barleys. Front Genet 12:809988. https:\/\/doi.org\/10.3389\/fgene.2021.809988","journal-title":"Front Genet"},{"key":"11949_CR36","doi-asserted-by":"publisher","first-page":"229","DOI":"10.1046\/j.1365-313X.2003.01871.x","volume":"36","author":"MP Apse","year":"2003","unstructured":"Apse MP, Sottosanto JB, Blumwald E (2003) Vacuolar cation\/H homeostasis, and leaf development are altered in a T-DNA insertional mutant of AtNHX1, the Arabidopsis vacuolar Na+\/H\u2009+\u2009antiporter. Plant J 36:229\u2013239","journal-title":"Plant J"}],"container-title":["Molecular Biology Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s11033-026-11949-x.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s11033-026-11949-x","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s11033-026-11949-x.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,5,12]],"date-time":"2026-05-12T08:29:52Z","timestamp":1778574592000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s11033-026-11949-x"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,5,12]]},"references-count":36,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2026,12]]}},"alternative-id":["11949"],"URL":"https:\/\/doi.org\/10.1007\/s11033-026-11949-x","relation":{},"ISSN":["0301-4851","1573-4978"],"issn-type":[{"value":"0301-4851","type":"print"},{"value":"1573-4978","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,5,12]]},"assertion":[{"value":"2 March 2026","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"7 May 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"12 May 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare no competing interests.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}},{"value":"Not applicable.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethical approval"}},{"value":"All authors agreed to the manuscript sent for publication.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"Not applicable.","order":5,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent to participate"}}],"article-number":"747"}}