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Natl. Acad. Sci. U.S.A."],"published-print":{"date-parts":[[2025,11,25]]},"abstract":"\n The conventional assumption is that stomatal conductance (\n \n \n \n g<\/mml:mi>\n s<\/mml:mi>\n <\/mml:msub>\n <\/mml:math>\n <\/jats:inline-formula>\n ) dominates the regulation of water and carbon dioxide fluxes between leaves and the atmosphere. Here, a nanoreporter of water status at the mesophyll cell surface and local xylem within intact maize leaves documents significant undersaturation of water vapor in the outside-xylem zone (OXZ) and a large loss of conductance of this zone (\n \n \n \n g<\/mml:mi>\n \n oxz<\/mml:mi>\n <\/mml:mrow>\n <\/mml:msub>\n <\/mml:math>\n <\/jats:inline-formula>\n ) at moderate xylem water stress, without stomatal closure or turgor loss. The ratio of the resistances\n \n \n \n \n (<\/mml:mo>\n 1<\/mml:mn>\n \/<\/mml:mo>\n \n g<\/mml:mi>\n \n oxz<\/mml:mi>\n <\/mml:mrow>\n <\/mml:msub>\n )<\/mml:mo>\n <\/mml:mrow>\n \/<\/mml:mo>\n \n (<\/mml:mo>\n 1<\/mml:mn>\n \/<\/mml:mo>\n \n g<\/mml:mi>\n s<\/mml:mi>\n <\/mml:msub>\n )<\/mml:mo>\n <\/mml:mrow>\n <\/mml:mrow>\n <\/mml:math>\n <\/jats:inline-formula>\n serves as a predictive phenotype of undersaturation, nonstomatal regulation of transpiration, errors in standard gas exchange analysis, and an increase of intrinsic water use efficiency (\n \n \n \n iWUE<\/mml:mi>\n <\/mml:mrow>\n <\/mml:math>\n <\/jats:inline-formula>\n ). Cell-scale access to water status reveals symplasmic-apoplasmic disequilibrium and informs a biophysical model that can explain experimental observations quantitatively based on localization of variable conductance to the plasma membrane. This work opens paths of inquiry into the molecular basis and functional consequences of nonstomatal regulation of transpiration.\n <\/jats:p>","DOI":"10.1073\/pnas.2504862122","type":"journal-article","created":{"date-parts":[[2025,11,19]],"date-time":"2025-11-19T17:23:32Z","timestamp":1763573012000},"update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":11,"title":["Loss of conductance between mesophyll symplasm and intercellular air spaces explains nonstomatal control of transpiration"],"prefix":"10.1073","volume":"122","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-2852-5285","authenticated-orcid":false,"given":"Piyush","family":"Jain","sequence":"first","affiliation":[{"name":"Sibley School of Mechanical and Aerospace Engineering, Cornell University","place":["Ithaca"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1015-5316","authenticated-orcid":false,"given":"Sabyasachi","family":"Sen","sequence":"additional","affiliation":[{"name":"Sibley School of Mechanical and Aerospace Engineering, Cornell University","place":["Ithaca"]},{"name":"Kavli Institute at Cornell for Nanoscale Science, Cornell University","place":["Ithaca"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2527-6033","authenticated-orcid":false,"given":"Fulton E.","family":"Rockwell","sequence":"additional","affiliation":[{"name":"Department of Organismic and Evolutionary Biology, Harvard University","place":["Cambridge"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0004-1824-3366","authenticated-orcid":false,"suffix":"III","given":"Robert J.","family":"Twohey","sequence":"additional","affiliation":[{"name":"Department of Crop Sciences, University of Illinois Urbana-Champaign","place":["Urbana"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0937-2324","authenticated-orcid":false,"given":"Annika E.","family":"Huber","sequence":"additional","affiliation":[{"name":"Smith School of Chemical and Biomolecular Engineering, Cornell University","place":["Ithaca"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9201-4626","authenticated-orcid":false,"given":"Sahil A.","family":"Desai","sequence":"additional","affiliation":[{"name":"Smith School of Chemical and Biomolecular Engineering, Cornell University","place":["Ithaca"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-4765-5142","authenticated-orcid":false,"given":"I-Feng","family":"Wu","sequence":"additional","affiliation":[{"name":"Smith School of Chemical and Biomolecular Engineering, Cornell University","place":["Ithaca"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8338-7786","authenticated-orcid":false,"given":"Tom","family":"De Swaef","sequence":"additional","affiliation":[{"name":"Plant Sciences Unit, Flanders Research Institute for Agriculture, Fisheries and Food","place":["Merelbeke, Belgium"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7664-5217","authenticated-orcid":false,"given":"Mehmet M.","family":"Ilman","sequence":"additional","affiliation":[{"name":"Sibley School of Mechanical and Aerospace Engineering, Cornell University","place":["Ithaca"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8571-3143","authenticated-orcid":false,"given":"Anthony J.","family":"Studer","sequence":"additional","affiliation":[{"name":"Department of Crop Sciences, University of Illinois Urbana-Champaign","place":["Urbana"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3325-5395","authenticated-orcid":false,"given":"N. Michele","family":"Holbrook","sequence":"additional","affiliation":[{"name":"Department of Organismic and Evolutionary Biology, Harvard University","place":["Cambridge"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8145-9977","authenticated-orcid":false,"given":"Abraham D.","family":"Stroock","sequence":"additional","affiliation":[{"name":"Kavli Institute at Cornell for Nanoscale Science, Cornell University","place":["Ithaca"]},{"name":"Smith School of Chemical and Biomolecular Engineering, Cornell University","place":["Ithaca"]},{"name":"School of Integrative Plant Science, Cornell University","place":["Ithaca"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"341","published-online":{"date-parts":[[2025,11,19]]},"reference":[{"key":"e_1_3_4_1_2","volume-title":"Physicochemical and Environmental Plant Physiology","author":"Nobel P. S.","year":"2009","unstructured":"P. S. 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J. Twohey Investigating the role of stomatal dynamics on agronomic traits using a slac1-2 Zea mays mutant. bioXriv [Preprint] (2025). https:\/\/doi.org\/10.1101\/2025.01.21.634166 (Accessed 27 January 2025).","DOI":"10.1101\/2025.01.21.634166"},{"key":"e_1_3_4_36_2","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/erm270"},{"key":"e_1_3_4_37_2","doi-asserted-by":"publisher","DOI":"10.1104\/pp.115.3.1089"},{"key":"e_1_3_4_38_2","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-313X.2011.04576.x"},{"key":"e_1_3_4_39_2","unstructured":"P. Jain . Data for \u201cLoss of conductance between mesophyll symplasm and intercellular air spaces explains nonstomatal control of transpiration.\u201d GitHub. https:\/\/github.com\/sabyasachi-sen\/Isothermal-model-Non-stomatal-control-of-transpiration. 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