Hourly and seasonal variation in photosynthesis and stomatal conductance of soybean grown at future CO(2) and ozone concentrations for 3 years under fully open-air field conditions

It is anticipated that enrichment of the atmosphere with CO(2) will increase photosynthetic carbon assimilation in C3 plants. Analysis of controlled environment studies conducted to date indicates that plant growth at concentrations of carbon dioxide ([CO(2)]) anticipated for 2050 ( approximately 550 micromol mol(-1)) will stimulate leaf photosynthetic carbon assimilation (A) by 20 to 40%. Simultaneously, concentrations of tropospheric ozone ([O(3)]) are expected to increase by 2050, and growth in controlled environments at elevated [O(3)] significantly reduces A. However, the simultaneous effects of both increases on a major crop under open-air conditions have never been tested. Over three consecutive growing seasons > 4700 individual measurements of A, photosynthetic electron transport (J(PSII)) and stomatal conductance (g(s)) were measured on Glycine max (L.) Merr. (soybean). Experimental treatments used free-air gas concentration enrichment (FACE) technology in a fully replicated, factorial complete block design. The mean A in the control plots was 14.5 micromol m(-2) s(-1). At elevated [CO(2)], mean A was 24% higher and the treatment effect was statistically significant on 80% of days. There was a strong positive correlation between daytime maximum temperatures and mean daily integrated A at elevated [CO(2)], which accounted for much of the variation in CO(2) effect among days. The effect of elevated [CO(2)] on photosynthesis also tended to be greater under water stress conditions. The elevated [O(3)] treatment had no statistically significant effect on mean A, g(s) or J(PSII) on newly expanded leaves. Combined elevation of [CO(2)] and [O(3)] resulted in a slightly smaller increase in average A than when [CO(2)] alone was elevated, and was significantly greater than the control on 67% of days. Thus, the change in atmospheric composition predicted for the middle of this century will, based on the results of a 3 year open-air field experiment, have smaller effects on photosynthesis, g(s) and whole chain electron transport through photosystem II than predicted by the substantial literature on relevant controlled environment studies on soybean and likely most other C3 plants.     

To what extent do current and projected increases in surface ozone affect photosynthesis and stomatal conductance of trees? A meta‐analytic review of the last 3 decades of experiments

The surface concentration of ozone ([O(3)]) has risen from less than 10 ppb prior to the industrial revolution to a day-time mean concentration of approximately 40 ppb over much of the northern temperate zone. If current global emission trends continue, surface [O(3)] is projected to rise a further 50% over this century, with larger increases in many locations including Northern Hemisphere forests. This review uses statistical meta-analysis to determine mean effects, and their confidence limits, of both the current and projected elevations of [O(3)] on light-saturated photosynthetic CO(2) uptake (A(sat)) and stomatal conductance (g(s)) in trees. In total, 348 measurements of A(sat) from 61 studies and 266 measures of g(s) from 55 studies were reviewed. Results suggested that the elevation of [O(3)] that has occurred since the industrial revolution is depressing A(sat) and g(s) by 11% (CI 9-13%) and 13% (CI 11-15%), respectively, where CI is the 95% confidence interval. In contrast to angiosperms, gymnosperms were not significantly affected. Both drought and elevated [CO(2)] significantly decreased the effect of ambient [O(3)]. Younger trees (<4 years) were affected less than older trees. Elevation of [O(3)] above current levels caused progressively larger losses of A(sat) and g(s), including gymnosperms. Results are consistent with the expectation that damage to photosynthesis depends on the cumulative uptake of ozone (O(3)) into the leaf. Thus, factors that lower g(s) lessen damage. Where both g(s) and [O(3)] were recorded, an overall decline in A(sat) of 0.21% per mmol m(-2) of estimated cumulative O(3) uptake was calculated. These findings suggest that rising [O(3)], an often overlooked aspect of global atmospheric change, is progressively depressing the ability of temperate and boreal forests to assimilate carbon and transfer water vapour to the atmosphere, with significant potential effects on terrestrial carbon sinks and regional hydrologies.     

Chronic ozone exposure affects leaf senescence of adult beech trees: a chlorophyll fluorescence approach

Accelerated leaf senescence is one of the harmful effects of elevated tropospheric ozone concentrations ([O(3)]) on plants. The number of studies dealing with mature forest trees is scarce however. Therefore, five 66-year-old beech trees (Fagus sylvatica L.) have been exposed to twice-ambient (2xambient) [O(3)] levels by means of a free-air canopy O(3) exposure system. During the sixth year of exposure, the hypothesis of accelerated leaf senescence in 2xambient [O(3)] compared with ambient [O(3)] trees was tested for both sun and shade leaves. Chlorophyll (chl) fluorescence was used to assess the photosynthetic quantum yield, and chl fluorescence images were processed to compare functional leaf homogeneity and the proportion of O(3)-injured leaf area (stipples) under ambient and 2xambient [O(3)] regimes. Based on the analysis of chl fluorescence images, sun leaves of both ambient and 2xambient [O(3)] trees had apparently developed typical necrotic O(3) stipples during high O(3) episodes in summer, while accelerated senescence was only observed with sun leaves of 2xambient [O(3)] trees. This latter effect was indicated along with a faster decrease of photosynthetic quantum yield, but without evidence of changes in non-photochemical quenching. Overall, treatment effects were small and varied among trees. Therefore, compared with ambient [O(3)], the consequence of the observed O(3)-induced accelerated leaf senescence for the carbon budget is likely limited.     

The effects of O3 fumigation during leaf development on photosynthesis of wheat and pea: An in vivo analysis

Previous studies have shown that short exposure of plants to high doses of ozone decreases subsequent photosynthesis; initially by reducing carboxylation capacity. This study tests the hypothesis that this is also the primary cause of loss of photosynthetic capacity in leaves affected by development under a low level of ozone. Triticum aestivum and Pisum sativum plants were exposed from germination to ozone in air (80 nmol mol^-1 for 7 hours per day, for 18 days. Leaves that had completed lamina expansion at this time were free of visible injury and light absorptance was unaffected. However, some significant changes in photosynthetic gas exchange were evident. Photosynthetic CO2 uptake at light saturation was decreased significantly by 35% in T. aestivum but was unchanged in P. sativum. The reduction in photosynthesis of T. aestivum was accompanied by a 31% decline in the maximum velocity of carboxylation measured in vivo. Decreased stomatal conductance did not contribute to this reduction of photosynthesis because there was no significant change in the stomatal limitation to CO2. Processes directly dependent upon photochemical reactions; that is, the quantum yield of CO2 uptake and capacity for regeneration of ribulose 1,5-bisphosphate were not affected by O3 fumigation in either species. This suggests that for wheat, the quantitative cause of decreased photosynthetic rate in vivo is a decrease in the quantity of active ribulose-1,5- bisphosphate carboxylase-oxygenase.     

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

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

Relationships between photosynthetic activity and silica accumulation with ages of leaf in Sasa veitchii (Poaceae, Bambusoideae)

BACKGROUND AND AIMS: Bamboos have long-lived, evergreen leaves that continue to accumulate silica throughout their life. Silica accumulation has been suggested to suppress their photosynthetic activity. However, nitrogen content per unit leaf area (N(area)), an important determinant of maximum photosynthetic capacity per unit leaf area (P(max)), decreases as leaves age and senescence. In many species, P(max) decreases in parallel with the leaf nitrogen content. It is hypothesized that if silica accumulation affects photosynthesis, then P(max) would decrease faster than N(area), leading to a decrease in photosynthetic rate per unit leaf nitrogen (photosynthetic nitrogen use efficiency, PNUE) with increasing silica content in leaves. METHODS: The hypothesis was tested in leaves of Sasa veitchii, which have a life span of 2 years and accumulate silica up to 41 % of dry mass. Seasonal changes in P(max), stomatal conductance, N(area) and silica content were measured for leaves of different ages. KEY RESULTS: Although P(max) and PNUE were negatively related with silica content across leaves of different ages, the relationship between PNUE and silica differed depending on leaf age. In second-year leaves, PNUE was almost constant although there was a large increase in silica content, suggesting that leaf nitrogen was a primary factor determining the variation in P(max) and that silica accumulation did not affect photosynthesis. PNUE was strongly and negatively correlated with silica content in third-year leaves, suggesting that silica accumulation affected photosynthesis of older leaves. CONCLUSIONS: Silica accumulation in long-lived leaves of bamboo did not affect photosynthesis when the silica concentration of a leaf was less than 25 % of dry mass. Silica may be actively transported to epidermal cells rather than chlorenchyma cells, avoiding inhibition of CO2 diffusion from the intercellular space to chloroplasts. However, in older leaves with a larger silica content, silica was also deposited in chlorenchyma cells, which may relate to the decrease in PNUE.     

Senescence and hyperspectral reflectance of cotton leaves exposed to ultraviolet-B radiation and carbon dioxide

The objectives of this study were to determine the effects of UV-B radiation and atmospheric carbon dioxide concentrations ([CO(2)]) on leaf senescence of cotton by measuring leaf photosynthesis and chlorophyll content and to identify changes in leaf hyperspectral reflectance occurring due to senescence and UV-B radiation. Plants were grown in controlled-environment growth chambers at two [CO(2)] (360 and 720 micro mol mol(-1)) and three levels of UV-B radiation (0, 7.7 and 15.1 kJ m(-2) day(-1)). Photosynthesis, chlorophyll, carotenoids and phenolic compounds along with leaf hyperspectral reflectance were measured on three leaves aged 12, 21 and 30 days in each of the treatments. No interaction was detected between [CO(2)] and UV-B for any of the measured parameters. Significant interactions were observed between UV-B and leaf age for photosynthesis and stomatal conductance. Elevated [CO(2)] enhanced leaf photosynthesis by 32%. On exposure to 0, 7.7 and 15.1 kJ of UV-B, the photosynthetic rates of 30-day-old leaves compared with 12-day-old leaves were reduced by 52, 76 and 86%, respectively. Chlorophyll pigments were not affected by leaf age at UV-B radiation of 0 and 7.7 kJ, but UV-B of 15.1 kJ reduced the chlorophylls by 20, 60 and 80% in 12, 21 and 30-day-old leaves, respectively. The hyperspectral reflectance between 726 and 1142 nm showed interaction for UV-B radiation and leaf age. In cotton, leaf photosynthesis can be used as an indicator of leaf senescence, as it is more sensitive than photosynthetic pigments on exposure to UV-B radiation. This study revealed that, cotton leaves senesced early on exposure to UV-B radiation as indicated by leaf photosynthesis, and leaf hyperspectral reflectance can be used to detect changes caused by UV-B and leaf ageing.     

Rising ozone concentrations decrease soybean evapotranspiration and water use efficiency whilst increasing canopy temperature

? Here, we investigated the effects of increasing concentrations of ozone ([O(3)]) on soybean canopy-scale fluxes of heat and water vapor, as well as water use efficiency (WUE), at the Soybean Free Air Concentration Enrichment (SoyFACE) facility. ? Micrometeorological measurements were made to determine the net radiation (R(n)), sensible heat flux (H), soil heat flux (G(0)) and latent heat flux (λET) of a commercial soybean (Glycine max) cultivar (Pioneer 93B15), exposed to a gradient of eight daytime average ozone concentrations ranging from approximately current (c. 40 ppb) to three times current (c. 120 ppb) levels. ? As [O(3)] increased, soybean canopy fluxes of λET decreased and H increased, whereas R(n) and G(0) were not altered significantly. Exposure to increased [O(3)] also resulted in warmer canopies, especially during the day. The lower λET decreased season total evapotranspiration (ET) by c. 26%. The [O(3)]-induced relative decline in ET was half that of the relative decline in seed yield, driving a 50% reduction in seasonal WUE. ? These results suggest that rising [O(3)] will alter the canopy energy fluxes that drive regional climate and hydrology, and have a negative impact on productivity and WUE, key ecosystem services.     

Studies on the Characteristics of Chlorophyll Fluorescence of Winter Wheat Flag Leaves at Different Developing Stages

The parameters of fluorescence induction kinetics and the maximal light-saturated net CO2 assimilation rate ( P sat ) of the flag leaves of four cultivars of winter wheat （Triticum aestivum L.） were compared at three different developing stages for the first time. From the blooming stage to the milky stage, the quantum efficiency of PSⅡ photochemistry ( Fv/Fm ) declined slightly only at the milk stage. The photochemical quenching co-efficient ( qP ), actual quantum yield of photosystem Ⅱ（PSⅡ）electron transport ( Φ PSⅡ) and P sat decreased substantially (>15%), while the non-photochemical quenching co-efficient ( qN ) increased significantly (>100%). There existed a linear correlation between the Φ PSⅡ and the P sat ( r =0.918). The results indicate that with the senescence of the flag leaves of winter wheat the photosynthetic efficiency including that of the energy transport and the CO2 assimilation significantly decreased.     

Elevated ozone reduces photosynthetic carbon gain by accelerating leaf senescence of inbred and hybrid maize in a genotype‐specific manner

Exposure to elevated tropospheric ozone concentration ([O₃]) accelerates leaf senescence in many C₃ crops. However, the effects of elevated [O₃] on C₄ crops including maize (Zea mays L.) are poorly understood in terms of physiological mechanism and genetic variation in sensitivity. Using free air gas concentration enrichment, we investigated the photosynthetic response of 18 diverse maize inbred and hybrid lines to season‐long exposure to elevated [O₃] (~100 nl L^?1) in the field. Gas exchange was measured on the leaf subtending the ear throughout the grain filling period. On average over the lifetime of the leaf, elevated [O₃] led to reductions in photosynthetic CO₂ assimilation of both inbred (?22%) and hybrid (?33%) genotypes. There was significant variation among both inbred and hybrid lines in the sensitivity of photosynthesis to elevated [O₃], with some lines showing no change in photosynthesis at elevated [O₃]. Based on analysis of inbred line B73, the reduced CO₂ assimilation at elevated [O₃] was associated with accelerated senescence decreasing photosynthetic capacity and not altered stomatal limitation. These findings across diverse maize genotypes could advance the development of more O₃ tolerant maize and provide experimental data for parameterization and validation of studies modeling how O₃ impacts crop performance.     

A role for brassinosteroids in the regulation of photosynthesis in Cucumis sativus

The effects of 24-epibrassinolide (EBR) spray application on gas-exchange, chlorophyll fluorescence characteristics, Rubisco activity, and carbohydrate metabolism were investigated in cucumber (Cucumis sativus L. cv. Jinchun No. 3) plants grown in a greenhouse. EBR significantly increased the light-saturated net CO(2) assimilation rate (A(sat)) from 3 h to 7d after spraying, with 0.1 mg l(-1) EBR proving most effective. Increased A(sat) in EBR-treated leaves was accompanied by increases in the maximum carboxylation rate of Rubisco (V(c,max)) and in the maximum rate of RuBP regeneration (J(max)). EBR-treated leaves also had a higher quantum yield of PSII electron transport (phi(PSII)) than the controls, which was mainly due to a significant increase in the photochemical quenching (q(P)), with no change in the efficiency of energy capture by open PSII reaction centres (F'(v)/F'(m)). EBR did not influence photorespiration. In addition, significant increases in the initial activity of Rubisco and in the sucrose, soluble sugars, and starch contents were observed followed by substantial increases in sucrose phosphate synthase (SPS), sucrose synthase (SS), and acid invertase (AI) activities after EBR treatment. It was concluded that EBR increases the capacity of CO(2) assimilation in the Calvin cycle, which was mainly attributed to an increase in the initial activity of Rubisco.     

Effect of solar ultraviolet-B radiation during springtime ozone depletion on photosynthesis and biomass production of Antarctic vascular plants

We assessed the influence of springtime solar UV-B radiation that was naturally enhanced during several days due to ozone depletion on biomass production and photosynthesis of vascular plants along the Antarctic Peninsula. Naturally growing plants of Colobanthus quitensis (Kunth) Bartl. and Deschampsia antarctica Desv. were potted and grown under filters that absorbed or transmitted most solar UV-B. Plants exposed to solar UV-B from mid-October to early January produced 11% to 22% less total, as well as above ground biomass, and 24% to 31% less total leaf area. These growth reductions did not appear to be associated with reductions in photosynthesis per se: Although rates of photosynthetic O(2) evolution were reduced on a chlorophyll and a dry-mass basis, on a leaf area basis they were not affected by UV-B exposure. Leaves on plants exposed to UV-B were denser, probably thicker, and had higher concentrations of photosynthetic and UV-B absorbing pigments. We suspect that the development of thicker leaves containing more photosynthetic and screening pigments allowed these plants to maintain their photosynthetic rates per unit leaf area. Exposure to UV-B led to reductions in quantum yield of photosystem II, based on fluorescence measurements of adaxial leaf surfaces, and we suspect that UV-B impaired photosynthesis in the upper mesophyll of leaves. Because the ratio of variable to maximal fluorescence, as well as the initial slope of the photosynthetic light response, were unaffected by UV-B exposure, we suggest that impairments in photosynthesis in the upper mesophyll were associated with light-independent enzymatic, rather than photosystem II, limitations.     

Maintenance of C sinks sustains enhanced C assimilation during long-term exposure to elevated [CO2] in Mojave Desert shrubs

During the first few years of elevated atmospheric [CO(2)] treatment at the Nevada Desert FACE Facility, photosynthetic downregulation was observed in desert shrubs grown under elevated [CO(2)], especially under relatively wet environmental conditions. Nonetheless, those plants maintained increased A (sat) (photosynthetic performance at saturating light and treatment [CO(2)]) under wet conditions, but to a much lesser extent under dry conditions. To determine if plants continued to downregulate during long-term exposure to elevated [CO(2)], responses of photosynthesis to elevated [CO(2)] were examined in two dominant Mojave Desert shrubs, the evergreen Larrea tridentata and the drought-deciduous Ambrosia dumosa, during the eighth full growing season of elevated [CO(2)] treatment at the NDFF. A comprehensive suite of physiological processes were collected. Furthermore, we used C labeling of air to assess carbon allocation and partitioning as measures of C sink activity. Results show that elevated [CO(2)] enhanced photosynthetic performance and plant water status in Larrea, especially during periods of environmental stress, but not in Ambrosia. δ(13)C analyses indicate that Larrea under elevated [CO(2)] allocated a greater proportion of newly assimilated C to C sinks than Ambrosia. Maintenance by Larrea of C sinks during the dry season partially explained the reduced [CO(2)] effect on leaf carbohydrate content during summer, which in turn lessened carbohydrate build-up and feedback inhibition of photosynthesis. δ(13)C results also showed that in a year when plant growth reached the highest rates in 5 years, 4% (Larrea) and 7% (Ambrosia) of C in newly emerging organs were remobilized from C that was assimilated and stored for at least 2 years prior to the current study. Thus, after 8 years of continuous exposure to elevated [CO(2)], both desert perennials maintained their photosynthetic capacities under elevated [CO(2)]. We conclude that C storage, remobilization, and partitioning influence the responsiveness of these desert shrubs during long-term exposure to elevated [CO(2)].     

Effects of drought on photosynthesis in Mediterranean plants grown under enhanced UV-B radiation

The effects of drought on the photosynthetic characteristics of three Mediterranean plants (olive, Olea europea L.; rosemary, Rosmarinus officinalis L.; lavender, Lavandula stoechas L.) exposed to elevated UV-B irradiation in a glasshouse were investigated over a period of weeks. Drought conditions were imposed on 2-year-old plants by withholding water. During the onset of water stress, analyses of the response of net carbon assimilation of leaves to their intercellular CO2 concentration were used to examine the potential limitations imposed by stomata, carboxylation velocity and capacity for regeneration of ribulose 1,5-bisphosphate on photosynthesis. Measurements of chlorophyll fluorescence were used to determine changes in the efficiency of light utilization for electron transport, the occurrence of photoinhibition of photosystem II photochemistry and the possibility of stomatal patchiness across leaves. The first stages of water stress produced decreases in the light-saturated rate of CO2 assimilation which were accompanied by decreases in the maximum carboxylation velocity and the capacity for regeneration of ribulose 1,5-bisphosphate in the absence of any significant photodamage to photosystem II. Leaves of rosemary and lavender were more sensitive than those of olive during the first stages of the drought treatment and also exhibited increases in stomatal limitation. With increasing water stress, significant decreases in the maximum quantum efficiency of photosystem II photochemistry occurred in lavender and rosemary, and stomatal limitation was increased in olive. No indication of any heterogeneity of photosynthesis was found in any leaves. Drought treatment significantly decreased leaf area in all species, an important factor in drought-induced decreases in photosynthetic productivity. Exposure of plants to elevated UV-B radiation (0.47 W m(-2)) prior to and during the drought treatment had no significant effects on the growth or photosynthetic activities of the plants. Consequently, it is predicted that increasing UV-B due to future stratospheric ozone depletion is unlikely to have any significant impact on the photosynthetic productivity of olive, lavender and rosemary in the field.     

Response of photosynthesis and respiration of resurrection plants to desiccation and rehydration

Using non-invasive techniques (CO₂ gas exchange, light scattering, light absorption, chlorophyll fluorescence, chlorophyll luminescence), we have analysed the response of respiration and photosynthesis to dehydration and rehydration of leaves of the resurrection plants Craterostigma plantagineum Hochst., Ramonda mykoni Reichb. and Ceterach officinarum Lam. et DC. and of the drought-sensitive mesophyte spinach (Spinacia oleracea L.). The following observations were made: (i) The rate of water loss during wilting of detached leaves of drought-tolerant resurrection plants was similar to that for leaves of the sensitive mesophyte, spinach. Leaves of Mediterranean xerophytes lost water much more slowly. (ii) Below a residual water content of about 20%, leaves of spinach did not recover turgor on rewatering, whereas leaves of the resurrection plants did. (iii) Respiration was less sensitive to the loss of water during wilting in the resurrection plants than in spinach. (iv) The sensitivity of photosynthesis to dehydration was similar in spinach and the resurrection plants. Up to a water loss of 50% from the leaves, photosynthesis was limited by stomatal closure, not by inhibition of reactions of the photosynthetic apparatus. Photosynthesis was inhibited and stomates reopened when loss of water became excessive. (v) After the leaves had lost 80% of their water or more, the light-dependent reactions of photosynthetic membranes were further inhibited by rewatering in spinach; they recovered in the resurrection plants. (vi) In desiccated leaves of the resurrection plants, slow rehydration reactivated mitochondrial gas exchange faster than photosynthetic membrane reactions. Photosynthetic carbon assimilation recovered only slowly.     

Effects of high light and ozone fumigation on photosynthesis in Phaseolus vulgaris

In the present work, the aim was to study the combined effects of high irradiance (about 1 000 μmol·m^–2·s^–1) on the effects of O₃ (150 nL·L^–1 for 2 h) on the photosynthetic process of primary pinto (Phaseolus vulgaris L.) leaves. The CO₂ assimilation rate significantly decreased in high light and/or O₃-treated leaves, as did the stomatal conductance. The present work shows that changes occur in photosynthesis-related parameters due to light stress. Photoinhibition was detected as a decrease in the F_v/F_m ratio. This indicates that the light treatment considerably exceeded the photoprotective capacity and resulted in significant photon damage. Moreover O₃ fumigation alone determined an impairment of the photosynthetic process, that in this case is related to the dark reactions of photosynthesis as also showed by the strong increase in (1-q_P). The data presented in this paper illustrate the complexity of photosynthetic response to high light intensities and ozone, which are two stress factors that often occur simultaneously under natural conditions. High light can modify the reaction of leaves to ozone treatment by reducing the chloroplastic electron transport rate and the quantum yield for PSII photochemistry. Thus, high light seems to enhance the detrimental effects of ozone on photosynthesis.     

Interactive effects of elevated CO2, warming, and drought on photosynthesis of Deschampsia flexuosa in a temperate heath ecosystem

Global change factors affect plant carbon uptake in concert. In order to investigate the response directions and potential interactive effects, and to understand the underlying mechanisms, multifactor experiments are needed. The focus of this study was on the photosynthetic response to elevated CO(2) [CO2; free air CO(2) enrichment (FACE)], drought (D; water-excluding curtains), and night-time warming (T; infrared-reflective curtains) in a temperate heath. A/C(i) curves were measured, allowing analysis of light-saturated net photosynthesis (P(n)), light- and CO(2)-saturated net photosynthesis (P(max)), stomatal conductance (g(s)), the maximal rate of Rubisco carboxylation (V(cmax)), and the maximal rate of ribulose bisphosphate (RuBP) regeneration (J(max)) along with leaf δ(13)C, and carbon and nitrogen concentration on a monthly basis in the grass Deschampsia flexuosa. Seasonal drought reduced P(n) via g(s), but severe (experimental) drought decreased P(n) via a reduction in photosynthetic capacity (P(max), J(max), and V(cmax)). The effects were completely reversed by rewetting and stimulated P(n) via photosynthetic capacity stimulation. Warming increased early and late season P(n) via higher P(max) and J(max). Elevated CO(2) did not decrease g(s), but stimulated P(n) via increased C(i). The T×CO2 synergistically increased plant carbon uptake via photosynthetic capacity up-regulation in early season and by better access to water after rewetting. The effects of the combination of drought and elevated CO(2) depended on soil water availability, with additive effects when the soil water content was low and D×CO2 synergistic stimulation of P(n) after rewetting. The photosynthetic responses appeared to be highly influenced by growth pattern. The grass has opportunistic water consumption, and a biphasic growth pattern allowing for leaf dieback at low soil water availability followed by rapid re-growth of active leaves when rewetted and possibly a large resource allocation capability mediated by the rhizome. This growth characteristic allowed for the photosynthetic capacity up-regulations that mediated the T×CO2 and D×CO2 synergistic effects on photosynthesis. These are clearly advantageous characteristics when exposed to climate changes. In conclusion, after 1 year of experimentation, the limitations by low soil water availability and stimulation in early and late season by warming clearly structure and interact with the photosynthetic response to elevated CO(2) in this grassland species.