Abstract.Various aspects of the biochemistry of photosynthetic carbon assimilation in C3 plants are integrated into a form compatible with studies of gas exchange in leaves. These aspects include the kinetic properties of ribulose bisphosphate carboxylaseoxygenase; the requirements of the photosynthetic carbon reduction and photorespiratory carbon oxidation cycles for reduced pyridine nucleotides; the dependence of electron transport on photon flux and the presence of a temperature dependent upper limit to electron transport. The measurements of gas exchange with which the model outputs may be compared include those of the temperature and partial pressure of CO2(p(CO2) ) dependencies of quantum yield, the variation of compensation point with temperature and partial pressure of O2(p(O2)), the dependence of net CO 2 assimilation rate on p(CO2) and irradiance, and the influence of p(CO2) and irradiance on the temperature dependence of assimilation rate.

A series of experiments is presented investigating short term and long term changes of the nature of the response of rate of CO2 assimilation to intercellular p(CO2). The relationships between CO2 assimilation rate and biochemical components of leaf photosynthesis, such as ribulose-bisphosphate (RuP2) carboxylase-oxygenase activity and electron transport capacity are examined and related to current theory of CO2 assimilation in leaves of C3 species. It was found that the response of the rate of CO2 assimilation to irradiance, partial pressure of O2, p(O2), and temperature was different at low and high intercellular p(CO2), suggesting that CO2 assimilation rate is governed by different processes at low and high intercellular p(CO2). In longer term changes in CO2 assimilation rate, induced by different growth conditions, the initial slope of the response of CO2 assimilation rate to intercellular p(CO2) could be correlated to in vitro measurements of RuP2 carboxylase activity. Also, CO2 assimilation rate at high p(CO2) could be correlated to in vitro measurements of electron transport rate. These results are consistent with the hypothesis that CO2 assimilation rate is limited by the RuP2 saturated rate of the RuP2 carboxylase-oxygenase at low intercellular p(CO2) and by the rate allowed by RuP2 regeneration capacity at high intercellular p(CO2).

CO 2 transfer conductance from the intercellular airspaces of the leaf into the chloroplast, defined as mesophyll conductance (g m ), is finite. Therefore, it will limit photosynthesis when CO 2 is not saturating, as in C3 leaves in the present atmosphere. Little is known about the processes that determine the magnitude of g m . The process dominating g m is uncertain, though carbonic anhydrase, aquaporins, and the diffusivity of CO 2 in water have all been suggested. The response of g m to temperature (10°C-40°C) in mature leaves of tobacco (Nicotiana tabacum L. cv W38) was determined using measurements of leaf carbon dioxide and water vapor exchange, coupled with modulated chlorophyll fluorescence. These measurements revealed a temperature coefficient (Q 10 ) of approximately 2.2 for g m , suggesting control by a protein-facilitated process because the Q 10 for diffusion of CO 2 in water is about 1.25. Further, g m values are maximal at 35°C to 37.5°C, again suggesting a protein-facilitated process, but with a lower energy of deactivation than Rubisco. Using the temperature response of g m to calculate CO 2 at Rubisco, the kinetic parameters of Rubisco were calculated in vivo from 10°C to 40°C. Using these parameters, we determined the limitation imposed on photosynthesis by g m . Despite an exponential rise with temperature, g m does not keep pace with increased capacity for CO 2 uptake at the site of Rubisco. The fraction of the total limitations to CO 2 uptake within the leaf attributable to g m rose from 0.10 at 10°C to 0.22 at 40°C. This shows that transfer of CO 2 from the intercellular air space to Rubisco is a very substantial limitation on photosynthesis, especially at high temperature.In C3 plants, the diffusion of CO 2 from the atmosphere to the active site of Rubisco follows a complex pathway involving as many as eight discrete conductance components (Nobel, 1999). Most commonly, this pathway is simplified into three main components: boundary layer, stomatal conductance, and mesophyll conductance (g m ; Farquhar and Sharkey, 1982). Boundary layer conductance depends on several leaf physical and environmental properties, in particular, size, surface structures, stomatal location, and air movement around the leaf, whereas stomatal conductance is primarily influenced by stomatal pore numbers and dimensions. The flexible and dynamic qualities of the stomatal pores provide the leaf with physiological control of CO 2 influx and water efflux (Farquhar and Sharkey, 1982). Estimates of boundary layer and stomatal conductances to CO 2 are based on water vapor released from the leaf because water and CO 2 share the same gaseous diffusion pathway (e.g. von Caemmerer and Farquhar, 1981). As a result, it has long been known that limitations of diffusion through the stomata and boundary layer are purely physical (Penman and Schofield, 1951). g m , defined as the conductance of CO 2 transfer from the intercellular leaf airspaces to the site of carboxylation, was initially assumed large enough to have a negligible...

The world's crop productivity is stagnating whereas population growth, rising affluence, and mandates for biofuels put increasing demands on agriculture. Meanwhile, demand for increasing cropland competes with equally crucial global sustainability and environmental protection needs. Addressing this looming agricultural crisis will be one of our greatest scientific challenges in the coming decades, and success will require substantial improvements at many levels. We assert that increasing the efficiency and productivity of photosynthesis in crop plants will be essential if this grand challenge is to be met. Here, we explore an array of prospective redesigns of plant systems at various scales, all aimed at increasing crop yields through improved photosynthetic efficiency and performance. Prospects range from straightforward alterations, already supported by preliminary evidence of feasibility, to substantial redesigns that are currently only conceptual, but that may be enabled by new developments in synthetic biology. Although some proposed redesigns are certain to face obstacles that will require alternate routes, the efforts should lead to new discoveries and technical advances with important impacts on the global problem of crop productivity and bioenergy production.light capture/conversion | carbon capture/conversion | smart canopy | enabling plant biotechnology tools | sustainable crop production Increasing demands for global food production over the next several decades portend a huge burden on the world's shrinking farmlands. Increasing global affluence, population growth, and demands for a bioeconomy (including livestock feed, bioenergy, chemical feedstocks, and biopharmaceuticals) will all require increased agricultural productivity, perhaps by as much as 60-120% over 2005 levels (e.g., refs. 1 and 2), putting increased productivity on a collision course with environmental and sustainability goals (3). The 45 y from 1960 to 2005 saw global food production grow ∼160%, mostly (135%) by improved production on

SummaryAccurate representation of photosynthesis in terrestrial biosphere models (TBMs) is essential for robust projections of global change. However, current representations vary markedly between TBMs, contributing uncertainty to projections of global carbon fluxes. Here we compared the representation of photosynthesis in seven TBMs by examining leaf and canopy level responses of photosynthetic CO 2 assimilation (A) to key environmental variables: light, temperature, CO 2 concentration, vapor pressure deficit and soil water content. We identified research areas where limited process knowledge prevents inclusion of physiological phenomena in current TBMs and research areas where data are urgently needed for model parameterization or evaluation. We provide a roadmap for new science needed to improve the representation of photosynthesis in the next generation of terrestrial biosphere and Earth system models.

Measurements of CO2 and water vapour exchange by leaves were combined with measurements of carbon isotope composition (13C/12C) of CO2 in the air passing over the leaf. Carbon isotope discrimination during CO2 uptake was determined from the difference in carbon isotope composition of the air leaving the leaf chamber with or without a leaf enclosed. Leaves of wheat plants grown with different nitrogen nutrition and leaves of several other species were examined. The measurements, made at different irradiances for a given leaf, showed that carbon isotope discrimination was strongly correlated with the rate of CO2 assimilation as well as the ratio of intercellular to ambient partial pressure of CO2, pI/pa. A function relating carbon isotope discrimination to the rate of CO2 assimilation was used to estimate the CO2 transfer conductance, gw, from the substomatal cavities to the sites of carboxylation for individual leaves. The photosynthetic capacity correlated with the CO2 transfer conductance, gw, and the average ratio of chloroplastic to intercellular partial pressure of CO2, pI/pa, was 0.7. This means that in general under high irradiance, the ratio of chloroplastic to ambient partial pressure of CO2 is about 0.5. In wheat, variation in gw was correlated with the chloroplast surface area appressing intercellular airspaces.

1077I.1078II.1079III.1080IV.1081V.1084VI.1087VII.10881089References1089 Summary The rate of CO2 assimilation by plants is directly influenced by the concentration of CO2 in the atmosphere, ca. As an environmental variable, ca also has a unique global and historic significance. Although relatively stable and uniform in the short term, global ca has varied substantially on the timescale of thousands to millions of years, and currently is increasing at seemingly an unprecedented rate. This may exert profound impacts on both climate and plant function. Here we utilise extensive datasets and models to develop an integrated, multi‐scale assessment of the impact of changing ca on plant carbon dioxide uptake and water use. We find that, overall, the sensitivity of plants to rising or falling ca is qualitatively similar across all scales considered. It is characterised by an adaptive feedback response that tends to maintain 1 − ci/ca, the relative gradient for CO2 diffusion into the leaf, relatively constant. This is achieved through predictable adjustments to stomatal anatomy and chloroplast biochemistry. Importantly, the long‐term response to changing ca can be described by simple equations rooted in the formulation of more commonly studied short‐term responses.

The partial pressure of CO2 at the sites of carboxylation within chloroplasts depends on the conductance to CO2 diffusion from intercellular airspace to the sites of carboxylation, termed mesophyll conductance (gm). We investigated the temperature response of gm in tobacco (Nicotiana tabacum) by combining gas exchange in high light, ambient CO2 in either 2 or 21% O2 with carbon isotope measurements using tuneable diode laser spectroscopy. The gm increased linearly with temperature in 2 or 21% O2. In 21% O2, isotope discrimination associated with gm decreased from 5.0 Ϯ 0.2 to 1.8 Ϯ 0.2‰ as temperature increased from 15 to 40°C, but the photorespiratory contribution to the isotopic signal is significant. While the fractionation factor for photorespiration ( f = 16.2 Ϯ 0.7‰) was independent of temperature between 20 and 35°C, discrimination associated with photorespiration increased from 1.1 Ϯ 0.01 to 2.7 Ϯ 0.02‰ from 15 to 40°C. Other mitochondrial respiration contributed around 0.2 Ϯ 0.03‰. The drawdown in CO2 partial pressure from ambient air to intercellular airspaces was nearly independent of leaf temperature. By contrast, the increase in gm with increasing leaf temperature resulted in the drawdown in CO2 partial pressure between intercellular airspaces and the sites of carboxylation decreasing substantially at high temperature.