Acclimation to temperature of the response of photosynthesis to increased carbon dioxide concentration in Taraxacum officinale

The relative stimulation of photosynthesis by elevated carbon dioxide in C₃ species normally increases strongly with increasing temperature. This results from the kinetic characteristics of Rubisco, and has potentially important implications for responses of vegetation to increasing atmospheric carbon dioxide. It is often assumed that because Rubisco characteristics are conservative, all C₃ species have the same temperature dependence of the response of photosynthesis to elevated carbon dioxide. However, in this field study of Taraxacum officinale, there were no significant differences in the relative stimulation of photosynthesis by elevated carbon dioxide among days with temperatures ranging from 15 to 34 °C. Nevertheless, short-term measurements indicated a strong temperature dependence of the stimulation. This suggested that acclimation to temperature caused the lack of variation in the seasonal data. Experiments in controlled environments indicated that complete acclimation of the relative stimulation of photosynthesis by elevated carbon dioxide occurred for growth temperatures of 10 – 25 °C. The apparent specificity of Rubisco for carbon dioxide relative to oxygen at 15 °C, as assayed in vivo by measurements of the carbon dioxide concentration at which carboxylation equalled oxygenation, also varied with growth temperature. Changes in the apparent specificity of Rubisco accounted for the acclimation of the temperature dependence of the relative stimulation of photosynthesis by elevated carbon dioxide. It is premature to conclude that low temperatures will necessarily reduce the relative stimulation of photosynthesis caused by rising atmospheric carbon dioxide. This revised version was published online in June 2006 with corrections to the Cover Date.     

Diurnal and Seasonal Changes in Photosynthesis and Photosystem 2 Photochemical Efficiency in Prosopis juliflora Leaves Subjected to Natural Environmental Stress

The plants of Prosopis juliflora growing in northern India are exposed to large variations of temperature, vapour pressure deficits (VPD), and photosynthetic photon flux density (PPFD) throughout the year. Under these conditions P. juliflora had two short periods of leaf production, one after the winter season and second after summer, which resulted in two distinct even aged cohorts of leaves. In winter with cold nights (2–8 °C) and moderate temperatures during the day, the plants showed high rates of photosynthesis. In summer the midday temperatures often reached <45 °C and plants showed severe inhibition of photosynthesis. The leaves of second cohort appeared in July and showed typical midday depression of photosynthesis. An analysis of diurnal partitioning of the absorbed excitation energy into photochemistry showed that a smaller fraction of the energy was utilised for photochemistry and a greater fraction was dissipated thermally, further the photon utilisation for photochemistry and thermal dissipation is largely affected by the interaction of irradiance and temperature. The plants showed high photochemical efficiency of photosystem 2 (PS2) at predawn and very little photoinhibition in all seasons except in summer. The photoinhibition in summer was pronounced with very poor recovery during night. Since P. juliflora exhibited distinct pattern of senescence and production of new leaves after winter and summer stress period, it appeared that the ontogenic characteristic together with its ability for safe dissipation of excess radiant energy in P. juliflora contributes to its growth and survival.     

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 elevated CO2 and temperature on photosynthesis and Rubisco in rice and soybean

Rice (Oryza sativa L. cv. IR-72) and soybean (Glycine max L. Merr. cv. Bragg), which have been reported to differ in acclimation to elevated CO₂, were grown for a season in sunlight at ambient and twice-ambient [CO₂], and under daytime temperature regimes ranging from 28 to 40°C. The objectives of the study were to test whether CO₂ enrichment could compensate for adverse effects of high growth temperatures on photosynthesis, and whether these two C₃ species differed in this regard. Leaf photosynthetic assimilation rates (A) of both species, when measured at the growth [CO₂], were increased by CO₂ enrichment, but decreased by supraoptimal temperatures. However, CO₂ enrichment more than compensated for the temperature-induced decline in A. For soybean, this CO₂ enhancement of A increased in a linear manner by 32–95% with increasing growth temperatures from 28 to 40°C, whereas with rice the degree of enhancement was relatively constant at about 60%, from 32 to 38°C. Both elevated CO₂ and temperature exerted coarse control on the Rubisco protein content, but the two species differed in the degree of responsiveness. CO₂ enrichment and high growth temperatures reduced the Rubisco content of rice by 22 and 23%, respectively, but only by 8 and 17% for soybean. The maximum degree of Rubisco down-regulation appeared to be limited, as in rice the substantial individual effects of these two variables, when combined, were less than additive. Fine control of Rubisco activation was also influenced by both elevated [CO₂] and temperature. In rice, total activity and activation were reduced, but in soybean only activation was lowered. The apparent catalytic turnover rate (K_cat) of rice Rubisco was unaffected by these variables, but in soybean elevated [CO₂] and temperature increased the apparent K_cat by 8 and 22%, respectively. Post-sunset declines in Rubisco activities were accelerated by elevated [CO₂] in rice, but by high temperature in soybean, suggesting that [CO₂] and growth temperature influenced the metabolism of 2-carboxyarabinitol-1-phosphate, and that the effects might be species-specific. The greater capacity of soybean for CO₂ enhancement of A at supraoptimal temperatures was probably not due to changes in stomatal conductance, but may be partially attributed to less down-regulation of Rubisco by elevated [CO₂] in soybean than in rice. However, unidentified species differences in the temperature optimum for photosynthesis also appeared to be important. The responses of photosynthesis and Rubisco in rice and soybean suggest that among C₃ plants species-specific differences will be encountered as a result of future increases in global [CO₂] and air temperatures.     

Photoprotective role of rhodoxanthin during cold acclimation in Cryptomeria japonica

To examine the role of rhodoxanthin in long‐term acclimation to low temperatures, we monitored seasonal changes in pigment composition, photosynthesis, chlorophyll fluorescence and the level of ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco) in needles of wild‐type and mutant forms of Cryptomeria japonica. In winter, rhodoxanthin accumulated in sun‐exposed needles of wild‐type plants, but not in those of the mutant. The level of chlorophyll decreased in both types of plant in winter. In contrast, the level of the xanthophyll cycle pool increased in both cases. The level of the pool in the mutant was twice that in the wild type in winter, on a Chl basis, even though the levels in both were similar in summer. The synthesis of rhodoxanthin might be triggered by photo‐inhibitory conditions, as suggested by the sustained elevated levels of zeaxanthin (Z) and antheraxanthin (A). In the wild type and the mutant, the quantum yield of CO₂ fixation (φ), the photosynthetic capacity, the photochemical efficiency of photosystem II (PSII), the photochemical quenching and the level of Rubisco in summer were similar. However, all these values for the wild type were higher than those for the mutant in winter. The non‐photochemical quenching (NPQ) in the mutant in winter increased rapidly even under low light conditions due to the high sustained levels of Z and A. In contrast, in the wild type, the conversion of Z via A to rhodoxanthin prevented the rapid increase in NPQ to maintain the relatively high level of φ. These findings suggest that rhodoxanthin might play an important photoprotective role in long‐term acclimation to cold. The dynamic regulation of the amount of rhodoxanthin relative to the level of the xanthophyll cycle pool might act to maintain an appropriate balance between light absorption, photosynthesis and the thermal dissipation of energy due to excess absorbed light in winter.     

The temperature response of CO2 assimilation, photochemical activities and Rubisco activation in Camelina sativa, a potential bioenergy crop with limited capacity for acclimation to heat stress

The temperature optimum of photosynthesis coincides with the average daytime temperature in a species’ native environment. Moderate heat stress occurs when temperatures exceed the optimum, inhibiting photosynthesis and decreasing productivity. In the present study, the temperature response of photosynthesis and the potential for heat acclimation was evaluated for Camelina sativa, a bioenergy crop. The temperature optimum of net CO₂ assimilation rate (A) under atmospheric conditions was 30–32?°C and was only slightly higher under non-photorespiratory conditions. The activation state of Rubisco was closely correlated with A at supra-optimal temperatures, exhibiting a parallel decrease with increasing leaf temperature. At both control and elevated temperatures, the modeled response of A to intercellular CO₂ concentration was consistent with Rubisco limiting A at ambient CO₂. Rubisco activation and photochemical activities were affected by moderate heat stress at lower temperatures in camelina than in the warm-adapted species cotton and tobacco. Growth under conditions that imposed a daily interval of moderate heat stress caused a 63?% reduction in camelina seed yield. Levels of cpn60 protein were elevated under the higher growth temperature, but acclimation of photosynthesis was minimal. Inactivation of Rubisco in camelina at temperatures above 35?°C was consistent with the temperature response of Rubisco activase activity and indicated that Rubisco activase was a prime target of inhibition by moderate heat stress in camelina. That photosynthesis exhibited no acclimation to moderate heat stress will likely impact the development of camelina and other cool season Brassicaceae as sources of bioenergy in a warmer world.     

Elevation provenance affects photosynthesis and its acclimation to temperature in the high-Andes alpine herb Phacelia secunda

• We analysed whether Phacelia secunda populations from different elevations exhibit intrinsic traits associated with diffusive and biochemical components of photosynthesis, and if they differ in acclimation of photosynthesis to warmer temperatures. We hypothesized that P. secunda will have similar photosynthetic performance regardless of altitudinal provenance and that plants from high elevations will have a lower photosynthetic acclimation capacity to higher temperature than plants from low elevations.
• Plants from 1600, 2800 and 3600 m a.s.l. in the central Chilean Andes were collected and grown under two temperature regimes (20/16 °C and 30/26 °C day/night). The following photosynthetic traits were measured in each plant for the two temperature regimes: A_N, g_s, g_m, J_max, V_cmax, Rubisco carboxylation k_cat^c.
• Under a common growth environment, plants from the highest elevation had slightly lower CO₂ assimilation rates compared to lower elevation plants. While diffusive components of photosynthesis increased with elevation provenance, the biochemical component decreased, suggesting compensation that explains the similar rates of photosynthesis among elevation provenances. Plants from high elevations had lower photosynthetic acclimation to warmer temperatures compared to plants from lower elevations, and these responses were related to elevational changes in diffusional and biochemical components of photosynthesis.
• Plants of P. secunda from different elevations maintain photosynthetic traits when grown in a common environment, suggesting low plasticity to respond to future climate changes. The fact that high elevation plants had lower photosynthetic acclimation to warmer temperature suggests higher susceptibility to increases in temperature associated with global warming.

     

Photosynthesis and water relations of well-watered orange plants as affected by winter and summer conditions

The aim of this study was to evaluate how the summer and winter conditions affect the photosynthesis and water relations of well-watered orange trees, considering the diurnal changes in leaf gas exchange, chlorophyll (Chl) fluorescence, and leaf water potential (Ψ) of potted-plants growing in a subtropical climate. The diurnal pattern of photosynthesis in young citrus trees was not significantly affected by the environmental changes when compared the summer and winter seasons. However, citrus plants showed higher photosynthetic performance in summer, when plants fixed 2.9 times more CO₂ during the diurnal period than in the winter season. Curiously, the winter conditions were more favorable to photosynthesis of citrus plants, when considering the air temperature (< 29 °C), leaf-to-air vapor pressure difference (< 2.4 kPa) and photon flux density (maximum values near light saturation) during the diurnal period. Therefore, low night temperature was the main environmental element changing the photosynthetic performance and water relations of well-watered plants during winter. Lower whole-plant hydraulic conductance, lower shoot hydration and lower stomatal conductance were noticed during winter when compared to the summer season. In winter, higher ratio between the apparent electron transport rate and leaf CO₂ assimilation was verified in afternoon, indicating reduction in electron use efficiency by photosynthesis. The high radiation loading in the summer season did not impair the citrus photochemistry, being photoprotective mechanisms active. Such mechanisms were related to increases in the heat dissipation of excessive light energy at the PSII level and to other metabolic processes consuming electrons, which impede the citrus photoinhibition under high light conditions.     

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     

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.     

The influence of temperature,sexual condition,and season on the metabolic rate of the lizardPsammodromus hispanicus

Summary Male and femalePsammodromus hispanicus from southern Europe were acclimated to four seasonal conditions of photoperiod and night time temperature. During the dark period, the lizards' body temperatures fell to ambient air temperature but during the light period the lizards were allowed to thermoregulate behaviourally and at such times the lizards' mean body temperature varied from 29.0°C to 32.6°C. The resting metabolic rate of these lizards was measured in 5°C steps from 5°C to 30°C or 35°C. Sexual condition had little effect on resting metabolic rate, but at low temperatures lizards acclimated to winter or spring seasonal conditions had lower resting metabolic rates than those acclimated to summer or autumn conditions. At temperatures above 20°C seasonal acclimation had no effect on resting metabolic rate. It is considered that the reduction in low temperature metabolic rate in spring and winter is induced by low night time temperatures and serves to conserve energy during those seasons when lizards must spend long periods at low temperature without being able to feed.     

Comparative effect of live food and manured treatments on water quality and production of ornamental carp, Cyprinus carpio var. koi L., during winter, summer, monsoon and post-monsoon growout experiments in concrete tanks

Four growth trials (each of 11 weeks duration) were conducted during different seasons throughout 1 year, namely, winter, summer, monsoon and post‐monsoon to assess the seasonal influence on growth and production of koi carp, Cyprinus carpio var. koi L. larvae in concrete tanks maintained under different management regimes. Individual weight gain, survival rate, fish deformities and number of marketable fish in each season were compared among four management regimes: (i) carp larvae fed with live zooplankton (LF); (ii) direct fertilization with poultry manure (PM); (iii) direct fertilization with cow dung (CD); and (iv) a control system (C), where a commercial diet containing 32% crude protein was applied. There were three replicates for each treatment. The LF treatment produced significantly higher weight gain, survival rate and number of marketable fish in all seasons (P < 0.05), compared with the other treatments, through maintenance of better water quality and greater abundance of zooplankton in the system. Fish deformities were highest in the C treatment in every season (P < 0.05). Water temperatures averaged 18.6°C (winter), 29.7°C (summer), 28.3°C (monsoon), and 26.5°C in the post‐monsoon trial. Average weight gain of koi carp achieved during the winter trial (2.01 g in C to 4.44 g in LF) was considerably lower than that achieved in the summer, monsoon or post‐monsoon trials. Survival rates of carp ranged from 70.5% (C) to 95.5% (LF) in the winter trial, and were considerably lower than the other seasons. The number of marketable fish was also lowest during the winter trial.     

An isoleucine residue acts as a thermal and regulatory switch in wheat Rubisco activase

The regulation of Rubisco, the gatekeeper of carbon fixation into the biosphere, by its molecular chaperone Rubisco activase (Rca) is essential for photosynthesis and plant growth. Using energy from ATP hydrolysis, Rca promotes the release of inhibitors and restores catalytic competence to Rubisco‐active sites. Rca is sensitive to moderate heat stress, however, and becomes progressively inhibited as the temperature increases above the optimum for photosynthesis. Here, we identify a single amino acid substitution (M159I) that fundamentally alters the thermal and regulatory properties of Rca in bread wheat (Triticum aestivum L.). Using site‐directed mutagenesis, we demonstrate that the M159I substitution extends the temperature optimum of the most abundant Rca isoform by 5°C in vitro, while maintaining the efficiency of Rubisco activation by Rca. The results suggest that this single amino acid substitution acts as a thermal and regulatory switch in wheat Rca that can be exploited to improve the climate resilience and efficiency of carbon assimilation of this cereal crop as temperatures become warmer and more volatile.     

Will rising CO2 protect plants from the midday sun? A study of photoinhibition of Quercus myrtifolia in a scrub‐oak community in two seasons

Over a large part of the photoperiod, light energy absorbed by upper canopy leaves saturates photosynthesis and exceeds the energetic requirements for light‐saturated linear electron flow through photosystem II (J_PSII), so that photoinhibition results. From a theoretical consideration of the response of light‐saturated photosynthesis to elevated atmospheric CO₂ partial pressure (pCO₂) it may be predicted that, where light‐saturated photosynthesis is Rubisco‐limited, an increase in pCO₂ will stimulate J_PSII. Therefore, the proportion of absorbed quanta dissipated photochemically will increase and the potential for photoinhibition of photosynthesis will decrease. This was tested by measuring modulated chlorophyll a fluorescence from Quercus myrtifolia Willd. growing in the field in open‐top chambers, at either current ambient or elevated (ambient + 35 Pa) pCO₂ on Merritt Island, Florida, USA. During spring and summer, light‐saturated photosynthesis at current ambient pCO₂ was Rubisco‐limited. Consistent with theoretical prediction, J_PSII was increased and photoinhibition decreased by elevated pCO₂ in spring. In the summer, when growth had largely ceased, an acclimatory decrease in the maximum Ribulose 1,5 bisphosphate saturated carboxylation capacity (V_{c max}) removed the stimulation of J_PSII seen in the spring, and photoinhibition was increased in elevated pCO₂. It is concluded that, for Q. myrtifolia growing in the field, the effects of elevated pCO₂ on J_PSII and photoinhibition will reflect seasonal differences in photosynthetic acclimation to elevated pCO₂ in a predictable manner.     

Symbiosis regulation in a facultatively symbiotic temperate coral: zooxanthellae division and expulsion

Zooxanthellae mitotic index (MI) and expulsion rates were measured in the facultatively symbiotic scleractinian Astrangia poculata during winter and summer off the southern New England coast, USA. While MI was significantly higher in summer than in winter, mean expulsion rates were comparable between seasons. Corals therefore appear to allow increases in symbiont density when symbiosis is advantageous during the warm season, followed by a net reduction during the cold season when zooxanthellae may draw resources from the coral. Given previous reports that photosynthesis in A. poculata symbionts does not occur below approximately 6°C, considerable zooxanthellae division at 3°C and in darkness suggests that zooxanthellae are heterotrophic at low seasonal temperatures. Finally, examination of expulsion as a function of zooxanthellae density revealed that corals with very low zooxanthellae densities export a significantly greater proportion of their symbionts, apparently allowing them to persist in a stable azooxanthellate state.     

Effects of long-term experimental night-time warming and drought on photosynthesis, <Emphasis Type="Italic">F</Emphasis>v/<Emphasis Type="Italic">F</Emphasis>m and stomatal conductance in the dominant species of a Mediterranean shrubland

We conducted a night-time warming and drought field experiment for 7 years (1999–2005) in a Mediterranean shrubland. We focused on the two dominant shrub species, Erica multiflora L. and Globularia alypum L. and the tree Pinus halepensis L. and the final years to study the effects of the experimental night-time warming and drought on Fv/Fm, photosynthesis, and stomatal conductance. Warming treatment increased mean air temperature and mean soil temperature through the years by an average of 0.7 and 0.9°C respectively, and drought treatment reduced soil moisture through the years by an average of 19%. Warming tended to increase photosynthetic rates in E. multiflora, G. alypum and P. halepensis mostly in the cold seasons, when plants were more limited by temperature, as shown by the lowest values of Fv/Fm being detected in winter in the three studied species. A negative effect of warming was only detected for E. multiflora in summer 2003. Drought treatment generated different responses of net photosynthetic rates depending on the species, season and year. Stomatal conductance showed the same pattern as photosynthesis for the three studied species, displaying seasonal and inter-annual variability, although with an overall negative effect of drought for P. halepensis. Photosynthetic rates decreased significantly in the dry winter 2005 and spring 2005 in comparison to the same seasons of 2003 and 2004. There were positive correlations between the photosynthetic rates in different seasons for E. multiflora, G. alypum and P. halepensis and the soil moisture of the week prior to measurements. The great variation in the photosynthetic rates was thus explained in a significant part by soil moisture levels. The lowest Fv/Fm values usually corresponded with lowest stomatal conductances suggesting that drought stress could be associated to stress by low temperatures in winter.     

Thylakoid membrane responses to moderately high leaf temperature in Pima cotton

Photosynthesis is inhibited by high temperatures that plants are likely to experience under natural conditions. Both increased thylakoid membrane ionic conductance and ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco) deactivation have been suggested as the primary cause. The moderately heat‐tolerant crop Pima S‐6 cotton (Gossypium barbadense) was used to examine heat stress‐induced inhibition of photosynthesis. Previous field‐work indicated that moderate heat stress (T = 35–45 °C) is associated with very rapid leaf temperature changes. Therefore, a system was devised for rapidly heating intact, attached leaves to mimic natural field heat‐stress conditions and monitored Rubisco activation, carbon‐cycle metabolites, thylakoid ionic conductance, and photosystem I activity. As a proxy for NADPH and stromal redox status the activation state of NADP‐malate dehydrogenase (NADP‐MDH) was measured. In dark‐adapted cotton leaves, heating caused an increase in thylakoid permeability at temperatures as low as 36 °C. The increased permeability did not cause a decline in adenosine 5′‐triphosphate (ATP) levels during steady‐state or transient heating. Rapid heating caused a transient decline in ribulose 1,5‐bisphosphate without a decrease in Rubisco activation. Sustained heating caused a decline in Rubisco activation and also oxidized the stroma as judged by NADP‐MDH activation and this is hypothesized to result from increased cyclic photophosphorylation, explaining the maintenance of ATP content in the face of increased thylakoid membrane ion leakiness.     

Acclimation of snow gum (Eucalyptus pauciflora) leaf respiration to seasonal and diurnal variations in temperature: the importance of changes in the capacity and temperature sensitivity of respiration

We investigated the relationship between daily and seasonal temperature variation and dark respiratory CO₂ release by leaves of snow gum (Eucalyptus pauciflora Sieb. ex Spreng) that were grown in their natural habitat or under controlled‐environment conditions. The open grassland field site in SE Australia was characterized by large seasonal and diurnal changes in air temperature. On each measurement day, leaf respiration rates in darkness were measured in situ at 2–3 h intervals over a 24 h period, with measurements being conducted at the ambient leaf temperature. The rate of respiration at a set measuring temperature (i.e. apparent ‘respiratory capacity’) was greater in seedlings grown under low average daily temperatures (i.e. acclimation occurred), both in the field and under controlled‐environment conditions. The sensitivity of leaf respiration to diurnal changes in temperature (i.e. the Q₁₀ of leaf respiration) exhibited little seasonal variation over much of the year. However, Q₁₀ values were significantly greater on cold winter days (i.e. when daily average and minimum air temperatures were below 6° and –1 °C, respectively). These differences in Q₁₀ values were not due to bias arizing from the contrasting daily temperature amplitudes in winter and summer, as the Q₁₀ of leaf respiration was constant over a wide temperature range in short‐term experiments. Due to the higher Q₁₀ values in winter, there was less difference between winter and summer leaf respiration rates measured at 5 °C than at 25 °C. The net result of these changes was that there was relatively little difference in total daily leaf respiratory CO₂ release per unit leaf dry mass in winter and summer. Under controlled‐environment conditions, acclimation of respiration to growth temperature occurred in as little as 1–3 d. Acclimation was associated with a change in the concentration of soluble sugars under controlled conditions, but not in the field. Our data suggest that acclimation in the field may be associated with the onset of cold‐induced photo‐inhibition. We conclude that cold‐acclimation of dark respiration in snow gum leaves is characterized by changes in both the temperature sensitivity and apparent ‘capacity’ of the respiratory apparatus, and that such changes will have an important impact on the carbon economy of snow gum plants.     

Differential responses of thermal tolerance to acclimation in the sub-Antarctic rove beetle Halmaeusa atriceps

Abstract. Investigations of the responses to acclimation of upper and lower lethal limits and limits to activity in insects have focused primarily on Drosophila. In the present study, Halmaeusa atriceps (Staphylinidae) is examined for thermal tolerance responses to acclimation, and seasonal acclimatization. In summer and winter, lower lethal temperatures of adults and larvae are approximately −7.6 ± 0.03 and −11.1 ± 0.06 °C, respectively. Supercooling points (SCPs) are more variable, with winter SCPs of −5.4 ± 0.4 °C in larvae and −6.3 ± 0.8 °C in adults. The species appears to be chill susceptible in summer and moderately freeze tolerant in winter, thus showing seasonal acclimatization. Similar changes cannot be induced solely by acclimation to low temperatures in the laboratory. Upper lethal temperatures show a weaker response to acclimation. There are also significant responses to acclimation of critical thermal limits. Critical thermal minima vary between −3.6 ± 0.2 and −0.6 ± 0.2 °C in larvae, and from −4.1 ± 0.1 to −0.8 ± 0.2 °C in adults. By contrast, critical thermal maxima vary much less within adults and larvae. These findings are in keeping with the general pattern found in insects, although this species differs in several respects from others found on Marion Island.     

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.