Effects of elevated [CO2] on photosynthesis in European forest species: a meta-analysis of model parameters

The effects of elevated atmospheric CO₂ concentration on growth of forest tree species are difficult to predict because practical limitations restrict experiments to much shorter than the average life-span of a tree. Long-term, process-based computer models must be used to extrapolate from shorter-term experiments. A key problem is to ensure a strong flow of information between experiments and models. In this study, meta-analysis techniques were used to summarize a suite of photosynthetic model parameters obtained from 15 field-based elevated [CO₂] experiments on European forest tree species. The parameters studied are commonly used in modelling photosynthesis, and include observed light-saturated photosynthetic rates (A_max), the potential electron transport rate (J_max), the maximum Rubisco activity (V_cmax) and leaf nitrogen concentration on mass (N_m) and area (N_a) bases. Across all experiments, light-saturated photosynthesis was strongly stimulated by growth in elevated [CO₂]. However, significant down-regulation of photosynthesis was also observed; when measured at the same CO₂ concentration, photosynthesis was reduced by 10–20%. The underlying biochemistry of photosynthesis was affected, as shown by a down-regulation of the parameters J_max and V_cmax of the order of 10%. This reduction in J_max and V_cmax was linked to the effects of elevated [CO₂] on leaf nitrogen concentration. It was concluded that the current model is adequate to model photosynthesis in elevated [CO₂]. Tables of model parameter values for different European forest species are given.     

Acclimation response of spring wheat in a free-air CO2 enrichment (FACE) atmosphere with variable soil nitrogen regimes. 1. Leaf position and phenology determine acclimation response

We have examined the photosynthetic acclimation of wheat leaves grown at an elevated CO₂ concentration, and ample and limiting N supplies, within a field experiment using free-air CO₂ enrichment (FACE). To understand how leaf age and developmental stage affected any acclimation response, measurements were made on a vertical profile of leaves every week from tillering until maturity. The response of assimilation (A) to internal CO₂ concentration (C_i) was used to estimate the in vivo carboxylation capacity (Vc_max) and maximum rate of ribulose-1,5-bisphosphate limited photosynthesis (A _sat). The total activity of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), and leaf content of Rubisco and the Light Harvesting Chlorophyll a/b protein associated with Photosystem II (LHC II), were determined. Elevated CO₂ did not alter Vc_max in the flag leaf at either low or high N. In the older shaded leaves lower in the canopy, acclimatory decline in Vc_max and A _sat was observed, and was found to correlate with reduced Rubisco activity and content. The dependency of acclimation on N supply was different at each developmental stage. With adequate N supply, acclimation to elevated CO₂ was also accompanied by an increased LHC II/Rubisco ratio. At low N supply, contents of Rubisco and LHC II were reduced in all leaves, although an increased LHC II/Rubisco ratio under elevated CO₂ was still observed. These results underscore the importance of leaf position, leaf age and crop developmental stage in understanding the acclimation of photosynthesis to elevated CO₂ and nutrient stress. This revised version was published online in June 2006 with corrections to the Cover Date.     

Brassinosteroids promote photosynthesis and growth by enhancing activation of Rubisco and expression of photosynthetic genes in <Emphasis Type="Italic">Cucumis sativus</Emphasis>

Brassinosteroids (BRs) are a new group of plant growth substances that promote plant growth and productivity. We showed in this study that improved growth of cucumber (Cucumis sativus) plants after treatment with 24-epibrassinolide (EBR), an active BR, was associated with increased CO₂ assimilation and quantum yield of PSII (Φ_PSII). Treatment of brassinazole (Brz), a specific inhibitor for BR biosynthesis, reduced plant growth and at the same time decreased CO₂ assimilation and Φ_PSII. Thus, the growth-promoting activity of BRs can be, at least partly, attributed to enhanced plant photosynthesis. To understand how BRs enhance photosynthesis, we have analyzed the effects of EBR and Brz on a number of photosynthetic parameters and their affecting factors, including the contents and activity of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco). Northern and Western blotting demonstrated that EBR upregulated, while Brz downregulated, the expressions of rbcL, rbcS and other photosynthetic genes. In addition, EBR had a positive effect on the activation of Rubisco based on increased maximum Rubisco carboxylation rates (V _c,max), total Rubisco activity and, to a greater extent, initial Rubisco activity. The accumulation patterns of Rubisco activase (RCA) based on immunogold-labeling experiments suggested a role of RCA in BR-regulated activation state of Rubisco. Enhanced expression of genes encoding other Calvin cycle genes after EBR treatment may also play a positive role in RuBP regeneration (J _max), thereby increasing maximum carboxylation rate of Rubisco (V _c,max). Thus, BRs promote photosynthesis and growth by positively regulating synthesis and activation of a variety of photosynthetic enzymes including Rubisco in cucumber.     

The rapid A–Ci response: photosynthesis in the phenomic era

Phenotyping for photosynthetic gas exchange parameters is limiting our ability to select plants for enhanced photosynthetic carbon gain and to assess plant function in current and future natural environments. This is due, in part, to the time required to generate estimates of the maximum rate of ribulose‐1,5‐bisphosphate carboxylase oxygenase (Rubisco) carboxylation (V_c,max) and the maximal rate of electron transport (J_max) from the response of photosynthesis (A) to the CO₂ concentration inside leaf air spaces (C_i). To relieve this bottleneck, we developed a method for rapid photosynthetic carbon assimilation CO₂ responses [rapid A–C_i response (RACiR)] utilizing non‐steady‐state measurements of gas exchange. Using high temporal resolution measurements under rapidly changing CO₂ concentrations, we show that RACiR techniques can obtain measures of V_c,max and J_max in ~5 min, and possibly even faster. This is a small fraction of the time required for even the most advanced gas exchange instrumentation. The RACiR technique, owing to its increased throughput, will allow for more rapid screening of crops, mutants and populations of plants in natural environments, bringing gas exchange into the phenomic era.     

Contrasting acclimation responses to elevated CO2 and warming between an evergreen and a deciduous boreal conifer

Rising atmospheric carbon dioxide (CO₂) concentrations may warm northern latitudes up to 8°C by the end of the century. Boreal forests play a large role in the global carbon cycle, and the responses of northern trees to climate change will thus impact the trajectory of future CO₂ increases. We grew two North American boreal tree species at a range of future climate conditions to assess how growth and carbon fluxes were altered by high CO₂ and warming. Black spruce (Picea mariana, an evergreen conifer) and tamarack (Larix laricina, a deciduous conifer) were grown under ambient (407 ppm) or elevated CO₂ (750 ppm) and either ambient temperatures, a 4°C warming, or an 8°C warming. In both species, the thermal optimum of net photosynthesis (T_optA) increased and maximum photosynthetic rates declined in warm‐grown seedlings, but the strength of these changes varied between species. Photosynthetic capacity (maximum rates of Rubisco carboxylation, V_cmax, and of electron transport, J_max) was reduced in warm‐grown seedlings, correlating with reductions in leaf N and chlorophyll concentrations. Warming increased the activation energy for V_cmax and J_max (E_aV and E_aJ, respectively) and the thermal optimum for J_max. In both species, the T_optA was positively correlated with both E_aV and E_aJ, but negatively correlated with the ratio of J_max/V_cmax. Respiration acclimated to elevated temperatures, but there were no treatment effects on the Q₁₀ of respiration (the increase in respiration for a 10°C increase in leaf temperature). A warming of 4°C increased biomass in tamarack, while warming reduced biomass in spruce. We show that climate change is likely to negatively affect photosynthesis and growth in black spruce more than in tamarack, and that parameters used to model photosynthesis in dynamic global vegetation models (E_aV and E_aJ) show no response to elevated CO₂.     

Seasonal response of photosynthesis to elevated CO2 in loblolly pine (Pinus taeda L.) over two growing seasons

Trees growing in natural systems undergo seasonal changes in environmental factors that generate seasonal differences in net photosynthetic rates. To examine how seasonal changes in the environment affect the response of net photosynthetic rates to elevated CO₂, we grew Pinus taeda L. seedlings for three growing seasons in open-top chambers continuously maintained at either ambient or ambient + 30 Pa CO₂. Seedlings were grown in the ground, under natural conditions of light, temperature nd nutrient and water availability. Photosynthetic capacity was measured bimonthly using net photosynthetic rate vs. intercellular CO₂ partial pressure (A-C_i) curves. Maximum Rubisco activity (Vc_max) and ribulose 1,5-bisphosphate regeneration capacity mediated by electron transport (J_max) and phosphate regeneration (PiRC) were calculated from A-C_i curves using a biochemically based model. Rubisco activity, activation state and content, and leaf carbohydrate, chlorophyll and nitrogen concentrations were measured concurrently with photosynthesis measurements. This paper presents results from the second and third years of treatment. Mean leaf nitrogen concentrations ranged from 13.7 to 23.8 mg g^?1, indicating that seedlings were not nitrogen deficient. Relative to ambient CO₂ seedlings, elevated CO₂ increased light-saturated net photosynthetic rates 60–110% during the summer, but < 30% during the winter. A relatively strong correlation between leaf temperature and the relative response of net photosynthetic rates to elevated CO₂ suggests a strong effect of leaf temperature. During the third growing season, elevated CO₂ reduced Rubisco activity 30% relative to ambient CO₂ seedlings, nearly completely balancing Rubisco and RuBP-regeneration regulation of photosynthesis. However, reductions in Rubisco activity did not eliminate the seasonal pattern in the relative response of net photosynthetic rates to elevated CO₂. These results indicate that seasonal differences in the relative response of net photosynthetic rates to elevated CO₂ are likely to occur in natural systems.     

Mechanisms underlying leaf photosynthetic acclimation to warming and elevated CO2 as inferred from least‐cost optimality theory

The mechanisms responsible for photosynthetic acclimation are not well understood, effectively limiting predictability under future conditions. Least‐cost optimality theory can be used to predict the acclimation of photosynthetic capacity based on the assumption that plants maximize carbon uptake while minimizing the associated costs. Here, we use this theory as a null model in combination with multiple datasets of C₃ plant photosynthetic traits to elucidate the mechanisms underlying photosynthetic acclimation to elevated temperature and carbon dioxide (CO₂). The model‐data comparison showed that leaves decrease the ratio of the maximum rate of electron transport to the maximum rate of Rubisco carboxylation (J_max/V_cmax) under higher temperatures. The comparison also indicated that resources used for Rubisco and electron transport are reduced under both elevated temperature and CO₂. Finally, our analysis suggested that plants underinvest in electron transport relative to carboxylation under elevated CO₂, limiting potential leaf‐level photosynthesis under future CO₂ concentrations. Altogether, our results show that acclimation to temperature and CO₂ is primarily related to resource conservation at the leaf level. Under future, warmer, high CO₂ conditions, plants are therefore likely to use less nutrients for leaf‐level photosynthesis, which may impact whole‐plant to ecosystem functioning.     

A mechanistic evaluation of photosynthetic acclimation at elevated CO2

Plants grown at elevated pCO₂ often fail to sustain the initial stimulation of net CO₂ uptake rate (A). This reduced, acclimated, stimulation of A often occurs concomitantly with a reduction in the maximum carboxylation velocity (V_c,max) of Rubisco. To investigate this relationship we used the Farquhar model of C₃ photosynthesis to predict the minimum V_c,max capable of supporting the acclimated stimulation in A observed at elevated pCO₂. For a wide range of species grown at elevated pCO₂ under contrasting conditions we found a strong correlation between observed and predicted values of V_c,max. This exercise mechanistically and quantitatively demonstrated that the observed acclimated stimulation of A and the simultaneous decrease in V_c,max observed at elevated pCO₂ is mechanistically consistent. With the exception of plants grown at a high elevated pCO₂ (> 90 Pa), which show evidence of an excess investment in Rubisco, the failure to maintain the initial stimulation of A is almost entirely attributable to the decrease in V_c,max and investment in Rubisco is coupled to requirements.     

The response of photosynthetic model parameters to temperature and nitrogen concentration in Pinus radiata D. Don

Responses of photosynthesis (A) to intercellular CO₂ concentration (c_i) in 2-year-old Pinus radiata D. Don seedlings were measured at a range of temperatures in order to parametrize a biophysical model of leaf photosynthesis. Increasing leaf temperature from 8 to 30°C caused a 4-fold increase in V_cmax, the maximum rate of carboxylation (10.7–43.3 μol m^?2 s^?1 and a 3-fold increase in J_max, the maximum electron transport rate (20.5–60.2 μmol m ^?2 s^?1). The temperature optimum for J_max was lower than that for V_cmax, causing a decline in the ratio J_max:V_cmax from 2.0 to 1.4 as leaf temperature increased from 8 to 30°C. To determine the response of photosynthesis to leaf nitrogen concentration, additional measurements were made on seedlings grown under four nitrogen treatments. Foliar N concentrations varied between 0.36 and 1.27 mol kg^?1, and there were linear relationships between N concentration and both V_cmax and J_max. Measurements made throughout the crown of a plantation forest tree, where foliar N concentrations varied from 0.83 mol kg^?1 near the base to 1.54 mol kg^?1 near the leader, yielded similar relationships. These results will be useful in scaling carbon assimilation models from leaves to canopies.     

Co‐limitation of photosynthetic capacity by nitrogen and phosphorus in West Africa woodlands

Photosynthetic leaf traits were determined for savanna and forest ecosystems in West Africa, spanning a large range in precipitation. Standardized major axis fits revealed important differences between our data and reported global relationships. Especially for sites in the drier areas, plants showed higher photosynthetic rates for a given N or P when compared with relationships from the global data set. The best multiple regression for the pooled data set estimated V_cmax and J_max from N_DW and S. However, the best regression for different vegetation types varied, suggesting that the scaling of photosynthesis with leaf traits changed with vegetation types. A new model is presented representing independent constraints by N and P on photosynthesis, which can be evaluated with or without interactions with S. It assumes that limitation of photosynthesis will result from the least abundant nutrient, thereby being less sensitive to the allocation of the non‐limiting nutrient to non‐photosynthetic pools. The model predicts an optimum proportionality for N and P, which is distinct for V_cmax and J_max and inversely proportional to S. Initial tests showed the model to predict V_cmax and J_max successfully for other tropical forests characterized by a range of different foliar N and P concentrations.     

Diffusion limitations and metabolic factors associated with inhibition and recovery of photosynthesis from drought stress in a C3 perennial grass species

Stomatal closure and metabolic impairment under drought stress limits photosynthesis. The objective of this study was to determine major stomatal and metabolic factors involved in photosynthetic responses to drought and recovery upon re‐watering in a C₃ perennial grass species, Kentucky bluegrass (Poa pratensis L.). Two genotypes differing in drought resistance, ‘Midnight’ (tolerant) and ‘Brilliant’ (sensitive), were subjected to drought stress for 15 days and then re‐watered for 10 days in growth chambers. Single‐leaf net photosynthetic rate (A), stomatal conductance (g_s) and transpiration rate (Tr) decreased during drought, with a less rapid decline in ‘Midnight’ than in ‘Brilliant’. Photochemical efficiency, Rubisco activity and activation state declined during drought, but were significantly higher in ‘Midnight’ than in ‘Brilliant’. The relationship between A and internal leaf CO₂ concentration (A/Ci curve) during drought and re‐watering was analyzed to estimate the relative influence of stomatal and non‐stomatal components on photosynthesis. Stomatal limitation (Ls %), non‐stomatal limitation (Lns %), CO₂ compensation point (CP) and dark respiration (Rd) increased with stress duration in both genotypes, but to a lesser extent in ‘Midnight’. Maximum CO₂ assimilation rate (A_max), carboxylation efficiency (CE) and mesophyll conductance (g_m) declined, but ‘Midnight’ had significantly higher levels of A_max, CE and g_m than ‘Brilliant’. Maximum carboxylation rate of Rubisco (V_cmax) and ribulose‐1,5‐bisphospate (RuBP) regeneration capacity mediated by maximum electron transport rate (J_max) decreased from moderate to severe drought stress in both genotypes, but to a greater extent in ‘Brilliant’ than in ‘Midnight’. After re‐watering, RWC restored to about 90% of the control levels in both genotypes, whereas A, g_s, Tr and Fv/Fm was only partially recovered, with a higher recovery level in ‘Midnight’ than in ‘Brilliant’. Rubisco activity and activation state restored to the control level after re‐watering, with more rapid increase in ‘Midnight’ than in ‘Brilliant’. The values of Ls, Lns, CP and Rd declined, and A_max, CE, V_cmax, J_max and g_m increased after re‐watering, with more rapid change in all parameters in ‘Midnight’ than in ‘Brilliant’. These results indicated that the maintenance of higher A and A_max under drought stress in drought‐tolerant Kentucky bluegrass could be attributed to higher Rubico activation state, higher CE and less stomatal limitation. The ability to resume metabolic activity (A_max, CE, Fv/Fm and Rubisco) was observed in the drought‐tolerant genotype and is the most likely cause for the increased recuperative ability of photosynthesis. Incomplete recovery of photosynthesis upon re‐watering could be attributable to lasting stomatal limitations caused by severe drought damage in both genotypes. Promoting rapid stomatal recovery from drought stress may be critical for plants to resume full photosynthetic capacity in C₃ perennial grass species.     

Temperature response of parameters of a biochemically based model of photosynthesis. I. Seasonal changes in mature maritime pine (Pinus pinaster Ait.)

Responses of plant processes to temperature may vary according to the time scale on which they are measured. In this study, both short‐term and seasonal responses of photosynthesis to temperature were examined. A field study of seasonal changes in the temperature response of photosynthesis was conducted on two provenances, French and Moroccan, of mature maritime pine (Pinus pinaster Ait.). Measurements were made every 2 months over a 1‐year period and used to parameterize a mechanistic model of photosynthesis. Temperature responses of maximum Rubisco activity, V_cmax, and potential electron transport rate, J_max, were obtained for each measurement period, as was the response of stomatal conductance, g_s, to water vapour pressure deficit (VPD). Absolute values of V_cmax and J_max at 25 °C were related to needle nitrogen content, N_area.N_area, and thus V_cmax and J_max, were negatively correlated with the mean minimum temperature in the month preceding measurements. The ratio of J_max : V_cmax at 25 °C varied between 1 and 1·7 but did not show any seasonal trend. Nor was there any seasonal trend in the relative temperature response of V_cmax, which had an activation energy H_a of approximately 57 kJ mol^?1 throughout the experiment. The activation energy of J_max was also close to constant throughout the experiment, averaging 39 kJ mol^?1. For the French provenance, the optimal temperature of J_max was positively correlated with the maximum temperature of the previous day, but no such correlation was found for the Moroccan provenance. The response of g_s to VPD also varied seasonally, with much stronger stomatal closure in winter months. Taken together, these results implied a translational shift downwards of the photosynthetic temperature response curve with increasing T_prev, and a shift in the temperature optimum of photosynthesis of 5–10 °C between summer and winter. These results illustrate that the short‐term temperature response of photosynthesis varies significantly on a seasonal basis.     

Effect of leaf age and position on light-saturated CO<Subscript>2</Subscript> assimilation rate,photosynthetic capacity,and stomatal conductance in rubber trees

Shoots of the tropical latex-producing tree Hevea brasiliensis (rubber tree) grow according to a periodic pattern, producing four to five whorls of leaves per year. All leaves in the same whorl were considered to be in the same leaf-age class, in order to assess the evolution of photosynthesis with leaf age in three clones of rubber trees, in a plantation in eastern Thailand. Light-saturated CO₂ assimilation rate (A _max) decreased more with leaf age than did photosynthetic capacity (maximal rate of carboxylation, V _cmax , and maximum rate of electron transport, J _max), which was estimated by fitting a biochemical photosynthesis model to the CO₂-response curves. Nitrogen-use efficiency (A _max/N_a, N_a is nitrogen content per leaf area) decreased also with leaf age, whereas J _max and V _cmax did not correlate with N _a. Although measurements were performed during the rainy season, the leaf gas exchange parameter that showed the best correlation with A _max was stomatal conductance (g _s). An asymptotic function was fitted to the A _max-g _s relationship, with R ² = 0.85. A _max, V _cmax, J _max and g _s varied more among different whorls in the same clone than among different clones in the same whorl. We concluded that leaf whorl was an appropriate parameter to characterize leaves for the purpose of modelling canopy photosynthesis in field-grown rubber trees, and that stomatal conductance was the most important variable explaining changes in A _max with leaf age in rubber trees.     

The growth of soybean under free air [CO<Subscript>2</Subscript>] enrichment (FACE) stimulates photosynthesis while decreasing in vivo Rubisco capacity

Down-regulation of light-saturated photosynthesis (A_sat) at elevated atmospheric CO₂ concentration, [CO₂], has been demonstrated for many C₃ species and is often associated with inability to utilize additional photosynthate and/or nitrogen limitation. In soybean, a nitrogen-fixing species, both limitations are less likely than in crops lacking an N-fixing symbiont. Prior studies have used controlled environment or field enclosures where the artificial environment can modify responses to [CO₂]. A soybean free air [CO₂] enrichment (FACE) facility has provided the first opportunity to analyze the effects of elevated [CO₂] on photosynthesis under fully open-air conditions. Potential ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) carboxylation (V_c,max) and electron transport through photosystem II (J_max) were determined from the responses of A_sat to intercellular [CO₂] (C_i) throughout two growing seasons. Mesophyll conductance to CO₂ (g_m) was determined from the responses of A_sat and whole chain electron transport (J) to light. Elevated [CO₂] increased A_sat by 15–20% even though there was a small, statistically significant, decrease in V_c,max. This differs from previous studies in that V_c,max/J_max decreased, inferring a shift in resource investment away from Rubisco. This raised the C_i at which the transition from Rubisco-limited to ribulose-1,5-bisphosphate regeneration-limited photosynthesis occurred. The decrease in V_c,max was not the result of a change in g_m, which was unchanged by elevated [CO₂]. This first analysis of limitations to soybean photosynthesis under fully open-air conditions reveals important differences to prior studies that have used enclosures to elevate [CO₂], most significantly a smaller response of A_sat and an apparent shift in resources away from Rubisco relative to capacity for electron transport.Abbreviations FACE Free air [CO₂] enrichment - Rubisco Ribulose-1,5-bisphosphate carboxylase/oxygenase - RuBP Ribulose-1,5-bisphosphate - SoyFACE Soybean free air [CO₂] enrichment - VPD Vapor pressure deficit     

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.     

Comparison of the A–Cc curve fitting methods in determining maximum ribulose 1·5-bisphosphate carboxylase/oxygenase carboxylation rate, potential light saturated electron transport rate and leaf dark respiration

A review of the literature revealed that a variety of methods are currently used for fitting net assimilation of CO₂–chloroplastic CO₂ concentration (A–C_c) curves, resulting in considerable differences in estimating the A–C_c parameters [including maximum ribulose 1·5‐bisphosphate carboxylase/oxygenase (Rubisco) carboxylation rate (V_cmax), potential light saturated electron transport rate (J_max), leaf dark respiration in the light (R_d), mesophyll conductance (g_m) and triose‐phosphate utilization (TPU)]. In this paper, we examined the impacts of fitting methods on the estimations of V_cmax, J_max, TPU, R_d and g_m using grid search and non‐linear fitting techniques. Our results suggested that the fitting methods significantly affected the predictions of Rubisco‐limited (A_c), ribulose 1,5‐bisphosphate‐limited (A_j) and TPU‐limited (A_p) curves and leaf photosynthesis velocities because of the inconsistent estimate of V_cmax, J_max, TPU, R_d and g_m, but they barely influenced the J_max : V_cmax, V_cmax : R_d and J_max : TPU ratio. In terms of fitting accuracy, simplicity of fitting procedures and sample size requirement, we recommend to combine grid search and non‐linear techniques to directly and simultaneously fit V_cmax, J_max, TPU, R_d and g_m with the whole A–C_c curve in contrast to the conventional method, which fits V_cmax, R_d or g_m first and then solves for V_cmax, J_max and/or TPU with V_cmax, R_d and/or g_m held as constants.     

The relationship of leaf photosynthetic traits – Vcmax and Jmax – to leaf nitrogen,leaf phosphorus,and specific leaf area: a meta‐analysis and modeling study

Great uncertainty exists in the global exchange of carbon between the atmosphere and the terrestrial biosphere. An important source of this uncertainty lies in the dependency of photosynthesis on the maximum rate of carboxylation (V_cmax) and the maximum rate of electron transport (J_max). Understanding and making accurate prediction of C fluxes thus requires accurate characterization of these rates and their relationship with plant nutrient status over large geographic scales. Plant nutrient status is indicated by the traits: leaf nitrogen (N), leaf phosphorus (P), and specific leaf area (SLA). Correlations between V_cmax and J_max and leaf nitrogen (N) are typically derived from local to global scales, while correlations with leaf phosphorus (P) and specific leaf area (SLA) have typically been derived at a local scale. Thus, there is no global-scale relationship between V_cmax and J_max and P or SLA limiting the ability of global-scale carbon flux models do not account for P or SLA. We gathered published data from 24 studies to reveal global relationships of V_cmax and J_max with leaf N, P, and SLA. V_cmax was strongly related to leaf N, and increasing leaf P substantially increased the sensitivity of V_cmax to leaf N. J_max was strongly related to V_cmax, and neither leaf N, P, or SLA had a substantial impact on the relationship. Although more data are needed to expand the applicability of the relationship, we show leaf P is a globally important determinant of photosynthetic rates. In a model of photosynthesis, we showed that at high leaf N (3 gm⁻²), increasing leaf P from 0.05 to 0.22 gm⁻² nearly doubled assimilation rates. Finally, we show that plants may employ a conservative strategy of J_max to V_cmax coordination that restricts photoinhibition when carboxylation is limiting at the expense of maximizing photosynthetic rates when light is limiting.     

Is stimulation of leaf photosynthesis by elevated carbon dioxide concentration maintained in the long term? A test with Lolium perenne grown for 10 years at two nitrogen fertilization levels under Free Air CO2Enrichment (FACE)

Photosynthesis is commonly stimulated in grasslands with experimental increases in atmospheric CO₂ concentration ([CO₂]), a physiological response that could significantly alter the future carbon cycle if it persists in the long term. Yet an acclimation of photosynthetic capacity suggested by theoretical models and short‐term experiments could completely remove this effect of CO₂. Perennial ryegrass (Lolium perenne L. cv. Bastion) was grown under an elevated [CO₂] of 600 µmol mol^?1 for 10 years using Free Air CO₂Enrichment (FACE), with two contrasting nitrogen levels and abrupt changes in the source : sink ratio following periodic harvests. More than 3000 measurements characterized the response of leaf photosynthesis and stomatal conductance to elevated [CO₂] across each growing season for the duration of the experiment. Over the 10 years as a whole, growth at elevated [CO₂] resulted in a 43% higher rate of light‐saturated leaf photosynthesis and a 36% increase in daily integral of leaf CO₂ uptake. Photosynthetic stimulation was maintained despite a 30% decrease in stomatal conductance and significant decreases in both the apparent, maximum carboxylation velocity (V_c,max) and the maximum rate of electron transport (J_max). Immediately prior to the periodic (every 4–8 weeks) cuts of the L. perenne stands, V_c,max and J_max, were significantly lower in elevated than in ambient [CO₂] in the low‐nitrogen treatment. This difference was smaller after the cut, suggesting a dependence upon the balance between the sources and sinks for carbon. In contrast with theoretical expectations and the results of shorter duration experiments, the present results provide no significant change in photosynthetic stimulation across a 10‐year period, nor greater acclimation in V_c,max and J_max in the later years in either nitrogen treatment.     

Estimation of photosynthesis parameters for a modified Farquhar–von Caemmerer–Berry model using simultaneous estimation method and nonlinear mixed effects model

The aims of this paper was to modify the photosynthesis model of Farquhar, von Caemmerer and Berry (FvCB) to be able to predict light dependency of the carboxylation capacity (V_c) and to improve the prediction of temperature dependency of the maximum carboxylation capacity (V_cmax) and the maximum electron transport rate (J_max). The FvCB model was modified by adding a sub-model for Ribulose-1,5-bisphosphate carboxylase (Rubisco) activation and validating the parameters for temperature dependency of V_cmax and J_max. Values of parameters for temperature dependency of V_cmax and J_max were validated and adjusted based on data of the photosynthesis response to temperature. Parameter estimation was based on measurements under a wide range of environmental conditions, providing parameters with broad validity. The simultaneous estimation method and the nonlinear mixed effects model were applied to ensure the accuracy of the parameter estimation. The FvCB parameters, V_cmax, J_max, α (the efficiency of light energy conversion), θ (the curvature of light response of electron transport), and R_d (the non-photorespiratory CO₂ release) were estimated and validated on a dataset from two other years. Observations and predictions matched well (R² = 0.94). We conclude that incorporating a sub-model of Rubisco activation improved the FvCB model through predicting light dependency of carboxylation rate; and that estimating V_cmax, J_max, α, θ, and R_d requires data sets of both CO₂ and light response curves.     

Temperature response of parameters of a biochemically based model of photosynthesis. II. A review of experimental data

The temperature dependence of C₃ photosynthesis is known to vary with growth environment and with species. In an attempt to quantify this variability, a commonly used biochemically based photosynthesis model was parameterized from 19 gas exchange studies on tree and crop species. The parameter values obtained described the shape and amplitude of the temperature responses of the maximum rate of Rubisco activity (V_cmax) and the potential rate of electron transport (J_max). Original data sets were used for this review, as it is shown that derived values of V_cmax and its temperature response depend strongly on assumptions made in derivation. Values of J_max and V_cmax at 25 °C varied considerably among species but were strongly correlated, with an average J_max : V_cmax ratio of 1·67. Two species grown in cold climates, however, had lower ratios. In all studies, the J_max : V_cmax ratio declined strongly with measurement temperature. The relative temperature responses of J_max and V_cmax were relatively constant among tree species. Activation energies averaged 50 kJ mol^?1 for J_max and 65 kJ mol^?1 for V_cmax, and for most species temperature optima averaged 33 °C for J_max and 40 °C for V_cmax. However, the cold climate tree species had low temperature optima for both J_max(19 °C) and V_cmax (29 °C), suggesting acclimation of both processes to growth temperature. Crop species had somewhat different temperature responses, with higher activation energies for both J_max and V_cmax, implying narrower peaks in the temperature response for these species. The results thus suggest that both growth environment and plant type can influence the photosynthetic response to temperature. Based on these results, several suggestions are made to improve modelling of temperature responses.