Photosynthetic limitations in olive cultivars with different sensitivity to salt stress

Olive (Olea europea L) is one of the most valuable and widespread fruit trees in the Mediterranean area. To breed olive for resistance to salinity, an environmental constraint typical of the Mediterranean, is an important goal. The photosynthetic limitations associated with salt stress caused by irrigation with saline (200 mm ) water were assessed with simultaneous gas‐exchange and fluorescence field measurements in six olive cultivars. Cultivars were found to possess inherently different photosynthesis when non‐stressed. When exposed to salt stress, cultivars with inherently high photosynthesis showed the highest photosynthetic reductions. There was no relationship between salt accumulation and photosynthesis reduction in either young or old leaves. Thus photosynthetic sensitivity to salt did not depend on salt exclusion or compartmentalization in the old leaves of the olive cultivars investigated. Salt reduced the photochemical efficiency, but this reduction was also not associated with photosynthesis reduction. Salt caused a reduction of stomatal and mesophyll conductance, especially in cultivars with inherently high photosynthesis. Mesophyll conductance was generally strongly associated with photosynthesis, but not in salt‐stressed leaves with a mesophyll conductance higher than 50 mmol m^?2 s^?1. The combined reduction of stomatal and mesophyll conductances in salt‐stressed leaves increased the CO₂ draw‐down between ambient air and the chloroplasts. The CO₂ draw‐down was strongly associated with photosynthesis reduction of salt‐stressed leaves but also with the variable photosynthesis of controls. The relationship between photosynthesis and CO₂ draw‐down remained unchanged in most of the cultivars, suggesting no or small changes in Rubisco activity of salt‐stressed leaves. The present results indicate that the low chloroplast CO₂ concentration set by both low stomatal and mesophyll conductances were the main limitations of photosynthesis in salt‐stressed olive as well as in cultivars with inherently low photosynthesis. It is consequently suggested that, independently of the apparent sensitivity of photosynthesis to salt, this effect may be relieved if conductances to CO₂ diffusion are restored.     

Analysis of fast chlorophyll fluorescence rise (O‐K‐J‐I‐P) curves in green fruits indicates electron flow limitations at the donor side of PSII and the acceptor sides of both photosystems

Limited evidence up to now indicates low linear photosynthetic electron flow and CO₂ assimilation rates in non‐foliar chloroplasts. In this investigation, we used chlorophyll fluorescence techniques to locate possible limiting steps in photosystem function in exposed, non‐stressed green fruits (both pericarps and seeds) of three species, while corresponding leaves served as controls. Compared with leaves, fruit photosynthesis was characterized by less photon trapping and less quantum yields of electron flow, while the non‐photochemical quenching was higher and potentially linked to enhanced carotenoid/chlorophyll ratios. Analysis of fast chlorophyll fluorescence rise curves revealed possible limitations both in the donor (oxygen evolving complex) and the acceptor (Q_A^?→ intermediate carriers) sides of photosystem II (PSII) indicating innately low PSII photochemical activity. On the other hand, PSI was characterized by faster reduction of its final electron acceptors and their small pool sizes. We argue that the fast reductive saturation of final PSI electron acceptors may divert electrons back to intermediate carriers facilitating a cyclic flow around PSI, while the partial inactivation of linear flow precludes strong reduction of plastoquinone. As such, the photosynthetic attributes of fruit chloroplasts may act to replenish the ATP lost because of hypoxia usually encountered in sink organs with high diffusive resistance to gas exchange.     

Photosynthetic Responses of a Temperate Liana to Xylella fastidiosa Infection and Water Stress

Xylella fastidiosa is a xylem‐limited bacterial plant pathogen that causes bacterial leaf scorch in its hosts. Our previous work showed that water stress enhances leaf scorch symptom severity and progression along the stem of a liana, Parthenocissus quinquefolia, infected by X. fastidiosa. This paper explores the photosynthetic gas exchange responses of P. quinquefolia, with the aim to elucidate mechanisms behind disease expression and its interaction with water stress. We used a 2 × 2‐complete factorial design, repeated over two growing seasons, with high and low soil moisture levels and infected and non‐infected plants. In both years, low soil moisture levels reduced leaf water potentials, net photosynthesis and stomatal conductance at all leaf positions, while X. fastidiosa‐infection reduced these parameters at basally located leaves only. Intercellular CO₂ concentrations were reduced in apical leaves, but increased at the most basal leaf location, implicating a non‐stomatal reduction of photosynthesis in leaves showing the greatest disease development. This result was supported by measured reductions in photosynthetic rates of basal leaves at high CO₂ concentrations, where stomatal limitation was eliminated. Repeated measurements over the summer of 2000 showed that the effects of water stress and infection were progressive over time, reaching their greatest extent in September. By reducing stomatal conductances at moderate levels of water stress, P. quinquefolia maintained relatively high leaf water potentials and delayed the onset of photosynthetic damage due to pathogen and drought‐induced water stress. In addition, chlorophyll fluorescence measurements showed that P. quinquefolia has an efficient means of dissipating excess light energy that protects the photosynthetic machinery of leaves from irreversible photoinhibitory damage that may occur during stress‐induced stomatal limitation of photosynthesis. However, severe stress induced by disease and drought eventually led to non‐stomatal decreases in photosynthesis associated with leaf senescence.     

Elevated CO<Subscript>2</Subscript> reduces stomatal and metabolic limitations on photosynthesis caused by salinity in <Emphasis Type="Italic">Hordeum vulgare</Emphasis>

The future environment may be altered by high concentrations of salt in the soil and elevated [CO₂] in the atmosphere. These have opposite effects on photosynthesis. Generally, salt stress inhibits photosynthesis by stomatal and non-stomatal mechanisms; in contrast, elevated [CO₂] stimulates photosynthesis by increasing CO₂ availability in the Rubisco carboxylating site and by reducing photorespiration. However, few studies have focused on the interactive effects of these factors on photosynthesis. To elucidate this knowledge gap, we grew the barley plant, Hordeum vulgare (cv. Iranis), with and without salt stress at either ambient or elevated atmospheric [CO₂] (350 or 700 μmol mol⁻¹ CO₂, respectively). We measured growth, several photosynthetic and fluorescence parameters, and carbohydrate content. Under saline conditions, the photosynthetic rate decreased, mostly because of stomatal limitations. Increasing salinity progressively increased metabolic (photochemical and biochemical) limitation; this included an increase in non-photochemical quenching and a reduction in the PSII quantum yield. When salinity was combined with elevated CO₂, the rate of CO₂ diffusion to the carboxylating site increased, despite lower stomatal and internal conductance. The greater CO₂ availability increased the electron sink capacity, which alleviated the salt-induced metabolic limitations on the photosynthetic rate. Consequently, elevated CO₂ partially mitigated the saline effects on photosynthesis by maintaining favorable biochemistry and photochemistry in barley leaves.     

Understanding the Low Photosynthetic Rates of Sun and Shade Coffee Leaves: Bridging the Gap on the Relative Roles of Hydraulic,Diffusive and Biochemical Constraints to Photosynthesis

It has long been held that the low photosynthetic rates (A) of coffee leaves are largely associated with diffusive constraints to photosynthesis. However, the relative limitations of the stomata and mesophyll to the overall diffusional constraints to photosynthesis, as well as the coordination of leaf hydraulics with photosynthetic limitations, remain to be fully elucidated in coffee. Whether the low actual A under ambient CO₂ concentrations is associated with the kinetic properties of Rubisco and high (photo)respiration rates also remains elusive. Here, we provide a holistic analysis to understand the causes associated with low A by measuring a variety of key anatomical/hydraulic and photosynthetic traits in sun- and shade-grown coffee plants. We demonstrate that leaf hydraulic architecture imposes a major constraint on the maximisation of the photosynthetic gas exchange of coffee leaves. Regardless of the light treatments, A was mainly limited by stomatal factors followed by similar limitations associated with the mesophyll and biochemical constraints. No evidence of an inefficient Rubisco was found; rather, we propose that coffee Rubisco is well tuned for operating at low chloroplastic CO₂ concentrations. Finally, we contend that large diffusive resistance should lead to large CO₂ drawdown from the intercellular airspaces to the sites of carboxylation, thus favouring the occurrence of relatively high photorespiration rates, which ultimately leads to further limitations to A.     

Microbial activity and organic matter dynamics during 4 years of irrigation with treated wastewater

The global changes in rainfall frequency and quantity have subjected arid and semi-arid regions to long periods of drought. As this phenomenon corresponds to increasing trend of water shortage, the use of treated wastewater (TWW) has been suggested as an alternative for irrigation of agricultural crops in these areas. The aim of the study was to investigate the short- and middle-term effects of TWW irrigation on the soil microbial activities and organic carbon content. The microbial community activity was measured every 1–3 months for 4 years in a persimmon (Diospyros kaki) orchard. These activities were used here as an indicator for the soil health. The hydrolysis activity (detected by fluorescein diacetate hydrolysis (FDA) assay) increased during the irrigation season and was significantly higher in soils irrigated with TWW compared to those irrigated with freshwater (FW). This activity was also negatively correlated with dissolved organic carbon (DOC) concentrations during the irrigation season, suggesting that the community degraded the DOC in the soils regardless of its origin. The irrigation season was also characterized by an increase in nitrification potential in both TWW- and FW-irrigated soils, which coincided with high concentrations of nitrate (50 mg kg⁻¹ soil). Overall, there was an increase in all measured activities during the irrigation season, and they were higher in the TWW soils. However, it appears that after each irrigation season, the potential activity of the community returned to levels similar to or even slightly lower than those of FW-irrigated soil during the wet season, suggesting that the periodic irrigation did not significantly change the soil microbial activity.     

Moderate water stress causes different stomatal and non‐stomatal changes in the photosynthetic functioning of Phaseolus vulgaris L. genotypes

The impact of moderate water deficit on the photosynthetic apparatus of three Phaseolus vulgaris L. cultivars, Plovdiv 10 (P10), Dobrudjanski Ran (DR) and Prelom (Prel), was investigated. Water shortage had less impact on leaf hydration, RWC (predawn and midday) and predawn water potential in Prel. RWC and Ψ_p were more reduced in P10, while there was no osmotic adjustment in any cultivar. Although drought drastically reduced stomatal opening in P10 and DR, reduced A_max indicated non‐stomatal limitations that contributed to the negligible P_n. These limitations were on potential thylakoid electron transport rates of PSI and II, pointing to photosystem functioning as a major limiting step in photosynthesis. This agrees with decreases in actual photochemical efficiency of PSII (F_v′/F_m′), quantum yield of photosynthetic non‐cyclic electron transport (?_e) and energy‐driven photochemical events (q_P), although the impact on these parameters would also include down‐regulation processes. When compared to DR, Prel retained a higher functional state of the photosynthetic machinery, justifying reduced need for photoprotective mechanisms (non‐photochemical quenching, zeaxanthin, lutein, β‐carotene) and maintenance of the balance between energy capture and dissipative pigments. The highest increases in fructose, glucose, arabinose and sorbitol in Prel might be related to tolerance to a lower oxidative state. All cultivars had reduced A_max due to daytime stomatal closure in well‐watered conditions. Under moderate drought, Prel had highest tolerance, higher leaf hydration and maintenance of important photochemical use of energy. However, water shortage caused appreciable non‐stomatal limitations to photosynthesis linked to regulation/imbalance at the metabolic level (and growth) in all cultivars. This included damage, as reflected in decreased potential photosystem functioning, pointing to higher sensitivity of photosynthesis to drought than is commonly assumed.     

Seasonal and diurnal changes in photosynthetic limitation of young sweet orange trees

This study tests the hypothesis that potted sweet orange plants show a significant variation in photosynthesis over seasonal and diurnal cycles, even in well-hydrated conditions. This hypothesis was tested by measuring diurnal variations in leaf gas exchange, chlorophyll fluorescence, leaf water potential, and the responses of CO₂ assimilation to increasing air CO₂ concentrations in 1-year-old ‘Valência’ sweet orange scions grafted onto ‘Cleopatra’ mandarin rootstocks during the winter and summer seasons in a subtropical climate. In addition, diurnal leaf gas exchange was evaluated under controlled conditions, with constant environmental conditions during both winter and summer. In relation to our hypothesis, a greater rate of photosynthesis is found during the summer compared to the winter. Reduced photosynthesis during winter was induced by cool night conditions, as the diurnal fluctuation of environmental conditions was not limiting. Low air and soil temperatures caused decreases in the stomatal conductance and in the rates of the biochemical reactions underlying photosynthesis (ribulose-1,5-bisphosphate (RuBP) carboxylation and RuBP regeneration) during the winter compared to the values obtained for those markers in the summer. Citrus photosynthesis during the summer was not impaired by biochemical or photochemical reactions, as CO₂ assimilation was only limited by stomatal conductance due to high leaf-to-air vapor pressure difference (VPD) during the afternoon. During the winter, the reduction in photosynthesis during the afternoon was caused by decreases in RuBP regeneration and stomatal conductance, which are both precipitated by low night temperature.     

Asymmetric responses of adaxial and abaxial stomata to elevated CO2: impacts on the control of gas exchange by leaves

The response of adaxial and abaxial stomatal conductance in Rumex obtusifolius to growth at elevated atmospheric concentrations of CO₂ (250 μmol mol^?1 above ambient) was investigated over two growing seasons. The conductance of both the adaxial and abaxial leaf surfaces was found to be reduced by elevated concentrations of CO₂. Elevated CO₂ caused a much greater reduction in conductance for the adaxial surface than for the abaxial surface. The absence of effects upon stomatal density indicated that the reductions were probably the result of changes in stomatal aperture. Partitioning of gas exchange between the leaf surfaces revealed that increased concentrations of CO₂ caused increased rates of photosynthesis only via the abaxial surface. Additionally, leaf thickness was found to increase during growth at elevated concentrations of CO₂. The tendency for these amphistomatous leaves to develop a distribution of conductance approaching that of hypostomatous leaves clearly reduced their maximum photosynthetic potential. This conclusion was supported by measurements of stomatal limitation, which showed greater values for the adaxial surfaces, and greater values at elevated CO₂. This reduction in photosynthesis may in part be caused by higher diffusive limitations imposed because of increased leaf thickness. In an uncoupled canopy, asymmetrical stomatal responses of the kind identified here may appreciably reduce transpiration. Species which show symmetrical responses are less likely to show reduced transpirational rates, and a redistribution of water loss between species may occur. The implications of asymmetrical stomatal responses for photosynthesis and canopy transpiration are discussed.     

Sinks for photosynthetic electron flow in green petioles and pedicels of Zantedeschia aethiopica: evidence for innately high photorespiration and cyclic electron flow rates

A combination of gas exchange and various chlorophyll fluorescence measurements under varying O₂ and CO₂ partial pressures were used to characterize photosynthesis in green, stomata-bearing petioles of Zantedeschia aethiopica (calla lily) while corresponding leaves served as controls. Compared to leaves, petioles displayed considerably lower CO₂ assimilation rates, limited by both stomatal and mesophyll components. Further analysis of mesophyll limitations indicated lower carboxylating efficiencies and insufficient RuBP regeneration but almost similar rates of linear electron transport. Accordingly, higher oxygenation/carboxylation ratios were assumed for petioles and confirmed by experiments under non-photorespiratory conditions. Higher photorespiration rates in petioles were accompanied by higher cyclic electron flow around PSI, the latter being possibly linked to limitations in electron transport from intermediate electron carriers to end acceptors and low contents of PSI. Based on chlorophyll fluorescence methods, similar conclusions can be drawn for green pedicels, although gas exchange in these organs could not be applied due to their bulky size. Since our test plants were not subjected to stress we argue that higher photorespiration and cyclic electron flow rates are innate attributes of photosynthesis in stalks of calla lily. Active nitrogen metabolism may be inferred, while increased cyclic electron flow may provide the additional ATP required for the enhanced photorespiratory activity in petiole and pedicel chloroplasts and/or the decarboxylation of malate ascending from roots.     

Photosynthesis and Transpiration as a Function of Gaseous Diffusive Resistances in Orange Leaves

The CO₂ and H₂O vapour exchange of single attached orange, Citrus sinensis (L.), leaves was measured under laboratory conditions using infrared gas analysis. Gaseous diffusive resistances were derived from measurements at a saturating irradiance and at a leaf temperature optimum for photosynthesis. Variation in leaf resistance (within the range 1.6 to 60 s cm^-1) induced by moisture status, or by cyclic oscillations in stomatal aperture, was associated with changes in both photosynthesis and transpiration. At low leaf resistance (ri less than 10 s cm^-1) the ratio of transpiration to photosynthesis declined with reduced stomatal aperture, indicating a tighter stomatal control over H₂O vapour loss than over CO₂ assimilation. At higher leaf resistance (ri greater than 10 s cm^-1) changes in transpiration and photosynthesis were linearly related, but leaf resistance and mesophyll resistance were also positively correlated, so that strictly stomatal control of photosynthesis became more apparent than real. This evidence, combined with direct measurements of CO₂ diffusive resistances (in a -O₂ gas stream) emphasised the presence of a significant mesophyll resistance; i.e., an additional and rate limiting resistance to CO₂ assimilation over and above that encountered by H₂O vapour escaping from the leaf.     

Contributions of diffusional limitation, photoinhibition and photorespiration to midday depression of photosynthesis in Arisaema heterophyllum in natural high light

Diurnal changes in photosynthetic gas exchange and chlorophyll fluorescence were measured under full sunlight to reveal diffusional and non‐diffusional limitations to diurnal assimilation in leaves of Arisaema heterophyllum Blume plants grown either in a riparian forest understorey (shade leaves) or in an adjacent deforested open site (sun leaves). Midday depressions of assimilation rate (A) and leaf conductance of water vapour were remarkably deeper in shade leaves than in sun leaves. To evaluate the diffusional (i.e. stomatal and leaf internal) limitation to assimilation, we used an index [1–A/A₃₅₀], in which A₃₅₀ is A at a chloroplast CO₂ concentration of 350 μ mol mol ^{? 1}. A₃₅₀ was estimated from the electron transport rate (J_T), determined fluorometrically, and the specificity factor of Rubisco (S), determined by gas exchange techniques. In sun leaves under saturating light, the index obtained after the ‘peak’ of diurnal assimilation was 70% greater than that obtained before the ‘peak’, but in shade leaves, it was only 20% greater. The photochemical efficiency of photosystem II ( Δ F/F_m ′ ) and thus J_T was considerably lower in shade leaves than in sun leaves, especially after the ‘peak’. In shade leaves but not in sun leaves, A at a photosynthetically active photon flux density (PPFD) > 500 μ mol m ^{? 2} s ^{? 1} depended positively on J_T throughout the day. Electron flows used by the carboxylation and oxygenation (J_O) of RuBP were estimated from A and J_T. In sun leaves, the J_O/J_T ratio was significantly higher after the ‘peak’, but little difference was found in shade leaves. Photorespiratory CO₂ efflux in the absence of atmospheric CO₂ was about three times higher in sun leaves than in shade leaves. We attribute the midday depression of assimilation in sun leaves to the increased rate of photorespiration caused by stomatal closure, and that in shade leaves to severe photoinhibition. Thus, for sun leaves, increased capacities for photorespiration and non‐photochemical quenching are essential to avoid photoinhibitory damage and to tolerate high leaf temperatures and water stress under excess light. The increased Rubisco content in sun leaves, which has been recognized as raising photosynthetic assimilation capacity, also contributes to increase in the capacity for photorespiration.     

Flavescence dorée phytoplasma deregulates stomatal control of photosynthesis in Vitis vinifera

Flavescence dorée (FD) is among the major grapevine diseases causing high management costs; curative methods against FD are unavailable. In FD‐infected plants, decrease in photosynthesis is usually recorded, but deregulation in stomatal control of leaf gas exchange during FD infection and recovery is unknown. We measured the seasonal time course of gas exchange rates in two cultivars (‘Barbera’ and ‘Nebbiolo’) during the term of 1 year when grapevines experienced a water stress and another with no drought, with difference in gas exchange rates in response to FD infection and recovery as assessed by symptom observation and phytoplasma detection through PCR analysis. Chlorophyll fluorescence was also evaluated at the time of maximum symptom severity in ‘Barbera’, the cultivar showing the most severe stress response to FD infection, causing the highest damage in vineyards of north‐western Italy. In FD‐infected plants, net photosynthesis and transpiration gradually decreased during the season, more during the no drought year than during drought. During recovery, healthy (PCR negative) plants infected 2 years before, but not those infected an year before, regained the gas exchange performances to the level as measured before infection. The relationships between stomatal conductance and the residual leaf intercellular CO₂ concentration (c_i) discriminated healthy versus FD‐infected and recovered plants; at the same c_i, FD‐infected leaves had higher non‐photochemical quenching than healthy ones. We conclude that metabolic, not stomatal, leaf gas exchange limitation in FD‐infected and recovered grapevines is the basis of plant response to FD disease. In addition, we also suggest that such response is dependent upon water stress, by showing that water stress superimposes on FD infection in terms of stomatal and metabolic non‐stomatal limitations to carbon assimilation.     

Effect of leaf surface wetness and wettability on photosynthesis in bean and pea

Leaf surface wetness that occurs frequently in natural environments has a significant impact on leaf photosynthesis. However, the physiological mechanisms for the photosynthetic responses to wetness are not well understood. The responses of leaf CO₂ assimilation rate (A) to 72 h of artificial mist of a wettable (bean; Phaseolus vulgaris) and a non‐wettable species (pea; Pisum sativum) were compared. Stomatal and non‐stomatal limitations to A were investigated. A 28% inhibition of A was observed in the bean leaves as a result of a 16% decrease in stomatal conductance and a 55% reduction in the amount of Rubisco. The decrease of Rubisco was mainly due to its partial degradation. In contrast to the bean leaves, a 22% stimulation of A was obtained in the 72 h mist‐treated pea leaves. Mist treatment increased stomatal conductance by 12.5% and had no effect on the amount of Rubisco. These results indicated that a positive photosynthetic response to wetness occurred only in non‐wettable species and is due to the change in stomatal regulation.     

The use of low [CO2] to estimate diffusional and non-diffusional limitations of photosynthetic capacity of salt-stressed olive saplings

In this study it has been shown that increased diffusional resistances caused by salt stress may be fully overcome by exposing attached leaves to very low [CO₂] (～ 50 µmol mol^?1), and, thus a non‐destructive‐in vivo method to correctly estimate photosynthetic capacity in stressed plants is reported. Diffusional (i.e. stomatal conductance, g_s, and mesophyll conductance to CO₂, g_m) and biochemical limitations to photosynthesis (A) were measured in two 1‐year‐old Greek olive cultivars (Chalkidikis and Kerkiras) subjected to salt stress by adding 200 mm NaCl to the irrigation water. Two sets of A–C_i curves were measured. A first set of standard A–C_i curves (i.e. without pre‐conditioning plants at low [CO₂]), were generated for salt‐stressed plants. A second set of A–C_i curves were measured, on both control and salt‐stressed plants, after pre‐conditioning leaves at [CO₂] of ～ 50 µmol mol^?1 for about 1.5 h to force stomatal opening. This forced stomata to be wide open, and g_s increased to similar values in control and salt‐stressed plants of both cultivars. After g_s had approached the maximum value, the A–C_i response was again measured. The analysis of the photosynthetic capacity of the salt‐stressed plants based on the standard A–C_i curves, showed low values of the J_max (maximum rate of electron transport) to V_cmax (RuBP‐saturated rate of Rubisco) ratio (1.06), that would implicate a reduced rate of RuBP regeneration, and, thus, a metabolic impairment. However, the analysis of the A–C_i curves made on pre‐conditioned leaves, showed that the estimates of the photosynthetic capacity parameters were much higher than in the standard A–C_i responses. Moreover, these values were similar in magnitude to the average values reported by Wullschleger (Journal of Experimental Botany 44, 907–920, 1993) in a survey of 109 C₃ species. These findings clearly indicates that: (1) salt stress did affect g_s and g_m but not the biochemical capacity to assimilate CO₂ and therefore, in these conditions, the sum of the diffusional resistances set the limit to photosynthesis rates; (2) there was a linear relationship (r² = 0.68) between g_m and g_s, and, thus, changes of g_m can be as fast as those of g_s; (3) the estimates of photosynthetic capacity based on A–C_i curves made without removing diffusional limitations are artificially low and lead to incorrect interpretations of the actual limitations of photosynthesis; and (4) the analysis of the photosynthetic properties in terms of stomatal and non‐stomatal limitations should be replaced by the analysis of diffusional and non‐diffusional limitations of photosynthesis. Finally, the C₃ photosynthesis model parameterization using in vitro‐measured and in vivo‐measured kinetics parameters was compared. Applying the in vivo‐measured Rubisco kinetics parameters resulted in a better parameterization of the photosynthesis model.     

Variation in the carbon and oxygen isotope composition of plant biomass and its relationship to water‐use efficiency at the leaf‐ and ecosystem‐scales in a northern Great Plains grassland

Measurements of the carbon (δ¹³C_m) and oxygen (δ¹⁸O_m) isotope composition of C₃ plant tissue provide important insights into controls on water‐use efficiency. We investigated the causes of seasonal and inter‐annual variability in water‐use efficiency in a grassland near Lethbridge, Canada using stable isotope (leaf‐scale) and eddy covariance measurements (ecosystem‐scale). The positive relationship between δ¹³C_m and δ¹⁸O_m values for samples collected during 1998–2001 indicated that variation in stomatal conductance and water stress‐induced changes in the degree of stomatal limitation of net photosynthesis were the major controls on variation in δ¹³C_m and biomass production during this time. By comparison, the lack of a significant relationship between δ¹³C_m and δ¹⁸O_m values during 2002, 2003 and 2006 demonstrated that water stress was not a significant limitation on photosynthesis and biomass production in these years. Water‐use efficiency was higher in 2000 than 1999, consistent with expectations because of greater stomatal limitation of photosynthesis and lower leaf c_i/ca during the drier conditions of 2000. Calculated values of leaf‐scale water‐use efficiency were 2–3 times higher than ecosystem‐scale water‐use efficiency, a difference that was likely due to carbon lost in root respiration and water lost during soil evaporation that was not accounted for by the stable isotope measurements.     

Enhanced tolerance of photosynthesis against high temperature damage in salt-adapted halophyte Atriplex centralasiatica plants

Thermotolerance of photosynthesis in salt‐adapted Atriplex centralasiatica plants (100–400 mm NaCl) was evaluated in this study after detached leaves and whole plants were exposed to high temperature stress (30–48 °C) either in the dark or under high light (1200 mol m^?2 s^?1). In parallel with the decrease in stomatal conductance, intercellular CO₂ concentration and CO₂ assimilation rate decreased significantly with increasing salt concentration. There was no change in the maximal efficiency of PSII photochemistry (F_v/F_m) with increasing salt concentration, suggesting that there was no damage to PSII in salt‐adapted plants. On the other hand, there was a striking difference in the response of PSII and CO₂ assimilation capacity to heat stress in non‐salt‐adapted and salt‐adapted leaves. Leaves from salt‐adapted plants maintained significantly higher F_v/F_m values than those from non‐salt‐adapted leaves at temperatures higher than 42 °C. The F_v/F_m differences between non‐salt‐adapted and salt‐adapted plants persisted for at least 24 h following heat stress. Leaves from salt‐adapted plants also maintained a higher net CO₂ assimilation rate than those in non‐salt‐adapted plants at temperatures higher than 42 °C. This increased thermotolerance was independent of the degree of salinity since no significant changes in F_v/F_m and net CO₂ assimilation rate were observed among the plants treated with different concentrations of NaCl. The increased thermotolerance of PSII induced by salinity was still evident when heat treatments were carried out under high light. Given that photosynthesis is considered to be the physiological process most sensitive to high temperature damage, increased thermotolerance of photosynthesis may be of significance since A. centralasiatica, a typical halophyte, grows in the high salinity regions in the north of China, where the temperature in the summer is often as high as 45 °C.     

Photorespiration and photoinhibition in the bracts of cotton under water stress

Gas exchange and chlorophyll fluorescence parameters of PSII were analyzed in the bracts and leaves of cotton plants after anthesis. Photosynthetic activity and photorespiration were measured in the leaves and bracts of cotton grown under either normal or reduced water-saving drip irrigation. The photosynthetic performance, amount of chlorophyll and Rubisco, and net photosynthesis were greater in the bracts than that in the leaves under water stress. The actual photochemical efficiency of PSII decreased in both the bracts and leaves after anthesis under reduced irrigation. However, the decrease was smaller in the bracts than in the leaves, indicating that the bracts experienced less severe photoinhibition compared to the leaves. The greater drought tolerance of bracts could be related to differences in relative water content, instantaneous water-use efficiency, and photorespiration rate. The ratio of photorespiration to net photosynthesis was much higher in the bracts than in leaves. Furthermore, water deficiency (due to the water-saving drip irrigation) had no significant effect on that ratio in the bracts. We hypothesized that photorespiration in the bracts alleviated photoinhibition and maintained photosynthetic activity.     

Water availability affects seasonal CO2‐induced photosynthetic enhancement in herbaceous species in a periodically dry woodland

Elevated atmospheric CO₂ (eCO₂) is expected to reduce the impacts of drought and increase photosynthetic rates via two key mechanisms: first, through decreased stomatal conductance (g_s) and increased soil water content (V_SWC) and second, through increased leaf internal CO₂ (C_i) and decreased stomatal limitations (S_lim). It is unclear if such findings from temperate grassland studies similarly pertain to warmer ecosystems with periodic water deficits. We tested these mechanisms in three important C₃ herbaceous species in a periodically dry Eucalyptus woodland and investigated how eCO₂‐induced photosynthetic enhancement varied with seasonal water availability, over a 3 year period. Leaf photosynthesis increased by 10%–50% with a 150 μmol mol^?1 increase in atmospheric CO₂ across seasons. This eCO₂‐induced increase in photosynthesis was a function of seasonal water availability, given by recent precipitation and mean daily V_SWC. The highest photosynthetic enhancement by eCO₂ (>30%) was observed during the most water‐limited period, for example, with V_SWC <0.07 in this sandy surface soil. Under eCO₂ there was neither a significant decrease in g_s in the three herbaceous species, nor increases in V_SWC, indicating no “water‐savings effect” of eCO₂. Periods of low V_SWC showed lower g_s (less than ≈ 0.12 mol m^?2 s^?1), higher relative S_lim (>30%) and decreased C_i under the ambient CO₂ concentration (aCO₂), with leaf photosynthesis strongly carboxylation‐limited. The alleviation of S_lim by eCO₂ was facilitated by increasing C_i, thus yielding a larger photosynthetic enhancement during dry periods. We demonstrated that water availability, but not eCO₂, controls g_s and hence the magnitude of photosynthetic enhancement in the understory herbaceous plants. Thus, eCO₂ has the potential to alter vegetation functioning in a periodically dry woodland understory through changes in stomatal limitation to photosynthesis, not by the “water‐savings effect” usually invoked in grasslands.     

Metabolism of <Superscript>14</Superscript>C-glycine as a substrate for photorespiration of the leaf at different developmental stages of ephemeroides

The metabolism of ¹⁴C-glycine (a substrate for photorespiration) was studied in the light and in darkness under natural CO₂ concentration (0.03%) in the leaves of ephemeroides Scilla sibirica Haw. and Ficaria verna Huds. at different developmental stages. Using one and the same sample, potential photosynthesis (at 1% CO₂), true photosynthesis (at 0.03% CO₂), and leaf respiratory capacity were measured by the radiometric and manometric methods, respectively. All measurements were performed at 15°C, an average temperature during ephemer growth. It was found that, in the white zone of the Scilla leaf, the rate of CO₂ evolution resulting from metabolization of exogenous ¹⁴C-glycine was similar in the light and in darkness. In the green zone of the Scilla leaf and in the green leaf of Ficaria, both ¹⁴C-glycine absorption and ¹⁴CO₂ evolution were lower in the light as compared with darkness, which is explained by CO₂ reassimilation. In all treatments of both plant species, a specific inhibitor of glycine decarboxylase complex (GDC), aminoacetonitrile (5 mM) suppressed CO₂ evolution by 20–40%. It was concluded that in ephemeroides mitochondrial GDC, responsible for CO₂ evolution in photorespiration, is formed at the earliest stage of leaf development. This indicates that photorespiration can occur simultaneously with the development of the leaf photosynthetic activity. On the basis of the assumption that carbon losses in the form of CO₂ evolved during photorespiration comprise 25% of true photosynthesis, it was calculated that, in ephemer leaves, the highest rates of photorespiration and photosynthesis were attained during flowering when the leaf area was the largest and the rate of dark respiration was reduced by 1.5–2.0 times. The highest rates of dark respiration were observed in the beginning of growth. In senescing leaves by the end of the plant vegetation, potential photosynthesis and true photosynthesis were reduced, whereas dark respiration remained essentially unchanged. It is concluded that the high rates of potential and true photosynthesis are characteristic of ephemeroides when they complete their short developmental program in early spring (at 15°C); theoretically, photorespiration also occurs at a high rate during this period, when this process provides for a defense against the threat of photoinhibition at low temperature and high insolation.