Acclimation of photosynthesis to temperature in <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis> and <Emphasis Type="Italic">Brassica oleracea</Emphasis>

Plants differ in how much the response of net photosynthetic rate (P _N) to temperature (T) changes with the T during leaf development, and also in the biochemical basis of such changes in response. The amount of photosynthetic acclimation to T and the components of the photosynthetic system involved were compared in Arabidopsis thaliana and Brassica oleracea to determine how well A. thaliana might serve as a model organism to study the process of photosynthetic acclimation to T. Responses of single-leaf gas exchange and chlorophyll fluorescence to CO₂ concentration measured over the range of 10–35 °C for both species grown at 15, 21, and 27 °C were used to determine the T dependencies of maximum rates of carboxylation (V_Cmax), photosynthetic electron transport (J_max), triose phosphate utilization rate (TPU), and mesophyll conductance to carbon dioxide (g’_m). In A. thaliana, the optimum T of P _N at air concentrations of CO₂ was unaffected by this range of growth T, and the T dependencies of V_Cmax, J_max, and g’_m were also unaffected by growth T. There was no evidence of TPU limitation of P _N in this species over the range of measurement conditions. In contrast, the optimum T of P _N increased with growth T in B. oleracea, and the T dependencies of V_Cmax, J_max, and g’_m, as well as the T at which TPU limited P _N all varied significantly with growth T. Thus B. oleracea had much a larger capacity to acclimate photosynthetically to moderate T than did A. thaliana.     

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.     

PHOTOSYNTHETIC SENSITIVITY TO TEMPERATURE IN POPULATIONS OF TWO C4 BOUTELOUA (POACEAE) SPECIES NATIVE TO DIFFERENT ALTITUDES

The relationships between photosynthesis, flowering, and growth temperatures were examined experimentally in four populations of the C₄ grass genus Bouteloua. Field-collected plants were grown under two temperature regimes, cool (20 C day/6 C night) and warm (30/16), representative of the extreme populations. Populations collected from the warm climates had significantly lower photosynthetic capacity when grown in the cool chamber relative to the warm chamber, while photosynthetic capacity in the cool climate populations did not differ between the growth conditions. Additionally, exposure to a 2-day cold temperature treatment (10/-2), representative of late-season frosts in high altitude sites, resulted in further reductions in photosynthesis in the warm climate plants, but not in the cool climate plants. This effect was greater for plants grown in the cool growth chamber. Flowering was reduced by 70% in the warm climate plants grown in the cool chamber, and was correlated with photosynthetic inhibition following the short-term cold temperature treatment. These results indicate that genetic differentiation for photosynthetic temperature sensitivity has occurred in the cool climate populations, and that long-term exposure to cool temperatures coupled with short-term relatively extreme low temperatures results in greater photosynthetic inhibition in nontolerant populations.     

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.     

Optimum leaf size predicted by a novel leaf energy balance model incorporating dependencies of photosynthesis on light and temperature

To clarify relationships between leaf size and the environment variables, we constructed an energy balance model for a single leaf incorporating Leuning’s stomatal conductance model and Farquhar’s leaf photosynthesis model. We ran this model for various environmental conditions paying particular attention to the leaf boundary layer. The leaf size maximizing the rate of photosynthesis per unit leaf area (A) at a high irradiance differed depending on the air temperature. In warm environments, A increased with decrease in leaf size, whereas in cool environments, there was the leaf size maximizing A. With the increase in leaf size, the CO₂ concentration inside the leaf (C _i) decreased and the leaf temperature increased, both due to lower boundary layer conductance. At low air temperatures, the negative effect of low C _i on A in large leaves was compensated by the increase in leaf temperature towards the optimum temperature for A. This balance determined the optimum leaf size for A at low air temperatures. With respect to water use efficiency, large leaves tended to be advantageous, especially in cool environments at low-to-medium irradiances. Some temperature-dependent trends in leaf size observed in nature are discussed based on the present results.     

Temperature-dependent feedback inhibition of photosynthesis in peanut

Arachis hypogaea L. is a tropical crop that is slow-growing at temperatures below 25°C. Unadapted CO₂-assimilation rate (A) showed insufficient variation between 15 and 30°C in the short term (hours) to explain this marked reduction in growth. However, at longer periods (12 d), A was depressed as were growth rate and leafproduction rate. To examine the possible relationship between growth, A and sink demand plants were transferred from 30°C, which is near the optimum for growth, to a suboptimal temperature (19°C). In the first 2 d of cooling, A decreased by 50–70%, the stomata stayed open, and the intercellular CO₂ concentration (c_i) rose, i.e. the decrease in A of the cooled plants was the result of non-stomatal factors. Changes in dark respiration did not account for the decline in A.Clear evidence was obtained of sink control of A by independently manipulating the temperature of different leaves on the plant. Cooling (to 19°C) most of the plant (the sink) led to a 70% decline in A of the remaining leaves at 30°C after 3 d, whereas the converse treatments (30°C sink, 19°C source) resulted in small changes (17%). In plants at 19°C which were exposed to low CO₂ concentration to prevent photosynthesis, A was not reduced when measured at normal CO₂ concentrations, indicating that carbohydrate accumulation was responsible for the decline in A. Dry-matter build-up at suboptimal temperature was also consistent with end-product inhibition of photosynthesis.Abbreviations and symbols A (mol·m^-2·s^-1) rate of net CO₂ assimilation - C_i (l·l^-1) substomatal CO₂ concentration - DW (g) dry weight - g (mol·m^-2·s^-1) stomatal conductance to diffusion of water vapour - PFD (mol·m^-2·s^-1) photon flux density     

Surprising lack of sensitivity of biochemical limitation of photosynthesis of nine tree species to open‐air experimental warming and reduced rainfall in a southern boreal forest

Photosynthetic biochemical limitation parameters (i.e., V_cmax, J_max and J_max:V_cmax ratio) are sensitive to temperature and water availability, but whether these parameters in cold climate species at biome ecotones are positively or negatively influenced by projected changes in global temperature and water availability remains uncertain. Prior exploration of this question has largely involved greenhouse based short‐term manipulative studies with mixed results in terms of direction and magnitude of responses. To address this question in a more realistic context, we examined the effects of increased temperature and rainfall reduction on the biochemical limitations of photosynthesis using a long‐term chamber‐less manipulative experiment located in northern Minnesota, USA. Nine tree species from the boreal‐temperate ecotone were grown in natural neighborhoods under ambient and elevated (+3.4°C) growing season temperatures and ambient or reduced (≈40% of rainfall removed) summer rainfall. Apparent rubisco carboxylation and RuBP regeneration standardized to 25°C (V_cmax25°C and J_max25°C, respectively) were estimated based on AC_i curves measured in situ over three growing seasons. Our primary objective was to test whether species would downregulate V_cmax25°C and J_max25°C in response to warming and reduced rainfall, with such responses expected to be greatest in species with the coldest and most humid native ranges, respectively. These hypotheses were not supported, as there were no overall main treatment effects on V_cmax25°C or J_max25°C (p > .14). However, J_max:V_cmax ratio decreased significantly with warming (p = .0178), whereas interactions between warming and rainfall reduction on the J_max25°C to V_cmax25°C ratio were not significant. The insensitivity of photosynthetic parameters to warming contrasts with many prior studies done under larger temperature differentials and often fixed daytime temperatures. In sum, plants growing in relatively realistic conditions under naturally varying temperatures and soil moisture levels were remarkably insensitive in terms of their J_max25°C and V_cmax25°C when grown at elevated temperatures, reduced rainfall, or both combined.     

In situ temperature relationships of biochemical and stomatal controls of photosynthesis in four lowland tropical tree species

Net photosynthetic carbon uptake of Panamanian lowland tropical forest species is typically optimal at 30–32 °C. The processes responsible for the decrease in photosynthesis at higher temperatures are not fully understood for tropical trees. We determined temperature responses of maximum rates of RuBP‐carboxylation (V_CMax) and RuBP‐regeneration (J_Max), stomatal conductance (G_s), and respiration in the light (R_Light) in situ for 4 lowland tropical tree species in Panama. G_s had the lowest temperature optimum (T_Opt), similar to that of net photosynthesis, and photosynthesis became increasingly limited by stomatal conductance as temperature increased. J_Max peaked at 34–37 °C and V_CMax ~2 °C above that, except in the late‐successional species Calophyllum longifolium, in which both peaked at ~33 °C. R_Light significantly increased with increasing temperature, but simulations with a photosynthesis model indicated that this had only a small effect on net photosynthesis. We found no evidence for Rubisco‐activase limitation of photosynthesis. T_Opt of V_CMax and J_Max fell within the observed in situ leaf temperature range, but our study nonetheless suggests that net photosynthesis of tropical trees is more strongly influenced by the indirect effects of high temperature—for example, through elevated vapour pressure deficit and resulting decreases in stomatal conductance—than by direct temperature effects on photosynthetic biochemistry and respiration.     

Low-temperature photosynthetic performance of a C4 grass and a co-occurring C3 grass native to high latitudes

The photosynthetic performance of C₄ plants is generally inferior to that of C₃ species at low temperatures, but the reasons for this are unclear. The present study investigated the hypothesis that the capacity of Rubisco, which largely reflects Rubisco content, limits C₄ photosynthesis at suboptimal temperatures. Photosynthetic gas exchange, chlorophyll a fluorescence, and the in vitro activity of Rubisco between 5 and 35 °C were measured to examine the nature of the low‐temperature photosynthetic performance of the co‐occurring high latitude grasses, Muhlenbergia glomerata (C₄) and Calamogrostis canadensis (C₃). Plants were grown under cool (14/10 °C) and warm (26/22 °C) temperature regimes to examine whether acclimation to cool temperature alters patterns of photosynthetic limitation. Low‐temperature acclimation reduced photosynthetic rates in both species. The catalytic site concentration of Rubisco was approximately 5.0 and 20 µmol m^?2 in M. glomerata and C. canadensis, respectively, regardless of growth temperature. In both species, in vivo electron transport rates below the thermal optimum exceeded what was necessary to support photosynthesis. In warm‐grown C. canadensis, the photosynthesis rate below 15 °C was unaffected by a 90% reduction in O₂ content, indicating photosynthetic capacity was limited by the capacity of P_i‐regeneration. By contrast, the rate of photosynthesis in C. canadensis plants grown at the cooler temperatures was stimulated 20–30% by O₂ reduction, indicating the P_i‐regeneration limitation was removed during low‐temperature acclimation. In M. glomerata, in vitro Rubisco activity and gross CO₂ assimilation rate were equivalent below 25 °C, indicating that the capacity of the enzyme is a major rate limiting step during C₄ photosynthesis at cool temperatures.     

Effects of irradiance and temperature on photosynthesis in C3, C4 and C3/C4 Panicum species

Species in the Laxa and Grandia groups of the genus Panicum are adapted to low, wet areas of tropical and subtropical America. Panicum milioides is a species with C₃ photosynthesis and low apparent photorespiration and has been classified as a C₃/C₄ intermediate. Other species in the Laxa group are C₃ with normal photorespiration. Panicum prionitis is a C₄ species in the Grandia group. Since P. milioides has some leaf characteristics intermediate to C₃ and C₄ species, its photosynthetic response to irradiance and temperature was compared to the closely related C₃ species, P. laxum and P. boliviense and to P. prionitis. The response of apparent photosynthesis to irradiance and temperature was similar to that of P. laxum and P. boliviense, with saturation at a photosynthetic photo flux density of about 1 mmol m^-2 s^-1 at 30°C and temperature optimum near 30°C. In contrast, P. prionitis showed no light saturation up to 2 mmol m^-2 s^-1 and an optimum temperature near 40°C. P. milioides exhibited low CO₂ loss into CO₂-free air in the light and this loss was nearly insensitive to temperature. Loss of CO₂ in the light in the C₃ species, P. laxum and P. boliviense, was several-fold higher than in P. milioides and increased 2- to 5-fold with increases in temperature from 10 to 40°C. The level of dark respiration and its response to temperature were similar in all four Panicum species examined. It is concluded that the low apparent photorespiration in P. milioides does not influence its response of apparent photosynthesis to irradiance and temperature in comparison to closely related C₃ Panicum species.Abbreviations AP apparent photosynthesis - I CO₂ compensation point - gl leaf conductance; gm, mesophyll conductance - PPFD photosynthetic photon flux density - PR apparent photorespiration rate - RuBPC sibulose bisphosphate carboxylase     

Dynamic photo-inhibition and carbon gain in a C4 and a C3 grass native to high latitudes

C₄ plants are rare in the cool climates characteristic of high latitudes and altitudes, perhaps because of an enhanced susceptibility to photo‐inhibition at low temperatures relative to C₃ species. In the present study we tested the hypothesis that low‐temperature photo‐inhibition is more detrimental to carbon gain in the C₄ grass Muhlenbergia glomerata than the C₃ species Calamogrostis Canadensis. These grasses occur together in boreal fens in northern Canada. Plants were grown under cool (14/10 °C day/night) and warm (26/22 °C) temperatures before measurement of the light responses of photosynthesis and chlorophyll fluorescence at different temperatures. Cool growth temperatures led to reduced rates of photosynthesis in M. glomerata at all measurement temperatures, but had a smaller effect on the C₃ species. In both species the amount of xanthophyll cycle pigments increased when plants were grown at 14/10 °C, and in M. glomerata the xanthophyll epoxidation state was greatly reduced. The detrimental effect of low growth temperature on photosynthesis in M. glomerata was almost completely reversed by a 24‐h exposure to the warm‐temperature regime. These data indicate that reversible dynamic photo‐inhibition is a strategy by which C₄ species may tolerate cool climates and overcome the Rubisco limitation that is prevalent at low temperatures in C₄ plants.     

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.     

Temperature and salinity regulation of growth and gas exchange of Salicornia fruticosa (L.) L.

Summary Salicornia fruticosa was collected from a salt marsh on the Mediterranean sea coast in Libya. Growth and gas exchange of this C₃ species were monitered in plants pretreated at various NaCl concentrations (0, 171, 342, 513 and 855 mM). Maximum growth was at 171 mM NaCl under cool growth conditions (20/10° C) and at 342 mM NaCl under warm growth conditions (30/15° C) with minimum growth at 0 mM NaCl (control). Net photosynthesis (Pn) was greatest in plants grown in 171 mM NaCl with plants grown at 513 and 855 mM having lowest rates. Maximum Pn was at 20–25° C shoot temperatures with statistically significant reductions at 30° C in control plants while salt treated plants showed such reductions at 35° C. Salt treatments increased dark respiration over the control at 171 and 342 mM but reduced it at higher concentrations. Photorespiration was reduced by salt treatment and increased by increasing shoot temperature. Greatest transpiration was in 171 mM NaCl treated plants and increasing shoot temperature increased transpiration in all treatments. Stomatal resistance to CO₂ influx was influenced only moderately by temperature while increasing salinity resulted in increased stomatal resistance. In general both temperature and salinity increased the mesophyll resistance to CO₂ influx. The species seems adapted to the warm saline habitat along the Mediterranean sea coast, at least partially, by its ability to maintain relatively high Pn at moderate NaCl concentrations over a broad range of shoot temperatures.     

C4 photosynthesis in Spartina townsendii at low and high temperatures

The metabolism of ¹⁴CO₂ in the cool temperate saltmarsh grass Spartina townsendii was investigated in plants grown in their natural habitats at two temperatures. Both in the spring at 10°C and in the late summer at 25°C radioactivity was initially incorporated into the organic acids malate and aspartate and then transferred to 3-phosphoglycerate in the manner characteristic of the C₄ pathway of photosynthesis. Metabolism was not disrupted at the lower temperature as in some C₄ plants. Radioactivity was transferred more slowly from malate into alanine, glycine and serine at 10°C, but sugars were labelled equally at both temperatures.     

Contrasting responses of photosynthesis at low temperatures in different annual legume species

Growth, net photosynthetic rate (P _N), chlorophyll fluorescence induction kinetics, and stromal fructose-1,6-bisphosphatase (sFBPase) in annual legumes native to the Mediterranean region, two clovers (Trifolium subterraneum L. ssp. oxaloides Nyman cv. Clare and T. michelianum Savi cv. Giorgia) and two Medicago species (M. polymorpha L. cv. Anglona and M. truncatula Gaertn. cv. Paraggio), shifted from 20 to 10 °C for 1 d or developed at 10 °C were compared with controls kept at 20 °C. Cold development produced a larger stimulation of growth in the clover cv. Giorgia and the Medicago cv. Paraggio. Transferring plants to low temperatures affected P _N relatively less in clovers than in Medicago plants. Development at 10 °C relieved the inhibition of photosynthesis in Giorgia and Paraggio, but not in Clare and Anglona, which correlated with increases in the maximum rate of carboxylation by ribulose-1,5-bisphosphate carboxylase/oxygenase, RuBPCO (V_cmax), and the photon-saturated rate of electron transport (J_max). In Medicago, transfer from high to low temperature inhibited photosynthesis in a lesser extent in Anglona than in Paraggio, which showed severe limitations at level of V_cmax and J_max. Development at 10 °C in Paraggio produced an efficient photosynthetic cold acclimation, this being associated with a two-fold increase of quantum yield of photosystem 2 electron transport (F/F'_m) and with the activity of sFBPase. By contrast, Anglona showed an irreversible inhibition of P _N coupled with the reduction of carbon metabolism by impairment of Calvin cycle enzyme activities such as RuBPCO and sFBPase, resulting in a poor cold acclimation of photosynthesis in this cultivar.     

Field measurements of photosynthesis,water-use efficiency,and growth inAgropyron smithii (C3) andBouteloua gracilis (C4) in the Colorado shortgrass steppe

Summary Field measurements of gas exchange and growth were conducted on a C₃ grass,Agropyron smithii, and a C₄ grass,Bouteloua gracilis, in order to further establish the adaptive significance of the C₄ pathway under natural conditions. Maximum rates of leaf area expansion in tillers and maximum seasonal photosynthesis rates of both species occurred during the cool, early summer month of June. The occurrence of maximum growth and photosynthesis inB. gracilis during this cool period was apparently related to its occupation of warm microenvironments next to the ground surface. As temperatures increased during the midsummer, photosynthesis rates decreased to 47% and 55% of the seasonal maximum inB. gracilis andA. smithii, respectively. Water-use efficiencies in both species were similar or slightly higher forB. gracilis during June, the period of maximum growth. By mid-July, however, leaves of the C₃ grass,A. smithii, exhibited water-use efficiencies approximately half as high asB. gracilis. These differences in water-use efficiency were the result of differences in stomatal conductance, rather than differences in daily CO₂ uptake rates which were similar in both species. The results demonstrate that in certain environments there are no offset periods of growth and maximum photosynthesis during the growing season in these C₃ and C₄ species. The greater amounts of daily water use inA. smithii during the midsummer might contribute to its much greater abundance in lowland sites in the shortgrass steppe. The C₄ grass,B. gracilis, occurs in dry upland sites in addition to the more mesic lowland sites.     

Modelled net carbon gain responses to climate change in boreal trees: Impacts of photosynthetic parameter selection and acclimation

Boreal forests are crucial in regulating global vegetation‐atmosphere feedbacks, but the impact of climate change on boreal tree carbon fluxes is still unclear. Given the sensitivity of global vegetation models to photosynthetic and respiration parameters, we determined how predictions of net carbon gain (C‐gain) respond to variation in these parameters using a stand‐level model (MAESTRA). We also modelled how thermal acclimation of photosynthetic and respiratory temperature sensitivity alters predicted net C‐gain responses to climate change. We modelled net C‐gain of seven common boreal tree species under eight climate scenarios across a latitudinal gradient to capture a range of seasonal temperature conditions. Physiological parameter values were taken from the literature together with different approaches for thermally acclimating photosynthesis and respiration. At high latitudes, net C‐gain was stimulated up to 400% by elevated temperatures and CO₂ in the autumn but suppressed at the lowest latitudes during midsummer under climate scenarios that included warming. Modelled net C‐gain was more sensitive to photosynthetic capacity parameters (V_cmax, J_max, Arrhenius temperature response parameters, and the ratio of J_max to V_cmax) than stomatal conductance or respiration parameters. The effect of photosynthetic thermal acclimation depended on the temperatures where it was applied: acclimation reduced net C‐gain by 10%–15% within the temperature range where the equations were derived but decreased net C‐gain by 175% at temperatures outside this range. Thermal acclimation of respiration had small, but positive, impacts on net C‐gain. We show that model simulations are highly sensitive to variation in photosynthetic parameters and highlight the need to better understand the mechanisms and drivers underlying this variability (e.g., whether variability is environmentally and/or biologically driven) for further model improvement.     

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     

Acclimation of photosynthesis in a boreal grass (Phalaris arundinacea L.) under different temperature, CO2, and soil water regimes

The aim of this work was to study the acclimation of photosynthesis in a boreal grass (Phalaris arundinacea L.) grown in controlled environment chambers under elevated temperature (ambient + 3.5°C) and CO₂ (700 μmol mol⁻¹) with varying soil water regimes. More specifically, we studied, during two development stages (early: heading; late: florescence completed), how the temperature response of light-saturated net photosynthetic rate (P _sat), maximum rate of ribulose-1,5-bisphosphate carboxylase/oxygenase activity (V _cmax) and potential rate of electron transport (J _max) acclimatized to the changed environment. During the early growing period, we found a greater temperature-induced enhancement of P _sat at higher measurement temperatures, which disappeared during the late stage. Under elevated growth temperature, V _cmax and J _max at lower measurement temperatures (5–15°C) were lower than those under ambient growth temperature during the early period. When the measurements were done at 20–30°C, the situation was the opposite. During the late growing period, V _cmax and J _max under elevated growth temperature were consistently lower across measurement temperatures. CO₂ enrichment significantly increased P _sat with higher intercellular CO₂ compared to ambient CO₂ treatment, however, elevated CO₂ slightly decreased V _cmax and J _max across measurement temperatures, probably due to down-regulation acclimation. For two growing periods, soil water availability affected the variation in photosynthesis and biochemical parameters much more than climatic treatment did. Over two growing periods, V _cmax and J _max were on average 36.4 and 30.6%, respectively, lower with low water availability compared to high water availability across measurement temperatures. During the late growing period, elevated growth temperature further reduced the photosynthesis under low water availability. V _cmax and J _max declined along with the decrease in nitrogen content of leaves as growing period progressed, regardless of climatic treatment and water regime. We suggest that, for grass species, seasonal acclimation of the photosynthetic parameters under varying environmental conditions needed to be identified to fairly estimate the whole-life photosynthesis.     

A correlation between photosynthetic temperature adaptation and seasonal phenology patterns in the shortgrass prairie

Summary The temperatures at which chlorophyll fluorescence yield is substantially increased and the temperatures at which the quantum yield for CO₂ uptake is irreversibly inhibited were measured for three shortgrass prairie species. The experimental taxa include, a cool season species (Agropyron smithii), a warm season species (Bouteloua gracilis), and a species which grows throughout the cool and warm seasons (Carex stenophylla). Agropyron smithii exhibited lower high temperature damage thresholds (43°C in cool grown plants, 46°C in warm grown plants), relative to the other two species. Bouteloua gracilis exhibited the highest tolerance to high temperature, with threshold values being 44–49°C for cool grown plants and 53–55°C for warm grown plants. Carex stenophylla exhibited threshold values which were intermediate to the other two species (43–47°C for cool grown plants, and 51–53°C for warm grown plants). Seasonal patterns in the fluorescence rise temperatures of field grown plants indicated acclimation to increased temperatures in all three species. The results demonstrate a correlation between the high temperature thresholds for damage to the photosynthetic apparatus, and in situ seasonal phenology patterns for the three species.