Leaf temperature effects on gas exchange in Quercus ilex L. growing under field conditions

ABSTRACT

Gas exchange temperature dependence in Quercus ilex shrubs growing in the Mediterranean maquis was analysed. The gas exchange trend was monitored during the year: photosynthetic activity (A _net) reached the highest average rates in early spring and autumn (12.5 µmol m^-2s^-1 was the absolute maximum A _net measured) and the lowest rates were monitored in the middle of June. There was a good correlation (r = 0.72) between A _net and g _s (A _net = 4.1246 ln g _s + 4316; P < 0.01), indicating that stomatal control of CO₂ diffusion plays an important role in controlling photosynthetic activity. Leaf temperature allowing the highest photosynthetic and stomatal conductance rates of Quercus ilex were in the range 17.5 – 29°C. A _net and g_s dropped below half its maximum value when leaf temperatures were below 11.5°C and above 35.7°C. Transpiration rates (E) were strongly related to leaf temperature; E increased as leaf temperature increased and the highest E rates were monitored in June, despite a 46% decrease in g _s. Leaf water loss from transpiration, during the drought period, could result in leaf water stress which would exacerbate heat effects on photosynthesis. During summer, the increase in leaf temperatures decreased g _s which in turn decreased A _net. Consequently, stomatal control in Quercus ilex may be considered as an adaptive strategy during drought.     

The influence of elevated temperature,elevated atmospheric CO2 concentration and water stress on net photosynthesis of loblolly pine (Pinus taeda L.) at northern,central and southern sites in its native range

We investigated the effect of elevated [CO₂] (700 μmol mol^?1), elevated temperature (+2 °C above ambient) and decreased soil water availability on net photosynthesis (A_net) and water relations of one‐year old potted loblolly pine (Pinus taeda L.) seedlings grown in treatment chambers with high fertility at three sites along a north‐south transect covering a large portion of the species native range. At each location (Blairsville, Athens and Tifton, GA) we constructed four treatment chambers and randomly assigned each chamber one of four treatments: ambient [CO₂] and ambient temperature, elevated [CO₂] and ambient temperature, ambient [CO₂] and elevated temperature, or elevated [CO₂] and elevated temperature. Within each chamber half of the seedlings were well watered and half received much less water (1/4 that of the well watered). Measurements of net photosynthesis (A_net), stomatal conductance (g_s), leaf water potential and leaf fluorescence were made in June and September, 2008. We observed a significant increase in A_net in response to elevated [CO₂] regardless of site or temperature treatment in June and September. An increase in air temperature of over 2 °C had no significant effect on A_net at any of the sites in June or September despite over a 6 °C difference in mean annual temperature between the sites. Decreased water availability significantly reduced A_net in all treatments at each site in June. The effects of elevated [CO₂] and temperature on g_s followed a similar trend. The temperature, [CO₂] and water treatments did not significantly affect leaf water potential or chlorophyll fluorescence. Our findings suggest that predicted increases in [CO₂] will significantly increase A_net, while predicted increases in air temperature will have little effect on A_net across the native range of loblolly pine. Potential decreases in precipitation will likely cause a significant reduction in A_net, though this may be mitigated by increased [CO₂].     

Differences in leaf gas exchange strategies explain Quercus rubra and Liriodendron tulipifera intrinsic water use efficiency responses to air pollution and climate change

Trees continuously regulate leaf physiology to acquire CO₂ while simultaneously avoiding excessive water loss. The balance between these two processes, or water use efficiency (WUE), is fundamentally important to understanding changes in carbon uptake and transpiration from the leaf to the globe under environmental change. While increasing atmospheric CO₂ (iCO₂) is known to increase tree intrinsic water use efficiency (iWUE), less clear are the additional impacts of climate and acidic air pollution and how they vary by tree species. Here, we couple annually resolved long-term records of tree-ring carbon isotope signatures with leaf physiological measurements of Quercus rubra (Quru) and Liriodendron tulipifera (Litu) at four study locations spanning nearly 100 km in the eastern United States to reconstruct historical iWUE, net photosynthesis (A_net), and stomatal conductance to water (g_s) since 1940. We first show 16%–25% increases in tree iWUE since the mid-20th century, primarily driven by iCO₂, but also document the individual and interactive effects of nitrogen (NO_x) and sulfur (SO₂) air pollution overwhelming climate. We find evidence for Quru leaf gas exchange being less tightly regulated than Litu through an analysis of isotope-derived leaf internal CO₂ (C_i), particularly in wetter, recent years. Modeled estimates of seasonally integrated A_net and g_s revealed a 43%–50% stimulation of A_net was responsible for increasing iWUE in both tree species throughout 79%–86% of the chronologies with reductions in g_s attributable to the remaining 14%–21%, building upon a growing body of literature documenting stimulated A_net overwhelming reductions in g_s as a primary mechanism of increasing iWUE of trees. Finally, our results underscore the importance of considering air pollution, which remains a major environmental issue in many areas of the world, alongside climate in the interpretation of leaf physiology derived from tree rings.     

Effects of extreme high temperature,drought and elevated CO2 on photosynthesis of the Mojave Desert evergreen shrub,Larrea tridentata

The interaction of extreme temperature events with future atmospheric CO₂ concentrations may have strong impacts on physiological performance of desert shrub seedlings, which during the critical establishment phase often endure temperature extremes in conjunction with pronounced drought. To evaluate the interaction of drought and CO₂ on photosynthesis during heat stress, one-year-old Larrea tridentata[DC] Cov. seedlings were exposed to nine days of heat with midday air temperature maxima reaching 53 °C under three atmospheric CO₂ concentrations (360, 550 and 700 mol mol^–1) and two water regimes (well-watered and droughted). Photosynthetic gas exchange, chlorophyll fluorescence and water potential responses were measured prior to, during and one week following the high temperature stress event. Heat stress markedly decreased net photosynthetic rate (A _net), stomatal conductance (g _s), and the photochemical efficiency of photosystem II (F _v/F _m) in all plants except for well-watered L. tridentata grown in 700 mol mol^–1 CO₂. A _net and g _s remained similar to pre-stress levels in these plants. In droughted L. tridentata, A _net was ca. 2× (in 550 mol mol^–1 CO₂) to 3× (in 700 mol mol^–1 CO₂) higher than in ambient-CO₂-grown plants, while g _s and F _v/F _m were similar and low in all CO₂ treatments. Following heat stress, g _s in all well-watered plants rose dramatically, exceeding pre-stress levels by up to 100%. In droughted plants, g _s and A _net rose only in plants grown at elevated CO₂ following release from heat. This recovery response was strongest at 700 mol mol^–1 CO₂, which returned to A _net and g _s values similar to pre-heat following several days of recovery. Extreme heat diminished the photosynthetic down-regulation response to growth at elevated CO₂ under well-watered conditions, similar to the action of drought. Ambient-CO₂-grown L. tridentata did not show significant recovery of photosynthetic capacity (A _\max and CE) after alleviation of temperature stress, especially when exposed to drought, while plants exposed to elevated CO₂ appeared to be unaffected. These findings suggest that elevated CO₂ could promote photosynthetic activity during critical periods of seedling establishment, and enhance the potential for L. tridentata to survive extreme high temperature events.     

Environmental and stomatal control of photosynthetic enhancement in the canopy of a sweetgum (Liquidambar styraciflua L.) plantation during 3 years of CO2 enrichment

Light‐saturated photosynthetic and stomatal responses to elevated CO₂ were measured in upper and mid‐canopy foliage of a sweetgum (Liquidambar styraciflua L) plantation exposed to free‐air CO₂ enrichment (FACE) for 3 years, to characterize environmental interactions with the sustained CO₂ effects in an intact deciduous forest stand. Responses were evaluated in relation to one another, and to seasonal patterns and natural environmental stresses, including high  temperatures, vapour pressure deficits (VPD), and drought. Photosynthetic CO₂ assimilation (A) averaged 46% higher in the +200 µmol mol^?1 CO₂ treatment, in mid‐ and upper canopy foliage. Stomatal conductance (g_s) averaged 14% (mid‐canopy) and 24% (upper canopy) lower under CO₂ enrichment. Variations in the relative responses of A and g_s were linked, such that greater relative stimulation of A was observed on dates when relative reductions in g_s were slight. Dry soils and high VPD reduced g_s and A in both treatments, and tended to diminish treatment differences. The absolute effects of CO₂ on A and g_s were minimized whenever g_s was low (<0·15 mol m^?2 s^?1), but relative effects, as the ratio of elevated to ambient rates, varied greatly under those conditions. Both stomatal and non‐stomatal limitations of A were involved during late season droughts. Leaf temperature had a limited influence on A and g_s, and there was no detectable relationship between prevailing temperature and CO₂ effects on A or g_s. The responsiveness of A and g_s to elevated CO₂, both absolute and relative, was maintained through time and within the canopy of this forest stand, subject to seasonal constraints and variability associated with limiting air and soil moisture.     

Optimal stomatal conductance in relation to photosynthesis in climatically contrasting Eucalyptus species under drought

Models of stomatal conductance (g_s) are based on coupling between g_s and CO₂ assimilation (A_net), and it is often assumed that the slope of this relationship (‘g₁’) is constant across species. However, if different plant species have adapted to different access costs of water, then there will be differences in g₁ among species. We hypothesized that g₁ should vary among species adapted to different climates, and tested the theory and its linkage to plant hydraulics using four Eucalyptus species from different climatic origins in a common garden. Optimal stomatal theory predicts that species from sub‐humid zones have a lower marginal water cost of C gain, hence lower g₁ than humid‐zone species. In agreement with the theory that g₁ is related to tissue carbon costs for water supply, we found a relationship between wood density and g₁ across Eucalyptus species of contrasting climatic origins. There were significant reductions in the parameter g₁ during drought in humid but not sub‐humid species, with the latter group maintaining g₁ in drought. There are strong differences in stomatal behaviour among related tree species in agreement with optimal stomatal theory, and these differences are consistent with the economics involved in water uptake and transport for carbon gain.     

Effects of elevated CO2 on leaf gas exchange in beech and oak at two levels of nutrient supply: consequences for sensitivity to drought in beech

Beech (Fagus sylvatica L.) and pedunculate oak (Quercus robur L.) were grown from seed for two whole seasons at two CO₂ concentrations (ambient and ambient + 250 μmol mol^?1) with two levels of soil nutrient supply. Measurements of net leaf photosynthetic rate (A) and stomatal conductance (g_s) of well-watered plants were taken over both seasons; a drought treatment was applied in the middle of the second growing season to a separate sample of beech drawn from the same population. The net leaf photosynthetic rate of well-watered plants was stimulated in elevated CO₂ by an average of 75% in beech and 33% in oak; the effect continued through both growing seasons at both nutrient levels. There were no interactive effects of CO₂ concentration and nutrient level on A or g_s in beech or oak. Stomatal conductance was reduced in elevated CO₂ by an average of 34% in oak, but in beech there were no significant reductions in g_s except under cloudy conditions (–22% in elevated CO₂). During drought, there was no effect of CO₂ concentration on g_s in beech grown with high nutrients, but for beech grown with low nutrients, g_s was significantly higher in elevated CO₂, causing more rapid soil drying. With high nutrient supply, soil drying was more rapid at elevated CO₂ due to increased leaf area. It appears that beech may substantially increase whole-plant water consumption in elevated CO₂, especially under conditions of high temperature and irradiance when damage due to high evaporative demand is most likely to occur, thereby putting itself at risk during periods of drought.     

Drought‐introduced variability of mesophyll conductance in Gossypium and its relationship with leaf anatomy

Mesophyll conductance (g_m) is one of the major determinants of photosynthetic rate, for which it has an impact on crop yield. However, the regulatory mechanisms behind the decline in g_m of cotton (Gossypium. spp) by drought are unclear. An upland cotton (Gossypium hirsutum) genotype and a pima cotton (Gossypium barbadense) genotype were used to determine the gas exchange parameters, leaf anatomical structure as well as aquaporin and carbonic anhydrase gene expression under well‐watered and drought treatment conditions. In this study, the decrease of net photosynthetic rate (A_N) under drought conditions was related to a decline in g_m and in stomatal conductance (g_s). g_m and g_s coordinate with each other to ensure optimum state of CO₂ diffusion and achieve the balance of water and CO₂ demand in the process of photosynthesis. Meanwhile, mesophyll limitations to photosynthesis are equally important to the stomatal limitations. Considering g_m, its decline in cotton leaves under drought was mostly regulated by the chloroplast surface area exposed to leaf intercellular air spaces per leaf area (S_c/S) and might also be regulated by the expression of leaf CARBONIC ANHYDRASE (CA1). Meanwhile, cotton leaves can minimize the decrease in g_m under drought by maintaining cell wall thickness (T_cw). Our results indicated that modification of chloroplasts might be a target trait in future attempts to improve cotton drought tolerance.     

Will intra‐specific differences in transpiration efficiency in wheat be maintained in a high CO2 world? A FACE study

This study evaluates whether the target breeding trait of superior leaf level transpiration efficiency is still appropriate under increasing carbon dioxide levels of a future climate using a semi‐arid cropping system as a model. Specifically, we investigated whether physiological traits governing leaf level transpiration efficiency, such as net assimilation rates (A_net), stomatal conductance (g_s) or stomatal sensitivity were affected differently between two Triticum aestivum L. cultivars differing in transpiration efficiency (cv. Drysdale, superior; cv. Hartog, low). Plants were grown under Free Air Carbon dioxide Enrichment (FACE, approximately 550 µmol mol^?1 or ambient CO₂ concentrations (approximately 390 µmol mol^?1). Mean A_net (approximately 15% increase) and g_s (approximately 25% decrease) were less affected by elevated [CO₂] than previously found in FACE‐grown wheat (approximately 25% increase and approximately 32% decrease, respectively), potentially reflecting growth in a dry‐land cropping system. In contrast to previous FACE studies, analyses of the Ball et al. model revealed an elevated [CO₂] effect on the slope of the linear regression by 12% indicating a decrease in stomatal sensitivity to the combination of [CO₂], photosynthesis rate and humidity. Differences between cultivars indicated greater transpiration efficiency for Drysdale with growth under elevated [CO₂] potentially increasing the response of this trait. This knowledge adds valuable information for crop germplasm improvement for future climates.     

Elevated CO2 affects photosynthetic responses in canopy pine and subcanopy deciduous trees over 10 years: a synthesis from Duke FACE

Leaf responses to elevated atmospheric CO₂ concentration (C_a) are central to models of forest CO₂ exchange with the atmosphere and constrain the magnitude of the future carbon sink. Estimating the magnitude of primary productivity enhancement of forests in elevated C_a requires an understanding of how photosynthesis is regulated by diffusional and biochemical components and up‐scaled to entire canopies. To test the sensitivity of leaf photosynthesis and stomatal conductance to elevated C_a in time and space, we compiled a comprehensive dataset measured over 10 years for a temperate pine forest of Pinus taeda, but also including deciduous species, primarily Liquidambar styraciflua. We combined over one thousand controlled‐response curves of photosynthesis as a function of environmental drivers (light, air C_a and temperature) measured at canopy heights up to 20 m over 11 years (1996–2006) to generate parameterizations for leaf‐scale models for the Duke free‐air CO₂ enrichment (FACE) experiment. The enhancement of leaf net photosynthesis (A_net) in P. taeda by elevated C_a of +200 μmol mol^?1 was 67% for current‐year needles in the upper crown in summer conditions over 10 years. Photosynthetic enhancement of P. taeda at the leaf‐scale increased by two‐fold from the driest to wettest growing seasons. Current‐year pine foliage A_net was sensitive to temporal variation, whereas previous‐year foliage A_net was less responsive and overall showed less enhancement (+30%). Photosynthetic downregulation in overwintering upper canopy pine needles was small at average leaf N (N_area), but statistically significant. In contrast, co‐dominant and subcanopy L. styraciflua trees showed A_net enhancement of 62% and no A_net–N_area adjustments. Various understory deciduous tree species showed an average A_net enhancement of 42%. Differences in photosynthetic responses between overwintering pine needles and subcanopy deciduous leaves suggest that increased C_a has the potential to enhance the mixed‐species composition of planted pine stands and, by extension, naturally regenerating pine‐dominated stands.     

Contrasting stomatal sensitivity to temperature and soil drought in mature alpine conifers

Conifers growing at high elevations need to optimize their stomatal conductance (g_s) for maximizing photosynthetic yield while minimizing water loss under less favourable thermal conditions. Yet the ability of high‐elevation conifers to adjust their g_s sensitivity to environmental drivers remains largely unexplored. We used 4 years of sap flow measurements to elucidate intraspecific and interspecific variability of g_s in Larix decidua Mill. and Picea abies (L.) Karst along an elevational gradient and contrasting soil moisture conditions. Site‐ and species‐specific g_s response to main environmental drivers were examined, including vapour pressure deficit, air temperature, solar irradiance, and soil water potential. Our results indicate that maximum g_s of L. decidua is >2 times higher, shows a more plastic response to temperature, and down‐regulates g_s stronger during atmospheric drought compared to P. abies. These differences allow L. decidua to exert more efficient water use, adjust to site‐specific thermal conditions, and reduce water loss during drought episodes. The stronger plasticity of g_s sensitivity to temperature and higher conductance of L. decidua compared to P. abies provide new insights into species‐specific water use strategies, which affect species' performance and should be considered when predicting terrestrial water dynamics under future climatic change.     

Effects of elevated CO2 and temperature on photosynthesis and leaf traits of an understory dwarf bamboo in subalpine forest zone,China

The dwarf bamboo (Fargesia rufa Yi), growing understory in subalpine dark coniferous forest, is one of the main foods for giant panda, and it influences the regeneration of subalpine coniferous forests in southwestern China. To investigate the effects of elevated CO₂, temperature and their combination, the dwarf bamboo plantlets were exposed to two CO₂ regimes (ambient and double ambient CO₂ concentration) and two temperatures (ambient and +2.2°C) in growth chambers. Gas exchange, leaf traits and carbohydrates concentration were measured after the 150‐day experiment. Elevated CO₂ significantly increased the net photosynthetic rate (A_net), intrinsic water‐use efficiency (WUE_i) and carbon isotope composition (δ¹³C) and decreased stomatal conductance (g_s) and total chlorophyll concentration based on mass (Chl_m) and area (Chl_a). On the other hand, elevated CO₂ decreased specific leaf area (SLA), which was increased by elevated temperature. Elevated CO₂ also increased foliar carbon concentration based on mass (C_m) and area (C_a), nitrogen concentration based on area (N_a), carbohydrates concentration (i.e. sucrose, sugar, starch and non‐structural carbohydrates) and the slope of the A_net–N_a relationship. However, elevated temperature decreased C_m, C_a and N_a. The combination of elevated CO₂ and temperature hardly affected SLA, C_m, C_a, N_m, N_a, Chl_m and Chl_a. Variables A_net and N_a had positive linear relationships in all treatments. Our results showed that photosynthetic acclimation did not occur in dwarf bamboo at elevated CO₂ and it could adjust physiology and morphology to enable the capture of more light, to increase WUE and improve nutritional conditions.     

Decoupling between stomatal conductance and photosynthesis occurs under extreme heat in broadleaf tree species regardless of water access

High air temperatures increase atmospheric vapor pressure deficit (VPD) and the severity of drought, threatening forests worldwide. Plants regulate stomata to maximize carbon gain and minimize water loss, resulting in a close coupling between net photosynthesis (A_net) and stomatal conductance (g_s). However, evidence for decoupling of g_s from A_net under extreme heat has been found. Such a response both enhances survival of leaves during heat events but also quickly depletes available water. To understand the prevalence and significance of this decoupling, we measured leaf gas exchange in 26 tree and shrub species growing in the glasshouse or at an urban site in Sydney, Australia on hot days (maximum T_air > 40°C). We hypothesized that on hot days plants with ample water access would exhibit reduced A_net and use transpirational cooling leading to stomatal decoupling, whereas plants with limited water access would rely on other mechanisms to avoid lethal temperatures. Instead, evidence for stomatal decoupling was found regardless of plant water access. Transpiration of well-watered plants was 23% higher than model predictions during heatwaves, which effectively cooled leaves below air temperature. For hotter, droughted plants, the increase in transpiration during heatwaves was even more pronounced—g_s was 77% higher than model predictions. Stomatal decoupling was found for most broadleaf evergreen and broadleaf deciduous species at the urban site, including some wilted trees with limited water access. Decoupling may simply be a passive consequence of the physical effects of high temperature on plant leaves through increased cuticular conductance of water vapor, or stomatal decoupling may be an adaptive response that is actively regulated by stomatal opening under high temperatures. This temperature response is not yet included in any land surface model, suggesting that model predictions of evapotranspiration may be underpredicted at high temperature and high VPD.     

基于FvCB模型的叶片光合生理对环境因子的响应研究进展

为提高叶片光合速率并更好地理解叶片光合生理对环境因子变化的响应机制,FvCB模型(C_3植物光合生化模型)常用于分析不同环境条件下CO_2响应曲线并预测叶片活体内光合系统的内在变化状况。系统介绍了FvCB模型的建立、发展过程和拟合方法等基本理论,综述了该模型在叶片光合生理对光、CO_2、水、温度和N营养等环境因子变化的响应机制中的应用研究。为进一步完善FvCB模型并更好地理解叶片活体内光合系统对环境因子变化的响应机制,未来拟加强以下研究:1)羧化速率与光合电子传递速率之间的联系;2)叶肉导度的具体组分及其对FvCB模型参数估计的影响;3)叶片气孔导度和叶肉导度对环境因子变化的调控机制。     

Late-stage canopy tree species with extremely low δ13C and high stomatal sensitivity to seasonal soil drought in the tropical rainforest of French Guiana

We assessed the daily time‐courses of CO₂ assimilation rate (A), leaf transpiration rate (E), stomatal conductance for water vapour (g_s), leaf water potential ( Ψ _w) and tree transpiration in a wet and a dry season for three late‐stage canopy rainforest tree species in French Guiana differing in leaf carbon isotope composition ( δ ¹³C). The lower sunlit leaf δ ¹³C values found in Virola surinamensis ( ? 29·9‰) and in Diplotropis purpurea ( ? 30·9‰), two light‐demanding species, as compared to Eperua falcata ( ? 28·6‰), a shade‐semi‐tolerant species, were clearly associated with higher maximum g_s values of sunlit leaves in the two former species. These two species were also characterized by a high sensitivity of g_s, sap flow density (Ju) and canopy conductance (g_c) to seasonal soil drought, allowing maintenance of high midday Ψ _w values in the dry season. The data for Diplotropis provided an original picture of increasing midday Ψ _w with increasing soil drought. In Virola, stomata were extremely sensitive to seasonal soil drought, leading to a dramatic decrease in leaf and tree transpiration in the dry season, whereas midday Ψ _w remained close to ? 0·3 MPa. The mechanisms underlying such an extremely high sensitivity of stomata to soil drought remain unknown. In Eperua, g_s of sunlit leaves was non‐responsive to seasonal drought, whereas Ju and g_c were lower in the dry season. This suggests a higher stomatal sensitivity to seasonal drought in shaded leaves than in sunlit ones in this species.     

Seasonal photosynthetic gas exchange and water-use efficiency in a constitutive CAM plant, the giant saguaro cactus (Carnegiea gigantea)

Crassulacean acid metabolism (CAM) and the capacity to store large quantities of water are thought to confer high water use efficiency (WUE) and survival of succulent plants in warm desert environments. Yet the highly variable precipitation, temperature and humidity conditions in these environments likely have unique impacts on underlying processes regulating photosynthetic gas exchange and WUE, limiting our ability to predict growth and survival responses of desert CAM plants to climate change. We monitored net CO₂ assimilation (A _net), stomatal conductance (g _s), and transpiration (E) rates periodically over 2 years in a natural population of the giant columnar cactus Carnegiea gigantea (saguaro) near Tucson, Arizona USA to investigate environmental and physiological controls over carbon gain and water loss in this ecologically important plant. We hypothesized that seasonal changes in daily integrated water use efficiency (WUE_day) in this constitutive CAM species would be driven largely by stomatal regulation of nighttime transpiration and CO₂ uptake responding to shifts in nighttime air temperature and humidity. The lowest WUE_day occurred during time periods with extreme high and low air vapor pressure deficit (D _a). The diurnal with the highest D _a had low WUE_day due to minimal net carbon gain across the 24 h period. Low WUE_day was also observed under conditions of low D _a; however, it was due to significant transpiration losses. Gas exchange measurements on potted saguaro plants exposed to experimental changes in D _a confirmed the relationship between D _a and g _s. Our results suggest that climatic changes involving shifts in air temperature and humidity will have large impacts on the water and carbon economy of the giant saguaro and potentially other succulent CAM plants of warm desert environments.     

On the Use of Leaf Spectral Indices to Assess Water Status and Photosynthetic Limitations in Olea europaea L. during Water-Stress and Recovery

Diffusional limitations to photosynthesis, relative water content (RWC), pigment concentrations and their association with reflectance indices were studied in olive (Olea europaea) saplings subjected to water-stress and re-watering. RWC decreased sharply as drought progressed. Following rewatering, RWC gradually increased to pre-stress values. Photosynthesis (A), stomatal conductance (g_s), mesophyll conductance (g_m), total conductance (g_t), photochemical reflectance index (PRI), water index (WI) and relative depth index (RDI) closely followed RWC. In contrast, carotenoid concentration, the carotenoid to chlorophyll ratio, water content reflectance index (WCRI) and structural independent pigment index (SIPI) showed an opposite trend to that of RWC. Photosynthesis scaled linearly with leaf conductance to CO₂; however, A measured under non-photorespiratory conditions (A_1%O2) was approximately two times greater than A measured at 21% [O₂], indicating that photorespiration likely increased in response to drought. A_1%O2 also significantly correlated with leaf conductance parameters. These relationships were apparent in saturation type curves, indicating that under non-photorespiratory conditions, CO₂ conductance was not the major limitations to A. PRI was significant correlated with RWC. PRI was also very sensitive to pigment concentrations and photosynthesis, and significantly tracked all CO₂ conductance parameters. WI, RDI and WCRI were all significantly correlated with RWC, and most notably to leaf transpiration. Overall, PRI correlated more closely with carotenoid concentration than SIPI; whereas WI tracked leaf transpiration more effectively than RDI and WCRI. This study clearly demonstrates that PRI and WI can be used for the fast detection of physiological traits of olive trees subjected to water-stress.     

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.     

Effects of elevated CO2, drought and temperature on the water relations and gas exchange of groundnut (Arachis hypogaea) stands grown in controlled environment glasshouses

Stands of groundnut (Arachis hypogaea L. cv. Kadiri‐3) were grown in controlled environment glasshouses at mean atmospheric CO₂ concentrations of 375 or 700 μmol mol^?1 and daily mean air temperatures of 28 or 32°C on irrigated or drying soil profiles. Leaf water (Ψ_l) and solute potential (Ψ_s), relative water content (RWC), stomatal conductance (g_l) and net photosynthesis (P_n) were measured at midday for the youngest mature leaf throughout the growing season. Elevated CO₂ and temperature had no detectable effect on the water relations of irrigated plants, but higher values of RWC, Ψ_l and Ψ_s were maintained for longer under elevated CO₂ during progressive drought. Turgor potential (Ψ_p) reached zero when Ψ_l declined to ?1.6 to ?1.8 MPa in all treatments; turgor was lost sooner when droughted plants were grown under ambient CO₂. A 4°C increase in mean air temperature had no effect on Ψ_s in droughted plants, but elicited a small increase in Ψ_l; midday g_l values were lower under elevated than under ambient CO₂, and Ψ_l and g_l declined below ?1.5 MPa and 0.25 cm s^?1, respectively, as the soil dried. Despite the low g_l values recorded for droughted plants late in the season, P_n was maintained under elevated CO₂, but declined to zero 3 weeks before final harvest under ambient CO₂. Concurrent reductions in g_l and increases in water use efficiency under elevated CO₂ prolonged photosynthetic activity during drought and increased pod yields relative to plants grown under ambient CO₂. The implications of future increases in atmospheric CO₂ for the productivity of indeterminate C₃ crops grown in rainfed subsistence agricultural systems in the semi‐arid tropics are discussed.     

Impacts of leafroll‐associated viruses (GLRaV‐1 and ‐3) on the physiology of the Portuguese grapevine cultivar ‘Touriga Nacional’ growing under field conditions

The impact of mixed infection of grapevine leafroll‐associated virus 1 and 3 (GLRaV‐1&‐3) on physiological performance of the Portuguese grapevine variety ‘Touriga Nacional’ was evaluated during 3 years with the main purpose of understanding the drastic reduction in yield. Overall, gas exchange was negatively affected in leaves with these leafroll virus infections. Particularly at ripeness stage, the reduction in stomatal conductance (g_s) was higher than in net CO₂ assimilation rate (A), leading to higher intrinsic water use efficiency (A/g_s) in infected leaves. However, the decrease in g_s and A were not a consequence of the decrease in bulk water potential, as the water index/normalised difference vegetation index ratio suggested similar magnitude for both treatments. The maximum quantum efficiency of photosystem II was unaffected by GLRaV‐1&‐3, whereas quantum effective efficiency of PSII, apparent electron transport rate and photochemical quenching significantly decreased in infected leaves and these was paralleled by a significant increase of non‐photochemical quenching. Relative to carbon metabolism, the analyses of the net CO₂ assimilation rate/photosynthetic photon flux density (A/PPFD) and net CO₂ assimilation rate/internal CO₂ concentration (A/C_i) curves revealed that virus infection had a negative effect on light saturated rate of CO₂ fixation at high irradiances and carboxylation efficiency but, in contrast, apparent quantum yield of CO₂ fixation was significantly higher. Meanwhile, the presence of GLRaV‐1&‐3 resulted in a marked decrease in photosynthetic pigments, soluble sugars and soluble proteins contents, while starch and anthocyanins were significantly improved. N, P, Ca, S and Fe leaf concentrations significantly decreased, while K, Mg, B, Cu, Zn and Mn were unaffected by these two leafroll virus species. Infected plants showed a significant decrease in yield, mainly due to a lower cluster weight. These results emphasised the important role of GLRaV‐1&‐3 as a biotic stress for the grapevine physiology and consequently to yield attributes.