The response of the high altitude C(4) grass Muhlenbergia montana (Nutt.) A.S. Hitchc. to long- and short-term chilling.

The acclimation of C(4) photosynthesis to low temperature was studied in the montane grass Muhlenbergia montana in order to evaluate inherent limitations in the C(4) photosynthetic pathway following chilling. Plants were grown in growth cabinets at 26 degrees C days, but at night temperatures of either 16 degrees C (the control treatment), 4 degrees C for at least 28 nights (the cold-acclimated treatment), or 1 night (the cold-stress treatment). Below a measurement temperature of 25 degrees C, little difference in the thermal response of the net CO(2) assimilation rate (A) was observed between the control and cold-acclimated treatment. By contrast, above 30 degrees C, A in the cold-acclimated treatment was 10% greater than in the control treatment. The temperature responses of Rubisco activity and net CO(2) assimilation rate were similar below 22 degrees C, indicating high metabolic control of Rubisco over the rate of photosynthesis at cool temperatures. Analysis of the response of A to intercellular CO(2) level further supported a major limiting role for Rubisco below 20 degrees C. As temperature declined, the CO(2) saturated plateau of A exhibited large reductions, while the initial slope of the CO(2) response was little affected. This type of response is consistent with a Rubisco limitation, rather than limitations in PEP carboxylase capacity. Stomatal limitations at low temperature were not apparent because photosynthesis was CO(2) saturated below 23 degrees C at air levels of CO(2). In contrast to the response of photosynthesis to temperature and CO(2) in plants acclimated for 4 weeks to low night temperature, plants exposed to 4 degrees C for one night showed substantial reduction in photosynthetic capacity at temperatures above 20 degrees C. Because these reductions were at both high and low CO(2), enzymes associated with the C(4) carbon cycle were implicated as the major mechanisms for the chilling inhibition. These results demonstrate that C(4) plants from climates with low temperature during the growing season can fully acclimate to cold stress given sufficient time. This acclimation appears to involve reversal of injury to the C(4) cycle following initial exposure to low temperature. By contrast, carbon gain at low temperatures generally appears to be constrained by the carboxylation capacity of Rubisco, regardless of acclimation time. The inability to overcome the Rubisco limitation at low temperature may be an inherent limitation restricting C(4) photosynthetic performance in cooler climates.     

Future CO2 concentrations, though not warmer temperatures, enhance wheat photosynthesis temperature responses

The temperature dependence of C3 photosynthesis is known to vary according to the growth environment. Atmospheric CO2 concentration and temperature are predicted to increase with climate change. To test whether long-term growth in elevated CO2 and temperature modifies photosynthesis temperature response, wheat (Triticum aestivum L.) was grown in ambient CO2 (370 micromol mol(-1)) and elevated CO2 (700 micromol mol(-1)) combined with ambient temperatures and 4 degrees C warmer ones, using temperature gradient chambers in the field. Flag leaf photosynthesis was measured at temperatures ranging from 20 to 35 degrees C and varying CO2 concentrations between ear emergence and anthesis. The maximum rate of carboxylation was determined in vitro in the first year of the experiment and from the photosynthesis-intercellular CO2 response in the second year. With measurement CO2 concentrations of 330 micromol mol(-1) or lower, growth temperature had no effect on flag leaf photosynthesis in plants grown in ambient CO2, while it increased photosynthesis in elevated growth CO2. However, warmer growth temperatures did not modify the response of photosynthesis to measurement temperatures from 20 to 35 degrees C. A central finding of this study was that the increase with temperature in photosynthesis and the photosynthesis temperature optimum were significantly higher in plants grown in elevated rather than ambient CO2. In association with this, growth in elevated CO2 increased the temperature response (activation energy) of the maximum rate of carboxylation. The results provide field evidence that growth under CO2 enrichment enhances the response of Rubisco activity to temperature in wheat.     

Temperature response of photosynthesis in transgenic rice transformed with 'sense' or 'antisense' rbcS

The responses of chlorophyll fluorescence, gas exchange rate and Rubisco activation state to temperature were examined in transgenic rice plants with 130 and 35% of the wild-type (WT) Rubisco content by transformation with rbcS cDNA in sense and antisense orientations, respectively. Although the optimal temperatures of PSII quantum efficiency and CO(2) assimilation were found to be between 25 and 32 degrees C, the maximal activation state of Rubisco was found to be between 16 and 20 degrees C in all genotypes. The Rubisco flux control coefficient was also the highest between 16 and 20 degrees C in the WT and antisense lines [>0.88 at an intercellular CO(2) pressure (Ci) of 28 Pa]. Gross photosynthesis at Ci = 28 Pa per Rubisco content in the WT between 12 and 20 degrees C was close to that of the antisense lines where high Rubisco control is present. Thus, Rubisco activity most strongly limited photosynthesis at cool temperatures. These results indicated that a selective enhancement of Rubisco content can enhance photosynthesis at cool temperatures, but in the sense line with enhanced Rubisco content Pi regeneration limitation occurred. Above 20 degrees C, the Rubisco flux control coefficient declined. This decline was associated with a decline in Rubisco activation. The activation state of Rubisco measured at each temperature decreased with increasing Rubisco content, and the slope of activation to Rubisco content was independent of temperature. We discuss the possibility that the decline in Rubisco activation at intermediate and high temperatures is part of a regulated response to a limitation in other photosynthetic processes.     

Sensitivity of photosynthesis in a C4 plant,maize, to heat stress

Our objective was to determine the sensitivity of components of the photosynthetic apparatus of maize (Zea mays), a C4 plant, to high temperature stress. Net photosynthesis (Pn) was inhibited at leaf temperatures above 38 degrees C, and the inhibition was much more severe when the temperature was increased rapidly rather than gradually. Transpiration rate increased progressively with leaf temperature, indicating that inhibition was not associated with stomatal closure. Nonphotochemical fluorescence quenching (qN) increased at leaf temperatures above 30 degrees C, indicating increased thylakoid energization even at temperatures that did not inhibit Pn. Compared with CO(2) assimilation, the maximum quantum yield of photosystem II (F(v)/F(m)) was relatively insensitive to leaf temperatures up to 45 degrees C. The activation state of phosphoenolpyruvate carboxylase decreased marginally at leaf temperatures above 40 degrees C, and the activity of pyruvate phosphate dikinase was insensitive to temperature up to 45 degrees C. The activation state of Rubisco decreased at temperatures exceeding 32.5 degrees C, with nearly complete inactivation at 45 degrees C. Levels of 3-phosphoglyceric acid and ribulose-1,5-bisphosphate decreased and increased, respectively, as leaf temperature increased, consistent with the decrease in Rubisco activation. When leaf temperature was increased gradually, Rubisco activation acclimated in a similar manner as Pn, and acclimation was associated with the expression of a new activase polypeptide. Rates of Pn calculated solely from the kinetics of Rubisco were remarkably similar to measured rates if the calculation included adjustment for temperature effects on Rubisco activation. We conclude that inactivation of Rubisco was the primary constraint on the rate of Pn of maize leaves as leaf temperature increased above 30 degrees C.     

The effects of Rubisco activase on C4 photosynthesis and metabolism at high temperature

The activation of Rubisco in vivo requires the presence of the regulatory protein Rubisco activase. This enzyme facilitates the release of sugar phosphate inhibitors from Rubisco catalytic sites thereby influencing carbamylation. T(1) progeny of transgenic Flaveria bidentis (a C(4) dicot) containing genetically reduced levels of Rubisco activase were used to explore the role of the enzyme in C(4) photosynthesis at high temperature. A range of T(1) progeny was screened at 25 degrees C and 40 degrees C for Rubisco activase content, photosynthetic rate, Rubisco carbamylation, and photosynthetic metabolite pools. The small isoform of F. bidentis activase was expressed and purified from E. coli and used to quantify leaf activase content. In wild-type F. bidentis, the activase monomer content was 10.6+/-0.8 micromol m(-2) (447+/-36 mg m(-2)) compared to a Rubisco site content of 14.2+/-0.8 micromol m(-2). CO(2) assimilation rates and Rubisco carbamylation declined at both 25 degrees C and 40 degrees C when the Rubisco activase content dropped below 3 mumol m(-2) (125 mg m(-2)), with the status of Rubisco carbamylation at an activase content greater than this threshold value being 44+/-5% at 40 degrees C compared to 81+/-2% at 25 degrees C. When the CO(2) assimilation rate was reduced, ribulose-1,5-bisphosphate and aspartate pools increased whereas 3-phosphoglycerate and phosphoenol pyruvate levels decreased, demonstrating an interconnectivity of the C(3) and C(4) metabolites pools. It is concluded that during short-term treatment at 40 degrees C, Rubisco activase content is not the only factor modulating Rubisco carbamylation during C(4) photosynthesis.     

C4 photosynthesis at low temperature. A study using transgenic plants with reduced amounts of Rubisco

C(4) plants are rare in the cool climates characteristic of high latitudes and elevations, but the reasons for this are unclear. We tested the hypothesis that CO(2) fixation by Rubisco is the rate-limiting step during C(4) photosynthesis at cool temperatures. We measured photosynthesis and chlorophyll fluorescence from 6 degrees C to 40 degrees C, and in vitro Rubisco and phosphoenolpyruvate carboxylase activity from 0 degrees C to 42 degrees C, in Flaveria bidentis modified by an antisense construct (targeted to the nuclear-encoded small subunit of Rubisco, anti-RbcS) to have 49% and 32% of the wild-type Rubisco content. Photosynthesis was reduced at all temperatures in the anti-Rbcs plants, but the thermal optimum for photosynthesis (35 degrees C) did not differ. The in vitro turnover rate (kcat) of fully carbamylated Rubisco was 3.8 mol mol(-)(1) s(-)(1) at 24 degrees C, regardless of genotype. The in vitro kcat (Rubisco Vcmax per catalytic site) and in vivo kcat (gross photosynthesis per Rubisco catalytic site) were the same below 20 degrees C, but at warmer temperatures, the in vitro capacity of the enzyme exceeded the realized rate of photosynthesis. The quantum requirement of CO(2) assimilation increased below 25 degrees C in all genotypes, suggesting greater leakage of CO(2) from the bundle sheath. The Rubisco flux control coefficient was 0.68 at the thermal optimum and increased to 0.99 at 6 degrees C. Our results thus demonstrate that Rubisco capacity is a principle control over the rate of C(4) photosynthesis at low temperatures. On the basis of these results, we propose that the lack of C(4) success in cool climates reflects a constraint imposed by having less Rubisco than their C(3) competitors.     

Variation in the k(cat) of Rubisco in C(3) and C(4) plants and some implications for photosynthetic performance at high and low temperature

The capacity of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) to consume RuBP is a major limitation on the rate of net CO(2) assimilation (A) in C(3) and C(4) plants. The pattern of Rubisco limitation differs between the two photosynthetic types, as shown by comparisons of temperature and CO(2) responses of A and Rubisco activity from C(3) and C(4) species. In C(3) species, Rubisco capacity is the primary limitation on A at light saturation and CO(2) concentrations below the current atmospheric value of 37 Pa, particularly near the temperature optimum. Below 20 degrees C, C(3) photosynthesis at 37 and 68 Pa is often limited by the capacity to regenerate phosphate for photophosphorylation. In C(4) plants, the Rubisco capacity is equivalent to A below 18 degrees C, but exceeds the photosynthetic capacity above 25 degrees C, indicating that Rubisco is an important limitation at cool but not warm temperatures. A comparison of the catalytic efficiency of Rubisco (k(cat) in mol CO(2) mol(-1) Rubisco active sites s(-1)) from 17 C(3) and C(4) plants showed that Rubisco from C(4) species, and C(3) species originating in cool environments, had higher k(cat) than Rubisco from C(3) species originating in warm environments. This indicates that Rubisco evolved to improve performance in the environment that plants normally experience. In C(4) plants, and C(3) species from cool environments, Rubisco often operates near CO(2) saturation, so that increases in k(cat) would enhance A. In warm-habitat C(4) species, Rubisco often operates at CO(2) concentrations below the K(m) for CO(2). Because k(cat) and K(m) vary proportionally, the low k(cat) indicates that Rubisco has been modified in a manner that reduces K(m) and thus increases the affinity for CO(2) in C(3) species from warm climates.     

田间冬小麦叶片光合午休过程中的非气孔限制

1　引　　言作物的光合午休是近几年作物栽培生理研究的热点之一 ,研究大都从生态、生理的各个方面阐述光合午休的成因与机理 ,如土壤水分降低 ,ＣＯ2 浓度下降 ,ＶＰＤ过高 ,气孔限制与非气孔限制 ,羧化效率降低等[4 ,6,10 ] .其中有关作物光合午休过程中的非气孔限制问题 ,多数研究仅根据Ｆａｒｑｕｈａｒ判据[2 ] 判断发生了非气孔限制即止 ,至于非气孔限制的具体内容 ,多见于讨论 ,缺乏直接的实验证据 .本文对大田生态条件下 ,冬小麦生育后期旗叶光合午休过程中的非气孔限制问题进行了较深入的研究 .2　材料与方法2 1　供试材料选用冬…     

Development of C4 photosynthesis in sorghum leaves grown under free-air CO2 enrichment (FACE)

The developmental pattern of C4 expression has been well characterized in maize and other C4 plants. However, few reports have explored the possibility that the development of this pathway may be sensitive to changes in atmospheric CO2 concentrations. Therefore, both the structural and biochemical development of leaf tissue in the fifth leaf of Sorghum bicolor plants grown at elevated CO2 have been characterized. Ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) and phosphoenolpyruvate carboxylase (PEPC) activities accumulate rapidly as the leaf tissue differentiates and emerges from the surrounding whorl. Rubisco was not expressed in a cell-specific manner in the youngest tissue at the base of the leaf, but did accumulate before PEPC was detected. This suggests that the youngest leaf tissue utilizes a C3-like pathway for carbon fixation. However, this tissue was in a region of the leaf receiving very low light and so significant rates of photosynthesis were not likely. Older leaf tissue that had emerged from the surrounding whorl into full sunlight showed the normal C4 syndrome. Elevated CO2 had no effect on the cell-specific localization of Rubisco or PEPC at any stage of leaf development, and the relative ratios of Rubisco to PEPC remained constant during leaf development. However, in the oldest tissue at the tip of the leaf, the total activities of Rubisco and PEPC were decreased under elevated CO2 implying that C4 photosynthetic tissue may acclimate to growth under elevated CO2.     

Raising yield potential of wheat. II. Increasing photosynthetic capacity and efficiency

Past increases in yield potential of wheat have largely resulted from improvements in harvest index rather than increased biomass. Further large increases in harvest index are unlikely, but an opportunity exists for increasing productive biomass and harvestable grain. Photosynthetic capacity and efficiency are bottlenecks to raising productivity and there is strong evidence that increasing photosynthesis will increase crop yields provided that other constraints do not become limiting. Even small increases in the rate of net photosynthesis can translate into large increases in biomass and hence yield, since carbon assimilation is integrated over the entire growing season and crop canopy. This review discusses the strategies to increase photosynthesis that are being proposed by the wheat yield consortium in order to increase wheat yields. These include: selection for photosynthetic capacity and efficiency, increasing ear photosynthesis, optimizing canopy photosynthesis, introducing chloroplast CO(2) pumps, increasing RuBP regeneration, improving the thermal stability of Rubisco activase, and replacing wheat Rubisco with that from other species with different kinetic properties.     

Temperature response of mesophyll conductance. Implications for the determination of Rubisco enzyme kinetics and for limitations to photosynthesis in vivo

CO(2) transfer conductance from the intercellular airspaces of the leaf into the chloroplast, defined as mesophyll conductance (g(m)), is finite. Therefore, it will limit photosynthesis when CO(2) is not saturating, as in C3 leaves in the present atmosphere. Little is known about the processes that determine the magnitude of g(m). The process dominating g(m) is uncertain, though carbonic anhydrase, aquaporins, and the diffusivity of CO(2) in water have all been suggested. The response of g(m) to temperature (10 degrees C-40 degrees C) in mature leaves of tobacco (Nicotiana tabacum L. cv W38) was determined using measurements of leaf carbon dioxide and water vapor exchange, coupled with modulated chlorophyll fluorescence. These measurements revealed a temperature coefficient (Q(10)) of approximately 2.2 for g(m), suggesting control by a protein-facilitated process because the Q(10) for diffusion of CO(2) in water is about 1.25. Further, g(m) values are maximal at 35 degrees C to 37.5 degrees C, again suggesting a protein-facilitated process, but with a lower energy of deactivation than Rubisco. Using the temperature response of g(m) to calculate CO(2) at Rubisco, the kinetic parameters of Rubisco were calculated in vivo from 10 degrees C to 40 degrees C. Using these parameters, we determined the limitation imposed on photosynthesis by g(m). Despite an exponential rise with temperature, g(m) does not keep pace with increased capacity for CO(2) uptake at the site of Rubisco. The fraction of the total limitations to CO(2) uptake within the leaf attributable to g(m) rose from 0.10 at 10 degrees C to 0.22 at 40 degrees C. This shows that transfer of CO(2) from the intercellular air space to Rubisco is a very substantial limitation on photosynthesis, especially at high temperature.     

供氮和增温对倍增二氧化碳浓度下荫香叶片光合作用的影响

供给0～0.6 mg N的盆栽荫香(Cinnamomum burmannii)幼树分别生长在倍增CO ₂(+CO₂,731 μmol·mol^-1)和正常空气CO ₂浓度(CO ₂,365 μmol·mol^-1)的生长箱内,昼夜温度分别为25/23 ℃和32/25 ℃,自然光照下生长30 d.以生长在CO₂和25/23 ℃下的植株为对照研究增温和氮对+CO₂叶片光合作用的影响.结果表明,在+CO₂和25/23 ℃下无氮和氮处理植株的平均光合速率(Pnsat)较+CO₂和32/25 ℃下的叶片高5.1%,温度增高降低叶片Pnsat;而Pnsat随供氮而增高.在+CO₂条件下,生长在32/25 ℃下的叶片Rubisco最大羧化速率(Vcmax)和最大电子传递速率(Jmax)较25/23 ℃下的低(P＜0.05),温度增高降低+CO₂下叶片的Vcmax和Jmax在+CO2下叶片光合呼吸速率(Rp)较低,生长温度增高提升Rp.在CO₂下生长温度从25/23 ℃增至32/25 ℃,叶片的Rubisco含量(NR)和Rubisco活化中心浓度(M)降低,而供氮能增高NR和M.供氮能减缓温度增高对倍增CO₂下荫香叶片光合作用的限制.     

Effects of Growth Temperature on the Responses of Ribulose-1,5-Biphosphate Carboxylase,Electron Transport Components,and Sucrose Synthesis Enzymes to Leaf Nitrogen in Rice,and Their Relationships to Photosynthesis

Effects of growth temperature on the photosynthetic gas-exchange rates and their underlying biochemical properties were examined in young, fully expanded leaves of rice (Oryza sativa L.). The plants were grown hydroponically under day/night temperature regimes of 18/15[deg]C, 23/18[deg]C, and 30/23[deg]C and all photosynthetic measurements were made at a leaf temperature of 25[deg]C and an irradiance of 1800 [mu]mol quanta m-2 s-1. Growth temperature affected the photosynthetic CO2 response curve. The relative ratio of the initial slope to the CO2-saturated photosynthesis increased with rising growth temperature. This was caused mainly by an increase in CO2-limited photosynthesis for a given leaf nitrogen content with rising growth temperature. However, there was no difference in ribulose-1,5-bisphosphate carboxylase (Rubisco) content at any given leaf nitrogen content among temperature treatments. In addition, the activation state and catalytic turnover rate of Rubisco were not affected by growth temperature. The increase in CO2-limited photosynthesis with rising growth temperature was the result of an increase in the CO2 transfer conductance between the intercellular airspaces and the carboxylation sites. The amounts of total chlorophyll and light-harvesting chlorophyll a/b protein II increased for the same leaf nitrogen content with rising growth temperature, but the amounts of cytochrome f and coupling factor 1 and the activities of cytosolic fructose-1,6-bisphosphatase and sucrose-phosphate synthase were the same between plants grown at 23/18[deg]C and those grown at 30/23[deg]C. Similarly, CO2-saturated photosynthesis was not different for the same leaf nitrogen content between these treatments. For the 18/15[deg]C-grown plants, a slight decrease in the amounts of cytochrome f and coupling factor 1 and an increase in the activities of cytosolic fructose-1,6-bisphosphatase and sucrose-phosphate synthase were found, but these were not reflected in CO2-saturated photosynthesis.     

Rubisco, Rubisco activase, and global climate change

Global warming and the rise in atmospheric CO(2) will increase the operating temperature of leaves in coming decades, often well above the thermal optimum for photosynthesis. Presently, there is controversy over the limiting processes controlling photosynthesis at elevated temperature. Leading models propose that the reduction in photosynthesis at elevated temperature is a function of either declining capacity of electron transport to regenerate RuBP, or reductions in the capacity of Rubisco activase to maintain Rubisco in an active configuration. Identifying which of these processes is the principal limitation at elevated temperature is complicated because each may be regulated in response to a limitation in the other. Biochemical and gas exchange assessments can disentangle these photosynthetic limitations; however, comprehensive assessments are often difficult and, for many species, virtually impossible. It is proposed that measurement of the initial slope of the CO(2) response of photosynthesis (the A/C(i) response) can be a useful means to screen for Rubisco activase limitations. This is because a reduction in the Rubisco activation state should be most apparent at low CO(2) when Rubisco capacity is generally limiting. In sweet potato, spinach, and tobacco, the initial slope of the A/C(i) response shows no evidence of activase limitations at high temperature, as the slope can be accurately modelled using the kinetic parameters of fully activated Rubisco. In black spruce (Picea mariana), a reduction in the initial slope above 30 degrees C cannot be explained by the known kinetics of fully activated Rubisco, indicating that activase may be limiting at high temperatures. Because black spruce is the dominant species in the boreal forest of North America, Rubisco activase may be an unusually important factor determining the response of the boreal biome to climate change.     

Functional analysis of corn husk photosynthesis

The husk surrounding the ear of corn/maize (Zea mays) has widely spaced veins with a number of interveinal mesophyll (M) cells and has been described as operating a partial C(3) photosynthetic pathway, in contrast to its leaves, which use the C(4) photosynthetic pathway. Here, we characterized photosynthesis in maize husk and leaf by measuring combined gas exchange and carbon isotope discrimination, the oxygen dependence of the CO(2) compensation point, and photosynthetic enzyme activity and localization together with anatomy. The CO(2) assimilation rate in the husk was less than that in the leaves and did not saturate at high CO(2), indicating CO(2) diffusion limitations. However, maximal photosynthetic rates were similar between the leaf and husk when expressed on a chlorophyll basis. The CO(2) compensation points of the husk were high compared with the leaf but did not vary with oxygen concentration. This and the low carbon isotope discrimination measured concurrently with gas exchange in the husk and leaf suggested C(4)-like photosynthesis in the husk. However, both Rubisco activity and the ratio of phosphoenolpyruvate carboxylase to Rubisco activity were reduced in the husk. Immunolocalization studies showed that phosphoenolpyruvate carboxylase is specifically localized in the layer of M cells surrounding the bundle sheath cells, while Rubisco and glycine decarboxylase were enriched in bundle sheath cells but also present in M cells. We conclude that maize husk operates C(4) photosynthesis dispersed around the widely spaced veins (analogous to leaves) in a diffusion-limited manner due to low M surface area exposed to intercellular air space, with the functional role of Rubisco and glycine decarboxylase in distant M yet to be explained.     

The temperature response of C(3) and C(4) photosynthesis

We review the current understanding of the temperature responses of C(3) and C(4) photosynthesis across thermal ranges that do not harm the photosynthetic apparatus. In C(3) species, photosynthesis is classically considered to be limited by the capacities of ribulose 1.5-bisphosphate carboxylase/oxygenase (Rubisco), ribulose bisphosphate (RuBP) regeneration or P(i) regeneration. Using both theoretical and empirical evidence, we describe the temperature response of instantaneous net CO(2) assimilation rate (A) in terms of these limitations, and evaluate possible limitations on A at elevated temperatures arising from heat-induced lability of Rubisco activase. In C(3) plants, Rubisco capacity is the predominant limitation on A across a wide range of temperatures at low CO(2) (<300 microbar), while at elevated CO(2), the limitation shifts to P(i) regeneration capacity at suboptimal temperatures, and either electron transport capacity or Rubisco activase capacity at supraoptimal temperatures. In C(4) plants, Rubisco capacity limits A below 20 degrees C in chilling-tolerant species, but the control over A at elevated temperature remains uncertain. Acclimation of C(3) photosynthesis to suboptimal growth temperature is commonly associated with a disproportional enhancement of the P(i) regeneration capacity. Above the thermal optimum, acclimation of A to increasing growth temperature is associated with increased electron transport capacity and/or greater heat stability of Rubisco activase. In many C(4) species from warm habitats, acclimation to cooler growth conditions increases levels of Rubisco and C(4) cycle enzymes which then enhance A below the thermal optimum. By contrast, few C(4) species adapted to cooler habitats increase Rubisco content during acclimation to reduced growth temperature; as a result, A changes little at suboptimal temperatures. Global change is likely to cause a widespread shift in patterns of photosynthetic limitation in higher plants. Limitations in electron transport and Rubisco activase capacity should be more common in the warmer, high CO(2) conditions expected by the end of the century.     

Rubisco activase is a key regulator of non-steady-state photosynthesis at any leaf temperature and, to a lesser extent, of steady-state photosynthesis at high temperature

The role of Rubisco activase in steady-state and non-steady-state photosynthesis was analyzed in wild-type (Oryza sativa) and transgenic rice that expressed different amounts of Rubisco activase. Below 25°C, the Rubisco activation state and steady-state photosynthesis were only affected when Rubisco activase was reduced by more than 70%. However, at 40°C, smaller reductions in Rubisco activase content were linked to a reduced Rubisco activation state and steady-state photosynthesis. As a result, overexpression of maize Rubisco activase in rice did not lead to an increase of the Rubisco activation state, nor to an increase in photosynthetic rate below 25°C, but had a small stimulatory effect at 40°C. On the other hand, the rate at which photosynthesis approached the steady state following an increase in light intensity was rapid in Rubisco activase-overexpressing plants, intermediate in the wild-type, and slowest in antisense plants at any leaf temperature. In Rubisco activase-overexpressing plants, Rubisco activation state at low light was maintained at higher levels than in the wild-type. Thus, rapid regulation by Rubisco activase following an increase in light intensity and/or maintenance of a high Rubisco activation state at low light would result in a rapid increase in Rubisco activation state and photosynthetic rate following an increase in light intensity. It is concluded that Rubisco activase plays an important role in the regulation of non-steady-state photosynthesis at any leaf temperature and, to a lesser extent, of steady-state photosynthesis at high temperature.     

Interaction of drought and high temperature on photosynthesis and grain-filling of wheat

Drought and high temperature often occur simultaneously, but their effects on crops are usually investigated individually. Our objective was to compare effects of drought, high temperature, and their interactions on photosynthesis and grain-growth of wheat (Triticum aestivum L.). Plants (cv. Len) were grown uniformly in well-watered soil at 25/20 ± 2 °C day/night until anthesis, when they were subjected to regimes of no drought (soil at field capacity) and drought (plant water potential of –.0 to –2.4 MPa) at 15/10, 25/20, and 35/30 °C in controlled environments until physiological maturity. Drought decreased photosynthesis, stomatal conductance, viable leaf area, shoot and grain mass, and weight and soluble sugar content of kernels but increased plant water-use efficiency. High temperature hastened the decline in photosynthesis and leaf area, decreased shoot and grain mass as well as weight and sugar content of kernels, and reduced water-use efficiency. Interactions between the two stresses were pronounced, and consequences of drought on all physiological parameters were more severe at high temperature than low temperature. The synergistic interactions indicated that productivity of wheat is reduced considerably more by the combined stresses than by either stress alone, and that much of the effect is on photosynthetic processes.     

Limitations to photosynthesis of lettuce grown under tropical conditions: alleviation by root-zone cooling.

Aerial parts of lettuce plants were grown under natural tropical fluctuating ambient temperatures, but with their roots exposed to two different root-zone temperatures (RZTs): a constant 20 degrees C-RZT and a fluctuating ambient (A-) RZT from 23-40 degrees C. Plants grown at A-RZT showed lower photosynthetic CO2 assimilation (A), stomatal conductance (gs), midday leaf relative water content (RWC), and chlorophyll fluorescence ratio Fv/Fm than 20 degrees C-RZT plants on both sunny and cloudy days. Substantial midday depression of A and g(s) occurred on both sunny and cloudy days in both RZT treatments, although Fv/Fm did not vary diurnally on cloudy days. Reciprocal temperature transfer experiments investigated the occurrence and possible causes of stomatal and non-stomatal limitations of photosynthesis. For both temperature transfers, light-saturated stomatal conductance (gs sat) and photosynthetic CO2 assimilation (A(sat)) were highly correlated with each other and with midday RWC, suggesting that A was limited by water stress-mediated stomatal closure. However, prolonged growth at A-RZT reduced light- and CO2-saturated photosynthetic O2 evolution (Pmax), indicating non-stomatal limitation of photosynthesis. Tight temporal coupling of leaf nitrogen content and P(max) during both temperature transfers suggested that decreased nutrient status caused this non-stomatal limitation of photosynthesis.