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      A Microscale Model for Combined CO 2 Diffusion and Photosynthesis in Leaves

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          Abstract

          Transport of CO 2 in leaves was investigated by combining a 2-D, microscale CO 2 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 CO 2. 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 CO 2 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 CO 2 concentration or photon flux density on stomatal and mesophyll conductance. It represents an important step forward to study CO 2 diffusion coupled to photosynthesis at the leaf tissue level, taking into account the leaf's actual microstructure.

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          Most cited references40

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          Fitting photosynthetic carbon dioxide response curves for C(3) leaves.

          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.
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            Mesophyll conductance to CO2: current knowledge and future prospects.

            During photosynthesis, CO2 moves from the atmosphere (C(a)) surrounding the leaf to the sub-stomatal internal cavities (C(i)) through stomata, and from there to the site of carboxylation inside the chloroplast stroma (C(c)) through the leaf mesophyll. The latter CO2 diffusion component is called mesophyll conductance (g(m)), and can be divided in at least three components, that is, conductance through intercellular air spaces (g(ias)), through cell wall (g(w)) and through the liquid phase inside cells (g(liq)). A large body of evidence has accumulated in the past two decades indicating that g(m) is sufficiently small as to significantly decrease C(c) relative to C(i), therefore limiting photosynthesis. Moreover, g(m) is not constant, and it changes among species and in response to environmental factors. In addition, there is now evidence that g(liq) and, in some cases, g(w), are the main determinants of g(m). Mesophyll conductance is very dynamic, changing in response to environmental variables as rapid or even faster than stomatal conductance (i.e. within seconds to minutes). A revision of current knowledge on g(m) is presented. Firstly, a historical perspective is given, highlighting the founding works and methods, followed by a re-examination of the range of variation of g(m) among plant species and functional groups, and a revision of the responses of g(m) to different external (biotic and abiotic) and internal (developmental, structural and metabolic) factors. The possible physiological bases for g(m), including aquaporins and carbonic anhydrases, are discussed. Possible ecological implications for variable g(m) are indicated, and the errors induced by neglecting g(m) when interpreting photosynthesis and carbon isotope discrimination models are highlighted. Finally, a series of research priorities for the near future are proposed.
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              Theoretical Considerations when Estimating the Mesophyll Conductance to CO(2) Flux by Analysis of the Response of Photosynthesis to CO(2).

              The conductance for CO(2) diffusion in the mesophyll of leaves can limit photosynthesis. We have studied two methods for determining the mesophyll conductance to CO(2) diffusion in leaves. We generated an ideal set of photosynthesis rates over a range of partial pressures of CO(2) in the stroma and studied the effect of altering the mesophyll diffusion conductance on the measured response of photosynthesis to intercellular CO(2) partial pressure. We used the ideal data set to test the sensitivity of the two methods to small errors in the parameters used to determine mesophyll conductance. The two methods were also used to determine mesophyll conductance of several leaves using measured rather than ideal data sets. It is concluded that both methods can be used to determine mesophyll conductance and each method has particular strengths. We believe both methods will prove useful in the future.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, USA )
                1932-6203
                2012
                7 November 2012
                : 7
                : 11
                : e48376
                Affiliations
                [1 ]Flanders Center of Postharvest Technology/BIOSYST-MeBioS, Katholieke Universiteit Leuven, Leuven, Belgium
                [2 ]Centre for Crop Systems Analysis, Wageningen University, Wageningen, The Netherlands
                Colorado State University, United States of America
                Author notes

                Competing Interests: Wageningen based authors have contributed to this work within the programme BioSolar Cells. Quang Tri Ho is a postdoctoral fellow of the Research Fund Flanders (FWO Vlaanderen). This does not alter the authors‚ adherence to all the PLOS ONE policies on sharing data and materials.

                Conceived and designed the experiments: BMN QTH PV XY PCS. Performed the experiments: XY PCS. Analyzed the data: QTH PV BMN. Contributed reagents/materials/analysis tools: BMN QTH PV XY PCS. Wrote the paper: BMN QTH PV XY PCS.

                Article
                PONE-D-12-09782
                10.1371/journal.pone.0048376
                3492360
                23144870
                584a4f6c-1292-4961-b3a5-eae971d03ae7
                Copyright @ 2012

                This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 6 April 2012
                : 24 September 2012
                Page count
                Pages: 15
                Funding
                The authors wish to thank the Research Council of the K.U. Leuven (OT 08/023), the Research Fund Flanders (project G.0603.08), and the Institute for the Promotion of Innovation by Science and Technology in Flanders (project IWT-050633) for financial support. Wageningen based authors have contributed to this work within the programme BioSolar Cells. Quang Tri Ho is a postdoctoral fellow of the Research Fund Flanders (FWO Vlaanderen). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Research Article
                Biology
                Biochemistry
                Plant Biochemistry
                Ecology
                Ecological Metrics
                Photosynthetic Efficiency
                Plant Ecology
                Plant-Environment Interactions
                Plant Science
                Plant Biochemistry
                Photosynthesis
                Plant Ecology
                Plant-Environment Interactions
                Plants
                Leafs
                Plant Physiology

                Uncategorized
                Uncategorized

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