Nitrogen deposition limits photosynthetic response to elevated CO2 differentially in a dioecious species

Sexual dimorphisms of dioecious plants are important in controlling and maintaining sex ratios under changing climate environments. Yet, little is known about sex-specific responses to elevated CO₂ with soil nitrogen (N) deposition. To investigate sex-related physiological and biochemical responses to elevated CO₂ with N deposition, Populus cathayana Rehd. was employed as a model species. The cuttings were subjected to two CO₂ regimes (350 and 700???mol?mol^?1) with two N levels (0 and 5?g?N?m^?2?year^?1). Our results showed that elevated CO₂ and N deposition separately increased the total number of leaves, leaf area (LA), leaf mass, net photosynthetic rate (P _n), light saturated photosynthetic rate (P _max), chlorophyll a (Chl a), and chlorophyll a to chlorophyll b ratio (Chl a/b) in both males and females of P. cathayana. However, the effects on LA, leaf mass, P _n, P _max, Chl a and Chl a/b were weakened under the combined treatment of elevated CO₂ and N deposition. Males had higher leaf mass, P _n, P _max, apparent quantum yield (??), carboxylation efficiency (CE), Chl a, Chl a/b, leaf N, and root carbon to N ratio (C/N) than did females under elevated CO₂ with N deposition. In contrast to males, females had significantly higher levels of soluble sugars in leaves and greater starch accumulation in roots and stems under the same condition. The results of the present work imply that P. cathayana females are more responsive and suffer from greater negative effects on growth and photosynthetic capacity than do males when grown under elevated CO₂ with soil N deposition.     

CO2 environment,microclimate and photosynthetic characteristics of the moss Hylocomium splendens in a subarctic habitat

Summary In order to document the natural CO₂ environment of the moss Hylocomium splendens, and ascertain whether or not the moss was adapted to this, and its interactions with other microenvironmental factors, two studies were carried out. Firstly, the seasonal variations of CO₂ concentration, photosynthetically active radiation (PAR), tissue water content and temperature were measured in the natural microenvironment of H. splendens in a subarctic forest during the summer period (July–September). Secondly, the photosynthetic responses of the species to controlled CO₂ concentrations, PAR, temperature, and hydration were measured in the laboratory. CO₂ concentrations around the upper parts of the plant, when PAR was above the compensation point (30 mol m^–2 s^–1), were mostly between 400 and 450 ppm. They occasionally increased up to 1143 ppm for short periods. PAR flux densities below saturating light levels for photosynthesis (100 mol m^–2 s^–1), occurred during 65% (July), 76% (August) and 96% (September) of the hours of the summer period. The temperature optimum of photosynthesis was 20° C: this temperature coincided with PAR above the compensation point during 5%, 6% and 0% of the time in July, August and September, respectively. Optimal hydration of tissues was infrequent. Hence PAR, temperature and water limit CO₂ uptake for most of the growing season. Our data suggest that the higher than normal ambient CO₂ concentration in the immediate environment of the plant counteracts some of the limitations in PAR supply that it experiences in its habitat. This species already experiences concentrations of atmospheric CO₂ predicted to occur over the next 50 years.     

Evidence for a physiological role of CO2 in the regulation of photosynthetic electron transport in intact leaves

The effect of CO₂ upon photosynthetic electron transport was examined in wheat and maize leaves in order to establish whether CO₂ had a direct role in electron-transport regulation in vivo. When intercellular CO₂ was depleted a transient rise in chlorophyll fluorescence occurred which correlated with an increase in the reduction of the Photosystem II primary quinone acceptor, Q_A, and a decrease in CO₂ fixation rate. However, when intercellular CO₂ was reduced from an already low level (50 μmol·mol⁻¹) towards zero a substantial further reduction in Q_A occurred with little change in fluorescence or CO₂ fixation. In very low intercellular CO₂ when no measurable CO₂ fixation was sustained, an appreciable fraction of Q_A still remained oxidised, however, maximal reduction of Q_A occurred when O₂ was also removed. Q_A could then be substantially reoxidised by the readdition of small amounts of CO₂ (20–40 μmol) which only facilitated a very small increase in CO₂ fixation. Changes in the kinetics of the fast rise in fluorescence emission indicated that Q_A-to-Q_B electron transfer was decreased in a CO₂-free atmosphere and Q_B was poised in a more oxidised state. Electron transport that was independent of CO₂ fixation was measured in methyl viologen-treated leaf discs. In 1% O₂, but not in 21% O₂, light-dependent electron transport to methyl viologen was decreased significantly by the depletion of CO₂. It is concluded that CO₂ can modify the redox state of Photosystem II electron transport acceptors in vivo independently of carbon assimilation and that there is a complex interaction between CO₂ and O₂ in the regulation of photosynthetic electron transport. The possibility that CO₂ operates via the reversible binding to PS II and thereby acts as a cofactor for efficient PS II electron transport in the leaf is discussed.     

A biochemical model of photosynthetic CO2 assimilation in leaves of C3 species

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 carboxylase-oxygenase; 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 CO₂ assimilation rate on p(CO₂) and irradiance, and the influence of p(CO₂) and irradiance on the temperature dependence of assimilation rate.Abbreviations RuP₂ ribulose bisphosphate - PGA 3-phosphoglycerate - C=p(CO₂) partial pressure of CO₂ - O=p(O₂) partial pressure of O₂ - PCR photosynthetic carbon reduction - PCO photorespiratory carbon oxidation     

Interactive effects of elevated CO2 and drought on photosynthetic capacity and PSII performance in maize

Elevated atmospheric CO₂ concentration [CO₂] and the change of water distribution in arid and semiarid areas affect plant physiology and ecosystem processes. The interaction of elevated [CO₂] and drought results in the complex response such as changes in the energy flux of photosynthesis. The performance of photosystem (PS) II and the electron transport were evaluated by using OJIP induction curves of chlorophyll a fluorescence and the P _N-C _i curves in the two-factor controlled experiment with [CO₂] of 380 (AC) or 750 (EC) [μmol mol^?1] and water stress by 10% polyethylene glycol 6000. Compared to water-stressed maize (Zea mays L.) under AC, the EC treatment combined with water stress decreased the number of active reaction centers but it increased the antenna size and the energy flux (absorbed photon flux, trapping flux, and electron transport flux) of each reaction center in PSII. Thus, the electron transport rate was enhanced, despite the indistinctively changed quantum yield of the electron transport and energy dissipation. The combination of EC and the water-stress treatment resulted in the robust carboxylation rate without elevating the saturated photosynthetic rate (P _max). This study demonstrated that maize was capable of transporting more electrons into the carboxylation reaction, but this could not be used to increase P _max under EC.     

Diurnal changes in the response of canopy photosynthetic rate to elevated CO2 in a coupled temperature-light environment

The relative increase with elevated CO₂ of canopy CO₂ uptake rate (A), derived from continuous measurements during the day, was examined in full-cover vegetative Lolium perenne canopies after 17 days of regrowth. The stands were grown at ambient (358±50 mol mol^-1) and increased (626±50 mol mol^-1) CO₂ concentration in sunlit growth chambers. Over the entire range of temperature and light conditions (which were strongly coupled and increased simultaneously), A was on average twice as large in high compared to ambient CO₂. This response (called M=A in high CO₂/A in ambient CO₂) could not be explained by changes in canopy conductance for CO₂ diffusion (GC). In spite of interaction and strong coupling between temperature and light intensity, there was evidence that temperature rather than light determined M. Further, high CO₂ treatment was found to alleviate the afternoon depression in A observed in ambient CO₂. A temperature optimum shift or/and a larger carbohydrate sink capacity through altered root/shoot ratio are proposed in explanation.Abbreviations A CO₂ uptake rate - C350 ambient CO₂ treatment - C600 elevated CO₂ treatment - E canopy evapotranspiration rate - GC canopy conductance for CO₂ diffusion - M high CO₂ modification factor     

Expression and regulation of Bradyrhizobium japonicum and Xanthobacter flavus CO2 fixation genes in a photosynthetic bacterial host.

Calvin cycle carbon dioxide fixation genes encoded on DNA fragments from two nonphotosynthetic, chemolithoautotrophic bacteria, Bradyrhizobium japonicum and Xanthobacter flavus, were found to complement and support photosynthetic growth of a ribulose 1,5-bisphosphate carboxylase-oxygenase (RubisCO) deletion mutant of the purple nonsulfur bacterium Rhodobacter sphaeroides. The regulation of RubisCO expression was analyzed in the complemented R. sphaeroides RubisCO deletion mutant. Distinct differences in the regulation of RubisCO synthesis were revealed when the complemented R. sphaeroides strains were cultured under photolithoautotrophic and photoheterotrophic growth conditions, e.g., a reversal in the normal pattern of RubisCO gene expression. These studies suggest that sequences and molecular signals which regulate the expression of diverse RubisCO genes may be probed by using the R. sphaeroides complementation system.     

Nutrient availability and atmospheric CO2 partial pressure modulate the effects of nutrient heterogeneity on the size structure of populations in grassland species

BACKGROUND AND AIMS: Size-asymmetric competition occurs when larger plants have a disproportionate advantage in competition with smaller plants. It has been hypothesized that nutrient heterogeneity may promote it. Experiments testing this hypothesis are inconclusive, and in most cases have evaluated the effects of nutrient heterogeneity separately from other environmental factors. The aim of this study was to test, using populations of Lolium perenne, Plantago lanceolata and Holcus lanatus, two hypotheses: (a) nutrient heterogeneity promotes size-asymmetric competition; and (b) nutrient heterogeneity interacts with both atmospheric CO2 partial pressure (P(CO2)) and nutrient availability to determine the magnitude of this response. METHODS: Microcosms consisting of monocultures of the three species were grown for 90 d in a factorial experiment with the following treatments: P(CO2) (37.5 and 70 Pa) and nutrient availability (NA; 40 and 120 mg of N added as organic material) combined with different spatial distribution of the organic material (NH; homogeneous and heterogeneous). Differences in the size of individual plants within populations (size inequality) were quantified using the coefficient of variation of individual above-ground biomass and the combined biomass of the two largest individuals in each microcosm. Increases in size inequality were associated with size-asymmetric competition. KEY RESULTS: Size inequality increased when the nutrients were heterogeneously supplied in the three species. The effects of NH on this response were more pronounced under high nutrient supply in both Plantago and Holcus (significant NA x NH interactions) and under elevated P(CO2) in Plantago (significant P(CO2) x NA x NH interaction). No significant two- and three-way interactions were found for Lolium. CONCLUSIONS: Our first hypothesis was supported by our results, as nutrient heterogeneity promoted size-asymmetric competition in the three species evaluated. Nutrient supply and P(CO2) modified the magnitude of this effect in Plantago and Holcus, but not in Lolium. Thus, our second hypothesis was partially supported.     

Effects of temperature and CO2 concentration on photosynthetic CO2 fixation by Chlorella

The rates of photosynthetic ¹⁴CO₂ fixation by Chlorella vulgarisllh, grown under high CO₂, were determined between 4 to 37°Cwith air containing from 300 to 13,000 ppm ¹⁴CO₂. When the CO₂level was increased, both the rate of photosynthesis and theoptimum temperature for maximum photosynthesis increased. Themaximum photosynthetic rate was reached at 12°C with 300ppm ^l4CO₂. Among the photosynthetic products fromed at 300 ppm ¹⁴CO₂, glycolatedecreased greatly when the temperature was raised from 20 to30°C. At 3,000 ppm ¹⁴CO₂ an insignificant amount of glycolatewas formed at all temperatures, whereas ¹⁴C-incorporation intothe insoluble fraction, sucrose, and the lipid fraction wassignificantly higher than at 300 ppm ¹⁴CO₂. The ¹⁴C in sucrosewas greatly increased and the radioactivity in the insolublefraction decreased when the temperature was raised from 28 to36°C. (Received April 8, 1980; )     

Coordination between leaf CO2 diffusion and Rubisco properties allows maximizing photosynthetic efficiency in Limonium species

High photosynthetic efficiency intrinsically demands tight coordination between traits related to CO₂ diffusion capacity and leaf biochemistry. Although this coordination constitutes the basis of existing mathematical models of leaf photosynthesis, it has been barely explored among closely related species, which could reveal rapid adaptation clues in the recent past. With this aim, we characterized the photosynthetic capacity of 12 species of Limonium, possessing contrasting Rubisco catalytic properties, grown under optimal (WW) and extreme drought conditions (WD). The availability of CO₂ at the site of carboxylation (C_c) determined the photosynthetic capacity of Limonium under WD, while both diffusional and biochemical components governed the photosynthetic performance under WW. The variation in the in vivo caboxylation efficiency correlated with both the concentration of active Rubisco sites and the in vitro‐based properties of Rubisco, such as the maximum carboxylase turnover rate (k_cat^c) and the Michaelis–Menten constant for CO₂ (K_c). Notably, the results confirmed the hypothesis of coordination between the CO₂ offer and demand functions of photosynthesis: those Limonium species with high total leaf conductance to CO₂ have evolved towards increased velocity (i.e. higher k_cat^c), at the penalty of lower affinity for CO₂ (i.e. lower specificity factor, S_c/o).     

Engineering of photosynthetic mannitol biosynthesis from CO2 in a cyanobacterium

d-Mannitol (hereafter denoted mannitol) is used in the medical and food industry and is currently produced commercially by chemical hydrogenation of fructose or by extraction from seaweed. Here, the marine cyanobacterium Synechococcus sp. PCC 7002 was genetically modified to photosynthetically produce mannitol from CO₂ as the sole carbon source. Two codon-optimized genes, mannitol-1-phosphate dehydrogenase (mtlD) from Escherichia coli and mannitol-1-phosphatase (mlp) from the protozoan chicken parasite Eimeria tenella, in combination encoding a biosynthetic pathway from fructose-6-phosphate to mannitol, were expressed in the cyanobacterium resulting in accumulation of mannitol in the cells and in the culture medium. The mannitol biosynthetic genes were expressed from a single synthetic operon inserted into the cyanobacterial chromosome by homologous recombination. The mannitol biosynthesis operon was constructed using a novel uracil-specific excision reagent (USER)-based polycistronic expression system characterized by ligase-independent, directional cloning of the protein-encoding genes such that the insertion site was regenerated after each cloning step. Genetic inactivation of glycogen biosynthesis increased the yield of mannitol presumably by redirecting the metabolic flux to mannitol under conditions where glycogen normally accumulates. A total mannitol yield equivalent to 10% of cell dry weight was obtained in cell cultures synthesizing glycogen while the yield increased to 32% of cell dry weight in cell cultures deficient in glycogen synthesis; in both cases about 75% of the mannitol was released from the cells into the culture medium by an unknown mechanism. The highest productivity was obtained in a glycogen synthase deficient culture that after 12 days showed a mannitol concentration of 1.1 g mannitol L⁻¹ and a production rate of 0.15 g mannitol L⁻¹ day⁻¹. This system may be useful for biosynthesis of valuable sugars and sugar derivatives from CO₂ in cyanobacteria.     

Nutrient availability and management in the rhizosphere: exploiting genotypic differences

Crop nutrition is frequently inadequate as a result of the expansion of cropping into marginal lands, elevated crop yields placing increasing demands on soil nutrient reserves, and environmental and economic concerns about applying fertilizers. Plants exposed to nutrient deficiency activate a range of mechanisms that result in increased nutrient availability in the rhizosphere compared with the bulk soil. Plants may change their root morphology, increase the affinity of nutrient transporters in the plasma membrane and exude organic compounds (carboxylates, phenolics, carbohydrates, enzymes, etc.) and protons. Chemical changes in the rhizosphere result in altered abundance and composition of microbial communities. Nutrient-efficient genotypes are adapted to environments with low nutrient availability. Nutrient efficiency can be enhanced by targeted breeding through pyramiding efficiency mechanisms in a desirable genotype as well as by gene transfer and manipulation. Rhizosphere microorganisms influence nutrient availability; adding beneficial microorganisms may result in enhanced availability of nutrients to crops. Understanding the role of plant-microbe-soil interactions in governing nutrient availability in the rhizosphere will enhance the economic and environmental sustainability of crop production.     

Long-term effects of a doubled atmospheric CO2 concentration on the CAM species Agave deserti

To examine the effects of a doubled atmospheric CO₂ concentrationand other aspects of global climate change on a common CAM speciesnative to the Sonoran Desert, Agave deserti was grown under370 and 750 µmol CO₂ mol^–1 air and gas exchangewas measured under various environmental conditions. Doublingthe CO₂ concentration increased daily net CO₂ uptake by 49%throughout the 17 months and decreased daily transpiration by24%, leading to a 110% increase in water-use efficiency. Underthe doubled CO₂ concentration, the activity of ribulose-1,5-bisphosphatecarboxylase/oxygenase (Rubisco) was 11% lower, phosphoenolpyruvatecarboxylase was 34% lower, and the activated:total ratio forRubisco was 25% greater than under the current CO₂ concentration.Less leaf epicuticular wax occurred on plants under the doubledCO₂ concentration, which decreased the reflectance of photosyntheticphoton flux (PPF); the chlorophyll content per unit leaf areawas also less. The enhancement of daily net CO₂ uptake by doublingthe CO₂ concentration increased when the PPF was decreased below25 mol m^–2 d^–1 when water was withheld, and whenday/night temperatures were below 17/12 C. More leaves, eachwith a greater surface area, were produced per plant under thedoubled CO₂ concentration. The combination of increased totalleaf surface area and increased daily net CO₂ uptake led toan 88% stimulation of dry mass accumulation under the doubledCO₂ concentration. A rising atmospheric CO₂ concentration, togetherwith accompanying changes in temperature, precipitation, andPPF, should increase growth and productivity of native populationsof A. deserti. Key words: Crassulacean acid metabolism, gas exchange, global climate change, Sonoran Desert     

Temperature effects on the photosynthetic response of C3 plants to long-term CO2 enrichment

To assess the long-term effect of increased CO₂ and temperature on plants possessing the C₃ photosynthetic pathway, Chenopodium album plants were grown at one of three treatment conditions: (1) 23 °C mean day temperature and a mean ambient partial pressure of CO₂ equal to 350 bar; (2) 34 °C and 350 bar CO₂; and (3) 34 °C and 750 bar CO₂. No effect of the growth treatments was observed on the CO₂ reponse of photosynthesis, the temperature response of photosynthesis, the content of Ribulose-1,5-bisphosphate carboxylase (Rubisco), or the activity of whole chain electron transport when measurements were made under identical conditions. This indicated a lack of photosynthetic acclimation in C. album to the range of temperature and CO₂ used in the growth treatments. Plants from every treatment exhibited similar interactions between temperature and CO₂ on photosynthetic activity. At low CO₂ (< 300 bar), an increase in temperature from 25 to 35 °C was inhibitory for photosynthesis, while at elevated CO₂ (> 400 bar), the same increase in temperature enhanced photosynthesis by up to 40%. In turn, the stimulation of photosynthesis by CO₂ enrichment increased as temperature increased. Rubisco capacity was the primary limitation on photosynthetic activity at low CO₂ (195 bar). As a consequence, the temperature response of A was relatively flat, reflecting a low temperature response of Rubisco at CO₂ levels below its k_m for CO₂. At elevated CO₂ (750 bar), the temperature response of electron transport appeared to control the temperature dependency of photosynthesis above 18 °C. These results indicate that increasing CO₂ and temperature could substantially enhance the carbon gain potential in tropical and subtropical habitats, unless feedbacks at the whole plant or ecosystem level limit the long-term response of photosynthesis to an increase in CO₂ and temperature.Abbreviations A net CO₂ assimilation rate - C _a ambient partial pressure of CO₂ - C _i intercellular partial pressure of CO₂ - Rubisco Ribulose-1,5-bisphosphate carboxylase - VPD vapor pressure difference between leaf and air     

Flowering in grassland predicted by CO2 and resource effects on species aboveground biomass

Continuing enrichment of atmospheric CO₂ may change plant community composition, in part by altering the availability of other limiting resources including soil water, nutrients, or light. The combined effects of CO₂ enrichment and altered resource availability on species flowering remain poorly understood. We quantified flowering culm and ramet production and biomass allocation to flowering culms/ramets for 10 years in C₄‐dominated grassland communities on contrasting soils along a CO₂ concentration gradient spanning pre‐industrial to expected mid‐21st century levels (250–500 μl/L). CO₂ enrichment explained up to 77% of the variation in flowering culm count across soils for three of the five species, and was correlated with flowering culm count on at least one soil for four of five species. In contrast, allocation to flowering culms was only weakly correlated with CO₂ enrichment for two species. Flowering culm counts were strongly correlated with species aboveground biomass (AGB; R² = .34–.74), a measure of species abundance. CO₂ enrichment also increased soil moisture and decreased light levels within the canopy but did not affect soil inorganic nitrogen availability. Structural equation models fit across the soils suggested species‐specific controls on flowering in two general forms: (1) CO₂ effects on flowering culm count mediated by canopy light level and relative species AGB (species AGB/total AGB) or by soil moisture effects on flowering culm count; (2) effects of canopy light level or soil inorganic nitrogen on flowering and/or relative species AGB, but with no significant CO₂ effect. Understanding the heterogeneity in species responses to CO₂ enrichment in plant communities across soils in edaphically variable landscapes is critical to predict CO₂ effects on flowering and other plant fitness components, and species potential to adapt to future environmental changes.     

Effects of CO2 concentration during growth on subsequent photosynthetic CO2 fixation in Chlorella1

The maximum rate of photosynthetic ¹⁴CO₂ fixation (V_max) aswell as the concentration of CO₂ at which the rate of photosynthetic¹⁴CO₂ fixation attains one-half its maximum velocity (K_m) inChlorella vulgaris 11h cells was strongly dependent on the concentrationof CO₂ continuously provided during the algal growth. The V_max (µmoles ¹⁴CO₂ fixed/ml pcv?min) and K_m (% CO₂)of the algal cells which had been grown in air containing 4%CO₂ (by volume) were ca. 10 and 0.15–0.17, while thosein the cells which had been grown in ordinary air (containing0.04% CO₂) were 7 and 0.05–0.06, respectively. When the concentration of CO₂ in the bubbling gas was loweredfrom 4 to 0.04% during the algal growth, their photosynthetickinetics attained the respective lower steady levels after 5–10hr. On the other hand, when the photosynthetic kinetics weredetermined 24 hr after raising the concentration of CO₂ from0.04 to 4%, the V_max and K_m-values were found to have alreadyattained the respective higher levels. (Received October 15, 1976; )