Carbon gains by desiccation-tolerant plants at elevated CO2

1. There have been no reports of the long-term responses of the desiccation-tolerant (DT) plants to elevated CO₂. Xerophyta scabrida is a DT woody shrub, which loses chlorophylls and thylakoids during desiccation: a so-called poikilochlorophyllous desiccation-tolerant species (PDT). When the leaves of X. scabria are allowed to desiccate, the species shows many of the normal features of (P)DT plants.
2. However, the duration of photosynthesis in X. scabria is prolonged by 300% when the measurements are made at 700 as opposed to 350p.p.m. CO₂. The implication is that the carboxylating enzymes must still have been active at this time to enable appreciable photosynthetic activity. This response could have far-reaching implications for the success of such species in a future climate.
3. Lichens and mosses, representing the homoiochlorophyllous DTs (HDT), retain their chlorophyll content and photosynthetic apparatus during desiccation. We show the desiccation responses of two common HDT species ( Cladonia convoluta and Tortula ruralis ) to elevated CO₂ for comparison. Both HDT species showed increased net CO₂ uptake in the material grown at high CO₂ by more than 30% in moss and by more than 50% in lichen. It is concluded that desiccation-tolerant plants will be among the main beneficiaries of a high CO₂ future.     

Stomatal and non-stomatal contribution in the decline in leaf net CO2 uptake during rapid water stress

A simple method is proposed for quantitative evaluation of Stomatal and non-stomatal components of the decline in leaf CO₂ uptake during rapid water stress. The changes in leaf conductance were measured during the stress and were used to calculate the photosynthetic rate which would be observed if Stomatal closure were the only cause of the decline in photosynthesis. Photosynthesis-CO₂ response curves, determined just before the stress, were used for this calculation. The difference between the calculated and the actual rate is a measure of the non-stomatal effect of water stress.
This analysis was tested on Sinapis alba submitted to rapid and severe water stress by excising leaves or roots. Experiments were performed at saturating light conditions under high (61 Pa), normal (34 Pa) or low (11 Pa) ambient CO 2 pressure. The non-stomatal effect on de-rooted plants reaches a maximum at the beginning of the stress and is dependent on the CO 2 pressure: after 45 min its influence is still about 100°, 70° and 8°, respectively, at high, normal and low CO₂. In the excised leaf system in which desiccation was more rapid, the non-stomatal effect accounted for nearly 100° of the assimilation decline whatever the CO₂ pressure.     

LIMITATIONS OF PHOTOSYNTHESIS IN DIFFERENT REGIONS OF THE ZEA MAYS LEAF

The progressive development of the photosynthetic apparatus occurring along the length of the Zea mays leaf offers a convenient system with which to examine the limitations to photosynthetic CO₂ assimilation during biogenesis of a C₄ leaf. Changes in light-induced O₂ evolution and CO₂ assimilation, chlorophyll content, activity of PEP-carboxylase, NADP-malic enzyme and the 'R5P system' (consisting of d -ribose-5-phosphate-keto isomerase, ATP- d -ribulose-5 phosphate 1-phosphotransferase and d -ribulose-1,5-bisphosphate carboxylase) and fluorescence emission characteristics were examined along the length of the second leaf of 7-day-old plants grown under a diurnal light regime. The results suggest that the major limitation to CO₂ assimilation in the leaf sheath lies within the chlorenchyma and is either energy supply for carboxylation or the capacity of key photosynthetic enzymes. In the leaf blade stomatal resistance to CO₂ diffusion constitutes a major fraction of the total leaf resistance to CO₂ assimilation implicating the stoma as the major limiting factor to photosynthetic CO₂ assimilation.     

An integrated portable apparatus for the simultaneous field measurement of photosynthetic CO2 and water vapour exchange, light absorption and chlorophyll fluorescence emission of attached leaves

Abstract. A portable apparatus has been constructed to measure simultaneously the quantum yield of CO₂ assimilation, light absorption, chlorophyll fluorescence emission and water vapour exchange of attached intact leaves in the field. The core of the instrument is a light-integrating spherical leaf chamber which includes ports for a light source, photosynthetically active radiation sensor, fluorescence probes and gas inlet and outlet manifolds. Measurement of the quantum flux inside the empty chamber and with a leaf present allows determination of leaf absorptance. An open gas exchange system is employed using an infra-red analyser to measure leaf CO₂ exchange. Using a DC white light source the quantum yield of CO₂ assimilation based on absorbed light (φ_abs) may be determined rapidly in either ambient air or artificial gas mixtures. Inclusion of capacitance humidity probes into the gas inlet and outlet ports allows simultaneous determination of water vapour exchange and subsequent estimation of stomatal conductance to CO₂ and intercellular CO₂ concentration. Measurement of fluorescence emission by the sample leaf exposed to white light is achieved by a modulated fluorescence detection system. In addition to determination of the minimal, maximal and variable fluorescence levels, a further analysis allows the photochemical and non-photochemical components of fluorescence quenching, to be estimated. The theory and design of this apparatus is described in detail. The use of the apparatus in the field is demonstrated through a study of the photosynthetic performance of a maize and bean crop during the growing season and by analysis of the photosynthetic performance of crops subjected to nitrogen-stress and a herbicide treatment.     

Effects of manganese toxicity on leaf CO2 assimilation of contrasting common bean genotypes

Parameters related to leaf photosynthesis were evaluated in three genotypes of common bean ( Phaseolus vulgaris L.) with contrasting tolerance to Mn toxicity. Two short-term studies in solution culture were used to assess the effect of excess Mn on CO₂ assimilation in mature and immature leaves. Mn toxicity decreased total chlorophyll content only in immature leaves, with a consequent reduction of leaf CO₂ assimilation. Mature leaves that showed brown speckles characteristic of Mn toxicity, did not suffer any detriment in their capacity to assimilate CO₂, at least in a 4-day experiment. Stomatal conductance and transpiration were not affected by the presence of high levels of Mn in leaf tissue. Lower stomatal conductance and transpiration rates were observed only in leaves with advanced chlorosis. Differences among genotypes were detected as increased chlorosis in the more sensitive genotype ZPV-292, followed by A-283 and less chlorosis in the tolerant genotype CALIMA. Since CO₂ assimilation expressed per unit of chlorophyll was not different between high-Mn plants and control plants, we conclude that the negative effect of Mn toxicity on CO₂ assimilation can be explained by a reduction in leaf chlorophyll content.     

Effect of elevated CO2 on the stomatal distribution and leaf physiology of Alnus glutinosa

Variation in stomatal development and physiology of mature leaves from Alnus glutinosa plants grown under reference (current ambient, 360 μmol mol⁻¹ CO₂) and double ambient (720 μmol mol⁻¹ CO₂) carbon dioxide (CO₂) mole fractions is assessed in terms of relative plant growth, stomatal characters (i.e. stomatal index and density) and leaf photosynthetic characters. This is the first study to consider the effects of elevated CO₂ concentration on the distribution of stomata and epidermal cells across the whole leaf and to try to ascertain the cause of intraleaf variation. In general, a doubling of the atmospheric CO₂ concentration enhanced plant growth and significantly increased stomatal index. However, there was no significant change in relative stomatal density. Under elevated CO₂ concentration there was a significant decrease in stomatal conductance and an increase in assimilation rate. However, no significant differences were found for the maximum rate of carboxylation ( V _cmax) and the light saturated rate of electron transport ( J _max) between the control and elevated CO₂ treatment.     

Non-stomatal limitation of CO2 assimilation in three tree species during natural drought conditions

The effect of drought on CO₂ assimilation and leaf conductance was studied in three northern hardwood species: Quercus rubra L., Acer rubrum L. and Populus grandidentata Michx. Leaf gas exchange characteristics at two CO₂ levels (320 and 620 μl I⁻¹) and temperatures from 20 to 35°C were measured at the end of a dry period and shortly after 10 cm of rainfall. The effects of drought varied with species, temperature and CO₂ level. Calculated values of internal CO₂ concentration showed little or no decline during drought. Differences in assimilation, before vs after the rains, were most apparent at the higher CO₂ level. These latter two observations indicate nonstomatal disruption of CO₂ assimilation during the dry period. In P. grandidentata there was a substantial interaction between drought and temperature, with a resultant shift in the temperature for maximum assimilation to lower temperatures during drought. During drought, internal CO₂ concentrations increased sharply in all three species under the combined conditions of high temperatures and the higher CO₂ level.     

Yield Loss in Sweet Corn in Response to Defoliation or Infection by Exserohilum turcicum

Abstract Net CO₂ assimilation was reduced in sites of infection by Exserohilum turcicum in leaves of Seneca 60 sweet corn before lesions appeared. In leaf tissue adjacent to infected areas, there was an early small increase in CO₂ assimilation followed by a gradual decline to nearly 0 net CO₂ exchange by 7 days after inoculation. Translocation of photosynthates in to disease lesions from healthy tissue distal to the lesions was observed within 1 h after exposure of 1.2–cm² areas of the leaf blade to ¹⁴CO₂. No translocation from lesions to healthy leaf tissue was observed. The effects of defoliation at specific leaf positions on yield of sweet corn plants were accurately simulated by a model in which yield is expressed as a function of healthy leaf area absorption of incident insolation. Removal of leaves from the bottom third of the plants caused no yield loss, whereas removal of leaves above the ear caused significant losses. The model underes, timated the yield loss caused by infection by E. turcicum by approximately 22 %. The observed effects of infection by E. turcicum on photosynthetic efficiency in leaf tissue adjacent to lesions and on translocation of photosynthates into lesion from distal parts of the leaf show that the effect of northern leaf blight on yield is greater than can be accounted for by the direct loss of healthy leaf area through necrosis within disease lesions.     

Gas exchange parameters, water relations and carbohydrate partitioning in leaves of field-grown Prunus domestica following fruit removal

The effect of fruit removal on gas exchange, water relations, chlorophyll and non-structural carbohydrate content of leaves from mature, field-grown plum trees ( Prunus domestica L. cv. Stanley) was determined over 2 consecutive growing seasons. Removal of fruits during stage II of fruit development decreased CO₂ assimilation rate within 24 h from 12.6 to 8.5 μmol m^-2 s^-1 in 1986, and from 12.1 to 10.2 μmol m^-2 s^-1 in 1987. Depression of net photosynthesis persisted for at least 5 days and was greatest in the early afternoon. Recovery of the CO₂ assimilation rate to pretreatment levels coincided in defruited trees with vegetative growth that was more than 5-fold that of fruiting trees in the first 6 weeks after fruit removal in 1986. Estimated photorespiration was similar in both fruiting and defruited trees. The stomatal contribution to the decrease of CO₂ assimilation rate, calculated from assimilation/intercellular CO₂ curves, ranged from 31 to 46%. Defruiting did not affect leaf water potential, but decreased leaf osmotic potential. Leaf levels of chlorophyll, fructose, glucose, sorbitol and sucrose were not affected by defruiting, whereas starch content increased up to 51% in leaves of defruited trees within 24 h after fruit removal. However, because of the small starch pool present in plum leaves (<1.9% dry weight) it is unlikely that starch accumulation was responsible for the observed decline in CO₂ assimilation rate after fruit removal. The decrease of CO₂ assimilation rate is discussed in relation to the hypothesis of assimilate demand regulating photosynthesis through a feedback mechanism.     

Physiological effects of long-term exposure to low and moderate concentrations of atmospheric NH3 on poplar leaves

Abstract. Poplar shoots ( Populus euramericana L.) obtained from cuttings were exposed for 6 or 8 weeks to NH₃ concentrations of 50 and 100 μgm⁻³ or filtered air in fumigation chambers. After this exposure the rates of NH₃ uptake, transpiration, CO₂ assimilation and respiration of leaves were measured using a leaf chamber. During the long-term exposure also modulated chlorophyll fluorescence measurements were carried out to obtain information about the photosynthetic performance of individual leaves. Both fluorescence and leaf chamber measurements showed a higher photosynthetic activity of leaves exposed to 100 μg NH₃ m⁻³. These leaves showed also a larger leaf conductance and a larger uptake rate of NH₃ than leaves exposed to 50 μg m⁻³ NH₃ or filtered air. The long-term NH₃ exposure did not induce an internal resistance against NH₃ transport in the leaf, nor did it affect the leaf cuticle. So, not only at a short time exposure, but also at a long-term exposure NH₃ uptake into leaves can be calculated from data on the boundary layer and stomatal resistance for H₂O and ambient NH₃-concentration. Furthermore, the NH₃ exposure had no effect on the relation between CO₂-assimilation and stomatal conductance, indicating that NH₃ in concentrations up to 100 μg m⁻³ has no direct effect on stomatal behaviour; for example, by affecting the guard or contiguous cells of the stomata.     

Reconstitution of chlorophylls and photosynthetic CO2 assimilation upon rehydration of the desiccated poikilochlorophyllous plant Xerophyta scabrida (Pax) Th. Dur. et Schinz

Resynthesis of the photosynthetic apparatus and resumption of CO₂ assimilation upon rehydration is reported for the monocotyledonous and poikilochlorophyllous desiccation-tolerant (PDT) plant Xerophyta scabrida (Pax) Th. Dur. et Schinz (Velloziaceae). During desiccation there was a complete breakdown of chlorophylls whereas the total carotenoid content of air-dried leaves was reduced to about 22% of that of functional leaves. The prerequisites for the resynthesis of photosynthetic pigments and functional thylakoids were the reappearance of turgor and maximum leaf water content at 2 and 10 h after rehydration, respectively. The period of increased initial respiration after rewetting leaves (rehydration respiration) lasted up to 30 h and was thus 6 to 10 times longer than in homoiochlorophyllous desiccation-tolerant plants (HDTs) in which chlorophylls are retained during desiccation. Accumulation of chlorophylls a + b and total carotenoids (xanthophylls and carotene) started 10 h after rehydration. Normal levels of chlorophyll and carotenoids were obtained 72 h after rehydration. Values for the variable-fluorescence decrease ratio (Rfd690 values), an indicator of photochemical activity, showed that photochemical function started 10 h after rehydration, but normal values of 2.7 were reached only 72 h after rehydration. Net CO₂ assimilation started 24 h after rewetting and normal rates were reached after 72 h, at the same time as normal values of stomatal conductance were obtained. The increasing rates of net CO₂ assimilation were paralleled by decreasing values of the intercellular CO₂ concentration. All photosynthetic parameters investigated showed values normal for functional chloroplasts by 72 h after the onset of rehydration. Fully regreened leaves of the presumed C₃ plant X. scabrida exhibited a net CO₂ assimilation rate which was in the same range as that of other C₃ plants and higher than that of recovered HDT plants. The fundamental difference between air-dried PDT plants, such as X. scabrida, which have to resynthesize the photosynthetic pigment apparatus, and air-dried HDT plants, which only undergo a functional recovery, is discussed.Abbreviations c -carotene - c_i intercellular CO₂ concentration - Car x + c total carotenoid content x + c - Chl a + b total chlorophyll a + b content - g_s stomatal conductance - HDT homoiochlorophyllous desiccation tolerant - LWC leaf-water content - P_N net photosynthesis rate - PDT poikilochloro phyllous desiccation tolerant - R_d dark respiration - Rfd variable fluorescence decrease ratio (Rfd = fd/fs) - x xanthophylls The senior author thanks the Deutschem Akademischem Auslandsdienst (Bonn, Germany), Soros Foundation (Budapest, Hungary) and European Community (Brussels, Belgium) for providing fellowships for research periods at Karlsruhe. The research was also supported by the Hungarian Scientific Research Foundation (OTKA I/848, OTKA I/3.1545 and OTKA I/4.F.5359). We wish to thank Professor T. Pocs (Eger, Hungary — Morogoro, Tanzania) for collecting the plant material and to the linguist Mr. A. Jackson for correcting the English.     

Stomatal limitation of photosynthesis in winter production of greenhouse tomato plants

In May, greenhouse tomato ( Lycopersicon esculentum Mill.) plants near the end of their winter production cycle were shown to exhibit a diurnal photosynthetic decrease. In order to identify the physiological causes of this decline, we compared in May the photosynthetic characteristics of the fifth youngest leaves from tomato plants of different ages corresponding to a winter production (11-month-old plants) and to a spring production (5-month-old plants). Although the leaves were developed simultaneously under the same environmental conditions, only the ones from the winter production showed a diurnal decline of the in situ CO₂ assimilation rate (A _{CO 2}). This was accompanied by a decline of internal CO₂ and stomatal conductance and by large accumulations of hexoses. When stomatal closure was relieved under saturated CO₂ concentration (5%) using a leaf-disc electrode system, the fifth leaves of both tomato cultures had similar maximum quantum efficiency of O₂ evolution (Φ_max), light-saturated rate of O₂ evolution (P_max) and quantum efficiency of photosystem II (PSII) photochemistry (ΔF/F'_m, q _P and q _N ). We concluded that the diurnal decline of A _{CO 2} observed in winter tomato production during May originates from a stomatal limitation that is not dependent on environmental conditions but rather related to the developmental stage of the plants.     

Effects of manganese toxicity on photosynthesis of white birch (Betula platyphylla var. japonica) seedlings

The effects of manganese (Mn) toxicity on photosynthesis in white birch ( Betula platyphylla var. japonica ) leaves were examined by the measurement of gas exchange and chlorophyll fluorescence in hydroponically cultured plants. The net photosynthetic rate at saturating light and ambient CO₂ (C_a) of 35 Pa decreased with increasing leaf Mn concentrations. The carboxylation efficiency, derived from the difference in CO₂ assimilation rate at intercellular CO₂ pressures attained at C_a of 13 Pa and O Pa, decreased with greater leaf Mn accumulation. Net photosynthetic rate at saturating light and saturating CO₂ (5%) also declined with leaf Mn accumulation while the maximum quantum yield of O₂ evolution at saturating CO₂ was not affected. The maximum efficiency of PSII photochemistry (F_v/F_m) was little affected by Mn accumulation in white birch leaves over a wide range of leaf Mn concentrations (2–17 mg g₋₁ dry weight). When measured in the steady state of photosynthesis under ambient air at 430 μmol quanta m₋₂ s₋₁, the levels of photochemical quenching (qP) and the excitation capture efficiency of open PSII (F'^v/F'^m) declined with Mn accumulation in leaves. The present results suggest that excess Mn in leaves affects the activities of the CO² reduction cycle rather than the potential efficiency of photochemistry, leading to increases in Q_A reduction state and thermal energy dissipation, and a decrease in quantum yield of PSII in the steady state.     

The role of temperature in determining the stimulation of CO2 assimilation at elevated carbon dioxide concentration in soybean seedlings

Soybean ( Glycine max cv. Clark) was grown at both ambient (ca 350 μmol mol⁻¹) and elevated (ca 700 μmol mol⁻¹) CO₂ concentration at 5 growth temperatures (constant day/night temperatures of 20, 25, 30, 35 and 40°C) for 17–22 days after sowing to determine the interaction between temperature and CO₂ concentration on photosynthesis (measured as A, the rate of CO₂ assimilation per unit leaf area) at both the single leaf and whole plant level. Single leaves of soybean demonstrated increasingly greater stimulation of A at elevated CO₂ as temperature increased from 25 to 35°C (i.e. optimal growth rates). At 40°C, primary leaves failed to develop and plants eventually died. In contrast, for both whole plant A and total biomass production, increasing temperature resulted in less stimulation by elevated CO₂ concentration. For whole plants, increased CO₂ stimulated leaf area more as growth temperature increased. Differences between the response of A to elevated CO₂ for single leaves and whole plants may be related to increased self-shading experienced by whole plants at elevated CO₂ as temperature increased. Results from the present study suggest that self-shading could limit the response of CO₂ assimilation rate and the growth response of soybean plants if temperature and CO₂ increase concurrently, and illustrate that light may be an important consideration in predicting the relative stimulation of photosynthesis by elevated CO₂ at the whole plant level.     

The significance of differences in the mechanisms of photosynthetic acclimation to light, nitrogen and CO2 for return on investment in leaves

1. We report changes in photosynthetic capacity of leaves developed in varying photon flux density (PFD), nitrogen supply and CO₂ concentration. We determined the relative effect of these environmental factors on photosynthetic capacity per unit leaf volume as well as the volume of tissue per unit leaf area. We calculated resource-use efficiencies from the photosynthetic capacities and measurements of leaf dry mass, carbohydrates and nitrogen content.
2. There were clear differences between the mechanisms of photosynthetic acclimation to PFD, nitrogen supply and CO₂. PFD primarily affected volume of tissue per unit area whereas nitrogen supply primarily affected photosynthetic capacity per unit volume. CO₂ concentration affected both of these parameters and interacted strongly with the PFD and nitrogen treatments.
3. Photosynthetic capacity per unit carbon invested in leaves increased in the low PFD, high nitrogen and low CO₂ treatments. Photosynthetic capacity per unit nitrogen was significantly affected only by nitrogen supply.
4. The responses to low PFD and low nitrogen appear to function to increase the efficiency of utilization of the limiting resource. However, the responses to elevated CO₂ in the high PFD and high nitrogen treatments suggest that high CO₂ can result in a situation where growth is not limited by either carbon or nitrogen supply. Limitation of growth at elevated CO₂ appears to result from internal plant factors that limit utilization of carbohydrates at sinks and/or transport of carbohydrates to sinks.     

The interaction of high temperature and elevated CO2 on photosynthetic acclimation of single leaves of rice in situ

Rice ( Oryza sativa L. cv. IR72) was grown at three different CO₂ concentrations (ambient, ambient + 200 μmol mol⁻¹, ambient + 300 μmol mol⁻¹) at two different growth temperatures (ambient, ambient + 4°C) from sowing to maturity to determine longterm photosynthetic acclimation to elevated CO₂ with and without increasing temperature. Single leaves of rice showed a cooperative enhancement of photosynthetic rate with elevated CO₂ and temperature during tillering, relative to the elevated CO₂ condition alone. However, after flowering, the degree of photosynthetic stimulation by elevated CO₂ was reduced for the ambient + 4°C treatment. This increasing insensitivity to CO₂ appeared to be accompanied by a reduction in ribulose-1.5-bisphosphate carboxylase/oxygenase (Rubisco) activity and/or concentration as evidenced by the reduction in the assimilation (A) to internal CO₂ (C₁) response curve. The reproductive response (e.g. percent filled grains, panicle weight) was reduced at the higher growth temperature and presumably reflects a greater increase in floral sterility. Results indicate that while CO₂ and temperature could act synergistically at the biochemical level, the direct effect of temperature on floral development with a subsequent reduction in carbon utilization may change sink strength so as to limit photosynthetic stimulation by elevated CO₂ concentration.     

Effect of nitrogen application on growth and photosynthetic nitrogen use efficiency in two ecotypes of wild strawberry, Fragaria chiloensis

The relationships between increasing nitrogen fertilization and growth, maximum CO₂ assimilation and the initial slope of the CO₂ response curve were studied in 2 ecotypes of wild strawberry, Fragaria chiloensis (L.) Duchn. Nitrogen accumulation of CA11, an ecotype from a low-nutrient dune site, was greater at all nitrogen concentrations than that of RCP37, an ecotype from a higher-nutrient strand site. Maximum CO₂ assimilation, total Rubisco activity, dry weight, and initiation of leaves and crowns were higher in CAI1 than RCP37 as nitrogen treatment was increased from 0 to 200 mg ^l-1, whereas these parameters were lower in CAl1 when fertilized at 300 mg T¹, but not in RCP37. The mean leaf area of CA11 was greater than RCP37 when grown with no supplemental nitrogen, but mean leaf area of the 2 lines was similar under nitrogen fertilization. Maximum CO₂ assimilation and carboxylation efficiency increased with increasing leaf nitrogen in both clones. At equivalent concentrations of leaf nitrogen, RCP37 had higher CO₂ assimilation and carboxylation efficiency than CA11 and the difference between the 2 clones increased as ieaf nitrogen increased. Thus, RCP37 had a higher photosynthetic nitrogen use efficiency than CA11. However, at a given applied nitrogen level, CA11 allocated more nitrogen to a unit of leaf area so that photosynthetic rates were higher than RCP37, except at the highest application of 300 mg ^l-1. The high nitrogen accumulation capacity and resource allocation to fruiting structures (crowns) in CA11 leads us to suggest that this clone may possess genes that could increase fruit yield in cultivated strawberry.     

Modelling environmental controls on ecosystem photosynthesis and the carbon isotope composition of ecosystem-respired CO2 in a coastal Douglas-fir forest

We developed and applied an ecosystem-scale model that calculated leaf CO₂ assimilation, stomatal conductance, chloroplast CO₂ concentration and the carbon isotope composition of carbohydrate formed during photosynthesis separately for sunlit and shaded leaves within multiple canopy layers. The ecosystem photosynthesis model was validated by comparison to leaf-level gas exchange measurements and estimates of ecosystem-scale photosynthesis from eddy covariance measurements made in a coastal Douglas-fir forest on Vancouver Island. A good agreement was also observed between modelled and measured δ ¹³C values of ecosystem-respired CO₂ ( δ _R). The modelled δ _R values showed strong responses to variation in photosynthetic photon flux density (PPFD), air temperature, vapour pressure deficit (VPD) and available soil moisture in a manner consistent with leaf-level studies of photosynthetic ¹³C discrimination. Sensitivity tests were conducted to evaluate the effect of (1) changes in the lag between the time of CO₂ fixation and the conversion of organic matter back to CO₂; (2) shifts in the proportion of autotrophic and heterotrophic respiration; (3) isotope fractionation during respiration; and (4) environmentally induced changes in mesophyll conductance, on modelled δ _R values. Our results indicated that δ _R is a good proxy for canopy-level C _c/ C _a and ¹³C discrimination during photosynthetic gas exchange, and therefore has several applications in ecosystem physiology.     

Light dependent activation of CO2 assimilation and the ratio of Rubisco activase to Rubisco during leaf aging of rice (Oryza sativa)

The effects of the ratio of Rubisco activase to Rubisco (activase/Rubisco ratio) on light dependent activation of CO₂ assimilation were investigated during leaf aging of rice. Changes of photosynthetic CO₂ gas exchange rates in relation to step increases of light intensity from two photon flux densities of 60 µmol m⁻² s⁻¹ (low initial PFD) and 500 µmol m⁻² s⁻¹ (high initial PFD) to saturated PFD of 1 800 µmol m⁻² s⁻¹ were measured. These photosynthetic activation processes were considered to be limited by the Rubisco activation rate when analyzed by the relaxation method. The relaxation time of low initial PFD gradually declined from 3 to 33 days after leaf emergence and showed high and negative correlation to the activase/Rubisco ratio. The initial rate of Rubisco activation under low initial PFD linearly correlated to the amounts of Rubisco activase, whereas these were almost constant from 3 to 23 days after leaf emergence. But these correlations could not be recognized in the case of high initial PFD. Moreover, the relaxation times were more sensitive to intercellular CO₂ concentration (C_i) under high initial PFD than under low initial PFD, especially, at C_i below 300 µl l⁻¹. These results suggest the involvement of the activase/Rubisco ratio in the photosynthetic activation under relatively low initial PFD, and the limitation of photosynthetic activation under relatively high initial PFD by Rubisco carbamylation during leaf aging of rice.