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Allometric relationships among the dimensions of leaves and their cells hold across diverse eudicotyledons, but have remained untested in the leaves of grasses. We hypothesised that geometric (proportional) allometries of cell sizes across tissues and of leaf dimensions would arise due to the coordination of cell development and that of cell functions such as water, nutrient and energy transport, and that cell sizes across tissues would be associated with light‐saturated photosynthetic rate. We tested predictions across 27 globally distributed C3 and C4 grass species grown in a common garden. We found positive relationships among average cell sizes within and across tissues, and of cell sizes with leaf dimensions. Grass leaf anatomical allometries were similar to those of eudicots, with exceptions consistent with the fewer cell layers and narrower form of grass leaves, and the specialised roles of epidermis and bundle sheath in storage and leaf movement. Across species, mean cell sizes in each tissue were associated with light‐saturated photosynthetic rate per leaf mass, supporting the functional coordination of cell sizes. These findings highlight the generality of evolutionary allometries within the grass lineage and their interlinkage with coordinated development and function. Summary statement: Allometries among leaf cell sizes and of cell sizes with leaf dimensions and photosynthetic rate hold across grass species, supporting their basis in developmental and functional coordination. [ABSTRACT FROM AUTHOR]

Incorporating C4 photosynthetic traits into C3 crops is a rational approach for sustaining future demands for crop productivity. Using classical plant breeding, engineering this complex trait is unlikely to achieve its target. Therefore, it is critical and timely to implement novel biotechnological crop improvement strategies to accomplish this goal. However, a fundamental understanding of C3, C4, and C3–C4 intermediate metabolism is crucial for the targeted use of biotechnological tools. This review assesses recent progress towards engineering C4 photosynthetic traits in C3 crops. We also discuss lessons learned from successes and failures of recent genetic engineering attempts in C3 crops, highlighting the pros and cons of using rice as a model plant for short‐, medium‐ and long‐term goals of genetic engineering. This review provides an integrated approach towards engineering improved photosynthetic efficiency in C3 crops for sustaining food, fibre and fuel production around the globe. [ABSTRACT FROM AUTHOR]

Summary: C2 photosynthesis is a carbon concentrating mechanism that can increase net CO2 assimilation by capturing, concentrating and re‐assimilating CO2 released by photorespiration. Empirical and modelling studies indicate that C2 plants assimilate more carbon than C3 plants under high temperature, bright light, and low CO2 conditions. I argue that engineering C2 photosynthesis into C3 crops is a promising approach to improve photosynthetic performance under these – and temporally heterogeneous – environments, and review the modifications that may re‐create a C2 phenotype in C3 plants. Although a C2 engineering program would encounter many of the same challenges faced by C4 engineering programmes, the simpler leaf anatomical requirements make C2 engineering a feasible approach to improve crops in the medium term. [ABSTRACT FROM AUTHOR]

Membrane permeabilities to CO2 and HCO3− constrain the function of CO2 concentrating mechanisms that algae use to supply inorganic carbon for photosynthesis. In diatoms and green algae, plasma membranes are moderately to highly permeable to CO2 but effectively impermeable to HCO3−. Here, CO2 and HCO3− membrane permeabilities were measured using an 18O‐exchange technique on two species of haptophyte algae, Emiliania huxleyi and Calcidiscus leptoporus, which showed that the plasma membranes of these species are also highly permeable to CO2 (0.006–0.02 cm · s−1) but minimally permeable to HCO3−. Increased temperature and CO2 generally increased CO2 membrane permeabilities in both species, possibly due to changes in lipid composition or CO2 channel proteins. Changes in CO2 membrane permeabilities showed no association with the density of calcium carbonate coccoliths surrounding the cell, which could potentially impede passage of compounds. Haptophyte plasma‐membrane permeabilities to CO2 were somewhat lower than those of diatoms but generally higher than membrane permeabilities of green algae. One caveat of these measurements is that the model used to interpret 18O‐exchange data assumes that carbonic anhydrase, which catalyzes 18O‐exchange, is homogeneously distributed in the cell. The implications of this assumption were tested using a two‐compartment model with an inhomogeneous distribution of carbonic anhydrase to simulate 18O‐exchange data and then inferring plasma‐membrane CO2 permeabilities from the simulated data. This analysis showed that the inferred plasma‐membrane CO2 permeabilities are minimal estimates but should be quite accurate under most conditions. [ABSTRACT FROM AUTHOR]

The transition of C3, via C2 towards C4 photosynthesis is an important example of stepwise evolution of a complex genetic trait. A common feature that was gradually emphasized during this trajectory is the evolution of a CO2 concentration mechanism around Rubisco. In C2 plants, this mechanism is based on tissue- specific accumulation of glycine decarboxylase (GDC) in bundle sheath (BS) cells, relative to global expression in the cells of C3 leaves. This limits photorespiratory CO2 release to BS cells. Because BS cells are surrounded by photosynthetically active mesophyll cells, this arrangement enhances the probability of re-fixation of CO2. The restriction of GDC to BS cells was mainly achieved by confinement of its P-subunit (GLDP). Here, we provide a mechanism for the establishment of C2-type gene expression by studying the upstream sequences of C3 Gldp genes in Arabidopsis thaliana. Deletion of 59 bp in the upstream region of AtGldp1 restricted expression of a reporter gene to BS cells and the vasculature without affecting diurnal variation. This region was named the 'M box'. Similar results were obtained for the AtGldp2 gene. Fusion of the M box to endogenous or exogenous promoters supported mesophyll expression. Nucleosome densities at the M box were low, suggesting an open chromatin structure facilitating transcription factor binding. In silico analysis defined a possible consensus for the element that was conserved across the Brassicaceae, but not in Moricandia nitens, a C2 plant. Collective results provide evidence that a simple mutation is sufficient for establishment of C2-specific gene expression in a C3 plant. [ABSTRACT FROM AUTHOR]

Oxygenic photosynthesis would not be possible without photorespiration in the present day O2-rich atmosphere. It is now generally accepted that cyanobacteria-like prokaryotes first evolved oxygenic photosynthesis, which was later conveyed via endosymbiosis into a eukaryotic host, which then gave rise to the different groups of algae and streptophytes. For photosynthetic CO2 fixation, all these organisms use Rubis CO, which catalyses both the carboxylation and the oxygenation of ribulose 1,5-bisphosphate. One of the reaction products of the oxygenase reaction, 2-phosphoglycolate (2 PG), represents the starting point of the photorespiratory C2 cycle, which is considered largely responsible for recapturing organic carbon via conversion to the Calvin-Benson cycle ( CBC) intermediate 3-phosphoglycerate, thereby detoxifying critical intermediates. Here we discuss possible scenarios for the evolution of this process toward the well-defined 2 PG metabolism in extant plants. While the origin of the C2 cycle core enzymes can be clearly dated back towards the different endosymbiotic events, the evolutionary scenario that allowed the compartmentalised high flux photorespiratory cycle is uncertain, but probably occurred early during the algal radiation. The change in atmospheric CO2/ O2 ratios promoting the acquisition of different modes for inorganic carbon concentration mechanisms, as well as the evolutionary specialisation of peroxisomes, clearly had a dramatic impact on further aspects of land plant photorespiration. [ABSTRACT FROM AUTHOR]

1. The ecophysiological significance of Crassulacean acid metabolism (CAM) in the invasive aquatic macrophyte Crassula helmsii was studied in an English soft-water lake. The extent and the contribution of CAM to the carbon budget was examined in spring (April) and summer (July) along a depth gradient (0.5–2.2 m), covering the growth range of C. helmsii in the lake. 2. Significant in situ CAM activity (30–80 meq kg−1 FW) was present in all specimens, although it decreased with depth and hence correlated with the decline in photon irradiance. Potential CAM activity (60–161 meq kg−1 FW), measured after exposure to low concentrations of CO2 in the day and high concentrations at night, were on average 2.7-times greater than in situ CAM activity. Overall CAM activity increased from April to July, which is consistent with higher potential carbon limitation caused by increased temperature and light availability. 3. CAM activity in C. helmsii appeared to be carbon-limited at night because night-time carbon-fixation increased at raised, compared to ambient, concentrations of CO2. 4. The high in situ CAM activity in C. helmsii was reflected in the contribution of CAM to the total carbon budget which, independent of depth and season, ranged from 18% to 42%. The amount of CO2 taken up in the night via CAM was 0.74 to 2.94 times the amount of CO2 lost in respiration, thus emphasizing the importance of CAM in refixation of potentially lost respiratory CO2. 5. The onset of decarboxylation in the morning appeared to be under circadian control as there was a delay of up to 5.5 h between the start of the light period and a decline in cell acidity level. 6. There was little variation in δ13C content (−21.69 to 23.49‰) with season or depth suggesting, along with the estimated contribution to the carbon-budget, that CAM is important for the whole population of C. helmsii. CAM may confer a competitive advantage in relation to growth, which may be one of the reasons for the invasiveness of this species. [ABSTRACT FROM AUTHOR]

Estimates of the effect of increased global atmospheric CO2 levels on oceanic primary productivity depend on the physiological responses of contemporary phytoplankton populations. However, microalgal populations will possibly adapt to rising CO2 levels in such a way that they become genetically different from contemporary populations. The unknown properties of these future populations introduce an undefined error into predictions of C pool dynamics, especially the presence and size of the biological C pump. To address the bias in predictions introduced by evolution, we measured the kinetics of CO2 uptake in populations of Chlamydomonas reinhardtii that had been selected for growth at high CO2 for 1000 generations. Following selection at high CO2, the populations were unable to induce high-affinity CO2 uptake, and one line had a lower rate of net CO2 uptake. We attribute this to conditionally neutral mutations in genes affecting the C concentrating mechanism (CCM). Lower affinity CO2 uptake, in addition to smaller population sizes, results in a significant reduction in net CO2 uptake of about 38% relative to contemporary populations under the same conditions. This shows how predictions about the properties of communities in the future can be influenced by the effect of natural selection. [ABSTRACT FROM AUTHOR]

Estimates of the effect of increased global atmospheric CO2 levels on oceanic primary productivity depend on the physiological responses of contemporary phytoplankton populations. However, microalgal populations will possibly adapt to rising CO2 levels in such a way that they become genetically different from contemporary populations. The unknown properties of these future populations introduce an undefined error into predictions of C pool dynamics, especially the presence and size of the biological C pump. To address the bias in predictions introduced by evolution, we measured the kinetics of CO2 uptake in populations of Chlamydomonas reinhardtii that had been selected for growth at high CO2 for 1000 generations. Following selection at high CO2, the populations were unable to induce high-affinity CO2 uptake, and one line had a lower rate of net CO2 uptake. We attribute this to conditionally neutral mutations in genes affecting the C concentrating mechanism (CCM). Lower affinity CO2 uptake, in addition to smaller population sizes, results in a significant reduction in net CO2 uptake of about 38% relative to contemporary populations under the same conditions. This shows how predictions about the properties of communities in the future can be influenced by the effect of natural selection. [ABSTRACT FROM AUTHOR]

Cyanobacteria show high metabolic plasticity by re-allocating macromolecular resources in response to variations in both environmental inorganic carbon (Ci) and light. We grew cultures of the picoplanktonic cyanobacterium Synechococcus elongatus Nägeli across a 50-fold range of growth irradiance at either a dissolved [Ci] <0.1 mM, sufficient to induce strongly the carbon-concentrating mechanism (CCM) or a dissolved [Ci] of ∼4 mM, sufficient to strongly induce the CCM to basal constitutive activity. There was no detectable growth cost of acclimation to low Ci across the entire range of irradiance and growth was nearly light saturated at 50 l mol photons·m−2·s−1. Cells acclimated to low Ci significantly re-allocated macromolecular resources to support their CCM, while maintaining near homeostatis of metabolic flux per unit photosynthetic complex. Changing growth irradiance also drove re-organization of the photosynthetic machinery to balance excitation flux and metabolic demands, but flux per complex varied widely across the range of tolerable growth irradiances. Across the range of growth irradiance, low Ci cells had significantly less phycocyanin than high Ci cells, which corresponded to a lower PSII absorbance capacity. Furthermore, low Ci cells maintained more PSI per cell−1 than high Ci cells under high growth irradiance. Low Ci cells could therefore maintain more of their PSII reaction centers open at high growth irradiance than could high Ci cells, which experienced a significant PSII closure. Thus, acclimation to growth under high available Ci actually constrained acclimation to high light by restricting electron transport downstream from PSII in S. elongatus. [ABSTRACT FROM AUTHOR]

ABSTRACT Physiological properties of photosynthesis were determined in the marine diatom, Phaeodactylum tricornutum UTEX640, during acclimation from 5% CO2 to air and related to H2CO3 dissociation kinetics and equilibria in artificial seawater. The concentration of dissolved inorganic carbon at half maximum rate of photosynthesis (K0·5[DIC]) value in high CO2-grown cells was 1009 mmol m-3 but was reduced three-fold by the addition of bovine carbonic anhydrase (CA), whereas in air-grown cells K0·5[DIC] was 71 mmol m-3, irrespective of the presence of CA. The maximum rate of photosynthesis (Pmax) values varied between 300 and 500 μmol O2 mg Chl-1 h-1 regardless of growth pCO2. Bicarbonate dehydration kinetics in artificial seawater were re-examined to evaluate the direct HCO3- uptake as a substrate for photosynthesis. The uncatalysed CO2 formation rate in artificial seawater of 31·65°/oo of salinity at pH 8·2 and 25 °C was found to be 0·6 mmol m-3 min-1 at 100 mmol m-3 DIC, which is 53·5 and 7·3 times slower than the rates of photosynthesis exhibited in air- and high CO2-grown cells, respectively. These data indicate that even high CO2-grown cells of P. tricornutum can take up both CO2 and HCO3- as substrates for photosynthesis and HCO3- use improves dramatically when the cells are grown in air. Detailed time courses were obtained of changes in affinity for DIC during the acclimation of high CO2-grown cells to air. The development of high-affinity photosynthesis started after a 2–5 h lag period, followed by a steady increase over the next 15 h. This acclimation time course is the slowest to be described so far. High CO2-grown cells were transferred to controlled DIC conditions, at which the concentrations of each DIC species could be defined, and were allowed to acclimate for more than 36 h. The K0·5[DIC] values in acclimated cells appeared to be correlated only with [CO2(aq)] in the medium but not to HCO3-, CO32-, total [DIC] or the pH of the medium and indicate that the critical signal regulating the affinity of cells for DIC in the marine diatom, P. tricornutum, is [CO2(aq)] in the medium. [ABSTRACT FROM AUTHOR]