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Abstract PremiseThe adaptive significance of amphistomy (stomata on both upper and lower leaf surfaces) is unresolved. A widespread association between amphistomy and open, sunny habitats suggests the adaptive benefit of amphistomy may be greatest in these contexts, but this hypothesis has not been tested experimentally. Understanding amphistomy informs its potential as a target for crop improvement and paleoenvironment reconstruction. MethodsWe developed a method to quantify “amphistomy advantage” () as the log‐ratio of photosynthesis in an amphistomatous leaf to that of the same leaf but with gas exchange blocked through the upper surface (pseudohypostomy). Humidity modulated stomatal conductance and thus enabled comparing photosynthesis at the same total stomatal conductance. We estimated and leaf traits in six coastal (open, sunny) and six montane (closed, shaded) populations of the indigenous Hawaiian species ʻilima (Sida fallax). ResultsCoastal ʻilima leaves benefit 4.04 times more from amphistomy than montane leaves. Evidence was equivocal with respect to two hypotheses: (1) that coastal leaves benefit more because they are thicker and have lower CO₂conductance through the internal airspace and (2) that they benefit more because they have similar conductance on each surface, as opposed to most conductance being through the lower surface. ConclusionsThis is the first direct experimental evidence that amphistomy increases photosynthesis, consistent with the hypothesis that parallel pathways through upper and lower mesophyll increase CO₂supply to chloroplasts. The prevalence of amphistomatous leaves in open, sunny habitats can partially be explained by the increased benefit of amphistomy in “sun” leaves, but the mechanistic basis remains uncertain. 

Abstract 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–adaxial stomatal coordination are miniscule (≪1%) 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. 

Summary A prevailing hypothesis posits that achieving higher maximum rates of leaf carbon gain and water loss is constrained by geometry and/or selection to limit the allocation of epidermal area to stomata (f_S). Under this ‘stomatal‐area minimization hypothesis’, higherg_s,maxis associated with greater numbers of smaller stomata because this trait combination increasesg_s,maxwith minimal increase inf_S, leading to relative conservation off_Ssemi‐independent ofg_s,maxdue to coordination in stomatal size, density, and pore depth. An alternative hypothesis is that the evolution of higherg_s,maxcan be enabled by a greater epidermal area allocated to stomata, leading to positive covariation betweenf_Sandg_s,max; we call this the ‘stomatal‐area adaptation hypothesis’. Under this hypothesis, the interspecific scaling betweeng_s,max, stomatal density, and stomatal size is a by‐product of selection on a moving optimalg_s,max.We integrated biophysical and evolutionary quantitative genetic modeling with phylogenetic comparative analyses of a global data set of stomatal density and size from 2408 vascular forest species. The models present specific assumptions of both hypotheses and deduce predictions that can be evaluated with our empirical analyses of forest plants.There are three main results. First, neither the stomatal‐area minimization nor adaptation hypothesis is sufficient to be supported. Second, estimates of interspecific scaling from common regression methods cannot reliably distinguish between hypotheses when stomatal size is bounded. Third, we reconcile both hypotheses with the data by including an additional assumption that stomatal size is bounded by a wide range and under selection; we refer to this synthetic hypothesis as the ‘stomatal adaptation + bounded size’ hypothesis.This study advances our understanding of scaling between stomatal size and density by mathematically describing specific assumptions of competing hypotheses, demonstrating that existing hypotheses are inconsistent with observations, and reconciling these hypotheses with phylogenetic comparative analyses by postulating a synthetic model of selection ong_s,max,f_S, and stomatal size. 

Abstract Plant ecophysiology is founded on a rich body of physical and chemical theory, but it is challenging to connect theory with data in unambiguous, analytically rigorous and reproducible ways. Custom scripts written in computer programming languages (coding) enable plant ecophysiologists to model plant processes and fit models to data reproducibly using advanced statistical techniques. Since many ecophysiologists lack formal programming education, we have yet to adopt a unified set of coding principles and standards that could make coding easier to learn, use and modify. We identify eight principles to help in plant ecophysiologists without much programming experience to write resilient code: (i) standardized nomenclature, (ii) consistency in style, (iii) increased modularity/extensibility for easier editing and understanding, (iv) code scalability for application to large data sets, (v) documented contingencies for code maintenance, (vi) documentation to facilitate user understanding; (vii) extensive tutorials and (viii) unit testing and benchmarking. We illustrate these principles using a new R package, {photosynthesis}, which provides a set of analytical and simulation tools for plant ecophysiology. Our goal with these principles is to advance scientific discovery in plant ecophysiology by making it easier to use code for simulation and data analysis, reproduce results and rapidly incorporate new biological understanding and analytical tools. 

Summary Leaf optical properties impact leaf energy balance and thus leaf temperature. The effect of leaf development on mid‐infrared (MIR) reflectance, and hence thermal emissivity, has not been investigated in detail.We measured a suite of morphological characteristics, as well as directional‐hemispherical reflectance from ultraviolet to thermal infrared wavelengths (250 nm to 20 µm) of leaves from five temperate deciduous tree species over the 8 wk following spring leaf emergence.By contrast to reflectance at shorter wavelengths, the shape and magnitude of MIR reflectance spectra changed markedly with development. MIR spectral differences among species became more pronounced and unique as leaves matured. Comparison of reflectance spectra of intact vs dried and ground leaves points to cuticular development – and not internal structural or biochemical changes – as the main driving factor. Accompanying the observed spectral changes was a drop in thermal emissivity from about 0.99 to 0.95 over the 8 wk following leaf emergence.Emissivity changes were not large enough to substantially influence leaf temperature, but they could potentially lead to a bias in radiometrically measured temperatures of up to 3 K. Our results also pointed to the potential for using MIR spectroscopy to better understand species‐level differences in cuticular development and composition.