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Plant leaf temperatures can differ from ambient air temperatures. A temperature gradient in a gas mixture gives rise to a phenomenon known as thermodiffusion, which operates in addition to ordinary diffusion. Whilst transpiration is generally understood to be driven solely by the ordinary diffusion of water vapour along a concentration gradient, we consider the implications of thermodiffusion for transpiration. We develop a new modelling framework that introduces the effects of thermodiffusion on the transpiration rate, E. By applying this framework, we quantify the proportion of E attributable to thermodiffusion for a set of physiological and environmental conditions, varied over a wide range. Thermodiffusion is found to be most significant (in some cases > 30% of E) when a leaf-to-air temperature difference coincides with a relatively small water vapour concentration difference across the boundary layer; a boundary layer conductance that is large as compared to the stomatal conductance; or a relatively low transpiration rate. Thermodiffusion also alters the conditions required for the onset of reverse transpiration, and the rate at which this water vapour uptake occurs., (© 2024 The Authors. New Phytologist © 2024 New Phytologist Foundation.)

This work aims at developing an adequate theoretical basis for comparing assimilation of the ancestral C 3 pathway with CO 2 concentrating mechanisms (CCM) that have evolved to reduce photorespiratory yield losses. We present a novel model for C 3 , C 2 , C 2  + C 4 and C 4 photosynthesis simulating assimilatory metabolism, energetics and metabolite traffic at the leaf level. It integrates a mechanistic description of light reactions to simulate ATP and NADPH production, and a variable engagement of cyclic electron flow. The analytical solutions are compact and thus suitable for larger scale simulations. Inputs were derived with a comprehensive gas-exchange experiment. We show trade-offs in the operation of C 4 that are in line with ecophysiological data. C 4 has the potential to increase assimilation over C 3 at high temperatures and light intensities, but this benefit is reversed under low temperatures and light. We apply the model to simulate the introduction of progressively complex levels of CCM into C 3 rice, which feeds > 3.5 billion people. Increasing assimilation will require considerable modifications such as expressing the NAD(P)H Dehydrogenase-like complex and upregulating cyclic electron flow, enlarging the bundle sheath, and expressing suitable transporters to allow adequate metabolite traffic. The simpler C 2 rice may be a desirable alternative., (© 2019 The Authors. New Phytologist © 2019 New Phytologist Trust.)

The arrangement of mitochondria and chloroplasts, together with the relative resistances of cell wall and chloroplast, determine the path of diffusion out of the leaf for (photo)respired CO 2 . Traditional photosynthesis models have assumed a tight arrangement of chloroplasts packed together against the cell wall with mitochondria located behind the chloroplasts, deep inside the cytosol. Accordingly, all (photo)respired CO 2 must cross the chloroplast before diffusing out of the leaf. Different arrangements have recently been considered, where all or part of the (photo)respired CO 2 diffuses through the cytosol without ever entering the chloroplast. Assumptions about the path for the (photo)respiratory flux are particularly relevant for the calculation of mesophyll conductance (g m ). If (photo)respired CO 2 can diffuse elsewhere besides the chloroplast, apparent g m is no longer a mere physical resistance but a flux-weighted variable sensitive to the ratio of (photo)respiration to net CO 2 assimilation. We discuss existing photosynthesis models in conjunction with their treatment of the (photo)respiratory flux and present new equations applicable to the generalized case where (photo)respired CO 2 can diffuse both into the chloroplast and through the cytosol. Additionally, we present a new generalized Δ 13 C model that incorporates this dual diffusion pathway. We assess how assumptions about the fate of (photo)respired CO 2 affect the interpretation of photosynthetic data and the challenges it poses for the application of different models.

Stable carbon isotopes are a powerful tool to study photosynthesis. Initial applications consisted of determining isotope ratios of plant biomass using mass spectrometry. Subsequently, theoretical models relating C-isotope values to gas exchange characteristics were introduced and tested against instantaneous online measurements of 13 C photosynthetic discrimination. Beginning in the twenty-first century, tunable diode laser spectroscopes with sufficient precision for determining isotope mixing ratios became commercially available. This has allowed collection of large data sets, at low cost and with unprecedented temporal resolution. With more data and accompanying knowledge, it has become apparent that there is a need for increased complexity in models and calculations. This chapter describes instantaneous online measurements of 13 C photosynthetic discrimination, provides recommendations for experimental setup, and presents a thorough compilation of equations needed for different applications.

It was shown over 40 years ago that plants maximize carbon gain for a given rate of water loss if stomatal conductance, g s , varies in response to external and internal conditions such that the marginal carbon revenue of water, ∂A/∂E, remains constant over time. This theory has long held promise for understanding the physiological ecology of water use and for informing models of plant-atmosphere interactions. Full realization of this potential hinges on three questions: (i) Are analytical approximations adequate for applying the theory at diurnal time scales? (ii) At what time scale is it realistic and appropriate to apply the theory? (iii) How should g s vary to maximize growth over long time scales? We review the current state of understanding for each of these questions and describe future research frontiers. In particular, we show that analytical solutions represent the theory quite poorly, especially when boundary layer or mesophyll resistances are significant; that diurnal variations in hydraulic conductance may help or hinder maintenance of ∂A/∂E, and the matter requires further study; and that optimal diurnal responses are distinct from optimal long-term variations in g s , which emerge from optimal shifts in carbon partitioning at the whole-plant scale., (© 2016 John Wiley & Sons Ltd.)

Mitochondrial respiration often appears to be inhibited in the light when compared with measurements in the dark. This inhibition is inferred from the response of the net CO 2 assimilation rate (A) to absorbed irradiance (I), changing slope around the light compensation point (I c ). We suggest a model that provides a plausible mechanistic explanation of this 'Kok effect'. The model uses the mathematical description of photosynthesis developed by Farquhar, von Caemmerer and Berry; it involves no inhibition of respiration rate in the light. We also describe a fitting technique for quantifying the Kok effect at low I. Changes in the chloroplastic CO 2 partial pressure (C c ) can explain curvature of A vs I, its diminution in C 4 plants and at low oxygen concentrations or high carbon dioxide concentrations in C 3 plants, and effects of dark respiration rate and of temperature. It also explains the apparent inhibition of respiration in the light as inferred by the Laisk approach. While there are probably other sources of curvature in A vs I, variation in C c can largely explain the curvature at low irradiance, and suggests that interpretation of day respiration compared with dark respiration of leaves on the basis of the Kok effect needs reassessment., (© 2017 The Authors. New Phytologist © 2017 New Phytologist Trust.)

We present a combined three-dimensional (3-D) model of light propagation, CO2 diffusion and photosynthesis in tomato (Solanum lycopersicum L.) leaves. The model incorporates a geometrical representation of the actual leaf microstructure that we obtained with synchrotron radiation X-ray laminography, and was evaluated using measurements of gas exchange and leaf optical properties. The combination of the 3-D microstructure of leaf tissue and chloroplast movement induced by changes in light intensity affects the simulated CO2 transport within the leaf. The model predicts extensive reassimilation of CO2 produced by respiration and photorespiration. Simulations also suggest that carbonic anhydrase could enhance photosynthesis at low CO2 levels but had little impact on photosynthesis at high CO2 levels. The model confirms that scaling of photosynthetic capacity with absorbed light would improve efficiency of CO2 fixation in the leaf, especially at low light intensity., (© 2015 John Wiley & Sons Ltd.)

The two-pool and Péclet effect models represent two theories describing mechanistic controls underlying leaf water oxygen isotope composition at the whole-leaf level (δ(18) OL ). To test these models, we used a laser spectrometer coupled to a gas-exchange cuvette to make online measurements of δ(18) O of transpiration (δ(18) Otrans ) and transpiration rate (E) in 61 cotton (Gossypium hirsutum) leaves. δ(18) Otrans measurements permitted direct calculation of δ(18) O at the sites of evaporation (δ(18) Oe ) which, combined with values of δ(18) OL from the same leaves, allowed unbiased estimation of the proportional deviation of enrichment of δ(18) OL from that of δ(18) Oe (f) under both steady-state (SS) and non-steady-state (NSS) conditions. Among all leaves measured, f expressed relative to both δ(18) O of transpired water (ftrans ) and source water (fsw ) remained relatively constant with a mean ± SD of 0.11 ± 0.05 and 0.13 ± 0.05, respectively, regardless of variation in E spanning 0.8-9.1 mmol m(-2)  s(-1) . Neither ftrans nor fsw exhibited a significant difference between the SS and NSS leaves at the P < 0.05 level. Our results suggest that the simpler two-pool model is adequate for predicting cotton leaf water enrichment at the whole-leaf level. We discuss the implications of adopting a two-pool concept for isotopic applications in ecological studies., (© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.)

Seedlings of a conifer (Pinus radiata D. Don) and a broad leaf angiosperm (Eucalyptus globulus Labill.) were grown for 100 days in two growth cabinets at 45 or 65% relative humidity. The seedlings were exposed to treatments designed to modify carbon assimilation rates and capacities, stomatal conductance and transpiration to test conceptual models that attempt to clarify the interpretation of carbon isotope discrimination (Δ(13)C) by using oxygen isotope enrichment (Δ(18)O). Differences in relative humidity and within-cabinet treatments (including lower irradiance, lower nitrogen inputs, higher leaf temperature and lower moisture status than control seedlings) produced significant differences in assimilation rates, photosynthetic capacities, stomatal conductance, leaf transpiration rates and leaf evaporative enrichment. The dual-isotope approach accurately interpreted the cause of variation in wood cellulose Δ(13)C for some of the treatments, but not for others. We also tested whether we could use Δ(13)C variation to constrain the interpretation of δ(18)O variation. Carbon isotope discrimination appears to be influenced by transpiration (providing information on leaf evaporative enrichment), but the results did not provide a clear way to interpret such variation. The dual-isotope approach appears to be valid conceptually, but more work is needed to make it operational under different scenarios.

To estimate the relative contributions of woody and herbaceous vegetation to savanna productivity, we measured the 13C/12C isotopic ratios of leaves from trees, shrubs, grasses and the surface soil carbon pool for 22 savannas in Australia, Brazil and Ghana covering the full savanna spectrum ranging from almost pure grassland to closed woodlands on all three continents. All trees and shrubs sampled were of the C3 pathway and all grasses of the C4 pathway with the exception of Echinolaena inflexa (Poir.) Chase, a common C3 grass of the Brazilian cerrado. By comparing the carbon isotopic compositions of the plant and carbon pools, a simple model relating soil delta 13C to the relative abundances of trees + shrubs (woody plants) and grasses was developed. The model suggests that the relative proportions of a savanna ecosystem's total foliar projected cover attributable to grasses versus woody plants is a simple and reliable index of the relative contributions of grasses and woody plants to savanna net productivity. Model calibrations against woody tree canopy cover made it possible to estimate the proportion of savanna productivity in the major regions of the world attributable to trees + shrubs and grasses from ground-based observational maps of savanna woodiness. Overall, it was estimated that 59% of the net primary productivity (Np) of tropical savannas is attributable to C4 grasses, but that this proportion varies significantly within and between regions. The C4 grasses make their greatest relative contribution to savanna Np in the Neotropics, whereas in African regions, a greater proportion of savanna Np is attributable to woody plants. The relative contribution of C4 grasses in Australian savannas is intermediate between those in the Neotropics and Africa. These differences can be broadly ascribed to large scale differences in soil fertility and rainfall.

Photosynthetic responses to carbon dioxide concentration can provide data on a number of important parameters related to leaf physiology. Methods for fitting a model to such data are briefly described. The method will fit the following parameters: V(cmax), J, TPU, R(d) and g(m)[maximum carboxylation rate allowed by ribulose 1.5-bisphosphate carboxylase/oxygenase (Rubisco), rate of photosynthetic electron transport (based on NADPH requirement), triose phosphate use, day respiration and mesophyll conductance, respectively]. The method requires at least five data pairs of net CO(2) assimilation (A) and [CO(2)] in the intercellular airspaces of the leaf (C(i)) and requires users to indicate the presumed limiting factor. The output is (1) calculated CO(2) partial pressure at the sites of carboxylation, C(c), (2) values for the five parameters at the measurement temperature and (3) values adjusted to 25 degrees C to facilitate comparisons. Fitting this model is a way of exploring leaf level photosynthesis. However, interpreting leaf level photosynthesis in terms of underlying biochemistry and biophysics is subject to assumptions that hold to a greater or lesser degree, a major assumption being that all parts of the leaf are behaving in the same way at each instant.

We described advection and diffusion of water isotopologues in leaves in the non-steady state, applied specifically to amphistomatous leaves. This explains the isotopic enrichment of leaf water from the xylem to the mesophyll, and we showed how it relates to earlier models of leaf water enrichment in non-steady state. The effective length or tortuosity factor of isotopologue movement in leaves is unknown and, therefore, is a fitted parameter in the model. We compared the advection-diffusion model to previously published data sets for Lupinus angustifolius and Eucalyptus globulus. Night-time stomatal conductance was not measured in either data set and is therefore another fitted parameter. The model compared very well with the observations of bulk mesophyll water during the whole diel cycle. It compared well with the enrichment at the evaporative sites during the day but showed some deviations at night for E. globulus. It became clear from our analysis that night-time stomatal conductance should be measured in the future and that the temperature dependence of the tracer diffusivities should be accounted for. However, varying mesophyll water volume did not seem critical for obtaining a good prediction of leaf water enrichment, at least in our data sets. In addition, observations of single diurnal cycles do not seem to constrain the effective length that relates to the tortuosity of the water path in the mesophyll. Finally, we showed when simpler models of leaf water enrichment were suitable for applications of leaf water isotopes once weighted with the appropriate gas exchange flux. We showed that taking an unsuitable leaf water enrichment model could lead to large biases when cumulated over only 1 day.

The analysis of the temperature dependence of the rate of photosynthetic carbon assimilation by leaves of C3 plants has involved subdivision into a diffusional (mainly stomatal) component and a biochemical one. The latter has been further analysed in terms of Rubisco activity and of the capacity for regeneration of RubP. Rubisco activity has been modelled in terms of the number of active (carbamylated) enzyme sites and the kinetics of these sites. RubP regeneration capacity has been modelled in terms of photosynthetic electron transport and photophosphorylation capacity and the rate of supply of inorganic phosphate. The short-term (minutes to hours) responses of assimilation rate to changes in temperature of Eucalyptus pauciflora leaves, in the range 15 to 35 degrees C, are examined using this framework. It is argued that the above approach has some useful features, even if these are mainly pedagogical. Problems with the simplified framework are also discussed, including heterogeneity of the light intensity and CO2 concentration within the leaf. The latter is discussed in terms of the resistances to diffusion within the leaf and observations of non-uniformity ('patches') of stomatal opening. The effects of temperature, couched in the above terms, appear to be mainly those on the supply of carbon as a substrate for growth. Yet the model (or description of carbon assimilation) can accommodate certain effects of demand for carbon (sink limitation). These occur, notably, via the supply of inorganic phosphate, but potentially also via Rubisco activation and other mechanisms. A striking example of effects of reduced temperature on assimilation and growth being caused by changed demand for carbon, is given for the tropical crop, Arachis hypogaea. This example prompts an appraisal of the effects of temperature on components of growth other than photosynthesis, for example, of leaf production rate.