Effects of growth and measurement light intensities on temperature dependence of CO2 assimilation rate in tobacco leaves

Effects of growth light intensity on the temperature dependence of CO₂ assimilation rate were studied in tobacco (Nicotiana tabacum) because growth light intensity alters nitrogen allocation between photosynthetic components. Leaf nitrogen, ribulose 1·5‐bisphosphate carboxylase/oxygenase (Rubisco) and cytochrome f (cyt f) contents increased with increasing growth light intensity, but the cyt f/Rubisco ratio was unaltered. Mesophyll conductance to CO₂ diffusion (g_m) measured with carbon isotope discrimination increased with growth light intensity but not with measuring light intensity. The responses of CO₂ assimilation rate to chloroplast CO₂ concentration (C_c) at different light intensities and temperatures were used to estimate the maximum carboxylation rate of Rubisco (V_cmax) and the chloroplast electron transport rate (J). Maximum electron transport rates were linearly related to cyt f content at any given temperature (e.g. 115 and 179 µmol electrons mol^?1 cyt f s^?1 at 25 and 40 °C, respectively). The chloroplast CO₂ concentration (C_trans) at which the transition from RuBP carboxylation to RuBP regeneration limitation occurred increased with leaf temperature and was independent of growth light intensity, consistent with the constant ratio of cyt f/Rubisco. In tobacco, CO₂ assimilation rate at 380 µmol mol^?1 CO₂ concentration and high light was limited by RuBP carboxylation above 32 °C and by RuBP regeneration below 32 °C.     

Effects of Rubisco kinetics and Rubisco activation state on the temperature dependence of the photosynthetic rate in spinach leaves from contrasting growth temperatures

Recently, several studies reported that the optimum temperature for the initial slope [IS(Ci)] of the light-saturated photosynthetic rate (A) versus intercellular CO2 concentration (Ci) curve changed, depending on the growth temperature. However, few studies compare IS(Ci) with ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) properties. Here, we assessed Rubisco activation state and in vitro Rubisco kinetics, the main determinants of IS(Ci), in spinach leaves grown at 30/25 [high temperature (HT)] and 15/10 degrees C [low temperature (LT)]. We measured Rubisco activation state and A at a CO2 concentration of 360 microL L(-1) (A360) at various temperatures. In both HT and LT leaves, the Rubisco activation state decreased with increasing temperatures above the optimum temperatures for A360, while the activation state remained high at lower temperatures. To compare Rubisco characteristics, temperature dependences of the maximum rate of ribulose 1,5-bisphosphate (RuBP) carboxylation (Vcmax), specificity factor (Sc/o) and thermal stability were examined. We also examined Vcmax, and thermal stability in the leaves that were transferred from HT to LT conditions and were subsequently kept under LT conditions for 2 weeks (HL). Rubisco purified from HT, LT and HL leaves are called HT, LT and HL Rubisco, respectively. Thermal stabilities of LT and HL Rubisco were similar and lower than that of HT Rubisco. Both Vcmax and Sc/o in LT Rubisco were higher than those of HT Rubisco at low temperatures, while these were lower at high temperatures. Vcmax in HL Rubisco were similar to those of LT Rubisco at low temperatures, and to those of HT Rubisco at high temperatures. The predicted photosynthetic rates, taking account of the Rubisco kinetics and the Rubisco activation state, agreed well with A360 in both HT and LT leaves. This study suggests that photosynthetic performance is largely determined by the Rubisco kinetics at low temperature and by Rubisco Kinetics and the Rubisco activation state at high temperature.     

Ribulose-1,5-bisphosphate regeneration limitation in rice leaf photosynthetic acclimation to elevated CO2

Our previous study has demonstrated that both RuBP carboxylation limitation and RuBP regeneration limitation exist simultaneously in rice grown under free-air CO₂ enrichment (FACE, about 200 μmol mol⁻¹ above the ambient air CO₂ concentration) conditions [G.-Y. Chen, Z.-H. Yong, Y. Liao, D.-Y. Zhang, Y. Chen, H.-B. Zhang, J. Chen, J.-G. Zhu, D.-Q. Xu, Photosynthetic acclimation in rice leaves to free-air CO₂ enrichment related to both ribulose-1,5-bisphosphate carboxylase limitation and ribulose-1,5-bisphosphate regeneration limitation. Plant Cell Physiol. 46 (2005) 1036–1045]. To explore the mechanism for forming of RuBP regeneration limitation, we conducted the gas exchange measurements and some biochemical analyses in FACE-treated and ambient rice plants. Net CO₂ assimilation rate (A_net) in FACE leaves was remarkably lower than that in ambient leaves when measured at the same CO₂ concentration, indicating that photosynthetic acclimation to elevated CO₂ occurred. In the meantime the maximum electron transport rate (ETR) (J_max), maximum carboxylation rate (V_cmax) in vivo, and RuBP contents decreased significantly in FACE leaves. The whole chain electron transport rate and photophosphorylation rate reduced significantly while ETR of photosystem II (PSII) did not significantly decrease and ETR of photosystem I (PSI) was significantly increased in the chloroplasts from FACE leaves. Further, the amount of cytochrome (Cyt) f protein, a key component localized between PSII and PSI, was remarkably declined in FACE leaves. It appears that during photosynthetic acclimation the decline in the Cyt f amount is an important cause for the decreased RuBP regeneration capacity by decreasing the whole chain electron transport in FACE leaves.     

The temperature acclimation of photosynthetic responses to CO2 in Zea mays and its relationship to the activities of photosynthetic enzymes and the CO2-concentrating mechanism of C4 photosynthesis

Abstract Associations between photosynthetic responses to CO₂ at rate-saturating light and photosynthetic enzyme activities were compared for leaves of maize grown under constant air temperatures of 19, 25 and 31°C. Key photosynthetic enzymes analysed were ribulose bisphosphatc (RuBP) carboxylase, phosphoenolpyruvate (PEP) carboxylase, NADP-malic enzyme and pyruvate, P_i dikinasc. Rates of CO₂-saturated photosynthesis were similar in leaves developed at 19°C and 25°C but were decreased significantly by growth at 31°C. In contrast, carboxylation efficiency differed significantly between all three temperature regimes. Carboxylation efficiency was greatest in leaves developed at 19°C and decreased with increasing temperature during growth. The changes of carboxylation efficiency were highly correlated with changes in the activity of pyruvate, P_i dikinase (r= 0.95), but not with other photosynthetic enzyme activities. The activities of these latter enzymes, including that of RuBP carboxylase, were relatively insensitive to temperature during growth. The sensitivity of quantum yield to O₂ concentration was lower in leaves grown at 19°C than in leaves grown at 31°C. These observations support the novel hypothesis that variation in the capacity for CO₂ delivery to the bundle sheath by the C₄ cycle, relative to the capacity for net assimilation by the C₂ cycle, can be a principal determinant of C₄ photosynthetic responses to CO₂.     

Electron transport is the functional limitation of photosynthesis in field-grown Pima cotton plants at high temperature

Restrictions to photosynthesis can limit plant growth at high temperature in a variety of ways. In addition to increasing photorespiration, moderately high temperatures (35–42 °C) can cause direct injury to the photosynthetic apparatus. Both carbon metabolism and thylakoid reactions have been suggested as the primary site of injury at these temperatures. In the present study this issue was addressed by first characterizing leaf temperature dynamics in Pima cotton (Gossypium barbadense) grown under irrigation in the US desert south‐west. It was found that cotton leaves repeatedly reached temperatures above 40 °C and could fluctuate as much as 8 or 10 °C in a matter of seconds. Laboratory studies revealed a maximum photosynthetic rate at 30–33 °C that declined by 22% at 45 °C. The majority of the inhibition persisted upon return to 30 °C. The mechanism of this limitation was assessed by measuring the response of photosynthesis to CO₂ in the laboratory. The first time a cotton leaf (grown at 30 °C) was exposed to 45 °C, photosynthetic electron transport was stimulated (at high CO₂) because of an increased flux through the photorespiratory pathway. However, upon cooling back to 30 °C, photosynthetic electron transport was inhibited and fell substantially below the level measured before the heat treatment. In the field, the response of assimilation (A) to various internal levels of CO₂ (C_i) revealed that photosynthesis was limited by ribulose‐1,5‐bisphosphate (RuBP) regeneration at normal levels of CO₂ (presumably because of limitations in thylakoid reactions needed to support RuBP regeneration). There was no evidence of a ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco) limitation at air levels of CO₂ and at no point on any of 30 A–C_i curves measured on leaves at temperatures from 28 to 39 °C was RuBP regeneration capacity measured to be in substantial excess of the capacity of Rubisco to use RuBP. It is therefore concluded that photosynthesis in field‐grown Pima cotton leaves is functionally limited by photosynthetic electron transport and RuBP regeneration capacity, not Rubisco activity.     

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.     

Photosynthetic performance at low temperature of Bouteloua gracilis Lag., a high-altitude C4 grass from the Rocky Mountains, USA

The mechanisms controlling the photosynthetic performance of C₄ plants at low temperature were investigated using ecotypes of Bouteloua gracilis Lag. from high (3000 m) and low (1500 m) elevation sites in the Rocky Mountains of Colorado. Plants were grown in controlled‐environment cabinets at a photon flux density of 700 μ mol m^?2 s^?1 and day/night temperatures of 26/16 °C or 14/7 °C. The thermal response of the net CO₂ assimilation rate (A) was evaluated using leaf gas‐exchange analysis and activity assays of ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco), phosphoenolpyruvate carboxylase (PEPCase) and pyruvate,orthophosphate dikinase (PPDK). In both ecotypes, a reduction in measurement temperature caused the CO₂‐saturated rate of photosynthesis to decline to a greater degree than the initial slope of A versus the intercellular CO₂ response, thereby reducing the photosynthetic CO₂ saturation point. As a consequence, A in normal air was CO₂‐saturated at sub‐optimal temperatures. Ecotypic variation was low when grown at 26/16 °C, with the major difference between the ecotypes being that the low‐elevation plants had higher A; however, the ecotypes responded differently when grown at cool temperature. At temperatures below the thermal optimum, A in high‐elevation plants grown at 14/7 °C was enhanced relative to plants grown at 26/16 °C, while A in low‐elevation plants grown at 14/7 °C was reduced compared to 26/16 °C‐grown plants. Photoinhibition at low growth temperature was minor in both ecotypes as indicated by small reductions in dark‐adapted F_v/F_m. In both ecotypes, the activity of Rubisco was equivalent to A below 17 °C but well in excess of A above 25 °C. Activities of PEPCase and PPDK responded to temperature in a similar proportion relative to Rubisco, and showed no evidence for dissociation that would cause them to become principal limitations at low temperature. Because of the similar temperature response of Rubisco and A, we propose that Rubisco is a major limitation on C₄ photosynthesis in B. gracilis below 17 °C. Based on these results and for theoretical reasons associated with how C₄ plants use Rubisco, we further suggest that Rubisco capacity may be a widespread limitation upon C₄ photosynthesis at low temperature.     

Specificity factor of Rubisco: estimation in intact leaves by carboxylation at different CO2/O2 ratios

The specificity factor of Rubisco (S _f) was estimated in intact leaves from the carboxylation of ribulose-1,5-bisphosphate (RuBP) at various CO₂/O₂ ratios. As oxygenation is calculated by the difference of the ¹⁴CO₂ uptake by RuBP in the absence and presence of oxygen, it is important to choose the optimum CO₂/O₂ ratios. At high CO₂ concentration (1,000 cm³ m^?3 and higher) oxygenation consumes less than 50% RuBP but the difference of concentrations of CO₂ at cell walls (C _w) and at the carboxylation centers (C _c) is 2?C5% and the influence of mesophyll resistance (r _md) is of minor importance. To accumulate large endogenous pool of RuBP, the leaves were preilluminated in the CO₂- and O₂-free gas environments for 8 to 10 s. Thereafter the light was switched off and the leaves were flushed with the gas containing different concentrations of ¹⁴CO₂ and O₂. The specificity factor of Rubisco was calculated from the amount of the tracer taken up under different ¹⁴CO₂/O₂ ratios by the exhaustion of the RuBP pool. Application of ¹⁴CO₂ allowed us to discriminate between the CO₂ uptake and the concurrent respiratory CO₂ release which proceeded at the expense of unlabelled intermediates.     

The effects of development at sub-optimal growth temperatures on photosynthetic capacity and susceptibility to chilling-dependent photoinhibition in Zea mays

When plants of Zea mays L. cv. LG11 that have been grown at optimal temperatures are transferred to chilling temperatures (0–12°C) photoinhibition of photosynthetic CO₂ assimilation can occur. This study examines how growth at sub-optimal temperatures alters both photosynthetic capacity and resistance to chilling-dependent photoinhibition. Plants of Z. mays cv. LG11 were grown in controlled environments at 14, 17, 20 and 25°C. As a measure of the capacity for photosynthesis under light limiting conditions, the maximum quantum yields of CO₂ assimilation (φ^a.c) and O₂ evolution (φ^a.o) were determined for the laminae of the second leaves at photon fluxes of 50–150 μmol m^-2s^-1. To determine photosynthetic capacity at photon fluxes approaching light saturation, rates of CO₂ uptake (A₁₅₀₀) and O₂ evolution (A₁₅₀₀) were determined in a photon flux of 1500 μmol m^-2s^-1. In leaves developed at 14°C, φ and φ were 26 and 43%, respectively, of the values for leaves grown at 25°C. Leaves grown at 17°C showed intermediate reductions in φ and φ, whilst leaves developed at 20°C showed no significant differences from those grown at 25°C. Similar patterns of decrease were observed for A₁₅₀₀ and A_1500.0 with decreasing growth temperature. Leaves developed at 25°C showed higher rates of CO₂ assimilation at all light levels and measurement temperatures in comparison to leaves developed at 17 and 14°C. A greater reduction in A₁₅₀₀ relative to A_1500.0 with decreasing growth temperature was attributed to increased stomatal limitation. Exposure of leaves to 800–1000 μmol m^-2 s^-1 when plant temperature was depressed to ca 6.5°C produced a photoinhibition of photosynthetic CO₂ assimilation in all leaves. However, in leaves developed at 17°C the decrease in A₁₅₀₀ following this chilling treatment was only 25% compared to 90% in leaves developed at 25°C. Recovery following chilling was completed earlier in leaves developed at 17°C. The results suggest that growth at sub-optimal temperatures induces increased tolerance to exposure to high light at chilling temperatures. This is offset by the large loss in photosynthetic capacity imposed by leaf development at sub-optimal temperatures.     

Elevated growth temperatures reduce the carbon gain of black spruce [Picea mariana (Mill.) B.S.P.]

We explored the effect of high‐growth temperatures on a dominant North American boreal tree, black spruce [Picea mariana (Mill.) B.S.P.]. In 2004 and 2005, we grew black spruce at either 22 °C/16 °C day/night temperatures [low temperature (LT)] or 30°/24 °C [high temperature (HT)] and determined how temperature affected growth, leaf morphology, photosynthesis, respiration and thermotolerance. HT spruce were 20% shorter, 58% lighter, and had a 58% lower root : shoot ratio than LT trees. Mortality was negligible in the LT treatment, but up to 14% of HT seedlings died by the end of the growing season. HT seedlings had a higher photosynthetic temperature optimum, but net photosynthesis at growth temperatures was 19–35% lower in HT than LT trees. HT seedlings had both a lower apparent maximum ribulose‐1,5‐bisphosphate carboxylation capacity (V_cmax) and a lower apparent maximum electron transport rate (J_max) than LT trees, indicating reduced allocation to photosynthetic components. Consistently, HT needles had 26% lower leaf nitrogen content than LT needles. At each measurement temperature, HT seedlings had 20–25% lower respiration rates than LT trees; however, this did not compensate for reduced photosynthetic rates at growth temperature, leading to a greater ratio of dark respiration to net carbon dioxide assimilation rate in HT trees. HT needles had 16% lower concentrations of soluble sugars than LT needles, but similar starch content. Growth at high temperatures increased the thermotolerance of black spruce. HT trees showed less PSII inhibition than LT seedlings and no increase in electrolyte leakage when briefly exposed to 40–57 °C. While trees that develop at high temperatures have enhanced tolerance for brief, extreme heat events, the reduction in root allocation indicates that seedlings will be more susceptible to episodic soil drying and less competitive for belowground resources in future climates of the boreal region.     

The relative impacts of daytime and night-time warming on photosynthetic capacity in Populus deltoides

In order to investigate the relative impacts of increases in day and night temperature on tree carbon relations, we measured night‐time respiration and daytime photosynthesis of leaves in canopies of 4‐m‐tall cottonwood (Populus deltoides Bartr. ex Marsh) trees experiencing three daytime temperatures (25, 28 or 31 °C) and either (i) a constant nocturnal temperature of 20 °C or (ii) increasing nocturnal temperatures (15, 20 or 25 °C). In the first (day warming only) experiment, rates of night‐time leaf dark respiration (R_dark) remained constant and leaves displayed a modest increase (11%) in light‐saturated photosynthetic capacity (A_max) during the day (1000–1300 h) over the 6 °C range. In the second (dual night and day warming) experiment, R_dark increased by 77% when nocturnal temperatures were increased from 15 °C (0·36 µmol m^?2 s^?1) to 25 °C (0·64 µmol m^?2 s^?1). A_max responded positively to the additional nocturnal warming, and increased by 38 and 64% in the 20/28 and 25/31 °C treatments, respectively, compared with the 15/25 °C treatment. These increases in photosynthetic capacity were associated with strong increases in the maximum carboxylation rate of rubisco (V_cmax) and ribulose‐1,5‐bisphosphate (RuBP) regeneration capacity mediated by maximum electron transport rate (J_max). Leaf soluble sugar and starch concentration, measured at sunrise, declined significantly as nocturnal temperature increased. The nocturnal temperature manipulation resulted in a significant inverse relationship between A_max and pre‐dawn leaf carbohydrate status. Independent measurements of the temperature response of photosynthesis indicated that the optimum temperature (T_opt) acclimated fully to the 6 °C range of temperature imposed in the daytime warming. Our findings are consistent with the hypothesis that elevated night‐time temperature increases photosynthetic capacity during the following light period through a respiratory‐driven reduction in leaf carbohydrate concentration. These responses indicate that predicted increases in night‐time minimum temperatures may have a significant influence on net plant carbon uptake.     

Photosynthetic responses to chilling in a chilling‐tolerant and chilling‐sensitive Miscanthus hybrid

Miscanthus is a C₄ perennial grass being developed for bioenergy production in temperate regions where chilling events are common. To evaluate chilling effects on Miscanthus, we assessed the processes controlling net CO₂ assimilation rate (A) in Miscanthus x giganteus (M161) and a chilling‐sensitive Miscanthus hybrid (M115) before and after a chilling treatment of 12/5 °C. The temperature response of A and maximum Rubisco activity in vitro were identical below 20 °C in chilled and unchilled M161, demonstrating Rubisco capacity limits or co‐limits A at cooler temperatures. By contrast, A in M115 decreased at all measurement temperatures after growth at 12/5 °C. Rubisco activity in vitro declined in proportion to the reduction in A in chilled M115 plants, indicating Rubisco capacity is responsible in part for the decline in A. Pyruvate orthophosphate dikinase activities were also reduced by the chilling treatment when assayed at 28 °C, indicating this enzyme may also contribute to the reduction in A in M115. The maximum extractable activities of PEPCase and NADP‐ME remained largely unchanged after chilling. The carboxylation efficiency of the C₄ cycle was depressed in both genotypes to a similar extent after chilling. Φ_P:Φ_CO2 remained unchanged in both genotypes indicating the C₃ and C₄ cycles decline equivalently upon chilling.     

Strong thermal acclimation of photosynthesis in tropical and temperate wet‐forest tree species: the importance of altered Rubisco content

Understanding of the extent of acclimation of light‐saturated net photosynthesis (A_n) to temperature (T), and associated underlying mechanisms, remains limited. This is a key knowledge gap given the importance of thermal acclimation for plant functioning, both under current and future higher temperatures, limiting the accuracy and realism of Earth system model (ESM) predictions. Given this, we analysed and modelled T‐dependent changes in photosynthetic capacity in 10 wet‐forest tree species: six from temperate forests and four from tropical forests. Temperate and tropical species were each acclimated to three daytime growth temperatures (T_growth): temperate – 15, 20 and 25 °C; tropical – 25, 30 and 35 °C. CO₂ response curves of A_n were used to model maximal rates of RuBP (ribulose‐1,5‐bisphosphate) carboxylation (V_cmax) and electron transport (J_max) at each treatment's respective T_growth and at a common measurement T (25 °C). SDS‐PAGE gels were used to determine abundance of the CO₂‐fixing enzyme, Rubisco. Leaf chlorophyll, nitrogen (N) and mass per unit leaf area (LMA) were also determined. For all species and T_growth, A_n at current atmospheric CO₂ partial pressure was Rubisco‐limited. Across all species, LMA decreased with increasing T_growth. Similarly, area‐based rates of V_cmax at a measurement T of 25 °C (V_cmax²⁵) linearly declined with increasing T_growth, linked to a concomitant decline in total leaf protein per unit leaf area and Rubisco as a percentage of leaf N. The decline in Rubisco constrained V_cmax and A_n for leaves developed at higher T_growth and resulted in poor predictions of photosynthesis by currently widely used models that do not account for T_growth‐mediated changes in Rubisco abundance that underpin the thermal acclimation response of photosynthesis in wet‐forest tree species. A new model is proposed that accounts for the effect of T_growth‐mediated declines in V_cmax²⁵ on A_n, complementing current photosynthetic thermal acclimation models that do not account for T sensitivity of V_cmax²⁵.     

In vivo temperature response functions of parameters required to model RuBP-limited photosynthesis

The leaf model of C₃ photosynthesis of Farquhar, von Caemmerer & Berry (Planta 149, 78–90, 1980) provides the basis for scaling carbon exchange from leaf to canopy and Earth‐System models, and is widely used to project biosphere responses to global change. This scaling requires using the leaf model over a wider temperature range than that for which the model was originally parameterized. The leaf model assumes that photosynthetic CO₂ uptake within a leaf is either limited by the rate of ribulose‐1,5‐bisphosphate (RuBP) regeneration or the activity of RuBP carboxylase‐oxygenase (Rubisco). Previously we reported a re‐parameterization of the temperature responses of Rubisco activity that proved robust when applied to a range of species. Herein this is extended to re‐parameterizing the response of RuBP‐limited photosynthesis to temperature. RuBP‐limited photosynthesis is assumed to depend on the whole chain electron transport rate, which is described as a three‐parameter non‐rectangular hyperbolic function of photon flux. Herein these three parameters are determined from simultaneous measurement of chlorophyll fluorescence and CO₂ exchange of tobacco leaves, at temperatures from 10 to 40 °C. All varied significantly with temperature and were modified further with variation in growth temperature from 15 to 35 °C. These parameters closely predicted the response of RuBP‐limited photosynthesis to temperature measured in both lemon and poplar and showed a significant improvement over predictions based on earlier parameterizations. We provide the necessary equations for use of the model of Farquhar et al. (1980) with our newly derived temperature functions for predicting both Rubisco‐ and RuBP‐limited photosynthesis.     

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.     

Acclimation of photosynthesis and respiration to elevated atmospheric CO2 in two Scrub Oaks

Abstract For two species of oak, we determined whether increasing atmospheric CO₂ concentration (C_a) would decrease leaf mitochondrial respiration (R) directly, or indirectly owing to their growth in elevated C_a, or both. In particular, we tested whether acclimatory decreases in leaf‐Rubisco content in elevated C_a would decrease R associated with its maintenance. This hypothesis was tested in summer 2000 on sun and shade leaves of Quercus myrtifolia Willd. and Quercus geminata Small. We also measured R on five occasions between summer 1999 and 2000 on leaves of Q. myrtifolia. The oaks were grown in the field for 4 years, in either current ambient or elevated (current ambient + 350 µmol mol^?1) C_a, in open‐top chambers (OTCs). For Q. myrtifolia, an increase in C_a from 360 to 710 µmol mol^?1 had no direct effect on R at any time during the year. In April 1999, R in young Q. myrtifolia leaves was significantly higher in elevated C_a—the only evidence for an indirect effect of growth in elevated C_a. Leaf R was significantly correlated with leaf nitrogen (N) concentration for the sun and shade leaves of both the species of oak. Acclimation of photosynthesis in elevated C_a significantly reduced maximum RuBP‐saturated carboxylation capacity (V_{c max}) for both the sun and shade leaves of only Q. geminata. However, we estimated that only 11–12% of total leaf N was invested in Rubisco; consequently, acclimation in this plant resulted in a small effect on N and an insignificant effect on R. In this study measurements of respiration and photosynthesis were made on material removed from the field; this procedure had no effect on gas exchange properties. The findings of this study were applicable to R expressed either per unit leaf area or unit dry weight, and did not support the hypothesis that elevated C_a decreases R directly, or indirectly owing to acclimatory decreases in Rubisco content.     

The relationship between CO2 assimilation, photosynthetic electron transport and water–water cycle in chill-exposed cucumber leaves under low light and subsequent recovery

The effects of chilling under low light (9/7 °C, 100 µmol m^?2 s^?1) on the photosynthetic and antioxidant capacities and subsequent recovery were examined in two (one tolerant and one sensitive) cucumber genotypes. Chilling resulted in an irreversible inhibition of net CO₂ assimilation and growth for the sensitive genotype, which was accompanied by decreases in the maximum velocity of RuBP carboxylation by Rubisco (V_cmax), the capacity for ribulose‐1,5‐bisphosphate regeneration (J_max), Rubisco content and activity, and the quantum efficiency of photosystem II, in the absence of any stomatal limitation of CO₂ supply or inorganic phosphate limitation. In contrast, CO₂ assimilation for the tolerant genotype fully recovered after chill. The chill‐induced decrease in the proportion of electron flux for photosynthetic carbon reduction was mostly compensated by an O₂‐dependent alternative electron flux driven by the water–water cycle, especially in the sensitive genotype. Compared with the tolerant genotype, the sensitive genotype after chill showed reduced capacity for scavenging reactive oxygen species and increased accumulation of reactive oxygen species. The balance between O₂‐dependent alternative electron flux and the capacity for scavenging reactive oxygen species in response to chill plays a major role in determining the tolerance of cucumber leaves to this stress factor. It is concluded that the water–water cycle operates at high rates when CO₂ assimilation is restricted in cucumber leaves subjected to chill and low light conditions.     

Effects of nitrogen supply and high temperature on the growth and physiology of the chickpea

Chickpeas were grown with or without nitrate nitrogen feeding, or nodulated with Rhizobium leguminosarum. High [40°C day, 25°C night (HT)] and moderate [25°C day, 177°C night (LT)] temperature regimes were employed during growth. Growth rates, photosynthetic capacity and enzymes of carbon and nitrogen metabolism were monitored to assess the acclimatory capacity of the chickpea. Initial growth rates were stimulated by high temperatures, particularly in nitrate-fed and nodulated plants. Older HT plants had fewer laterals, smaller leaves, and fewer flowers were produced than in LT plants. There was some indication of an acclimation of photosynthesis to high temperatures and this was independent of nitrogen supply. Rubisco activity was increased by high growth temperatures. However, HT plants also had higher transpiration rates and lower water use efficiency than LT plants both in respective growth conditions and when compared in a common condition. High temperatures reduced shoot nitrate reductase activity but had little effect on root activity, which was the same if not greater than activity in LT roots. The amino acid, asparagine, was found at high concentrations in all treatments. Concentrations were maintained throughout growth in HT plants but declined with age in LT plants.     

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.     

Effect of heat stress on the inhibition and recovery of the ribulose-1,5-bisphosphate carboxylase/oxygenase activation state

Experiments were conducted to determine the relative contributions of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco; EC 4.1.1.39) activation state vis-à-vis Rubisco activase and metabolite levels to the inhibition of cotton (Gossypium hirsutum L.) photosynthesis by heat stress. Exposure of leaf tissue in the light to temperatures of 40 or 45 °C decreased the activation state of Rubisco to levels that were 65 or 10%, respectively, of the 28 °C control. Ribulose-1,5-bisphosphate (RuBP) levels increased in heat-stressed leaves, whereas the 3-phosphoglyceric acid pool was depleted. Heat stress did not affect Rubisco per se, as full activity could be restored by incubation with CO₂ and Mg²⁺. Inhibition and recovery of Rubisco activation state and carbon dioxide exchange rate (CER) were closely related under moderate heat stress (up to 42.5 °C). Moderate heat stress had negligible effect on Fv/Fm, the maximal quantum yield of photosystem II. In contrast, severe heat stress (45 °C) caused significant and irreversible damage to Rubisco activation, CER, and Fv/Fm. The rate of Rubisco activation after alleviating moderate heat stress was comparable to that of controls, indicating rapid reversibility of the process. However, moderate heat stress decreased both the rate and final extent of CER activation during dark-to-light transition. Treatment of cotton leaves with methyl viologen or an oxygen-enriched atmosphere reduced the effect of heat stress on Rubisco inactivation. Both treatments also reduced tissue RuBP levels, indicating that the amount of RuBP present during heat stress may influence the degree of Rubisco inactivation. Under both photorespiratory and non-photorespiratory conditions, the inhibition of the CER during heat stress could be completely reversed by increasing the internal partial pressure of CO₂ (Ci). However, the inhibition of the CER by nigericin, a K⁺ ionophore, was not reversible when the Ci was increased at ambient or high temperature. Our results indicate that inhibition of photosynthesis by moderate heat stress is not caused by inhibition of the capacity for RuBP regeneration. We conclude that heat stress inhibits Rubisco activation via a rapid and direct effect on Rubisco activase, possibly by perturbing Rubisco activase subunit interactions with each other or with Rubisco. Received: 25 February 2000 / Accepted: 13 May 2000