{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,26]],"date-time":"2026-09-26T09:45:29Z","timestamp":1790415929826,"version":"4.1.0"},"reference-count":82,"publisher":"Oxford University Press (OUP)","issue":"2","license":[{"start":{"date-parts":[[2024,3,20]],"date-time":"2024-03-20T00:00:00Z","timestamp":1710892800000},"content-version":"vor","delay-in-days":48,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2024,2,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n               <jats:p>Photosynthesis is co-limited by multiple factors depending on the plant and its environment. These include biochemical rate limitations, internal and external water potentials, temperature, irradiance and carbon dioxide ( CO2). Amphistomatous leaves have stomata on both abaxial and adaxial leaf surfaces. This feature is considered an adaptation to alleviate CO2 diffusion limitations in productive environments as the diffusion path length from stomate to chloroplast is effectively halved in amphistomatous leaves. Plants may also reduce CO2 limitations through other aspects of optimal stomatal anatomy: stomatal density, distribution, patterning and size. Some studies have demonstrated that stomata are overdispersed compared to a random distribution on a single leaf surface; however, despite their prevalence in nature and near ubiquity among crop species, much less is known about stomatal anatomy in amphistomatous leaves, especially the coordination between leaf surfaces. Here, we use novel spatial statistics based on simulations and photosynthesis modelling to test hypotheses about how amphistomatous plants may optimize CO2 diffusion in the model angiosperm Arabidopsis thaliana grown in different light environments. We find that (i) stomata are overdispersed, but not ideally dispersed, on both leaf surfaces across all light treatments; (ii) the patterning of stomata on abaxial and adaxial leaf surfaces is independent and (iii) the theoretical improvements to photosynthesis from abaxial\u2013adaxial stomatal coordination are miniscule (\u226a1%) across the range of feasible parameter space. However, we also find that (iv) stomatal size is correlated with the mesophyll volume that it supplies with CO2, suggesting that plants may optimize CO2 diffusion limitations through alternative pathways other than ideal, uniform stomatal spacing. We discuss the developmental, physical and evolutionary constraints that may prohibit plants from reaching this theoretical adaptive peak of uniform stomatal spacing and inter-surface stomatal coordination. These findings contribute to our understanding of variation in the anatomy of amphistomatous leaves.<\/jats:p>","DOI":"10.1093\/aobpla\/plae015","type":"journal-article","created":{"date-parts":[[2024,3,18]],"date-time":"2024-03-18T07:06:16Z","timestamp":1710745576000},"source":"Crossref","is-referenced-by-count":12,"title":["Does stomatal patterning in amphistomatous leaves minimize the CO2 diffusion path length within leaves?"],"prefix":"10.1093","volume":"16","author":[{"given":"Jacob L","family":"Watts","sequence":"first","affiliation":[{"name":"School of Life Sciences, University of Hawai\u2019i at M\u0101noa , Honolulu, HI 96822 , USA"},{"name":"Ecology and Evolutionary Biology, University of Colorado , Boulder, CO 80309 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Graham J","family":"Dow","sequence":"additional","affiliation":[{"name":"Department of Crop Science and Production Systems, NIAB , Cambridge CB3 0LE , UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Thomas N","family":"Buckley","sequence":"additional","affiliation":[{"name":"Department of Plant Sciences, University of California , Davis, CA 95616 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christopher D","family":"Muir","sequence":"additional","affiliation":[{"name":"School of Life Sciences, University of Hawai\u2019i at M\u0101noa , Honolulu, HI 96822 , USA"},{"name":"Department of Botany, University of Wisconsin, Madison , WI 53706 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2024,3,20]]},"reference":[{"key":"2024082904153874700_CIT0001","doi-asserted-by":"crossref","first-page":"1399","DOI":"10.1046\/j.0016-8025.2002.00906.x","article-title":"A three-dimensional model of CO2 transport in airspaces and mesophyll cells of a silver birch leaf","volume":"25","author":"Aalto","year":"2002","journal-title":"Plant, Cell & Environment"},{"key":"2024082904153874700_CIT0002","doi-asserted-by":"crossref","first-page":"794","DOI":"10.1007\/s004250100556","article-title":"Acclimation of Arabidopsis thaliana to the light environment: the existence of separate low light and high light responses","volume":"213","author":"Bailey","year":"2001","journal-title":"Planta"},{"key":"2024082904153874700_CIT0003","doi-asserted-by":"crossref","first-page":"1120","DOI":"10.1111\/nph.16341","article-title":"The developmental relationship between stomata and mesophyll airspace","volume":"225","author":"Baillie","year":"2020","journal-title":"New Phytologist"},{"key":"2024082904153874700_CIT0004","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1111\/j.2517-6161.1995.tb02031.x","article-title":"Controlling the false discovery rate: a practical and powerful approach to multiple testing","volume":"57","author":"Benjamini","year":"1995","journal-title":"Journal of the Royal Statistical Society. Series B (Methodological)"},{"key":"2024082904153874700_CIT0005","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1006\/dbio.1995.1058","article-title":"Arrest of stomatal initials in tradescantia is linked to the proximity of neighboring stomata and results in the arrested initials acquiring properties of epidermal cells","volume":"168","author":"Boetsch","year":"1995","journal-title":"Developmental Biology"},{"key":"2024082904153874700_CIT0006","doi-asserted-by":"crossref","first-page":"946","DOI":"10.1111\/nph.17971","article-title":"Structural organization of the spongy mesophyll","volume":"234","author":"Borsuk","year":"2022","journal-title":"New Phytologist"},{"key":"2024082904153874700_CIT0007","doi-asserted-by":"crossref","first-page":"881","DOI":"10.1111\/pce.12823","article-title":"Optimal plant water economy","volume":"40","author":"Buckley","year":"2017","journal-title":"Plant, Cell & Environment"},{"key":"2024082904153874700_CIT0008","doi-asserted-by":"crossref","DOI":"10.18637\/jss.v080.i01","article-title":"brms: an R package for Bayesian multilevel models using Stan","volume":"80","author":"B\u00fcrkner","year":"2017","journal-title":"Journal of Statistical Software"},{"key":"2024082904153874700_CIT0009","doi-asserted-by":"crossref","first-page":"395","DOI":"10.32614\/RJ-2018-017","article-title":"Advanced Bayesian multilevel modeling with the R package brms","volume":"10","author":"B\u00fcrkner","year":"2018","journal-title":"The R Journal"},{"key":"2024082904153874700_CIT0010","doi-asserted-by":"crossref","first-page":"1037","DOI":"10.1071\/FP06078","article-title":"Stomatal aperture can compensate altered stomatal density in Arabidopsis thaliana at growth light conditions","volume":"33","author":"B\u00fcssis","year":"2006","journal-title":"Functional Plant Biology"},{"key":"2024082904153874700_CIT0011","doi-asserted-by":"crossref","first-page":"1965","DOI":"10.1890\/0012-9658(2002)083[1965:SIPAIE]2.0.CO;2","article-title":"Shade-induced plasticity and its ecological significance in wild populations of Arabidopsis thaliana","volume":"83","author":"Callahan","year":"2002","journal-title":"Ecology"},{"key":"2024082904153874700_CIT0012","doi-asserted-by":"crossref","first-page":"448","DOI":"10.1111\/pce.12402","article-title":"Diffusional limitations explain the lower photosynthetic capacity of ferns as compared with angiosperms in a common garden study: photosynthetic comparison in ferns and angiosperms","volume":"38","author":"Carriqu\u00ed","year":"2015","journal-title":"Plant, Cell & Environment"},{"key":"2024082904153874700_CIT0013","doi-asserted-by":"crossref","first-page":"445","DOI":"10.2307\/1931034","article-title":"Distance to nearest neighbor as a measure of spatial relationships in populations","volume":"35","author":"Clark","year":"1954","journal-title":"Ecology"},{"key":"2024082904153874700_CIT0014","first-page":"471","article-title":"Stomatal function in relation to leaf metabolism and environment","volume":"31","author":"Cowan","year":"1977","journal-title":"Symposia of the Society for Experimental Biology"},{"key":"2024082904153874700_CIT0015","doi-asserted-by":"crossref","first-page":"1069","DOI":"10.2307\/2656643","article-title":"Stomatal patterning in angiosperms","volume":"87","author":"Croxdale","year":"2000","journal-title":"American Journal of Botany"},{"key":"2024082904153874700_CIT0016","doi-asserted-by":"crossref","first-page":"1219","DOI":"10.1111\/nph.13929","article-title":"Optimal allocation of leaf epidermal area for gas exchange","volume":"210","author":"de Boer","year":"2016","journal-title":"New Phytologist"},{"key":"2024082904153874700_CIT0017","doi-asserted-by":"crossref","first-page":"1116","DOI":"10.1038\/s41477-020-00760-6","article-title":"Optimization can provide the fundamental link between leaf photosynthesis, gas exchange and water relations","volume":"6","author":"Deans","year":"2020","journal-title":"Nature Plants"},{"key":"2024082904153874700_CIT0018","doi-asserted-by":"crossref","first-page":"1205","DOI":"10.1111\/nph.12586","article-title":"The physiological importance of developmental mechanisms that enforce proper stomatal spacing in Arabidopsis thaliana","volume":"201","author":"Dow","year":"2014","journal-title":"New Phytologist"},{"key":"2024082904153874700_CIT0019","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1111\/nph.14746","article-title":"Disruption of stomatal lineage signaling or transcriptional regulators has differential effects on mesophyll development, but maintains coordination of gas exchange","volume":"216","author":"Dow","year":"2017","journal-title":"New Phytologist"},{"key":"2024082904153874700_CIT0020","doi-asserted-by":"crossref","first-page":"1179","DOI":"10.1111\/nph.15652","article-title":"Two sides to every leaf: water and CO2 transport in hypostomatous and amphistomatous leaves","volume":"222","author":"Drake","year":"2019","journal-title":"New Phytologist"},{"key":"2024082904153874700_CIT0021","doi-asserted-by":"crossref","first-page":"1082","DOI":"10.1104\/pp.17.00223","article-title":"Excess diffuse light absorption in upper mesophyll limits CO2 drawdown and depresses photosynthesis","volume":"174","author":"Earles","year":"2017","journal-title":"Plant Physiology"},{"key":"2024082904153874700_CIT0022","doi-asserted-by":"crossref","first-page":"148","DOI":"10.1104\/pp.18.00550","article-title":"Beyond porosity: 3D leaf intercellular airspace traits that impact mesophyll conductance","volume":"178","author":"Earles","year":"2018","journal-title":"Plant Physiology"},{"key":"2024082904153874700_CIT0023","doi-asserted-by":"crossref","first-page":"2235","DOI":"10.1093\/jxb\/erp117","article-title":"Resistances along the CO2 diffusion pathway inside leaves","volume":"60","author":"Evans","year":"2009","journal-title":"Journal of Experimental Botany"},{"key":"2024082904153874700_CIT0024","author":"Evans","year":"2023"},{"key":"2024082904153874700_CIT0025","doi-asserted-by":"crossref","first-page":"78","DOI":"10.1007\/BF00386231","article-title":"A biochemical model of photosynthetic CO2 assimilation in leaves of C3 species","volume":"149","author":"Farquhar","year":"1980","journal-title":"Planta"},{"key":"2024082904153874700_CIT0026","doi-asserted-by":"crossref","first-page":"216","DOI":"10.1111\/nph.13577","article-title":"Transport efficiency through uniformity: organization of veins and stomata in angiosperm leaves","volume":"209","author":"Fiorin","year":"2016","journal-title":"New Phytologist"},{"key":"2024082904153874700_CIT0027","doi-asserted-by":"crossref","first-page":"537","DOI":"10.1098\/rstb.2011.0270","article-title":"Physiological framework for adaptation of stomata to CO2 from glacial to future concentrations","volume":"367","author":"Franks","year":"2012","journal-title":"Philosophical Transactions of the Royal Society B: Biological Sciences"},{"key":"2024082904153874700_CIT0028","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1111\/j.1469-8137.1975.tb01368.x","article-title":"The influence of light on stomatal density in the tomato","volume":"75","author":"Gay","year":"1975","journal-title":"New Phytologist"},{"key":"2024082904153874700_CIT0029","doi-asserted-by":"crossref","first-page":"2075","DOI":"10.1105\/tpc.12.11.2075","article-title":"Oriented asymmetric divisions that generate the stomatal spacing pattern in Arabidopsis are disrupted by the too many mouths mutation","volume":"12","author":"Geisler","year":"2000","journal-title":"The Plant Cell"},{"key":"2024082904153874700_CIT0030","doi-asserted-by":"crossref","first-page":"768","DOI":"10.1111\/tpj.14560","article-title":"The influence of stomatal morphology and distribution on photosynthetic gas exchange","volume":"101","author":"Harrison","year":"2020","journal-title":"The Plant Journal"},{"key":"2024082904153874700_CIT0031","article-title":"2021. Understanding airspace in leaves: 3D anatomy and directional tortuosity","author":"Harwood","year":"4079","journal-title":"Plant, Cell & Environment"},{"key":"2024082904153874700_CIT0032","author":"Hijmans","year":"2023"},{"key":"2024082904153874700_CIT0033","doi-asserted-by":"crossref","first-page":"168","DOI":"10.1016\/j.palaeo.2013.12.035","article-title":"Using fossil leaves as evidence for open vegetation","volume":"395","author":"Jordan","year":"2014","journal-title":"Palaeogeography, Palaeoclimatology, Palaeoecology"},{"key":"2024082904153874700_CIT0034","doi-asserted-by":"crossref","first-page":"608","DOI":"10.1111\/nph.13076","article-title":"Environmental adaptation in stomatal size independent of the effects of genome size","volume":"205","author":"Jordan","year":"2015","journal-title":"New Phytologist"},{"key":"2024082904153874700_CIT0035","doi-asserted-by":"crossref","first-page":"31252","DOI":"10.1038\/srep31252","article-title":"Metabolic and diffusional limitations of photosynthesis in fluctuating irradiance in Arabidopsis thaliana","volume":"6","author":"Kaiser","year":"2016","journal-title":"Scientific Reports"},{"key":"2024082904153874700_CIT0036","doi-asserted-by":"crossref","first-page":"76","DOI":"10.1007\/BF00386887","article-title":"Responses of stomata to changes in humidity","volume":"100","author":"Lange","year":"1971","journal-title":"Planta"},{"key":"2024082904153874700_CIT0037","doi-asserted-by":"crossref","first-page":"1381","DOI":"10.1093\/jxb\/50.337.1381","article-title":"Spatial and temporal variation in gas exchange over the lower surface of Phaseolus vulgaris L. primary leaves","volume":"50","author":"Lawson","year":"1999","journal-title":"Journal of Experimental Botany"},{"key":"2024082904153874700_CIT0038","doi-asserted-by":"crossref","first-page":"1387","DOI":"10.1093\/jxb\/49.325.1387","article-title":"The nature of heterogeneity in the stomatal behaviour of Phaseolus vulgaris L. primary leaves","volume":"49","author":"Lawson","year":"1998","journal-title":"Journal of Experimental Botany"},{"key":"2024082904153874700_CIT0039","doi-asserted-by":"crossref","first-page":"963","DOI":"10.1002\/j.1537-2197.1964.tb06725.x","article-title":"Diffusion resistance in leaves as related to their stomatal anatomy and micro-structure","volume":"51","author":"Lee","year":"1964","journal-title":"American Journal of Botany"},{"key":"2024082904153874700_CIT0040","doi-asserted-by":"crossref","first-page":"1015","DOI":"10.1111\/nph.13442","article-title":"Effects of stomata clustering on leaf gas exchange","volume":"207","author":"Lehmann","year":"2015","journal-title":"New Phytologist"},{"key":"2024082904153874700_CIT0041","doi-asserted-by":"crossref","first-page":"981","DOI":"10.1111\/tpj.13727","article-title":"Cell density and airspace patterning in the leaf can be manipulated to increase leaf photosynthetic capacity","volume":"92","author":"Lehmeier","year":"2017","journal-title":"The Plant Journal"},{"key":"2024082904153874700_CIT0042","author":"Lenth","year":"2023"},{"key":"2024082904153874700_CIT0043","article-title":"Scaling between stomatal size and density in forest plants","author":"Liu","year":"2021","journal-title":"Plant Biology"},{"key":"2024082904153874700_CIT0044","doi-asserted-by":"crossref","first-page":"550","DOI":"10.1104\/pp.17.00114","article-title":"Stomatal biology of CAM plants","volume":"174","author":"Males","year":"2017","journal-title":"Plant Physiology"},{"key":"2024082904153874700_CIT0045","doi-asserted-by":"crossref","first-page":"2581","DOI":"10.1093\/jxb\/erh260","article-title":"A\/Ci curve analysis across a range of woody plant species: influence of regression analysis parameters and mesophyll conductance","volume":"55","author":"Manter","year":"2004","journal-title":"Journal of Experimental Botany"},{"key":"2024082904153874700_CIT0046","doi-asserted-by":"crossref","first-page":"2008","DOI":"10.1104\/pp.16.00380","article-title":"Linking turgor with ABA biosynthesis: implications for stomatal responses to vapor pressure deficit across land plants","volume":"171","author":"McAdam","year":"2016","journal-title":"Plant Physiology"},{"key":"2024082904153874700_CIT0047","doi-asserted-by":"crossref","first-page":"94","DOI":"10.1111\/nph.13579","article-title":"Using modern plant trait relationships between observed and theoretical maximum stomatal conductance and vein density to examine patterns of plant macroevolution","volume":"209","author":"McElwain","year":"2016","journal-title":"New Phytologist"},{"key":"2024082904153874700_CIT0048","doi-asserted-by":"crossref","first-page":"561","DOI":"10.1104\/pp.16.01853","article-title":"Stomatal defense a decade later","volume":"174","author":"Melotto","year":"2017","journal-title":"Plant Physiology"},{"key":"2024082904153874700_CIT0049","doi-asserted-by":"crossref","first-page":"254","DOI":"10.1104\/pp.105.062950","article-title":"Lateral diffusion of CO2 in leaves is not sufficient to support photosynthesis","volume":"139","author":"Morison","year":"2005","journal-title":"Plant Physiology"},{"key":"2024082904153874700_CIT0050","doi-asserted-by":"crossref","first-page":"1072","DOI":"10.1111\/j.1365-3040.2007.01685.x","article-title":"Does lateral gas diffusion in leaves matter","volume":"30","author":"Morison","year":"2007","journal-title":"Plant, Cell & Environment"},{"key":"2024082904153874700_CIT0051","doi-asserted-by":"crossref","first-page":"76","DOI":"10.1002\/j.1537-2197.1991.tb12573.x","article-title":"Amphistomy as an adaptation to high light intensity in Ambrosia cordifolia (Compositae)","volume":"78","author":"Mott","year":"1991","journal-title":"American Journal of Botany"},{"key":"2024082904153874700_CIT0052","doi-asserted-by":"crossref","first-page":"455","DOI":"10.1111\/1365-3040.ep11611750","article-title":"The adaptive significance of amphistomatic leaves","volume":"5","author":"Mott","year":"1982","journal-title":"Plant, Cell & Environment"},{"key":"2024082904153874700_CIT0053","doi-asserted-by":"crossref","first-page":"242","DOI":"10.1111\/nph.14956","article-title":"Light and growth form interact to shape stomatal ratio among British angiosperms","volume":"218","author":"Muir","year":"2018","journal-title":"New Phytologist"},{"key":"2024082904153874700_CIT0054","doi-asserted-by":"crossref","first-page":"571","DOI":"10.1093\/icb\/icz085","article-title":"Is amphistomy an adaptation to high light? Optimality models of stomatal traits along light gradients","volume":"59","author":"Muir","year":"2019","journal-title":"Integrative and Comparative Biology"},{"key":"2024082904153874700_CIT0055","doi-asserted-by":"crossref","first-page":"794","DOI":"10.1086\/723780","article-title":"How important are functional and developmental constraints on phenotypic evolution? An empirical test with the stomatal anatomy of flowering plants","volume":"201","author":"Muir","year":"2023","journal-title":"The American Naturalist"},{"key":"2024082904153874700_CIT0056","doi-asserted-by":"crossref","first-page":"142","DOI":"10.1086\/706260","article-title":"Consistent relationship between field-measured stomatal conductance and theoretical maximum stomatal conductance in C3 woody angiosperms in four major biomes","volume":"181","author":"Murray","year":"2020","journal-title":"International Journal of Plant Sciences"},{"key":"2024082904153874700_CIT0057","doi-asserted-by":"crossref","first-page":"2309","DOI":"10.1093\/jxb\/erx072","article-title":"Stomatal clustering in Begonia associates with the kinetics of leaf gaseous exchange and influences water use efficiency","volume":"68","author":"Papanatsiou","year":"2017","journal-title":"Journal of Experimental Botany"},{"key":"2024082904153874700_CIT0058","doi-asserted-by":"crossref","first-page":"367","DOI":"10.2307\/2259142","article-title":"The adaptive significance of stomatal occurrence on one or both surfaces of leaves","volume":"66","author":"Parkhurst","year":"1978","journal-title":"Journal of Ecology"},{"key":"2024082904153874700_CIT0059","doi-asserted-by":"crossref","first-page":"449","DOI":"10.1111\/j.1469-8137.1994.tb04244.x","article-title":"Diffusion of CO2 and other gases inside leaves","volume":"126","author":"Parkhurst","year":"1994","journal-title":"New Phytologist"},{"key":"2024082904153874700_CIT0060","doi-asserted-by":"crossref","first-page":"1024","DOI":"10.1104\/pp.94.3.1024","article-title":"Intercellular diffusion limits to CO2 uptake in leaves: studies in air and helox","volume":"94","author":"Parkhurst","year":"1990","journal-title":"Plant Physiology"},{"key":"2024082904153874700_CIT0061","doi-asserted-by":"crossref","first-page":"857","DOI":"10.1093\/jxb\/eri072","article-title":"Lateral gas diffusion inside leaves","volume":"56","author":"Pieruschka","year":"2005","journal-title":"Journal of Experimental Botany"},{"key":"2024082904153874700_CIT0062","doi-asserted-by":"crossref","first-page":"779","DOI":"10.1111\/j.1469-8137.2005.01605.x","article-title":"Lateral diffusion of CO 2 from shaded to illuminated leaf parts affects photosynthesis inside homobaric leaves","volume":"169","author":"Pieruschka","year":"2006","journal-title":"New Phytologist"},{"key":"2024082904153874700_CIT0063","doi-asserted-by":"crossref","first-page":"591","DOI":"10.1146\/annurev-arplant-042811-105451","article-title":"Mechanisms of stomatal development","volume":"63","author":"Pillitteri","year":"2012","journal-title":"Annual Review of Plant Biology"},{"key":"2024082904153874700_CIT0064","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1007\/0-387-29362-0_15","article-title":"Multiple testing procedures: the multtest package and applications to genomics","volume-title":"Bioinformatics and computational biology solutions using R and bioconductor","author":"Pollard","year":"2005"},{"key":"2024082904153874700_CIT0065","doi-asserted-by":"crossref","first-page":"1073","DOI":"10.1111\/nph.15754","article-title":"A meta-analysis of plant responses to light intensity for 70 traits ranging from molecules to whole plant performance","volume":"223","author":"Poorter","year":"2019","journal-title":"New Phytologist"},{"key":"2024082904153874700_CIT0066","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1086\/706186","article-title":"The scaling of genome size and cell size limits maximum rates of photosynthesis with implications for ecological strategies","volume":"181","author":"Roddy","year":"2020","journal-title":"International Journal of Plant Sciences"},{"key":"2024082904153874700_CIT0067","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/S0034-6667(00)00074-9","article-title":"Stomatal density and stomatal index as indicators of paleoatmospheric CO2 concentration","volume":"114","author":"Royer","year":"2001","journal-title":"Review of Palaeobotany and Palynology"},{"key":"2024082904153874700_CIT0068","doi-asserted-by":"crossref","first-page":"314","DOI":"10.1086\/336767","article-title":"The developmental origin of stomata pattern in crinum","volume":"135","author":"Sachs","year":"1974","journal-title":"Botanical Gazette"},{"key":"2024082904153874700_CIT0069","doi-asserted-by":"crossref","first-page":"2358","DOI":"10.1104\/pp.16.00476","article-title":"The developmental basis of stomatal density and flux","volume":"171","author":"Sack","year":"2016","journal-title":"Plant Physiology"},{"key":"2024082904153874700_CIT0070","doi-asserted-by":"crossref","first-page":"671","DOI":"10.1038\/nmeth.2089","article-title":"NIH Image to ImageJ: 25 years of image analysis","volume":"9","author":"Schneider","year":"2012","journal-title":"Nature Methods"},{"key":"2024082904153874700_CIT0071","doi-asserted-by":"crossref","first-page":"1211","DOI":"10.1093\/jxb\/31.5.1211","article-title":"Dependence of the stomatal index on environmental factors during stomatal differentiation in leaves of Vigna sinensis L. 1. Effect of light intensity","volume":"31","author":"Schoch","year":"1980","journal-title":"Journal of Experimental Botany"},{"key":"2024082904153874700_CIT0072","doi-asserted-by":"crossref","first-page":"816","DOI":"10.1111\/pce.12852","article-title":"Predicting stomatal responses to the environment from the optimization of photosynthetic gain and hydraulic cost: a stomatal optimization model","volume":"40","author":"Sperry","year":"2017","journal-title":"Plant, Cell & Environment"},{"key":"2024082904153874700_CIT0073","author":"Stan Development Team","year":"2023"},{"key":"2024082904153874700_CIT0074","doi-asserted-by":"crossref","first-page":"18488","DOI":"10.1073\/pnas.1216223109","article-title":"Drift-barrier hypothesis and mutation-rate evolution","volume":"109","author":"Sung","year":"2012","journal-title":"Proceedings of the National Academy of Sciences"},{"key":"2024082904153874700_CIT0075","doi-asserted-by":"crossref","first-page":"1609","DOI":"10.1111\/nph.14687","article-title":"The bias of a two-dimensional view: comparing two-dimensional and three-dimensional mesophyll surface area estimates using noninvasive imaging","volume":"215","author":"Th\u00e9roux-Rancourt","year":"2017","journal-title":"New Phytologist"},{"key":"2024082904153874700_CIT0076","doi-asserted-by":"crossref","DOI":"10.1214\/20-BA1221","article-title":"Rank-normalization, folding, and localization: an improved R\u02c6 for assessing convergence of MCMC (with discussion)","volume":"16","author":"Vehtari","year":"2021","journal-title":"Bayesian Analysis"},{"key":"2024082904153874700_CIT0077","doi-asserted-by":"crossref","first-page":"1743","DOI":"10.1111\/nph.18257","article-title":"Stomata on the abaxial and adaxial leaf surfaces contribute differently to leaf gas exchange and photosynthesis in wheat","volume":"235","author":"Wall","year":"2022","journal-title":"New Phytologist"},{"key":"2024082904153874700_CIT0078","doi-asserted-by":"crossref","first-page":"311","DOI":"10.1111\/nph.16572","article-title":"A theoretical and empirical assessment of stomatal optimization modeling","volume":"227","author":"Wang","year":"2020","journal-title":"New Phytologist"},{"key":"2024082904153874700_CIT0079","first-page":"317","article-title":"Heterogeneity in stomatal characteristics","volume-title":"Advances in botanical research","author":"Weyers","year":"1997"},{"key":"2024082904153874700_CIT0080","doi-asserted-by":"crossref","first-page":"617","DOI":"10.1038\/327617a0","article-title":"Stomatal numbers are sensitive to increases in CO2 from pre-industrial levels","volume":"327","author":"Woodward","year":"1987","journal-title":"Nature"},{"key":"2024082904153874700_CIT0081","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1186\/1746-4811-6-17","article-title":"High-contrast three-dimensional imaging of the Arabidopsis leaf enables the analysis of cell dimensions in the epidermis and mesophyll","volume":"6","author":"Wuyts","year":"2010","journal-title":"Plant Methods"},{"key":"2024082904153874700_CIT0082","first-page":"325","article-title":"Stomatal clustering, a new marker for environmental perception and adaptation in terrestrial plants","volume":"51","author":"Yi","year":"2010","journal-title":"Botanical Studies"}],"container-title":["AoB PLANTS"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/aobpla\/advance-article-pdf\/doi\/10.1093\/aobpla\/plae015\/57027197\/plae015.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/aobpla\/article-pdf\/16\/2\/plae015\/58955348\/plae015.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/aobpla\/article-pdf\/16\/2\/plae015\/58955348\/plae015.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,8,29]],"date-time":"2024-08-29T04:16:24Z","timestamp":1724904984000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/aobpla\/article\/doi\/10.1093\/aobpla\/plae015\/7632359"}},"subtitle":[],"editor":[{"given":"William","family":"Salter","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2024,2,1]]},"references-count":82,"journal-issue":{"issue":"2","published-print":{"date-parts":[[2024,2,1]]}},"URL":"https:\/\/doi.org\/10.1093\/aobpla\/plae015","relation":{},"ISSN":["2041-2851"],"issn-type":[{"value":"2041-2851","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2024,2,1]]},"published":{"date-parts":[[2024,2,1]]},"article-number":"plae015"}}