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Oxygen isotopic composition (Δ18OLW) of leaf water can help improve our understanding of how anatomy interacts with physiology to influence leaf water transport. Leaf water isotope models of Δ18OLW such as the Péclet effect model have been developed to predict Δ18OLW, and it incorporates transpiration rate (E) and the mixing length between unenriched xylem water and enriched mesophyll water, which can occur in the mesophyll (Lm) or veins (Lv). Here we used two cell wall composition mutants grown under two light intensities and RH to evaluate the effect of cell wall composition on Δ18OLW. In maize (Zea mays), the compromised ultrastructure of the suberin lamellae in the bundle sheath of the ALIPHATIC SUBERIN FERULOYL TRANSFERASE mutant (Zmasft) reduced barriers to apoplastic water movement, resulting in higher E and Lv and, consequently, lower Δ18OLW. In cellulose synthase-like F6 (Cslf6) mutants and wildtype of rice (Oryza sativa), the difference in Δ18OLW in plants grown under high and low growth light intensity co-varied with their differences in stomatal density. These results show that cell wall composition and stomatal density influence Δ18OLW by altering the Péclet effect and that stable isotopes can facilitate the development of a physiologically and anatomically explicit water transport model.

In habitats with low water availability, a fundamental challenge for plants will be to maximize photosynthetic C-gain while minimizing transpirational water-loss. This trade-off between C-gain and water-loss can in part be achieved through the coordination of leaf-level photosynthetic and hydraulic traits. To test the relationship of photosynthetic C-gain and transpirational water-loss, we grew, under common growth conditions, 18 C4 grasses adapted to habitats with different mean annual precipitation (MAP) and measured leaf-level structural and anatomical traits associated with mesophyll conductance (gm ) and leaf hydraulic conductance (Kleaf ). The C4 grasses adapted to lower MAP showed greater mesophyll surface area exposed to intercellular air spaces (Smes ) and adaxial stomatal density (SDada ) which supported greater gm . These grasses also showed greater leaf thickness and vein-to-epidermis distance, which may lead to lower Kleaf . Additionally, grasses with greater gm and lower Kleaf also showed greater photosynthetic rates (Anet ) and leaf-level water-use efficiency (WUE). In summary, we identify a suite of leaf-level traits that appear important for adaptation of C4 grasses to habitats with low MAP and may be useful to identify C4 species showing greater Anet and WUE in drier conditions.

Photosynthesis in C(3) -C(4) intermediates reduces carbon loss by photorespiration through refixing photorespired CO(2) within bundle sheath cells. This is beneficial under warm temperatures where rates of photorespiration are high; however, it is unknown how photosynthesis in C(3) -C(4) plants acclimates to growth under cold conditions. Therefore, the cold tolerance of the C(3) -C(4) Salsola divaricata was tested to determine whether it reverts to C(3) photosynthesis when grown under low temperatures. Plants were grown under cold (15/10 °C), moderate (25/18 °C) or hot (35/25 °C) day/night temperatures and analysed to determine how photosynthesis, respiration and C(3) -C(4) features acclimate to these growth conditions. The CO(2) compensation point and net rates of CO(2) assimilation in cold-grown plants changed dramatically when measured in response to temperature. However, this was not due to the loss of C(3) -C(4) intermediacy, but rather to a large increase in mitochondrial respiration supported primarily by the non-phosphorylating alternative oxidative pathway (AOP) and, to a lesser degree, the cytochrome oxidative pathway (COP). The increase in respiration and AOP capacity in cold-grown plants likely protects against reactive oxygen species (ROS) in mitochondria and photodamage in chloroplasts by consuming excess reductant via the alternative mitochondrial respiratory electron transport chain.

Biochemical models are used to predict and understand the response of photosynthesis to rising temperatures and CO2 partial pressures. These models require the temperature dependency of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) kinetics and mesophyll conductance to CO2 (g(m)). However, it is not known how the temperature response of Rubisco kinetics differs between species, and comprehensive in vivo Rubisco kinetics that include gm have only been determined in the warm-adapted Nicotiana tabacum. Here, we measured the temperature response of Rubisco kinetics and gm in N. tabacum and the cold-adapted Arabidopsis thaliana using gas exchange and (13)CO2 isotopic discrimination on plants with genetically reduced levels of Rubisco. While the individual Rubisco kinetic parameters in N. tabacum and A. thaliana were similar across temperatures, they collectively resulted in significantly different modelled rates of photosynthesis. Additionally, gm increased with temperature in N. tabacum but not in A. thaliana. These findings highlight the importance of considering species-dependent differences in Rubisco kinetics and gm when modelling the temperature response of photosynthesis.

The efficiency of C(4) photosynthesis in Zea mays, Miscanthus x giganteus and Flaveria bidentis in response to light was determined using measurements of gas exchange, (13) CO(2) photosynthetic discrimination, metabolite pools and spectroscopic assays, with models of C(4) photosynthesis and leaf (13) CO(2) discrimination. Spectroscopic and metabolite assays suggested constant energy partitioning between the C(4) and C(3) cycles across photosynthetically active radiation (PAR). Leakiness (φ), modelled using C(4) light-limited photosynthesis equations (φ(mod)), matched values from the isotope method without simplifications (φ(is)) and increased slightly from high to low PAR in all species. However, simplifications of bundle-sheath [CO(2)] and respiratory fractionation lead to large overestimations of φ at low PAR with the isotope method. These species used different strategies to maintain similar φ. For example, Z. mays had large rates of the C(4) cycle and low bundle-sheath cells CO(2 ) conductance (g(bs)). While F. bidentis had larger g(bs) but lower respiration rates and M. giganteus had less C(4) cycle capacity but low g(bs), which resulted in similar φ. This demonstrates that low g(bs) is important for efficient C(4) photosynthesis but it is not the only factor determining φ. Additionally, these C(4) species are able to optimize photosynthesis and minimize φ over a range of PARs, including low light.

Photosynthetic carbon gain in plants using the C(3) photosynthetic pathway is substantially inhibited by photorespiration in warm environments, particularly in atmospheres with low CO(2) concentrations. Unlike C(4) plants, C(3) plants are thought to lack any mechanism to compensate for the loss of photosynthetic productivity caused by photorespiration. Here, for the first time, we demonstrate that the C(3) plants rice and wheat employ a specific mechanism to trap and reassimilate photorespired CO(2) . A continuous layer of chloroplasts covering the portion of the mesophyll cell periphery that is exposed to the intercellular air space creates a diffusion barrier for CO(2) exiting the cell. This facilitates the capture and reassimilation of photorespired CO(2) in the chloroplast stroma. In both species, 24-38% of photorespired and respired CO(2) were reassimilated within the cell, thereby boosting photosynthesis by 8-11% at ambient atmospheric CO(2) concentration and 17-33% at a CO(2) concentration of 200 µmol mol(-1) . Widespread use of this mechanism in tropical and subtropical C(3) plants could explain why the diversity of the world's C(3) flora, and dominance of terrestrial net primary productivity, was maintained during the Pleistocene, when atmospheric CO(2) concentrations fell below 200 µmol mol(-1) .

Differences in light quality penetration within a leaf and absorption by the photosystems alter rates of CO2 assimilation in C3 plants. It is also expected that light quality will have a profound impact on C4 photosynthesis due to disrupted coordination of the C4 and C3 cycles. To test this hypothesis, we measured leaf gas exchange, 13CO2 discrimination (Δ13C), photosynthetic metabolite pools and Rubisco activation state in Zea mays and Miscanthus × giganteus under steady-state red, green, blue and white light. Photosynthetic rates, quantum yield of CO2 assimilation, and maximum phosphoenolpyruvate carboxylase activity were significantly lower under blue light than white, red and green light in both species. However, similar leakiness under all light treatments suggests the C4 and C3 cycles were coordinated to maintain the photosynthetic efficiency. Measurements of photosynthetic metabolite pools also suggest coordination of C4 and C3 cycles across light treatments. The energy limitation under blue light affected both C4 and C3 cycles, as we observed a reduction in C4 pumping of CO2 into bundle-sheath cells and a limitation in the conversion of C3 metabolite phosphoglycerate to triose phosphate. Overall, light quality affects rates of CO2 assimilation, but not the efficiency of CO2 concentrating mechanism.

The photosynthetic efficiency of the CO(2)-concentrating mechanism in two forms of single-cell C(4) photosynthesis in the family Chenopodiaceae was characterized. The Bienertioid-type single-cell C(4) uses peripheral and central cytoplasmic compartments (Bienertia sinuspersici), while the Borszczowioid single-cell C(4) uses distal and proximal compartments of the cell (Suaeda aralocaspica). C(4) photosynthesis within a single-cell raises questions about the efficiency of this type of CO(2) -concentrating mechanism compared with the Kranz-type. We used measurements of leaf CO(2) isotope exchange (Δ(13) C) to compare the efficiency of the single-cell and Kranz-type forms of C(4) photosynthesis under various temperature and light conditions. Comparisons were made between the single-cell C(4) and a sister Kranz form, S. eltonica[NAD malic enzyme (NAD ME) type], and with Flaveria bidentis[NADP malic enzyme (NADP-ME) type with Kranz Atriplicoid anatomy]. There were similar levels of Δ(13) C discrimination and CO(2) leakiness (Φ) in the single-cell species compared with the Kranz-type. Increasing leaf temperature (25 to 30 °C) and light intensity caused a decrease in Δ(13) C and Φ across all C(4) types. Notably, B. sinuspersici had higher Δ(13) C and Φ than S. aralocaspica under lower light. These results demonstrate that rates of photosynthesis and efficiency of the CO(2) -concentrating mechanisms in single-cell C(4) plants are similar to those in Kranz-type.

Carbon and oxygen isotope discrimination of CO(2) during photosynthesis (Δ(13)C(obs) and Δ(18)O(obs)) were measured along a monocot leaf, triticale (Triticum × Secale). Both Δ(13)C(obs) and Δ(18)O(obs) increased towards the leaf tip. While this was expected for Δ(18)O(obs) , because of progressive enrichment of leaf water associated with the Peclet effect, the result was surprising for Δ(13) C(obs). To explore parameters determining this pattern, we measured activities of key photosynthetic enzymes [ribulose bis-phosphate carboxylase-oxygenase (Rubisco), phosphoenolpyruvate carboxylase (PEPC) and carbonic anhydrase) as well as maximum carboxylation and electron transport rates (V(cmax) and J(max)) along the leaf. Patterns in leaf internal anatomy along the leaf were also quantified. Mesophyll conductance (g(m)) is known to have a strong influence on Δ(13)C(obs) , so we used three commonly used estimation methods to quantify variation in g(m) along the leaf. Variation in Δ(13)C(obs) was correlated with g(m) and chloroplast surface area facing the intercellular air space, but unrelated to photosynthetic enzyme activity. The observed variation could cause errors at higher scales if the appropriate portion of a leaf is not chosen for leaf-level measurements and model parameterization. Our study shows that one-third of the way from the base of the leaf represents the most appropriate portion to enclose in the leaf chamber.

Measurements of CO 2 and O 2 gas exchange and chlorophyll a fluorescence were used to test the hypothesis that elevated atmospheric CO 2 inhibits nitrate (NO 3 ‐ ) photo-assimilation in the C 4 plant, maize ( Zea mays L.). The assimilatory quotient ( AQ ), the ratio of net CO 2 assimilation to net O 2 evolution, decreases as NO 3 ‐ photo-assimilation increases so that the difference in AQ between the ammonium- and nitrate-fed plants ( D D D AQ ) provided an in planta estimate of NO 3 ‐ photo-assimilation. In fully expanded maize leaves, NO 3 ‐ photo-assimilation was detectable only under high light and was not affected by CO 2 treatments. Furthermore, CO 2 assimilation and O 2 evolution were higher under NO 3 ‐ than ammonia (NH 4 + + + ) regardless of CO 2 levels. In conclusion, NO 3 ‐ photo-assimilation in maize primarily occurred at high light when reducing equivalents were presumably not limiting. Nitrate photo-assimilation enhanced C 4 photosynthesis, and in contrast to C 3 plants, elevated CO 2 did not inhibit foliar NO 3 ‐ photo-assimilation.

The present study was carried out to test the hypothesis that elevated atmospheric CO 2 (Ca) will alleviate over-excitation of the C 4 photosynthetic apparatus and decrease non-photochemical quenching (NPQ) during periods of limited water availability. Chlorophyll a fluorescence was monitored in Sorghum bicolor plants grown under a freeair carbon-dioxide enrichment (FACE) by water-stress (Dry) experiment. Under Dry conditions elevated Ca increased the quantum yield of photosystem II ( f PSII) throughout the day through increases in both photochemical quenching coefficient ( q p ) and the efficiency with which absorbed quanta are transferred to open PSII reaction centres ( F v ¢¢¢ ¢ / F m ¢¢ ). However, in the well-watered plants (Wets) FACE enhanced f PSII only at midday and was entirely attributed to changes in F v ¢¢¢ ¢ / F m ¢¢ . Under field conditions, decreases in f f f PSII under Dry treatments and ambient Ca corresponded to increases in NPQ but the de-epoxidation state of the xanthophyll pool (DPS) showed no effects. Water-stress did not lead to long-term damage to the photosynthetic apparatus as indicated by f PSII and carbon assimilation measured after removal of stress conditions. We conclude that elevated Ca enhances photochemical light energy usage in C 4 photosynthesis during drought and/ or midday conditions. Additionally, NPQ protects against photo-inhibition and photodamage. However, NPQ and the xanthophyll cycle were affected differently by elevated Ca and water-stress.