{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,14]],"date-time":"2026-09-14T13:09:00Z","timestamp":1789391340421,"version":"build-2803163510"},"reference-count":59,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,9,14]],"date-time":"2026-09-14T00:00:00Z","timestamp":1789344000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Plant Sci."],"abstract":"<jats:p>\n                    Drought severely limits wheat productivity, highlighting the need for tolerant genetic resources. Targeted allele mining and high-throughput phenotyping (HTP) were combined to assess natural variation in\n                    <jats:italic>TdBCAT<\/jats:italic>\n                    genes and drought response in 38 durum wheat genotypes. A cost-effective strategy integrating next-generation sequencing and Kompetitive Allele Specific PCR (KASP) genotyping identified 27 single nucleotide polymorphisms (SNPs) and 10 haplotypes across\n                    <jats:italic>TdBCAT-A<\/jats:italic>\n                    and\n                    <jats:italic>TdBCAT-B<\/jats:italic>\n                    , mainly in promoter regions. Under controlled drought, RGB imaging and the digital biovolume ratio (DBR) classified SSD 195, SSD 269, SSD 322, SSD 343, SSD 409, SSD 416, SSD 441, and Svevo as the best-performing genotypes, with SSD 195 and SSD 441 showing the greatest stability. Integrating genotypic and phenotypic data identified\n                    <jats:italic>TdBCAT<\/jats:italic>\n                    haplotype combinations associated with improved drought performance and valuable landraces for pre-breeding. This integrated approach efficiently characterized haplotypic diversity and identified drought-adaptive germplasm, providing valuable resources for pre-breeding and the sustainable selection of durum wheat under water-limited conditions.\n                  <\/jats:p>","DOI":"10.3389\/fpls.2026.1826616","type":"journal-article","created":{"date-parts":[[2026,9,14]],"date-time":"2026-09-14T12:53:06Z","timestamp":1789390386000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Uncovering natural variation in TdBCAT genes and phenotypic variation under drought in durum wheat"],"prefix":"10.3389","volume":"17","author":[{"given":"Valentina","family":"Buffagni","sequence":"first","affiliation":[{"name":"Department of Chemistry, Life Sciences and Environmental Sustainability, University of Parma","place":["Parma, Italy"]},{"name":"John Innes Centre","place":["Norwich, United Kingdom"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Donatella","family":"Danzi","sequence":"additional","affiliation":[{"name":"Institute of Biosciences and Bioresources (IBBR)-Consiglio Nazionale delle Ricerche (CNR)","place":["Bari, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ricardo Humberto","family":"Ramirez-Gonzalez","sequence":"additional","affiliation":[{"name":"John Innes Centre","place":["Norwich, United Kingdom"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cristobal","family":"Uauy","sequence":"additional","affiliation":[{"name":"John Innes Centre","place":["Norwich, United Kingdom"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mariolina","family":"Gull\u00ec","sequence":"additional","affiliation":[{"name":"Department of Chemistry, Life Sciences and Environmental Sustainability, University of Parma","place":["Parma, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"James","family":"Simmonds","sequence":"additional","affiliation":[{"name":"John Innes Centre","place":["Norwich, United Kingdom"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Angelo","family":"Petrozza","sequence":"additional","affiliation":[{"name":"ALSIA Centro Ricerche Metapontum Agrobios","place":["Metaponto, MT, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Francesco","family":"Cellini","sequence":"additional","affiliation":[{"name":"ALSIA Centro Ricerche Metapontum Agrobios","place":["Metaponto, MT, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stephan","family":"Summerer","sequence":"additional","affiliation":[{"name":"ALSIA Centro Ricerche Metapontum Agrobios","place":["Metaponto, MT, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Domenico","family":"Pignone","sequence":"additional","affiliation":[{"name":"Institute of Biosciences and Bioresources (IBBR)-Consiglio Nazionale delle Ricerche (CNR)","place":["Bari, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michela","family":"Janni","sequence":"additional","affiliation":[{"name":"Institute of Biosciences and Bioresources (IBBR)-Consiglio Nazionale delle Ricerche (CNR)","place":["Bari, Italy"]},{"name":"Institute of Materials for Electronics and Magnetism IMEM-Consiglio Nazionale delle Ricerche (CNR)","place":["Parma, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nelson","family":"Marmiroli","sequence":"additional","affiliation":[{"name":"Department of Chemistry, Life Sciences and Environmental Sustainability, University of Parma","place":["Parma, Italy"]},{"name":"CINSA, Interuniversity Consortium for Environmental Sciences","place":["Parma, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,9,14]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","DOI":"10.1016\/j.heliyon.2024.e31734","article-title":"Identifying plant traits to increase wheat yield under irrigated conditions","volume":"10","author":"Abidi","year":"2024","journal-title":"Heliyon"},{"key":"B2","doi-asserted-by":"publisher","first-page":"349","DOI":"10.1186\/s12864-020-6756-0","article-title":"Optimized breeding strategies to harness genetic resources with different performance levels","volume":"21","author":"Allier","year":"2020","journal-title":"BMC Genomics"},{"key":"B3","doi-asserted-by":"publisher","DOI":"10.20900\/cbgg20190004","article-title":"Genetic gain for yield and allelic diversity over 35 years of durum wheat breeding at ICARDA","volume":"1","author":"Bassi","year":"2019","journal-title":"Crop Breeding Genet. Genomics"},{"key":"B4","doi-asserted-by":"publisher","first-page":"484","DOI":"10.1186\/s12870-021-03272-3","article-title":"Association mapping of drought tolerance and agronomic traits in rice (Oryza sativa L.) landraces","volume":"21","author":"Beena","year":"2021","journal-title":"BMC Plant Biol."},{"key":"B5","doi-asserted-by":"publisher","DOI":"10.3389\/fgene.2016.00221","article-title":"Genomic selection in the era of next generation sequencing for complex traits in plant breeding","volume":"7","author":"Bhat","year":"2016","journal-title":"Front. Genet."},{"key":"B6","doi-asserted-by":"publisher","first-page":"248","DOI":"10.1016\/j.jbiotec.2020.11.010","article-title":"Harnessing high-throughput phenotyping and genotyping for enhanced drought tolerance in crop plants","volume":"324","author":"Bhat","year":"2020","journal-title":"J. Biotechnol."},{"key":"B7","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1007\/978-3-030-34163-3_4","article-title":"Contribution of genetic resources to grain storage protein composition and wheat quality","volume-title":"Wheat Quality for Improving Processing and Human Health","author":"Branlard","year":"2020"},{"key":"B8","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2020.00891","article-title":"Shaping durum wheat for the future: Gene expression analyses and metabolites profiling support the contribution of BCAT genes to drought stress response","volume":"11","author":"Buffagni","year":"2020","journal-title":"Front. Plant Sci."},{"key":"B9","doi-asserted-by":"publisher","first-page":"823","DOI":"10.1038\/s41586-024-07682-9","article-title":"Harnessing landrace diversity empowers wheat breeding","volume":"632","author":"Cheng","year":"2024","journal-title":"Nature"},{"key":"B10","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2018.00893","article-title":"Comparative aerial and ground based high throughput phenotyping for the genetic dissection of NDVI as a proxy for drought adaptive traits in durum wheat","volume":"9","author":"Condorelli","year":"2018","journal-title":"Front. Plant Sci."},{"key":"B11","doi-asserted-by":"publisher","first-page":"48","DOI":"10.1111\/eva.12773","article-title":"Linking genotype to phenotype to identify genetic variation relating to host susceptibility in the mountain pine beetle system","volume":"13","author":"Cullingham","year":"2020","journal-title":"Evol. Appl."},{"key":"B12","first-page":"1","article-title":"Wheat in the world","volume-title":"Bread Wheat Improvement and Production","author":"Curtis","year":"2002"},{"key":"B13","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2019.00015","article-title":"Can high throughput phenotyping help food security in the Mediterranean area","volume":"10","author":"Danzi","year":"2019","journal-title":"Front. Plant Sci."},{"key":"B14","doi-asserted-by":"publisher","DOI":"10.3390\/agriculture12040537","article-title":"The use of near-infrared imaging (NIR) as a fast non-destructive screening tool to identify drought-tolerant wheat genotypes","volume":"12","author":"Danzi","year":"2022","journal-title":"Agriculture"},{"key":"B15","doi-asserted-by":"publisher","first-page":"372","DOI":"10.1111\/nph.19619","article-title":"Back to the future for drought tolerance","volume":"242","author":"Guadarrama-Escobar","year":"2024","journal-title":"New Phytol."},{"key":"B16","doi-asserted-by":"publisher","first-page":"688","DOI":"10.1016\/j.cj.2020.04.005","article-title":"Harness the power of genomic selection and the potential of germplasm in crop breeding for global food security in the era with rapid climate change","volume":"8","author":"He","year":"2020","journal-title":"Crop J."},{"key":"B17","doi-asserted-by":"publisher","first-page":"7921","DOI":"10.1038\/s41598-022-11423-1","article-title":"Six decades of warming and drought in the world\u2019s top wheat-producing countries offset the benefits of rising CO2 to yield","volume":"12","author":"Helman","year":"2022","journal-title":"Sci. Rep."},{"key":"B18","doi-asserted-by":"publisher","first-page":"16394","DOI":"10.1038\/s41598-017-16006-z","article-title":"Genotyping-by-sequencing derived high-density linkage map and its application to QTL mapping of flag leaf traits in bread wheat","volume":"7","author":"Hussain","year":"2017","journal-title":"Sci. Rep."},{"key":"B19","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2021.766803","article-title":"Genetic variation in drought-tolerance traits and their relationships to growth in Pinus radiata D. Don under water stress","volume":"12","author":"Ismael","year":"2022","journal-title":"Front. Plant Sci."},{"key":"B20","doi-asserted-by":"publisher","first-page":"16929","DOI":"10.1038\/s41598-018-35251-4","article-title":"Gene-ecology of durum wheat HMW glutenin reflects their diffusion from the center of origin","volume":"8","author":"Janni","year":"2018","journal-title":"Sci. Rep."},{"key":"B21","doi-asserted-by":"publisher","first-page":"1","DOI":"10.34133\/2019\/6168209","article-title":"In vivo phenotyping for the early detection of drought stress in tomato","volume":"2019","author":"Janni","year":"2019","journal-title":"Plant Phenomics"},{"key":"B22","doi-asserted-by":"publisher","first-page":"3780","DOI":"10.1093\/jxb\/eraa034","article-title":"Molecular and genetic bases of heat stress responses in crop plants and breeding for increased resilience and productivity","volume":"71","author":"Janni","year":"2020","journal-title":"J. Exp. Bot"},{"key":"B23","doi-asserted-by":"publisher","first-page":"5085","DOI":"10.1093\/jxb\/erac286","article-title":"Plant phenotyping for a sustainable future","volume":"73","author":"Janni","year":"2022","journal-title":"J. Exp. Bot."},{"key":"B24","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2017.01277","article-title":"Genetic diversity within a global panel of durum wheat (Triticum durum) landraces and modern germplasm reveals the history of alleles exchange","volume":"8","author":"Kabbaj","year":"2017","journal-title":"Front. Plant Sci."},{"key":"B25","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2020.01149","article-title":"Recent progress in germplasm evaluation and gene mapping to enable breeding of drought-tolerant wheat","volume":"11","author":"Khadka","year":"2020","journal-title":"Front. Plant Sci."},{"key":"B26","doi-asserted-by":"publisher","first-page":"509","DOI":"10.1093\/jxb\/ert393","article-title":"Structural\u2013functional dissection and characterization of yield-contributing traits originating from a group 7 chromosome of the wheatgrass species Thinopyrum ponticum after transfer into durum wheat","volume":"65","author":"Kuzmanovi\u0107","year":"2014","journal-title":"J. Exp. Bot."},{"key":"B27","doi-asserted-by":"publisher","first-page":"486","DOI":"10.3390\/agronomy10040486","article-title":"Engineered durum wheat germplasm with multiple alien introgressions: Agronomic and quality performance","volume":"10","author":"Kuzmanovi\u0107","year":"2020","journal-title":"Agronomy"},{"key":"B28","doi-asserted-by":"publisher","first-page":"3477","DOI":"10.1093\/jxb\/erv122","article-title":"Exploiting genetic diversity from landraces in wheat breeding for adaptation to climate change","volume":"66","author":"Lopes","year":"2015","journal-title":"J. Exp. Bot."},{"key":"B29","doi-asserted-by":"publisher","first-page":"885","DOI":"10.1038\/s41588-019-0381-3","article-title":"Durum wheat genome highlights past domestication signatures and future improvement targets","volume":"51","author":"Maccaferri","year":"2019","journal-title":"Nat. Genet."},{"key":"B30","doi-asserted-by":"publisher","first-page":"409","DOI":"10.1093\/jxb\/erq287","article-title":"Association mapping in durum wheat grown across a broad range of water regimes","volume":"62","author":"Maccaferri","year":"2011","journal-title":"J. Exp. Bot."},{"key":"B31","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2020.569905","article-title":"The global durum wheat panel (GDP): an international platform to identify and exchange beneficial alleles","volume":"11","author":"Mazzucotelli","year":"2020","journal-title":"Front. Plant Sci."},{"key":"B32","doi-asserted-by":"publisher","first-page":"54","DOI":"10.3390\/crops1020007","article-title":"Diversity and adaptation of currently grown wheat landraces and modern germplasm in Afghanistan, Iran, and Turkey","volume":"1","author":"Morgounov","year":"2021","journal-title":"Crops"},{"key":"B33","doi-asserted-by":"publisher","DOI":"10.1016\/j.fcr.2021.108371","article-title":"A European perspective on opportunities and demands for field-based crop phenotyping","volume":"276","author":"Morisse","year":"2022","journal-title":"Field Crops Res."},{"key":"B34","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/s10681-011-0588-6","article-title":"Can Mediterranean durum wheat landraces contribute to improved grain quality attributes in modern cultivars","volume":"185","author":"Nazco","year":"2012","journal-title":"Euphytica"},{"key":"B35","doi-asserted-by":"publisher","DOI":"10.1016\/j.envexpbot.2024.105986","article-title":"From landraces to haplotypes, exploiting a genomic and phenomic approach to identify heat tolerant genotypes within durum wheat landraces","volume":"228","author":"Palermo","year":"2024","journal-title":"Environ. Exp. Bot."},{"key":"B36","doi-asserted-by":"publisher","DOI":"10.1016\/j.plantsci.2025.112930","article-title":"Durum wheat germplasm response to high temperatures, the role of small HSP26 in the defense response","volume":"364","author":"Palermo","year":"2026","journal-title":"Plant Sci."},{"key":"B37","doi-asserted-by":"publisher","first-page":"185","DOI":"10.1016\/j.scienta.2014.05.023","article-title":"Physiological responses to Megafol\u00ae treatments in tomato plants under drought stress: A phenomic and molecular approach","volume":"174","author":"Petrozza","year":"2014","journal-title":"Sci. Hortic."},{"key":"B38","doi-asserted-by":"publisher","first-page":"1029","DOI":"10.1007\/s10722-014-0206-2","article-title":"Single seed descent: a tool to exploit durum wheat (Triticum durum Desf.) genetic resources","volume":"62","author":"Pignone","year":"2015","journal-title":"Genet. Resour. Crop Evol."},{"key":"B39","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1079\/9781780646138.0047","article-title":"Capturing wild relative and landrace diversity for crop improvement using a new selection tool to exploit genetic resources in durum wheat","volume-title":"Enhancing Crop Genepool Use: Capturing Wild Relative and Landrace Diversity for Crop Improvement","author":"Pignone","year":"2016"},{"key":"B40","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1007\/978-3-642-37045-8_2","article-title":"Conservation, evaluation, and utilization of biodiversity","volume-title":"Genomics and Breeding for Climate-Resilient Crops: Vol. 1 Concepts and Strategies","author":"Pignone","year":"2013"},{"key":"B41","doi-asserted-by":"publisher","first-page":"781","DOI":"10.1111\/jipb.12198","article-title":"Digital imaging approaches for phenotyping whole plant nitrogen and phosphorus response in Brachypodium distachyon","volume":"56","author":"Poir\u00e9","year":"2014","journal-title":"Journal of Integrative Plant Biology"},{"key":"B42","doi-asserted-by":"publisher","DOI":"10.3390\/plants8020034","article-title":"Impact of climate change on crops adaptation and strategies to tackle its outcome: A review","volume":"8","author":"Raza","year":"2019","journal-title":"Plants"},{"key":"B43","doi-asserted-by":"publisher","DOI":"10.3390\/metabo12060511","article-title":"Advances in metabolomics-driven diagnostic breeding and crop improvement","volume":"12","author":"Razzaq","year":"2022","journal-title":"Metabolites"},{"key":"B44","doi-asserted-by":"publisher","DOI":"10.3390\/ijms20133137","article-title":"Drought stress tolerance in wheat and barley: Advances in physiology, breeding and genetics research","volume":"20","author":"Sallam","year":"2019","journal-title":"Int. J. Mol. Sci."},{"key":"B45","doi-asserted-by":"publisher","first-page":"4572","DOI":"10.1038\/s41467-020-18404-w","article-title":"Diversity analysis of 80,000 wheat accessions reveals consequences and opportunities of selection footprints","volume":"11","author":"Sansaloni","year":"2020","journal-title":"Nat. Commun."},{"key":"B46","doi-asserted-by":"publisher","DOI":"10.3390\/agronomy13030891","article-title":"Resilience to terminal drought, heat, and their combination stress in wheat genotypes","volume":"13","author":"Sareen","year":"2023","journal-title":"Agronomy"},{"key":"B47","doi-asserted-by":"publisher","DOI":"10.3390\/plants10061071","article-title":"Profiling of plant growth-promoting metabolites by phosphate-solubilizing bacteria in maize rhizosphere","volume":"10","author":"Shen","year":"2021","journal-title":"Plants"},{"key":"B48","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2019.00394","article-title":"High-throughput phenotyping enabled genetic dissection of crop lodging in wheat","volume":"10","author":"Singh","year":"2019","journal-title":"Front. Plant Sci."},{"key":"B49","doi-asserted-by":"publisher","first-page":"1098","DOI":"10.20546\/ijcmas.2020.905.121","article-title":"Potential of allele mining for improving drought tolerance in crops","volume":"9","author":"Sinha","year":"2020","journal-title":"Int. J. Curr. Microbiol. Appl. Sci."},{"key":"B50","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0206682","article-title":"Genetic basis of drought tolerance during seed germination in barley","volume":"13","author":"Thabet","year":"2018","journal-title":"PloS One"},{"key":"B51","volume-title":"World Population Prospects 2024: Summary of Results","year":"2024"},{"key":"B52","doi-asserted-by":"publisher","first-page":"631","DOI":"10.1016\/j.tplants.2021.03.010","article-title":"Designing future crops: Genomics-assisted breeding comes of age","volume":"26","author":"Varshney","year":"2021","journal-title":"Trends Plant Sci."},{"key":"B53","doi-asserted-by":"publisher","DOI":"10.1016\/j.pbi.2018.03.007","article-title":"Can genomics deliver climate-change ready crops","volume":"45","author":"Varshney","year":"2018","journal-title":"Curr. Opin. Plant Biol"},{"key":"B54","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2022.896985","article-title":"How does the environment affect wheat yield and protein content response to drought? A meta-analysis","volume":"13","author":"Wan","year":"2022","journal-title":"Front. Plant Sci."},{"key":"B55","doi-asserted-by":"publisher","DOI":"10.1155\/2013\/489583","article-title":"Establishing an efficient way to utilize the drought resistance germplasm population in wheat","volume":"2013","author":"Wang","year":"2013","journal-title":"Sci. World J."},{"key":"B56","doi-asserted-by":"publisher","DOI":"10.3390\/ijms221910265","article-title":"Exploitation of drought tolerance-related genes for crop improvement","volume":"22","author":"Wang","year":"2021","journal-title":"Int. J. Mol. Sci."},{"key":"B57","volume-title":"Strategic Research Agenda: Global Vision for Sustainable Wheat Production","year":"2023"},{"key":"B58","doi-asserted-by":"publisher","first-page":"496","DOI":"10.1111\/jipb.13451","article-title":"The battle of crops against drought: Genetic dissection and improvement","volume":"65","author":"Yang","year":"2023","journal-title":"J. Integr. Plant Biol."},{"key":"B59","doi-asserted-by":"publisher","DOI":"10.3390\/ijms23136929","article-title":"Reproductive-stage heat stress in cereals: Impact, plant responses and strategies for tolerance improvement","volume":"23","author":"Zenda","year":"2022","journal-title":"Int. J. Mol. Sci."}],"container-title":["Frontiers in Plant Science"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2026.1826616\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,14]],"date-time":"2026-09-14T12:53:08Z","timestamp":1789390388000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2026.1826616\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,14]]},"references-count":59,"alternative-id":["10.3389\/fpls.2026.1826616"],"URL":"https:\/\/doi.org\/10.3389\/fpls.2026.1826616","relation":{},"ISSN":["1664-462X"],"issn-type":[{"value":"1664-462X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,14]]},"article-number":"1826616"}}