Photosynthetic acclimation to CO2 enrichment related to ribulose-1,5-bisphosphate carboxylation limitation in wheat

Net photosynthetic rate (P _N) measured at the same CO₂ concentration, the maximum in vivo carboxylation rate, and contents of ribulose-1,5-bisphosphate (RuBP) carboxylase/oxygenase (RuBPCO) and RuBPCO activase were significantly decreased, but the maximum in vivo electron transport rate and RuBP content had no significant change in CO₂-enriched [EC, about 200 μmol mol⁻¹ above the ambient CO₂ concentration (AC)] wheat leaves compared with those in AC grown wheat leaves. Hence photosynthetic acclimation in wheat leaves to EC is largely due to RuBP carboxylation limitation.     

Photosynthetic acclimation in rice leaves to free-air CO2 enrichment related to both ribulose-1,5-bisphosphate carboxylation limitation and ribulose-1,5-bisphosphate regeneration limitation

Net photosynthetic rates (Pns) in leaves were compared between rice plants grown in ambient air control and free-air CO2 enrichment (FACE, about 200 micromol mol(-1) above ambient) treatment rings. When measured at the same CO2 concentration, the Pn of FACE leaves decreased significantly, indicating that photosynthetic acclimation to high CO2 occurs. Although stomatal conductance (Gs) in FACE leaves was markedly decreased, intercellular CO2 concentrations (Ci) were almost the same in FACE and ambient leaves, indicating that the photosynthetic acclimation is not caused by the decreased Gs. Furthermore, carboxylation efficiency and maximal Pn, both light and CO2-saturated Pn, were decreased in FACE leaves, as shown by the Pn-Ci curves. In addition, the soluble protein, Rubisco (ribulose-1,5-bisphosphate caboxylase/oxygenase), and its activase contents as well as the sucrose-phosphate synthase activity decreased significantly, while some soluble sugar, inorganic phosphate, chlorophyll and light-harvesting complex II (LHC II) contents increased in FACE leaves. It appears that the photosynthetic acclimation in rice leaves is related to both ribulose-1,5-bisphosphate (RuBP) carboxylation limitation and RuBP regeneration limitation.     

Is photosynthetic acclimation to free-air CO<Subscript>2</Subscript> enrichment (FACE) related to a strong competition for the assimilatory power between carbon assimilation and nitrogen assimilation in rice leaf?

Net photosynthetic rate (P _N) of leaves grown under free-air CO₂ enriched condition (FACE, about 200 μmol mol⁻¹ above ambient air) was significantly lower than P _N of leaves grown at ambient CO₂ concentration (AC) when measured at CO₂ concentration of 580 μmol mol⁻¹. This difference was found in rice plants grown at normal nitrogen supply (25 g m⁻²; NN-plants) but not in plants grown at low nitrogen supply (15 g m⁻²; LN-plants). Namely, photosynthetic acclimation to FACE was observed in NN-plants but not in LN-plants. Different from the above results measured in a period of continuous sunny days, such photosynthetic acclimation occurred in NN-plants, however, it was also observed in LN-plants when P _N was measured before noon of the first sunny day after rain. Hence strong competition for the assimilatory power between nitrogen (N) and carbon (C) assimilations induced by an excessive N supply may lead to the photosynthetic acclimation to FACE in NN-plants. The hypothesis is supported by the following facts: FACE induced significant decrease in both apparent photosynthetic quantum yield (Φ_c) and ribulose-1,5-bisphosphate (RuBP) content in NN-plants but not in LN-plants.     

Changes in leaf photosynthesis of transgenic rice with silenced <Emphasis Type="Italic">OsBP-73</Emphasis> gene

In comparison with its wild type (WT), the transgenic (TG) rice with silenced OsBP-73 gene had significantly lower plant height, grain number per panicle, and leaf net photosynthetic rate (P _N). Also, the TG rice showed significantly lower chlorophyll (Chl), ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO), RuBPCO activase, and RuBP contents, photosystem 2 (PS2) photochemical efficiency (F_v/F_m and ΔF/F_m′), apparent quantum yield of carbon assimilation (Φ_c), carboxylation efficiency (CE), photosynthetic electron transport and photophosphorylation rates as well as sucrose phosphate synthase activity, but higher intercellular CO₂ concentration, sucrose, fructose, and glycerate 3-phosphate contents, and non-photochemical quenching of Chl fluorescence (NPQ). Thus the decreased P _N in the TG rice leaves is related to both RuBP carboxylation and RuBP regeneration limitations, and the latter is a predominant limitation to photosynthesis.     

Influence of Etherel and Gibberellic Acid on Carbon Metabolism,Growth, and Essential Oil Accumulation in Spearmint (Mentha Spicata)

Controlled environment chamber and glasshouse studies were conducted on six herbaceous annual species grown at 350 (AC) and 700 (EC) mol(CO₂) mol^-1 to determine whether growth at EC resulted in acclimation of the apparent quantum yield of photosynthesis (QY) measured at limiting photosynthetic photon flux density (PPFD), or in acclimation of net photosynthetic rate (P _N) measured at saturating PPFD. It was also determined whether acclimation in P _N at limiting PPFD was correlated with acclimation of carboxylation efficiency or ribulose-1,5-bisphosphate (RuBP) regeneration rate measured at saturating PPFD. Growth at EC reduced both the QY and P _N at limiting PPFD in three of the six species. The occurrence of photosynthetic acclimation measured at a rate limiting PPFD was independent of whether photosynthetic acclimation was apparent at saturating measurement PPFD. At saturating measurement PPFD, acclimation to EC in the apparent carboxylation efficiency and RuBP regeneration capacity also occurred independently. Thus at least three components of the photosynthetic system may adjust independently when leaves are grown at EC. Estimates of photosynthetic acclimation at both high and low PPFD are necessary to accurately predict photosynthesis at the whole plant or canopy level as [CO₂] increases.     

The relationship between carbon-dioxide-limited photosynthetic rate and ribulose-1,5-bisphosphate-carboxylase content in two nuclear-cytoplasm substitution lines of wheat,and the coordination of ribulose-bisphosphate-carboxylation and electron-transport capacities

Photosynthesis in two cultivars of Triticum aestivum was compared with photosynthesis in two lines having the same nuclear genomes but with cytoplasms derived from T. boeoticum. The in-vitro specific activity of ribulose-1,5-bisphosphate carboxylase (RuBPCase; EC 4.1.1.39) isolated from lines with T. boeoticum cytoplasm was only 71% of that of normal T. aestivum. By contrast, the RuBPCase activities calculated from the CO₂-assimilation rate at low partial pressures of CO₂, p(CO₂), were the same for all lines for a given RuBPCase content. This indicates that both types of RuBPCase have the same turnover numbers in-vivo of 27.5 mol CO₂·(mol enzyme)^–1·s^–1 (23°). The rate of CO₂ assimilation measured at normal p(CO₂), p _a =340 bar, and high irradiance could be quantitatively predicted from the amount of RuBPCase protein. The maximum rate of RuBP regeneration could also predict the rate of CO₂ assimilation at normal ambient conditions. Therefore, the maximum capacities for RuBP carboxylation and RuBP regeneration appear to be well-balanced for normal ambient conditions. As photosynthetic capacity declined with increasing leaf age, the capacities for RuBP carboxylation and RuBP regeneration declined in parallel.Abbreviations PAR photosynthetically active radiation - RuBP(Case) ribulose-1,5-bisphosphate (carboxylase)     

Use of an analytical model to study limitations on net photosynthesis in Arbutus unedo under field conditions

Summary From field gas-exchange measurements on Arbutus unedo growing in Portugal, parameter values necessary to apply an analytical, physiologicallybased model of C ₃ photosynthesis were obtained. The model successfully simulated measured diurnal photosynthetic responses in Arbutus during periods without water stress, under both natural and CO₂-saturating conditions. The model was used to analyze those factors limiting primary productivity during each of the experimental days. Due to a large investment in ribulose bisphosphate (RuBP) regeneration capacity, irradiance was rarely limiting, even during cloudy periods, but the limitation imposed by stomatal conductance was quite large, averaging over 30%. The fact that experimental leaves were maintained in a horizontal position is at least partially responsible for these results. Possible other reasons for this apparent excess of RuBP regeneration capacity visa-vis RuBP carboxylase-oxygenase concentration are discussed.     

Cotton bracts are adapted to a microenvironment of concentrated CO2 produced by rapid fruit respiration

Background and Aims

Elucidation of the mechanisms by which plants adapt to elevated CO₂ is needed; however, most studies of the mechanisms investigated the response of plants adapted to current atmospheric CO₂. The rapid respiration rate of cotton (Gossypium hirsutum) fruits (bolls) produces a concentrated CO₂ microenvironment around the bolls and bracts. It has been observed that the intercellular CO₂ concentration of a whole fruit (bract and boll) ranges from 500 to 1300 µmol mol⁻¹ depending on the irradiance, even in ambient air. Arguably, this CO₂ microenvironment has existed for at least 1·1 million years since the appearance of tetraploid cotton. Therefore, it was hypothesized that the mechanisms by which cotton bracts have adapted to elevated CO₂ will indicate how plants will adapt to future increased atmospheric CO₂ concentration. Specifically, it is hypothesized that with elevated CO₂ the capacity to regenerate ribulose-1,5-bisphosphate (RuBP) will increase relative to RuBP carboxylation.

Methods

To test this hypothesis, the morphological and physiological traits of bracts and leaves of cotton were measured, including stomatal density, gas exchange and protein contents.

Key results

Compared with leaves, bracts showed significantly lower stomatal conductance which resulted in a significantly higher water use efficiency. Both gas exchange and protein content showed a significantly greater RuBP regeneration/RuBP carboxylation capacity ratio (J_max/V_cmax) in bracts than in leaves.

Conclusions

These results agree with the theoretical prediction that adaptation of photosynthesis to elevated CO₂ requires increased RuBP regeneration. Cotton bracts are readily available material for studying adaption to elevated CO₂.     

Photosynthetic acclimation in tomato plants grown in high CO2

The effects of prolonged CO₂ enrichment of tomato plants on photosynthetic performance and Calvin cycle enzymes, including the amount and activity of ribulose-1,5-bisphosphate carboxylase (RuBPco), were determined. Also the light-saturated rate of photosynthesis (P_max) of the 5th leaf throughout leaf development was predicted based on the amount and kinetics of RuBPco. With short-term CO₂ enrichment, i.e. only during the photosynthesis measurements, P_max of the young leaves did not increase while the leaves reaching full expansion more than doubled their net rate of CO₂ fixation. However, with longer-term CO₂ enrichment, i.e. growing the crop in high CO₂, the plants did not maintain this photosynthetic gain. Compared with leaves of plants grown in normal ambient CO₂ the high CO₂-grown leaves, when almost fully expanded, contained only about half as much RuBPco protein and P_max in 300 and 1000 vpm CO₂ was similarly reduced.The loss of RuBPco protein may be a factor associated with the accelerated fall in P_max since P_max was close to that predicted from the amount and kinetics of RuBPco assuming RuBP saturation. Acclimation to high CO₂ is fundamentally different from acclimation to high light. In contrast to acclimation to high light, acclimation to high CO₂ does not usually involve an increase in photosynthetic machinery so the synthesis and maintenance costs (as indicated by the dark respiration rate) are generally lower.     

Differences between wheat and rice in the enzymic properties of ribulose-1,5-bisphosphate carboxylase/oxygenase and the relationship to photosynthetic gas exchange

The kinetic parameters of ribulose-1,5-bisphosphate (RuBP) carboxylase/oxygenase (EC 4.1.1.39) in wheat (Triticum aestivum L.) and rice (Oryza sativa L.) were determined by rapidly assaying the leaf extracts. The respective K _m and V _max values for carboxylase and oxygenase activities were significantly higher for wheat than for rice. In particular, the differences in the V _max values between the two species were greater. When the net activity of CO₂ exchange was calculated at the physiological CO₂-O₂ concentration from these kinetic parameters, it was 22% greater in wheat than in rice. This difference in the in-vitro RuBP-carboxylase/oxygenase activity between the two species reflected a difference in the CO₂-assimilation rate per unit of RuBP-carboxylase protein. However, there was no apparent difference in the CO₂-assimilation rate for a given leaf-nitrogen content between the two species. When the RuBP-carboxylase/oxygenase activity was estimated at the intercellular CO₂ pressure from the enzyme content and kinetic parameters, these estimated enzyme activities in wheat and rice were similar to each other for the same rate of CO₂ assimilation. These results indicate that the difference in the kinetic parameters of RuBP carboxylase between the two species was offset by the differences in RuBP-carboxylase content and conductance for a given leaf-nitrogen content.Abbreviations DTT dithiothreitol - EDTA ethylenediamine-tetraacetic - PAR photosynthetically active radiation - RuBP ribulose-1,5-bisphosphate     

The occurrence of both C3 and C4 photosynthetic characteristics in a single Zea mays plant

The activities of the carboxylating enzymes ribulose-1,5-biphosphate (RuBP) carboxylase and phosphoenolpyruvate (PEP) carboxylase in leaves of three-week old Zea mays plants grown under phytotron conditions were found to vary according to leaf position. In the lower leaves the activity of PEP carboxylase was lower than that of RuBP carboxylase, while the upper leaves exhibited high levels of PEP carboxylase. Carbon dioxide compensation points and net photosynthetic rates also differed in the lower and upper leaves. Differences in the fine structure of the lowermost and uppermost leaves are shown. The existence of both the C₃ and C₄ photosynthetic pathways in the same plant, in this and other species, is discussed.Abbreviations PEP phosphoenolpyruvate - RuBP ribulose-1,5-biphosphate     

The temporal and species dynamics of photosynthetic acclimation in flag leaves of rice (Oryza sativa) and wheat (Triticum aestivum) under elevated carbon dioxide

In this study, we tested for the temporal occurrence of photosynthetic acclimation to elevated [CO₂] in the flag leaf of two important cereal crops, rice and wheat. In order to characterize the temporal onset of acclimation and the basis for any observed decline in photosynthetic rate, we characterized net photosynthesis, g_s, g_m, C_i/C_a, C_i/C_c, V_cmax, J_max, cell wall thickness, content of Rubisco, cytochrome (Cyt) f, N, chlorophyll and carbohydrate, mRNA expression for rbcL and petA, activity for Rubisco, sucrose phosphate synthase (SPS) and sucrose synthase (SS) at full flag expansion, mid‐anthesis and the late grain‐filling stage. No acclimation was observed for either crop at full flag leaf expansion. However, at the mid‐anthesis stage, photosynthetic acclimation in rice was associated with RuBP carboxylation and regeneration limitations, while wheat only had the carboxylation limitation. By grain maturation, the decline of Rubisco content and activity had contributed to RuBP carboxylation limitation of photosynthesis in both crops at elevated [CO₂]; however, the sharp decrease of Rubisco enzyme activity played a more important role in wheat. Although an increase in non‐structural carbohydrates did occur during these later stages, it was not consistently associated with changes in SPS and SS or photosynthetic acclimation. Rather, over time elevated [CO₂] appeared to enhance the rate of N degradation and senescence so that by late‐grain fill, photosynthetic acclimation to elevated [CO₂] in the flag leaf of either species was complete. These data suggest that the basis for photosynthetic acclimation with elevated [CO₂] may be more closely associated with enhanced rates of senescence, and, as a consequence, may be temporally dynamic, with significant species variation.     

Low concentration of bisulfite enhances photosynthesis in tea tree by promoting carboxylation efficiency in leaves

Tea tree (Melaleuca alternifolia) canopy was sprayed with low concentration of NaHSO₃ or mixture of NaHSO₃+ KH₂PO₄. The treatments significantly enhanced net photosynthetic rate (P _N), carboxylation efficiency (CE), and the maximum response of P _N to intercellular CO₂ concentration. The enhancement of P _N by foliar application of low concentrations of bisulfite was due to increasing CE relevant to ribulose-1,5-bisphosphate (RuBP) carboxylase/oxygenase activity and regeneration rate of RuBP depending on ATP formation.     

Growth and photosynthetic response of nine tropical species with long-term exposure to elevated carbon dioxide

Summary Seedlings of nine tropical species varying in growth and carbon metabolism were exposed to twice the current atmospheric level of CO₂ for a 3 month period on Barro Colorado Island, Panama. A doubling of the CO₂ concentration resulted in increases in photosynthesis and greater water use efficiency (WUE) for all species possessing C₃ metabolism, when compared to the ambient condition. No desensitization of photosynthesis to increased CO₂ was observed during the 3 month period. Significant increases in total plant dry weight were also noted for 4 out of the 5 C₃ species tested and in one CAM species, Aechmea magdalenae at high CO₂. In contrast, no significant increases in either photosynthesis or total plant dry weight were noted for the C₄ grass, Paspallum conjugatum. Increases in the apparent quantum efficiency (AQE) for all C₃ species suggest that elevated CO₂ may increase photosynthetic rate relative to ambient CO₂ over a wide range of light conditions. The response of CO₂ assimilation to internal C_i suggested a reduction in either the RuBP and/or Pi regeneration limitation with long term exposure to elevated CO₂. This experiment suggests that: (1) a global rise in CO₂ may have significant effects on photosynthesis and productivity in a wide variety of tropical species, and (2) increases in productivity and photosynthesis may be related to physiological adaptation(s) to increased CO₂.     

Limitation of photosynthesis by changes in temperature

The aim of this work was to examine the effect of abrupt changes in temperature in the range 5 to 30°C upon the rate of photosynthetic carbon assimilation in leaves of barley (Hordeum vulgare L.). Measurement of the CO₂-assimilation rate in relation to the intercellular partial pressure of CO₂ at different temperatures and O₂ concentrations and at saturating irradiance showed that as the temperature was decreased photosynthesis was saturated at progressively lower CO₂ partial pressures and that the transition between the CO₂-limited and ribulose-1,5-bisphosphate-regeneration-limited rate became more abrupt. Feeding of orthophosphate to leaves resulted in an increased rate of CO₂ assimilation at lower temperatures at around ambient or higher CO₂ partial pressures both in 20% O₂ and in 2% O₂ and it removed the abruptness in the transition between the CO₂-limited and ribulose-1,5-bisphosphate-regeneration-limited rates. Phosphate feeding tended to inhibit carbon assimilation at higher temperatures. The response of carbon assimilation to temperature was altered by feeding orthophosphate, by changing the concentrations of CO₂ or of O₂ or by leaving plants in the dark at 4°C for several hours. Similarly, the response of carbon assimilation to phosphate feeding or to changes in 2% O₂ was altered by leaving the plants in the dark at 4°C. The mechanism of limitation of photosynthesis by an abrupt lowering of temperature is discussed in the light of the results.Abbreviations A rate of CO₂ assimilation - P _i intercellular partial pressure of CO₂ - RuBP ribulose-1,5-bisphosphate     

Photosynthesis and ribulose-1,5-bisphosphate carboxylase/oxygenase in rice leaves from emergence through senescence. Quantitative analysis by carboxylation/oxygenation and regeneration of ribulose 1,5-bisphosphate

Changes in gas-exchange rates during the life span of the leaves of rice (Oryza sativa L.) were analyzed quantitatively by measuring changes in the carboxylation/oxygenation and regeneration of ribulose 1,5-bisphosphate (RuBP) at photon fluence rates of 2000 (saturating) and 500 (subsaturating) μmol quanta·m^-2·s^-1 under ambient air conditions. The RuBP levels were always higher than the active-site concentrations of RuBP carboxylase (EC 4.1.1.39), irrespective of the irradiance supplied. Analysis of the CO₂-assimilation rate as a function of intercellular CO₂ concentration indicated that RuBP regeneration does not limit CO₂ assimilation. The estimated RuBP-carboxylase/oxygenase activity in vivo was linearly correlated to the rate of CO₂ assimilation at each level of irradiance. This enzyme activity was just enough to account for the rate of CO₂ assimilation at the saturating irradiance and was 35% more than the rate of CO₂ assimilation at the subsaturating irradiance. Analysis of the assimilation rate at subsaturating irradiance as a function of intercellular CO₂ concentration indicated that a limitation caused by enzyme activation comes into play. The results indicate that the rate of CO₂ assimilation in rice leaves under ambient air conditions is limited during their entire life span by the RuBP-carboxylation/oxygenation capacity.     

The relationship between steady-state gas exchange of bean leaves and the levels of carbon-reduction-cycle intermediates

The relationship between the gas-exchange characteristics of attached leaves of Phaseolus vulgaris L. and the pool sizes of several carbon-reduction-cycle intermediates was examined. After determining the rate of CO₂ assimilation at known intercellular CO₂ pressure, O₂ pressure and light, the leaf was rapidly killed (<0.1 s) and the levels of ribulose-1,5-bisphosphate (RuBP), 3-phosphoglyceric acid (PGA), fructose-1,6-bisphosphate, fructose-6-phosphate, glucose-6-phosphate, glyceraldehyde-3-phosphate, and dihydroxyacetone phosphate were measured. In 210 mbar O₂, photosynthesis appeared RuBP-saturated at low CO₂ pressure and RuBP-limited at high CO₂ pressure. In 21 mbar (2%) O₂, the level of RuBP always appeared saturating. Very high levels of PGA and other phosphate-containing compounds were found with some conditions, especially under low oxygen.Abbreviations and symbols C₁ intercellular CO₂ pressure - PGA 3-phosphoglyceric acid - RuBP ribulose-1,5-bisphosphate - Rubisco ribulose-1,5-bisphosphate carboxylase-oxygenase     

Effect of growth conditions on carboxylating enzymes of Zea mays plants

Phosphoenolpyruvate (PEP) carboxylase (EC 4.1.1.31) and ribulose-1,5-bisphospate (RuBP) carboxylase (EC 4.1.1.39) activities in leaves of different maize hybrids grown under field conditions (high light intensity) and in a growth chamber (low light intensity) were determined. Light intensity and leaf age affected PEP carboxylase activity, whereas RuBP carboxylase was affected by leaf age only at low light intensity. PEP carboxylase/RuBP carboxylase activity ratio decreased according to light intensity and leaf age. Results demonstrate that Zea mays grown under field conditions is a typical C₄ species in all leaves independently from their position on the stem, whereas it may be a C₃ plant when it is grown in a growth chamber at low light intensityAbbreviations PEP phosphoenolpyruvate - RuBP ribulose-1,5-bisphosphate     

Seasonal Changes of Selected Parameters of CO2 Fixation Biochemistry of Norway Spruce Under the Long-Term Impact of Elevated CO2

Twelve-year-old Norway spruce (Picea abies [L.] Karst.) trees were exposed to ambient (AC) or elevated (EC) [ambient + 350 μmol(CO₂) mol^-1] CO₂ concentrations in open-top-chamber (OTC) experiment under the field conditions of a mountain stand. Short-term (4 weeks, beginning of the vegetation season) and long-term (4 growing seasons, end of the vegetation season) effects of this treatment on biochemical parameters of CO₂ assimilation were evaluated. A combination of gas exchange, fluorescence of chlorophyll a, and application of a mathematical model of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) activity was used. The analysis showed that the depression of photosynthetic activity by long-term impact of elevated CO₂ was mainly caused by decreased RuBPCO carboxylation rate. The electron transport rate as well as the rate of ribulose-1,5-bisphosphate (RuBP) formation were also modified. These modifications to photosynthetic assimilation depended on time during the growing season. Changes in the spring were caused mainly by local deficiency of nitrogen in the assimilating tissue. However, the strong depression of assimilation observed in the autumn months was the result of insufficient carbon sink capacity. This revised version was published online in June 2006 with corrections to the Cover Date.     

A Model Describing the Regulation of Ribulose-1,5-Bisphosphate Carboxylase, Electron Transport, and Triose Phosphate Use in Response to Light Intensity and CO(2) in C(3) Plants

A model of the regulation of the activity of ribulose-1,5-bisphosphate carboxylase, electron transport, and the rate of orthophosphate regeneration by starch and sucrose synthesis in response to changes in light intensity and partial pressures of CO₂ and O₂ is presented. The key assumption behind the model is that nonlimiting processes of photosynthesis are regulated to balance the capacity of limiting processes. Thus, at CO₂ partial pressures below ambient, when a limitation on photosynthesis by the capacity of rubisco is postulated, the activities of electron transport and phosphate regeneration are down-regulated in order that the rate of RuBP regeneration matches the rate of RuBP consumption by rubisco. Similarly, at subsaturating light intensity or elevated CO₂, when electron transport or Pi regeneration may limit photosynthesis, the activity of rubisco is downregulated to balance the limitation in the rate of RuBP regeneration. Comparisons with published data demonstrate a general consistency between modelled predictions and measured results.