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While iWUE is generally higher in C\n                    <jats:sub>4<\/jats:sub>\n                    than in C\n                    <jats:sub>3<\/jats:sub>\n                    plants, future climate scenarios, marked by increasing atmospheric CO\n                    <jats:sub>2<\/jats:sub>\n                    , temperatures, and aridity, present ongoing pressures on crop productivity in C\n                    <jats:sub>4<\/jats:sub>\n                    \u2010dominated environments. This review explores the physiological, anatomical, and environmental factors controlling iWUE in C\n                    <jats:sub>4<\/jats:sub>\n                    plants, with a focus on balancing productivity and resilience under climate change. Using process models of photosynthesis and stomatal behaviour, we assess how iWUE varies with photosynthetic limitations, highlight the central role of stomata in controlling iWUE, and predict the physiological adjustments needed to optimize carbon gain per unit water loss. Recent evidence points to leaf width as a useful trait influencing leaf energy balance, temperature, and transpiration. We propose integrated physiological and breeding strategies to optimize crop iWUE and yield under climate stress, supported by emerging sensing and selection technologies.\n                  <\/jats:p>","DOI":"10.1111\/nph.70660","type":"journal-article","created":{"date-parts":[[2025,10,23]],"date-time":"2025-10-23T11:32:53Z","timestamp":1761219173000},"page":"2656-2673","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":23,"title":["Opportunities for improving intrinsic water use efficiency in C\n                    <sub>4<\/sub>\n                    plants under climate change"],"prefix":"10.1111","volume":"248","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1341-0741","authenticated-orcid":false,"given":"Oula","family":"Ghannoum","sequence":"first","affiliation":[{"name":"Hawkesbury Institute for the Environment Western Sydney University  Locked Bag 1797 Penrith NSW 2751 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Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2025,10,23]]},"reference":[{"key":"e_1_2_11_2_1","doi-asserted-by":"publisher","DOI":"10.1111\/nph.19933"},{"key":"e_1_2_11_3_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/erae418"},{"key":"e_1_2_11_4_1","doi-asserted-by":"publisher","DOI":"10.1111\/pce.14592"},{"key":"e_1_2_11_5_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/erad063"},{"key":"e_1_2_11_6_1","doi-asserted-by":"publisher","DOI":"10.1111\/tpj.16346"},{"key":"e_1_2_11_7_1","doi-asserted-by":"publisher","DOI":"10.1111\/ele.13516"},{"key":"e_1_2_11_8_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.tplants.2013.09.008"},{"key":"e_1_2_11_9_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/erab097"},{"key":"e_1_2_11_10_1","doi-asserted-by":"publisher","DOI":"10.1093\/plphys\/kiad355"},{"key":"e_1_2_11_11_1","doi-asserted-by":"publisher","DOI":"10.1016\/S1360-1385(03)00136-5"},{"key":"e_1_2_11_12_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.jplph.2007.09.003"},{"key":"e_1_2_11_13_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00122-018-3193-4"},{"key":"e_1_2_11_14_1","doi-asserted-by":"publisher","DOI":"10.1111\/nph.20341"},{"key":"e_1_2_11_15_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-021-03370-0"},{"key":"e_1_2_11_16_1","doi-asserted-by":"publisher","DOI":"10.1016\/0168-1923(85)90070-X"},{"key":"e_1_2_11_17_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/erae389"},{"key":"e_1_2_11_18_1","doi-asserted-by":"publisher","DOI":"10.1111\/nph.14063"},{"key":"e_1_2_11_19_1","doi-asserted-by":"publisher","DOI":"10.1111\/nph.20167"},{"key":"e_1_2_11_20_1","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2019.00225"},{"key":"e_1_2_11_21_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.fcr.2009.03.009"},{"key":"e_1_2_11_22_1","doi-asserted-by":"publisher","DOI":"10.1111\/pce.12817"},{"key":"e_1_2_11_23_1","doi-asserted-by":"publisher","DOI":"10.1016\/B978-012614440-6\/50015-X"},{"key":"e_1_2_11_24_1","doi-asserted-by":"publisher","DOI":"10.1104\/pp.91.4.1543"},{"key":"e_1_2_11_25_1","doi-asserted-by":"publisher","DOI":"10.1104\/pp.16.01772"},{"key":"e_1_2_11_26_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41559-018-0643-y"},{"key":"e_1_2_11_27_1","doi-asserted-by":"publisher","DOI":"10.1104\/pp.100.1.191"},{"key":"e_1_2_11_28_1","doi-asserted-by":"publisher","DOI":"10.1071\/9780643103405"},{"key":"e_1_2_11_29_1","doi-asserted-by":"publisher","DOI":"10.1016\/B978-012614440-6\/50007-0"},{"key":"e_1_2_11_30_1","doi-asserted-by":"publisher","DOI":"10.1023\/A:1025830019591"},{"key":"e_1_2_11_31_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.pbi.2016.04.003"},{"key":"e_1_2_11_32_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/eru127"},{"key":"e_1_2_11_33_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-3040.2003.01157.x"},{"key":"e_1_2_11_34_1","doi-asserted-by":"publisher","DOI":"10.1111\/nph.18704"},{"key":"e_1_2_11_35_1","doi-asserted-by":"publisher","DOI":"10.1111\/nph.15344"},{"key":"e_1_2_11_36_1","doi-asserted-by":"publisher","DOI":"10.1111\/nph.15920"},{"key":"e_1_2_11_37_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/erq071"},{"key":"e_1_2_11_38_1","doi-asserted-by":"publisher","DOI":"10.1071\/FP08216"},{"key":"e_1_2_11_39_1","doi-asserted-by":"publisher","DOI":"10.1111\/nph.12423"},{"key":"e_1_2_11_40_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/erab039"},{"key":"e_1_2_11_41_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature09487"},{"key":"e_1_2_11_42_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrg3503"},{"key":"e_1_2_11_43_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1216777110"},{"key":"e_1_2_11_44_1","doi-asserted-by":"publisher","DOI":"10.1093\/molbev\/msn178"},{"key":"e_1_2_11_45_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/eraa002"},{"key":"e_1_2_11_46_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/ern001"},{"key":"e_1_2_11_47_1","doi-asserted-by":"publisher","DOI":"10.1104\/pp.002170"},{"key":"e_1_2_11_48_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00442-009-1364-3"},{"key":"e_1_2_11_49_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/erab464"},{"key":"e_1_2_11_50_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.tplants.2017.08.011"},{"key":"e_1_2_11_51_1","doi-asserted-by":"publisher","DOI":"10.1105\/tpc.16.00155"},{"key":"e_1_2_11_52_1","doi-asserted-by":"publisher","DOI":"10.1038\/s42003-021-01772-4"},{"key":"e_1_2_11_53_1","doi-asserted-by":"publisher","DOI":"10.5194\/bg-14-4435-2017"},{"key":"e_1_2_11_54_1","doi-asserted-by":"publisher","DOI":"10.1105\/tpc.112.100115"},{"key":"e_1_2_11_55_1","doi-asserted-by":"publisher","DOI":"10.1006\/anbo.1994.1029"},{"key":"e_1_2_11_56_1","doi-asserted-by":"publisher","DOI":"10.1016\/B978-012614440-6\/50006-9"},{"key":"e_1_2_11_57_1","doi-asserted-by":"publisher","DOI":"10.1111\/nph.12586"},{"key":"e_1_2_11_58_1","doi-asserted-by":"publisher","DOI":"10.1093\/j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