Farquhar-von Caemmerer-Berry Model Research Articles

The importance of species-specific and temperature-sensitive parameterisation of A/Ci models: A case study using cotton (Gossypium hirsutum L.) and the automated 'OptiFitACi' R-package.

Leaf gas exchange measurements are an important tool for inferring a plant's photosynthetic biochemistry. In most cases, the responses of photosynthetic CO2 assimilation to variable intercellular CO2 concentrations (A/Ci response curves) are used to model the maximum (potential) rate of carboxylation by ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco, Vcmax) and the rate of photosynthetic electron transport at a given incident photosynthetically active radiation flux density (PAR; JPAR). The standard Farquhar-von Caemmerer-Berry model is often used with default parameters of Rubisco kinetic values and mesophyll conductance to CO2 (gm) derived from tobacco that may be inapplicable across species. To study the significance of using such parameters for other species, here we measured the temperature responses of key in vitro Rubisco catalytic properties and gm in cotton (Gossypium hirsutum cv. Sicot 71) and derived Vcmax and J2000 (JPAR at 2000 µmol m-2 s-1 PAR) from cotton A/Ci curves incrementally measured at 15°C-40°C using cotton and other species-specific sets of input parameters with our new automated fitting R package 'OptiFitACi'. Notably, parameterisation by a set of tobacco parameters produced unrealistic J2000:Vcmax ratio of <1 at 25°C, two- to three-fold higher estimates of Vcmax above 15°C, up to 2.3-fold higher estimates of J2000 and more variable estimates of Vcmax and J2000, for our cotton data compared to model parameterisation with cotton-derived values. We determined that errors arise when using a gm,25 of 2.3 mol m-2 s-1 MPa-1 or less and Rubisco CO2-affinities in 21% O2 (KC 21%O2) at 25°C outside the range of 46-63 Pa to model A/Ci responses in cotton. We show how the A/Ci modelling capabilities of 'OptiFitACi' serves as a robust, user-friendly, and flexible extension of 'plantecophys' by providing simplified temperature-sensitivity and species-specificity parameterisation capabilities to reduce variability when modelling Vcmax and J2000.

Development of Modified Farquhar-von Caemmerer-Berry Model Describing Photodamage of Photosynthetic Electron Transport in C3 Plants under Different Temperatures.

Photodamage of photosynthetic electron transport is a key mechanism of disruption of photosynthesis in plants under action of stressors. This means that investigation of photodamage is an important task for basic and applied investigations. However, its complex mechanisms restrict using experimental methods of investigation for this process; the development of mathematical models of photodamage and model-based analysis can be used for overcoming these restrictions. In the current work, we developed the modified Farquhar-von Caemmerer-Berry model which describes photodamage of photosynthetic electron transport in C3 plants. This model was parameterized on the basis of experimental results (using an example of pea plants). Analysis of the model showed that combined inactivation of linear electron flow and Rubisco could induce both increasing and decreasing photodamage at different magnitudes of inactivation of these processes. Simulation of photodamage under different temperatures and light intensities showed that simulated temperature dependences could be multi-phase; particularly, paradoxical increases in the thermal tolerance of photosynthetic electron transport could be observed under high temperatures (37-42 °C). Finally, it was shown that changes in temperature optimums of linear electron flow and Rubisco could modify temperature dependences of the final activity of photosynthetic electron transport under photodamage induction; however, these changes mainly stimulated its photodamage. Thus, our work provides a new theoretical tool for investigation of photodamage of photosynthetic processes in C3 plants and shows that this photodamage can be intricately dependent on parameters of changes in activities of linear electron flow and Rubisco including changes induced by temperature.

Development of a semi-open chamber system for the gas exchange measurement of whole-canopy under steady and unsteady states in cucumber seedlings

BackgroundLarge-scale data on the photosynthetic characteristics of whole crop canopy is crucial for improving yield. However, current data collection methods remain challenging, and the time constraints associated with photosynthetic data collection further complicate matters. Developing a practical yet easy-to-use tool for collecting whole-canopy data is essential to address these challenges. Furthermore, it is necessary to obtain instantaneous measurements of photosynthetic rate over a wide range of CO2 concentrations under an unsteady state to enable faster data collection and obtain reliable biochemical limits of carbon assimilation. This study developed a semi-open chamber system with steady and unsteady state measurement techniques to collect biochemical photosynthetic data from an entire cucumber canopy, emphasizing the correction procedures for CO2 concentration of unsteady state measurements applicable regardless of chamber scale.ResultsAfter constructing a semi-open chamber system, we described how to correct measurement errors according to chamber volume. In order to assess the accuracy of the newly developed system, an analysis was conducted to determine the overall measurement error resulting from variations in the reference, sample CO2 concentration, and leakage flow rate. The total measurement error was accurate to no more than 10%. Furthermore, the difference between the photosynthetic rate of the single leaf and that of the whole-canopy was not significant in Rubisco activity-limited carboxylation range. In addition, the Farquhar–von Caemmerer–Berry (FvCB) model parameters and the photosynthetic rate estimation values were compared to evaluate the steady- and unsteady state measurement methods between the cucumber seedlings' single-leaf and whole-canopy. The average root mean square error of the FvCB model in the steady (standard A-Ci response) and unsteady states (800 to 400 ramp) of the chambers was 1.4 and 2.3, respectively. Results show that the developed system is suitable for measuring the gas exchange rate of the cucumber canopy.ConclusionsWe demonstrate the correction method for measurement errors to enable the gas exchange rate of the whole-canopy even in an unsteady state. The correction method of the measurement system of the gas exchange rate for the whole- canopy can be applied regardless of the volume of the chamber, and it can be applied simply to other chamber systems. In addition, an unsteady state measurement method for fast data collection was also applicable. However, it was deemed necessary to identify a more optimal measurement range by conducting measurements across a broader range of values.

A small dynamic leaf-level model predicting photosynthesis in greenhouse tomatoes.

The conversion of supplemental greenhouse light energy into biomass is not always optimal. Recent trends in global energy prices and discussions on climate change highlight the need to reduce our energy footprint associated with the use of supplemental light in greenhouse crop production. This can be achieved by implementing "smart" lighting regimens which in turn rely on a good understanding of how fluctuating light influences photosynthetic physiology. Here, a simple fit-for-purpose dynamic model is presented. It accurately predicts net leaf photosynthesis under natural fluctuating light. It comprises two ordinary differential equations predicting: 1) the total stomatal conductance to CO2 diffusion and 2) the CO2 concentration inside a leaf. It contains elements of the Farquhar-von Caemmerer-Berry model and the successful incorporation of this model suggests that for tomato (Solanum lycopersicum L.), it is sufficient to assume that Rubisco remains activated despite rapid fluctuations in irradiance. Furthermore, predictions of the net photosynthetic rate under both 400ppm and enriched 800ppm ambient CO2 concentrations indicate a strong correlation between the dynamic rate of photosynthesis and the rate of electron transport. Finally, we are able to indicate whether dynamic photosynthesis is Rubisco or electron transport rate limited.

Quantitative expression of mesophyll conductance temperature response in the FvCB model and impacts on plant gas exchange estimations

The way of quantitatively expressing mesophyll conductance (gm) in the Farquhar-von Caemmerer-Berry (FvCB) photosynthesis model and its impacts on plant gas exchange estimations have not been well explored, primarily due to huge uncertainties in gm parameterization. Here, a peaked Arrhenius function to depict gm temperature response was introduced into the FvCB model and parameterized through evaluating four different gm estimation methods in 19 C3 species at 31 experimental treatments. Results indicated that the FvCB model without explicitly considering gm cannot perform well in eight species/treatments, while the model that considers gm estimated by the chlorophyll fluorescence–gas exchange method and biochemical parameters estimated by the Bayesian retrieval algorithm was superior. Overall modeling accuracy was not further ameliorated when taking anatomy-based gm into consideration. The increasing Arrhenius function without considering the suboptimal stage of gm temperature response caused significant overestimations in photosynthesis under high leaf temperatures by 2–3 folds. The gm explicit expression had equally important effects on photosynthesis and transpiration estimations, which disagreed with “the asymmetric effects on photosynthesis and transpiration estimations” hypothesis proposed by Knauer et al. (2020). Literature survey plus our data indicated that observed variations of photosynthesis optimal temperature (ToptA) were primarily explained by the gm optimal temperature (Topt_gm) (58%) rather than biochemical limitations, which disagreed with “the JVr biochemical limitations” hypothesis proposed by Kumarathunge et al. (2019).

Vertical profile of photosynthetic CO&lt;sub&gt;2&lt;/sub&gt; response within rice canopy

Leaf-specific Farquhar-von Caemmerer-Berry (FvCB) model was fitted to characterize the vertical profile of photosynthetic CO2 response within rice canopy. Leaf-position-specific and canopy average FvCB models were fitted to study a suitable leaf representing photosynthetic parameters at the canopy scale. The results showed that leaf photosynthesis was limited by Rubisco activity or ribulose-1,5-bisphosphate regeneration under field conditions. The maximum rate of carboxylation, maximum rate of electron transport, rate of triose phosphates utilization, and light respiration rate in the FvCB model reached the highest values for the top second leaf and then decreased, while the mesophyll diffusion conductance kept decreased in downward leaves. The integrated photosynthetic CO2-response curves for the top fourth and fifth leaves were appropriate for estimating parameters in the FvCB model at the canopy scale.

The physiological basis for estimating photosynthesis from Chla fluorescence.

Solar-induced Chl fluorescence (SIF) offers the potential to curb large uncertainties in the estimation of photosynthesis across biomes and climates, and at different spatiotemporal scales. However, it remains unclear how SIF should be used to mechanistically estimate photosynthesis. In this study, we built a quantitative framework for the estimation of photosynthesis, based on a mechanistic light reaction model with the Chla fluorescence of Photosystem II (SIFPSII ) as an input (MLR-SIF). Utilizing 29 C3 and C4 plant species that are representative of major plant biomes across the globe, we confirmed the validity of this framework at the leaf level. The MLR-SIF model is capable of accurately reproducing photosynthesis for all C3 and C4 species under diverse light, temperature, and CO2 conditions. We further tested the robustness of the MLR-SIF model using Monte Carlo simulations, and found that photosynthesis estimates were much less sensitive to parameter uncertainties relative to the conventional Farquhar, von Caemmerer, Berry (FvCB) model because of the additional independent information contained in SIFPSII . Once inferred from direct observables of SIF, SIFPSII provides 'parameter savings' to the MLR-SIF model, compared to the mechanistically equivalent FvCB model, and thus avoids the uncertainties arising as a result of imperfect model parameterization. Our findings set the stage for future efforts to employ SIF mechanistically to improve photosynthesis estimates across a variety of scales, functional groups, and environmental conditions.

Evaluation of the light profile and carbon assimilation of tomato plants in greenhouses with respect to film diffuseness and regional solar radiation using ray-tracing simulation

Diffuse fraction, which can be increased by using diffuse films, has been considered to influence light interception and photosynthesis of crops in greenhouses. However, quantifying the influence of diffuse films is challenging owing to the complicated optical interactions between climatic factors inside and outside greenhouses. Thus, versatile methods for evaluating the effect of diffuse films are required. The objective of this study was to evaluate the effect of diffuse films on the improvement of the light profile and photosynthesis of tomatoes in greenhouses according to film diffuseness and regional solar radiation using ray-tracing simulation. The structural and optical properties of the greenhouse components were applied in a 3D-framework combined with a ray-tracing module. The light transmission patterns of diffuse films and solar radiation properties were incorporated. The reliability of the simulation was confirmed by comparing measured and estimated irradiances inside greenhouses covered with films having different haze factors. For scenarios, the diffuse film efficiency was assessed under typically different solar radiations, a low irradiance, high diffuse radiation fraction (LIHD) and a high irradiance, low diffuse radiation fraction (HILD). The light interception was estimated through the simulation and used to calculate the photosynthesis using the Farquhar-von Caemmerer-Berry model. The simulation was found to be reliable with R2 of 0.95 and 0.94 for the two greenhouses covered with different diffuse films. The light distribution on the tomato plants were less affected by film diffuseness under LIHD than HILD. With increasing film diffuseness, carbon uptake and light use efficiency increased by 5.30% and 4.58% under HILD, but did not change under LIHD. The light distribution and photosynthesis in diffuse film-covered greenhouses under different light environments could be reasonably estimated by the simulation. Thus, this method can be used to evaluate the applicability of diffuse films to various regions with diverse meteorological characteristics.

The maximum carboxylation rate of Rubisco affects CO2 refixation in temperate broadleaved forest trees

Mesophyll resistance to CO2 diffusion (rm) and the maximum carboxylation rate of Rubisco (Vcmax) affect photosynthetic rates, and can potentially also influence the percentage of respiratory and photorespiratory CO2 being refixated (Pr) by mesophyll cells. Here we investigated how various leaf anatomical traits (e.g. leaf mass per area [LMA] and leaf dry matter content [LDMC]) influenced rm in leaves of mature forest trees. We further explored how rm and Vcmax in turn affected Pr, and if these traits varied among species and leaves along a light gradient. Photosynthetic CO2 response of leaves grown in high-, medium-, and low-light environments was measured, from Pinus sylvestris [Scots pine], Picea abies [Norway spruce], Quercus robur [English oak], and Betula pendula [Silver birch] in southern Sweden. A modified version of the Farquhar-von Caemmerer-Berry model was fitted to the leaf gas exchange data to estimate Vcmax, rm and Pr. We found that of all leaf traits measured, only LMA for Q. robur was significantly higher in leaves from high-light environments. When comparing species, both rm and LMA were significantly higher in the conifers, and rm had a negative correlation with Vcmax. We found that Pr was similar between different species and functional groups, with an average of 73.2% (and SD of ±10.4) across all species. There was a strong, positive correlation between Pr and Vcmax in broadleaves, and we hypothesise that this effect might derive from a higher CO2 drawdown near Rubisco in leaves with high Vcmax.

Oxygen response of leaf CO2 compensation points used to determine Rubisco specificity factors of gymnosperm species

Rubisco specificity factor (Sc/o), a measure of the relative capacities of an enzyme to catalyze carboxylation and oxygenation of ribulose-1,5-bisphosphate, determines the extent of photosynthetic CO2 assimilation and photorespiratory CO2 release. The current model of C3 photosynthesis, the Farquhar-von Caemmerer-Berry (FvCB) model, requires a species-specific Sc/o. However, Sc/o values have never been reported in conifers, likely because in vitro kinetic analysis of conifer Rubisco presents difficulties. To estimate the Sc/o of conifers and compare it with angiosperm Sc/o, we measured changes in leaf CO2 compensation points (Γ) in response to O2 partial pressure for a variety of leaves, with different rates of day respiration (Rday) and maximum Rubisco carboxylation (Vcmax) in gymnosperms (Ginkgo biloba), conifers (Metasequoia glyptostroboides and Cryptomeria japonica), and angiosperms (Nicotiana tabacum and Phaseolus vulgaris). As predicted by the FvCB model, the slope of a linear function of Γ vs O2 partial pressure, d, increased alongside increasing Rday/Vcmax. The Sc/o was obtainable from this relationship between d and Rday/Vcmax, because the d values at Rday/Vcmax = 0 corresponded to α/Sc/o, where α was the photorespiratory CO2 release rate per Rubisco oxygenation rate (generally assumed to be 0.5). The calculated Sc/o values of N. tabacum and P. vulgaris exhibited good agreement with those reported by in vitro studies. The Sc/o values of both conifers were similar to those of the two angiosperm species. In contrast, the Sc/o value of G. biloba was significantly lower than those of the other four studied species. These results suggest that our new method for Sc/o estimation is applicable to C3 plants, including those for which in vitro kinetic analysis is difficult. Furthermore, results also suggest that conifer Sc/o does not differ significantly from that of C3 angiosperms, assuming α remains unchanged.

A dynamic model of RuBP-regeneration limited photosynthesis accounting for photoinhibition, heat and water stress

Plant physiological responses to stress and subsequent damage have not been successfully integrated into leaf, plant or land surface models. Water deficit leads to a series of plant responses: stomatal closure decreases evaporative cooling leading to higher leaf temperature, which it can affect photosynthetic dissipation of absorbed energy, especially under high irradiance. Excess absorbed energy by PSII results in photoinhibition, which is not typically included when modeling photosynthesis using the Farquhar-von Caemmerer-Berry (FvCB) model. We introduce a novel approach to represent plant-atmosphere interactions: a non-steady state model of photoinhibition linked to the RuBP-regeneration component of the FvCB model that accounts for the interactions between high temperature, irradiance, and water deficits. Gas exchange and chlorophyll fluorescence dynamics showed that, under severe water stress, the maximum quantum efficiency of PSII (Fv/Fm) quickly decreases, and it takes days to recover. We found a clear relationship between maximum electron transport rate and Fv/Fm and that short-term acclimation to irradiance affected photosynthetic FvCB model parameters. Fv/Fm functioned well as a coupling variable, able to scale RuBP-regeneration limited photosynthesis parameters proportional to photoinhibition. These results indicate that plant stress, photo-inhibitory damage, and recovery can be explicitly represented when modeling photosynthesis.

Sensitivity analysis and estimation using a hierarchical Bayesian method for the parameters of the FvCB biochemical photosynthetic model

Photosynthesis is a major process included in land surface models. Accurately estimating the parameters of the photosynthetic sub-models can greatly improve the ability of these models to accurately simulate the carbon cycle of terrestrial ecosystems. Here, we used a hierarchical Bayesian approach to fit the Farquhar-von Caemmerer-Berry model, which is based on the biochemistry of photosynthesis using 236 curves for the relationship between net CO2 assimilation and changes in the intercellular CO2 concentration. An advantage of the hierarchical Bayesian algorithm is that parameters can be estimated at multiple levels (plant, species, plant functional type, and population level) simultaneously. The parameters of the hierarchical strategy were based on the results of a sensitivity analysis. The Michaelis-Menten constant (Kc25), enthalpies of activation (EJ and EV), and two optical parameters (θ and α) demonstrated considerable variation at different levels, which suggests that this variation cannot be ignored. The maximum electron transport rate (Jmax25), maximum rate of Rubisco activity (Vcmax25), and dark respiration in the light (Rd25) were higher for broad-leaved plants than for needle-leaved plants. Comparison of the model's simulated outputs with observed data showed strong and significant positive correlations, particularly when the model was parameterized at the plant level. In summary, our study is the first effort to combine sensitivity analysis and hierarchical Bayesian parameter estimation. The resulting realistic parameter distributions for the four levels provide a reference for current and future land surface models. Furthermore, the observed variation in the parameters will require attention when using photosynthetic parameters in future models.

Parameter Estimation of the Farquhar—von Caemmerer—Berry Biochemical Model from Photosynthetic Carbon Dioxide Response Curves

The Farquhar—von Caemmerer—Berry (FvCB) biochemical model of photosynthesis, commonly used to estimate CO2 assimilation at various spatial scales from leaf to global, has been used to assess the impacts of climate change on crop and ecosystem productivities. However, it is widely known that the parameters in the FvCB model are difficult to accurately estimate. The objective of this study was to assess the methods of Sharkey et al. and Gu et al., which are often used to estimate the parameters of the FvCB model. We generated An/Ci datasets with different data accuracies, numbers of data points, and data point distributions. The results showed that neither method accurately estimated the parameters; however, Gu et al.’s approach provided slightly better estimates. Using Gu et al.’s approach and datasets with measurement errors and the same accuracy as a typical open gas exchange system (i.e., Li-6400), the majority of the estimated parameters—Vcmax (maximal Rubisco carboxylation rate), Kco (effective Michaelis-Menten coefficient for CO2), gm (internal (mesophyll) conductance to CO2 transport) and Γ* (chloroplastic CO2 photocompensation point)—were underestimated, while the majority of Rd (day respiration) and α (the non-returned fraction of the glycolate carbon recycled in the photorespiratory cycle) were overestimated. The distributions of Tp (the rate of triose phosphate export from the chloroplast) were evenly dispersed around the 1:1 line using both approaches. This study revealed that a high accuracy of leaf gas exchange measurements and sufficient data points are required to correctly estimate the parameters for the biochemical model. The accurate estimation of these parameters can contribute to the enhancement of food security under climate change through accurate predictions of crop and ecosystem productivities. A further study is recommended to address the question of how the measurement accuracies can be improved.

Plantecophys--An R Package for Analysing and Modelling Leaf Gas Exchange Data.

Here I present the R package 'plantecophys', a toolkit to analyse and model leaf gas exchange data. Measurements of leaf photosynthesis and transpiration are routinely collected with portable gas exchange instruments, and analysed with a few key models. These models include the Farquhar-von Caemmerer-Berry (FvCB) model of leaf photosynthesis, the Ball-Berry models of stomatal conductance, and the coupled leaf gas exchange model which combines the supply and demand functions for CO2 in the leaf. The 'plantecophys' R package includes functions for fitting these models to measurements, as well as simulating from the fitted models to aid in interpreting experimental data. Here I describe the functionality and implementation of the new package, and give some examples of its use. I briefly describe functions for fitting the FvCB model of photosynthesis to measurements of photosynthesis-CO2 response curves ('A-Ci curves'), fitting Ball-Berry type models, modelling C3 photosynthesis with the coupled photosynthesis-stomatal conductance model, modelling C4 photosynthesis, numerical solution of optimal stomatal behaviour, and energy balance calculations using the Penman-Monteith equation. This open-source package makes technically challenging calculations easily accessible for many users and is freely available on CRAN.

Modelling photosynthesis in flag leaves of winter wheat (Triticum aestivum) considering the variation in photosynthesis parameters during development.

The Farquhar-von Caemmerer-Berry (FvCB) model of photosynthesis has been widely used to estimate the photosynthetic C flux of plants under different growth conditions. However, the seasonal fluctuation of some photosynthesis parameters (e.g. the maximum carboxylation rate of Rubisco (Vcmax), the maximum electron transport rate (Jmax) and internal mesophyll conductance to CO2 transport (gm)) is not considered in the FvCB model. In this study, we investigated the patterns of the FvCB parameters during flag leaf development based on measured photosynthesis-intercellular CO2 curves in two cultivars of winter wheat (Triticum aestivum L.). Parameterised seasonal patterns of photosynthesis parameters in the FvCB model have subsequently been applied in order to predict the photosynthesis of flag leaves. The results indicate that the Gaussian curve characterises the dynamic patterns of Vcmax, Jmax and gm well. Compared with the model with fixed photosynthesis parameter values, updating the FvCB model by considering seasonal changes in Vcmax and Jmax during flag leaf development slightly improved predictions of photosynthesis. However, if the updated FvCB model incorporated the seasonal patterns of Vcmax and Jmax, and also of gm, predictions of photosynthesis was improved a lot, matching well with the measurements (R2=0.87, P<0.0001). This suggests that the dynamics of photosynthesis parameters, particularly gm, play an important role in estimating the photosynthesis rate of winter wheat.

Accounting for the decrease of photosystem photochemical efficiency with increasing irradiance to estimate quantum yield of leaf photosynthesis.

Maximum quantum yield for leaf CO2 assimilation under limiting light conditions (Φ CO2LL) is commonly estimated as the slope of the linear regression of net photosynthetic rate against absorbed irradiance over a range of low-irradiance conditions. Methodological errors associated with this estimation have often been attributed either to light absorptance by non-photosynthetic pigments or to some data points being beyond the linear range of the irradiance response, both causing an underestimation of Φ CO2LL. We demonstrate here that a decrease in photosystem (PS) photochemical efficiency with increasing irradiance, even at very low levels, is another source of error that causes a systematic underestimation of Φ CO2LL. A model method accounting for this error was developed, and was used to estimate Φ CO2LL from simultaneous measurements of gas exchange and chlorophyll fluorescence on leaves using various combinations of species, CO2, O2, or leaf temperature levels. The conventional linear regression method under-estimated Φ CO2LL by ca. 10-15%. Differences in the estimated Φ CO2LL among measurement conditions were generally accounted for by different levels of photorespiration as described by the Farquhar-von Caemmerer-Berry model. However, our data revealed that the temperature dependence of PSII photochemical efficiency under low light was an additional factor that should be accounted for in the model.

Improving the estimation of mesophyll conductance to CO₂: on the role of electron transport rate correction and respiration.

Mesophyll conductance (gm) can markedly limit photosynthetic CO2 assimilation and is required to estimate the parameters of the Farquhar–von Caemmerer–Berry (FvCB) model properly. The variable J (electron transport rate) is the most frequently used method for estimating gm, and the correct determination of J is one of its requirements. Recent evidence has shown that calibrating J can lead to some errors in estimating gm, but to what extent the parameterization of the FvCB model is affected by calibrations is not well known. In addition to determining the FvCB parameters, variants of the J calibration method were tested to address whether varying CO2 or light levels, possible alternative electron sinks, or contrasting leaf structural properties might play a role in determining differences in αβ, the product of the leaf absorptance (α) and the photosystem II optical cross-section (β). It was shown that differences in αβ were mainly attributed to the use of A/Ci or A/PPFD curves to calibrate J. The different αβ values greatly influenced gm, leading to a high number of unrealistic values in addition to affecting the estimates of the FvCB model parameters. A new approach was devised to retrieve leaf respiration in the light from combined A/Ci and A/Cc curves and a framework to understand the high variation in observed gm values. Overall, a background is provided to decrease the noise in gm, facilitating data reporting and allowing better retrieval of the information presented in A/Ci and A/Cc curves.

A microscale model for combined CO(2) diffusion and photosynthesis in leaves.

Transport of CO2 in leaves was investigated by combining a 2-D, microscale CO2 transport model with photosynthesis kinetics in wheat (Triticum aestivum L.) leaves. The biophysical microscale model for gas exchange featured an accurate geometric representation of the actual 2-D leaf tissue microstructure and accounted for diffusive mass exchange of CO2. The resulting gas transport equations were coupled to the biochemical Farquhar-von Caemmerer-Berry model for photosynthesis. The combined model was evaluated using gas exchange and chlorophyll fluorescence measurements on wheat leaves. In general a good agreement between model predictions and measurements was obtained, but a discrepancy was observed for the mesophyll conductance at high CO2 levels and low irradiance levels. This may indicate that some physiological processes related to photosynthesis are not incorporated in the model. The model provided detailed insight into the mechanisms of gas exchange and the effects of changes in ambient CO2 concentration or photon flux density on stomatal and mesophyll conductance. It represents an important step forward to study CO2 diffusion coupled to photosynthesis at the leaf tissue level, taking into account the leaf's actual microstructure.

Reliable estimation of biochemical parameters from C3 leaf photosynthesis–intercellular carbon dioxide response curves

The Farquhar-von Caemmerer-Berry (FvCB) model of photosynthesis is a change-point model and structurally overparameterized for interpreting the response of leaf net assimilation (A) to intercellular CO₂ concentration (Ci). The use of conventional fitting methods may lead not only to incorrect parameters but also several previously unrecognized consequences. For example, the relationships between key parameters may be fixed computationally and certain fits may be produced in which the estimated parameters result in contradictory identification of the limitation states of the data. Here we describe a new approach that is better suited to the FvCB model characteristics. It consists of four main steps: (1) enumeration of all possible distributions of limitation states; (2) fitting the FvCB model to each limitation state distribution by minimizing a distribution-wise cost function that has desirable properties for parameter estimation; (3) identification and correction of inadmissible fits; and (4) selection of the best fit from all possible limitation state distributions. The new approach implemented theoretical parameter resolvability with numerical procedures that maximally use the information content of the data. It was tested with model simulations, sampled A/Ci curves, and chlorophyll fluorescence measurements of different tree species. The new approach is accessible through the automated website leafweb.ornl.gov.