Drought responses in Aleppo pine seedlings from two wild provenances with different climatic features

Global warming will likely exacerbate the negative effects of limited water availability in the Mediterranean area. The Italian Aleppo pine (Pinus halepensis Mill.) provenances are distributed along the coasts except Otricoli provenance growing in an unusual location between 300 and 1,000 m a. s. l., in Umbria (central Italy). The aim of the present study was to investigate the photosynthetic response to a 28-day-long drought and to a subsequent reestablishment of water availability in Otricoli and North Euboea (Greece) provenances, representing different locations along a rainfall gradient in the natural range of this species. Six-month-old seedlings were used in this experiment since at this age Aleppo pine plants in Mediterranean climate face their first water stress potentially affecting plant survival. Water potential (ψ_w), net photosynthesis and stomatal conductance decreased during drought in both provenances and showed minimal values 28 days after beginning the treatment (DAT). Otricoli seedlings adjusted ψ_w gradually as the stress level increased and 21 DAT showed a lower ψ_w than North Euboea. In contrast, in North Euboea seedlings ψ_w that was not affected until 21 DAT rapidly dropped to a minimum of −3.81 MPa 28 DAT. At the onset of the stress the intercellular CO₂ concentration (C _i) was reduced, and the “instrinsic” water-use efficiency (WUE_i) was enhanced in both provenances, as stomatal conductance decreased more rapidly than photosynthesis. However, 28 DAT, C _i increased and WUEi decreased as stomatal conductance and photosynthesis declined to minimum levels, revealing nonstomatal limitations of photosynthesis. A rapid decrease in PSII maximal photochemical efficiency estimated by chlorophyll (Chl) fluorescence (F_v/F_m) was also observed when the stress became severe. At the final stage of water stress, North Euboea seedlings maintained significantly higher values of F_v/F_m than Otricoli seedlings. Upon rewatering, photosynthesis did not fully recover in Otricoli seedlings (41 DAT), while all other parameters recovered to control levels in both provenances. No drought-induced physiological differences were consistent with the regional climatic features of these two provenances. Our results suggest that phenotypic plasticity in drought response may help Otricoli provenance cope with global warming, but that recurrent drought episode may slow down the primary productivity of this provenance.     

Anatomical adjustments of the tree hydraulic pathway decrease canopy conductance under long-term elevated CO2

The cause of reduced leaf-level transpiration under elevated CO₂ remains largely elusive. Here, we assessed stomatal, hydraulic, and morphological adjustments in a long-term experiment on Aleppo pine (Pinus halepensis) seedlings germinated and grown for 22–40 months under elevated (eCO₂; c. 860 ppm) or ambient (aCO₂; c. 410 ppm) CO₂. We assessed if eCO₂-triggered reductions in canopy conductance (g_c) alter the response to soil or atmospheric drought and are reversible or lasting due to anatomical adjustments by exposing eCO₂ seedlings to decreasing [CO₂]. To quantify underlying mechanisms, we analyzed leaf abscisic acid (ABA) level, stomatal and leaf morphology, xylem structure, hydraulic efficiency, and hydraulic safety. Effects of eCO₂ manifested in a strong reduction in leaf-level g_c (−55%) not caused by ABA and not reversible under low CO₂ (c. 200 ppm). Stomatal development and size were unchanged, while stomatal density increased (+18%). An increased vein-to-epidermis distance (+65%) suggested a larger leaf resistance to water flow. This was supported by anatomical adjustments of branch xylem having smaller conduits (−8%) and lower conduit lumen fraction (−11%), which resulted in a lower specific conductivity (−19%) and leaf-specific conductivity (−34%). These adaptations to CO₂ did not change stomatal sensitivity to soil or atmospheric drought, consistent with similar xylem safety thresholds. In summary, we found reductions of g_c under elevated CO₂ to be reflected in anatomical adjustments and decreases in hydraulic conductivity. As these water savings were largely annulled by increases in leaf biomass, we do not expect alleviation of drought stress in a high CO₂ atmosphere.

Increases in atmospheric CO₂ can trigger structural modifications in leaves and branches, which contribute to lower leaf-level water loss, but do not affect the drought sensitivity in Aleppo pine.     

Daily and seasonal CO2 exchange in Scots pine grown under elevated O3 and CO2: experiment and simulation

Starting in early spring of 1994, naturally regenerated, 30-year-old Scots pine (Pinus sylvestris L.) trees were grown in open-top chambers and exposed in situ to doubled ambient O₃,doubled ambient CO₂ and a combination of O₃ and CO₂ from 15 April to 15 September. To investigate daily and seasonal responses of CO₂ exchange to elevated O₃ and CO₂, the CO₂ exchange of shoots was measured continuously by an automatic system for measuring gas exchange during the course of one year (from 1 Januray to 31 December 1996). A process-based model of shoot photosynthesis was constructed to quantify modifications in the intrinsic capacity of photosynthesis and stomatal conductance by simulating the daily CO₂ exchange data from the field. Results showed that on most days of the year the model simulated well the daily course of shoot photosynthesis. Elevated O₃ significantly decreased photosynthetic capacity and stomatal conductance during the whole photosynthetic period. Elevated O₃ also led to a delay in onset of photosynthetic recovery in early spring and an increase in the sensitivity of photosynthesis to environmental stress conditions. The combination of elevated O₃ and CO₂ had an effect on photosynthesis and stomatal conductance similar to that of elevated O₃ alone, but significantly reduced the O₃-induced depression of photosynthesis. Elevated CO₂ significantly increased the photosynthetic capacity of Scots pine during the main growing season but slightly decreased it in early spring and late autumn. The model calculation showed that, compared to the control treatment, elevated O₃ alone and the combination of elevated O₃ and CO₂ decreased the annual total of net photosynthesis per unit leaf area by 55% and 38%, respectively. Elevated CO₂ increased the annual total of net photosynthesis by 13%.     

A review of ozone responses in Scots pine (Pinus sylvestris)

The data on Scots pine responses to elevated ozone (O₃) mainly come from experimental studies with young seedlings and trees. Based on the 38 experiments reviewed here, Scots pine may be considered as an O₃-sensitive conifer species, with mature pines more sensitive than younger trees. This is due to their relatively small proportion of current (c) year needles with the highest photosynthetic capacity. Moreover, young seedlings and trees seem to acclimate to slightly elevated realistic O₃ exposures, and hence do not often exhibit growth and biomass reductions in spite of the visible and microscopic needle injuries and changes in needle chemistry. The O₃ sensitivity in Scots pine is thought to relate to impaired water status due to the malfunction of stomata and subsequent increase in transpiration. This may lead to reduced wood biomass in the long term, if Scots pines try to maximise the biomass of c needles and root biomass to maintain efficient water and nitrogen (N) supply to support the photosynthesis of c needles. Tree water status also contributes to the spring-time recovery of photosynthesis. We call especially for studies on atmosphere–needle surface interaction that would yield novel information on the impact of O₃ on epicuticular waxes and stomatal functioning, which both regulate O₃ flux and tree water status and hence also modify photosynthesis. The need for flux-based field studies is especially important in the light of future climatic change, since the risk presented by O₃ to Scots pine forests in Northern and Central Europe seems to be equal.     

Interactive effects of ozone and drought stress on pigments and activities of antioxidative enzymes in Pinus halepensis

The aim of this study was to investigate the interactive effects of ozone (O₃) and drought on pigments and antioxidant enzymes of Aleppo pine (Pinus halepensis). Two‐year‐old seedlings were exposed in open‐top chambers to charcoal‐filtered air or non‐filtered air plus an additional 40 nL L^?1 of ozone. After 20 months of O₃ exposure, a subset of plants was subjected to drought stress by withholding water supply for 11 d. Ozone induced higher guaiacol peroxidase, catalase and KCN‐resistant superoxide dismutase (SOD) activities in young needles, while drought stress increased glutathione reductase and CuZnSOD. One‐year‐old needles showed lower capacity to activate these enzymes in response to stress. Both ozone and drought activated the xanthophyll cycle pool and reduced chlorophyll contents in both current and 1‐year‐old needles. The combined effects of ozone and drought decreased antioxidant enzyme activities and the capacity of recovering after re‐watering. Similarly, interactive effects of O₃ and drought reduced xanthophyll‐mediated photoprotection capacity in 1‐year‐old needles but induced a higher conversion of the cycle in current‐year needles. These results showed that ozone modified the Aleppo pine response to drought stress, suggesting that this pollutant might be reducing the ability of this species to withstand other environmental stresses.     

Stomatal and nonstomatal limitations to net photosynthesis in seedlings of woody angiosperms

Comparative responses of net photosynthesis (A) to water stress in woody species from a variety of habitats were studied to assess the relationship between photosynthetic attributes and drought tolerance. Stomatal and nonstomatal limitations to A were compared in three-month-old white oak (Quercus alba L.), post oak (Quercus stellata Wangenh.), sugar maple (Acer saccharum Marsh.), and black walnut (Juglans nigra L.) seedlings during a drying cycle. Relative stomatal limitation of photosynthesis (I) was less than 50% in all species except for Q. stellata seedlings subjected to severe water stress. No significant changes in I were observed in Q. alba and J. nigra before, during, and after drought. In A. saccharum, I was generally low and decreased significantly under water stress. Under well-watered conditions, A was highest in Q. stellata, intermediate in Q. alba, and lower in A. saccharum and J. nigra. High A in well-watered Q. stellata was associated with high stomatal conductance and carboxylation efficiency, whereas low A was associated with low stomatal conductance and carboxylation efficiency in A. saccharum and low stomatal conductance, low carboxylation efficiency, and high CO₂ compensation point in J. nigra. Under severe water stress, A, carboxylation efficiency, and stomatal conductance decreased substantially in all species; however, Q. stellata had the highest carboxylation efficiency and lowest CO₂ compensation point under these conditions. After 5 days at high soil moisture after drought, stomatal and mesophyll components of A in A. saccharum and J. nigra had not recovered to predrought levels, whereas they had completely recovered in Q. stellata and Q. alba. The photosynthetic apparatus, especially mesophyll components, of drought-tolerant Quercus species showed either less inhibition under water stress, superior recovery to predrought capacity, or both. Exposure of the leaves to ¹⁴CO₂ indicated apparent asymmetric stomatal closure for mildly water-stressed seedlings, but not for leaves of well-watered, severely stressed, or rehydrated plants. These results suggest that patchy stomatal closure under mild water stress might be important for water stress-induced inhibition of photosynthesis, but not under the more severe water stress imposed in this study.     

Hydraulic control of stomatal conductance in Douglas fir [Pseudotsuga menziesii (Mirb.) Franco] and alder [Alnus rubra (Bong)] seedlings

Experiments were conducted on 1-year-old Douglas fir [Pseudotsuga menziesii (Mirb.) Franco] and 2- to 3-month-old alder [Alnus rubra (Bong)] seedlings growing in drying soils to determine the relative influence of root and leaf water status on stomatal conductance (g_c). The water status of shoots was manipulated independently of that of the roots using a pressure chamber that enclosed the root system. Pressurizing the chamber increases the turgor of cells in the shoot but not in the roots. Seedling shoots were enclosed in a whole-plant cuvette and transpiration and net photosynthesis rates measured continuously. In both species, stomatal closure in response to soil drying was progressively reversed with increasing pressurization. Responses occurred within minutes of pressurization and measurements almost immediately returned to pre-pressurization levels when the pressure was released. Even in wet soils there was a significant increase in g_c with pressurization. In Douglas fir, the stomatal response to pressurization was the same for seedlings grown in dry soils for up to 120 d as for those subjected to drought stress over 40 to 60 d. The stomatal conductance of both Douglas fir and alder seedlings was less sensitive to root chamber pressure at higher vapour pressure deficits (D), and stomatal closure in response to increasing D from 1.04 to 2.06 kPa was only partially reversed by pressurization. Our results are in contrast to those of other studies on herbaceous species, even though we followed the same experimental approach. They suggest that it is not always appropriate to invoke a ‘feedforward’ model of short-term stomatal response to soil drying, whereby chemical messengers from the roots bring about stomatal closure.     

Effect of drought and ABA on growth, photosynthesis and antioxidant system of Cotinus coggygria seedlings under two different light conditions

We exposed seedlings of Cotinus coggygria var. cinerea to drought and exogenous abscisic acid (ABA) under two different light conditions. Two watering regimes (well-watered and drought), two exogenous ABA applications (no ABA and with ABA) and two light regimes (full sunlight and shade) were employed. Compared with well-watered treatment, drought treatment significantly reduced the relative growth rate, relative water content (RWC), net photosynthesis rate (A) and transpiration (E), but increased chlorophyll a (chla), carbon isotope (δ¹³C), endogenous ABA, malondialdehyde (MDA) and hydrogen peroxide (H₂O₂) contents, and guaiacol peroxidase (POD) and catalase (CAT) activities. There was an apparent alleviation of drought effects by shade, as indicated by the lower relative growth rate, and chlorophyll, MDA and H₂O₂ contents, and increases in indoleacetic acid (IAA) and reduced glutathione (GSH) contents. On the other hand, the exogenous ABA application under shade induced protective effects on drought-stressed seedlings, as visible in RWC, MDA, A, stomatal conductance (g_s), E, δ¹³C, ABA and IAA values. In all, our results suggest that seedlings of C. coggygria are more sensitive to drought under full-light than under shade.     

Arbuscular Mycorrhizal Fungi Alter Fractal Dimension Characteristics of Robinia pseudoacacia L. Seedlings Through Regulating Plant Growth,Leaf Water Status,Photosynthesis, and Nutrient Concentration Under Drought Stress

The influence of arbuscular mycorrhizal fungi (AMF), Funneliformis mosseae and Rhizophagus intraradices, on plant growth, leaf water status, chlorophyll concentration, photosynthesis, nutrient concentration, and fractal dimension (FD) characteristics of black locust (Robinia pseudoacacia L.) seedlings was studied in pot culture under well-watered, moderate drought stress, and severe drought stress treatments. Mycorrhizal seedlings had higher dry biomass, leaf relative water content (RWC), and water use efficiency (WUE) compared with non-mycorrhizal seedlings. Under all treatments, AMF colonization notably enhanced net photosynthetic rate, stomatal conductance, and transpiration rate, but decreased intercellular CO₂ concentration. Leaf chlorophyll a and total chlorophyll concentrations were higher in AM seedlings than those in non-AM seedlings although there was no significant difference between AMF species. AMF colonization improved leaf C, N, and P concentrations, but decreased C:N, C:P, and N:P ratios. Mycorrhizal seedlings had a larger FD value than non-mycorrhizal seedlings. The FD value was positively and significantly correlated to the plant growth parameters, photosynthesis, RWC, WUE, and nutrient concentration but negatively correlated to leaf/stem ratio, C:N and C:P ratios, and intercellular CO₂ concentration. We conclude that AMF lead to an improvement of growth performance of black locust seedlings under all growth conditions, including drought stress via improving leaf water status, chlorophyll concentration, photosynthesis, and nutrient uptake. Moreover, FD technology proved to be a powerful non-destructive method to characterize the effect of AMF on the physiology of host plants during drought stress.     

A stomatal optimization theory to describe the effects of atmospheric CO2 on leaf photosynthesis and transpiration

Background and Aims

Global climate models predict decreases in leaf stomatal conductance and transpiration due to increases in atmospheric CO₂. The consequences of these reductions are increases in soil moisture availability and continental scale run-off at decadal time-scales. Thus, a theory explaining the differential sensitivity of stomata to changing atmospheric CO₂ and other environmental conditions must be identified. Here, these responses are investigated using optimality theory applied to stomatal conductance.

Methods

An analytical model for stomatal conductance is proposed based on: (a) Fickian mass transfer of CO₂ and H₂O through stomata; (b) a biochemical photosynthesis model that relates intercellular CO₂ to net photosynthesis; and (c) a stomatal model based on optimization for maximizing carbon gains when water losses represent a cost. Comparisons between the optimization-based model and empirical relationships widely used in climate models were made using an extensive gas exchange dataset collected in a maturing pine (Pinus taeda) forest under ambient and enriched atmospheric CO₂.

Key Results and Conclusion

In this interpretation, it is proposed that an individual leaf optimally and autonomously regulates stomatal opening on short-term (approx. 10-min time-scale) rather than on daily or longer time-scales. The derived equations are analytical with explicit expressions for conductance, photosynthesis and intercellular CO₂, thereby making the approach useful for climate models. Using a gas exchange dataset collected in a pine forest, it is shown that (a) the cost of unit water loss λ (a measure of marginal water-use efficiency) increases with atmospheric CO₂; (b) the new formulation correctly predicts the condition under which CO₂-enriched atmosphere will cause increasing assimilation and decreasing stomatal conductance.     

Developing a mathematical model for variation of physiological responses of Jatropha curcas leaves depending on leaf positions under soil flooding

The effect of soil flooding on photosynthesis, transpiration and stomatal conductance of Jatropha curcas seedlings were studied under natural environmental variables. Soil flooding reduced photosynthesis (P _N), transpiration (E) and stomatal conductance (gs) in response to leaf positions of Jatropha curcas plants. Based on the results, we conclude that decrease in stomatal opening and stomatal limitation of photosynthesis, followed by decrease in individual leaf area are the main causes of reductions in carbon uptake of flooded seedlings. A mathematical relationship was successfully developed to describe photosynthesis, transpiration and stomatal response of Jatropha under soil flooding stress.     

Limitations of Photosynthesis in Pinus taeda L. (Loblolly Pine) at Low Soil Temperatures

The relative importance of stomatal and nonstomatal limitations to net photosynthesis (A) and possible signals responsible for stomatal limitations were investigated in unhardened Pinus taeda seedlings at low soil temperatures. After 2 days at soil temperatures between 13 and 7°C, A was reduced by 20 to 50%, respectively. The reduction in A at these moderate root-chilling conditions appeared to be the result of stomatal limitations, based on the decrease in intercellular CO₂ concentrations (c_i). This conclusion was supported by A versus c_i analysis and measurements of O₂ evolution at saturating CO₂, which suggested increases in stomatal but not biochemical limitations at these soil temperatures. Nonuniform stomatal apertures, which were demonstrated with abscisic acid, were not apparent 2 days after root chilling, and results of our A versus c_i analysis appear valid. Bulk shoot water potential (ψ) declined as soil temperature dropped below 16°C. When half the root system of seedlings was chilled, shoot ψ and gas-exchange rates did not decline. Thus, nonhydraulic root-shoot signals were not implicated in stomatal limitations. The initial decrease in leaf conductance to water vapor after root chilling appeared to precede any detectable decrease in bulk fascicle ψ, but may be in response to a decrease in turgor of epidermal cells. These reductions in leaf conductance to water vapor, which occurred within 30 minutes of root chilling, could be delayed and temporarily reversed by reducing the leaf-to-air vapor-pressure deficit, suggesting that hydraulic signals may be involved in initiating stomatal closure. By independently manipulating the leaf-to-air vapor-pressure deficit of individual fascicles, we could induce uptake of water vapor through stomata, suggesting that nonsaturated conditions occur in the intercellular airspaces. There was an anomaly in our results on seedlings maintained for 2 days at soil temperatures below 7°C. Lower A appeared primarily the result of nonstomatal limitations, based on large increases in calculated c_i and A versus c_i analysis. In contrast, measurements of O₂ evolution at saturating CO₂ concentrations implied nonstomatal limitations per se did not increase at these temperatures. One explanation for this paradox is that calculations of c_i are unreliable at very low gas-exchange rates because of inadequate measurement resolution, and limitations of A are predominantly stomatal. An alternative interpretation is that increases in c_i are real and the results from O₂-evolution measurements are in error. The high CO₂ concentration used in O₂-evolution measurements (15%) may have overcome nonstomatal limitations by enzymes that were down-regulated by a feedback mechanism. In this scenario, carbohydrate feedback limitations may be responsible for nonstomatal reductions in A after 2 days at soil temperatures below 7°C.     

Recovery responses of photosynthesis,transpiration, and stomatal conductance in kidney bean following drought stress

Photosynthesis, transpiration, stomatal conductance and chlorophyll fluorescence characteristics were examined in kidney bean plants, with developing gradually water stress for several days after watering and then permitted to recover by re-watering. The photosynthetic rate, transpiration rate, and stomatal conductance decreased rapidly by withholding water for 2 days. The F_v/F_m of chlorophyll fluorescence characteristics slightly decreased when the water was withheld for 7 days. After re-watering the rate of recovery of photosynthesis, transpiration, and stomatal conductance decreased gradually as the days without watering became longer. The differences existed in rates of recovery of photosynthesis, transpiration, and stomatal conductance following drought stress. Among the fractional recoveries the highest was photosynthesis, and the lowest was stomatal conductance. Photosynthesis rate following drought stress was rapidly recovered until 2 days after re-watering, then recovered slowly. The critical time for the recovery of photosynthesis was recognized. The results show clearly a close correlation between the leaf water potential and the recovery level and speed of photosynthesis, transpiration, and stomatal conductance.     

干旱胁迫对降香黄檀幼苗光合生理特性的影响

采用温室盆栽方法,设置对照(CK)、轻度(LS)、中度(MS)和重度(HS)干旱胁迫4个水分条件,研究不同水分条件对降香黄檀幼苗光合和生理特性的影响。结果表明:(1)随着干旱胁迫程度增加,降香黄檀幼苗叶片叶绿素总含量总体呈现出下降趋势。(2)降香黄檀幼苗叶片净光合速率、气孔导度、胞间CO2浓度和蒸腾速率随着干旱胁迫强度增加均呈现出先增加后降低趋势,且MS和HS处理下的气孔导度和胞间CO2浓度同时降低,此时幼苗光合能力的下降主要受气孔因素限制。(3)随着干旱胁迫强度的增加,降香黄檀幼苗叶片细胞膜相对透性、丙二醛含量、游离脯氨酸含量和POD活性均呈现出增加趋势,而同期SOD和CAT活性呈现出先升高后降低趋势。可见,降香黄檀幼苗在轻度干旱胁迫下可通过增加叶片保护酶活性来清除活性氧对其组织造成的伤害,但胁迫超过一定程度后保护酶活性下降,表明降香黄檀幼苗的耐旱能力有限。     

The effect of CaCl<Subscript>2</Subscript> on calcium content,photosynthesis, and chlorophyll fluorescence of tung tree seedlings under drought conditions

The study investigated the effects of different CaCl₂ concentrations (2, 5, and 10 mM) on photosynthetic enzymatic activities, photosynthesis, and chlorophyll fluorescence of tung tree seedlings under drought conditions. Plants were sprayed with either CaCl₂ or distilled water until run-off. Irrigation was then withheld to induce drought stress. The strength of drought stress was evaluated by relative leaf water content and soil water content, which was 27.3 and 9.5% on day 0 and day 12, respectively. Drought stress decreased activities of ribulose-1,5-bisphosphate carboxylase/oxygenase and phosphoenolpyruvate carboxylase, chlorophyll (a+b) content, net photosynthetic rate, stomatal conductance, transpiration rate, electron transport rate, the maximal quantum yield of PSII photochemistry, and effective quantum yield of PSII in tung tree seedlings. The CaCl₂ pretreatments alleviated the negative effect of drought stress to some degree on all the parameters mentioned above.     

Improvement of growth conditions and gas exchange of<Emphasis Type="Italic"> Fagus sylvatica</Emphasis> L. seedlings planted below a recently thinned<Emphasis Type="Italic"> Pinus sylvestris</Emphasis> L. stand

Abstract Gas exchange and growth of beech seedlings planted in the understory of a recently thinned pinewood were recorded for 2 years. Relative irradiance was assessed by hemispherical photographs taken just after the thinning. Predawn water potential (_pd), daily gas exchange and chlorophyll fluorescence were measured several times during the two growing seasons. Maximum values of photosynthesis (A _max) and stomatal conductance to water vapour (g _wvmax) were established from daily data. Maximum quantum efficiency of PS II was recorded at dawn by taking the variable to maximum chlorophyll fluorescence ratio on dark adapted leaves (F _v/F _m). In the middle of each summer, leaf nitrogen content and leaf mass per area were evaluated, and height growth and basal area increment were recorded at the end of the season. The thinning treatment removed half the trees and generated around 10% more available relative irradiance (GLF). This was followed by an increase in net photosynthesis at saturating PPFD (A _sat) and in maximum stomatal conductance to water vapour (g _wvmax). Moreover, specific leaf mass (SLM) and mass based nitrogen content (N_m) showed higher values for seedlings in the thinned stand. In both years, a positive relationship was established between the area based nitrogen content (N_a) and maximum net photosynthesis (A _max). In 1998, a year with a dry summer, seedlings suffered a significant drop in daily A _max irrespective of the thinning regime. This was a response to an increase in stomatal limitation to net photosynthesis, g _wvmax reaching the lowest value on dates with the highest drought. A lack of decrease of Fv/Fm confirmed the absence of significant non-stomatal limitation to A as a consequence of photoinhibition after opening the pinewood. A higher maximum quantum efficiency of open PS II centres (Fv/Fm) was registered in seedlings in the thinned stand. The significance of the differences between the treatments was stronger in the second year after thinning. In 1999, a year with frequent summer storms, water availability increased for seedlings growing under the thinned pinewood. Overall, the reduced pine overstory had a positive effect on physiological responses of beech seedlings, which was translated into improved seedling growth.     

Photosynthetic responses of Monarch birch seedlings to differing timings of free air ozone fumigation

To study the effects of different periods of ozone (O₃) fumigation on photosynthesis in leaves of the Monarch birch (Betula maximowicziana), we undertook free air O₃ fumigation to Monarch birch seedlings at a concentration of 60 nmol mol^?1 during daytime. Plants were exposed to O₃ at early, late or both periods in the growing season. The light-saturated net photosynthetic rate (A _sat) in July and August was reduced by O₃ exposure through a reduction in the maximum rate of carboxylation (V _c,max). In early September, on the other hand, despite a reduction in V _c,max, A _sat was not reduced by O₃ due to a counteracting increase in the stomatal conductance. Through the experiment, there was no difference in sensitivity to O₃ between maturing and matured leaves. We analyzed the relationship between A _sat, V _c,max and accumulated stomatal O₃ flux (AF_st). Whereas V _c,max decreased with increasing AF_st, the correlation between A _sat and AF_st was weak because the response of stomatal conductance to O₃ was affected by season. We conclude photosynthetic response of Monarch birch to O₃ exposure changes with season. This is due to the inconstant stomatal response to O₃ but not due to the respose of biochemical assimilation capacity in chloroplasts.     

Photosynthesis in Encelia farinosa Gray in Response to Decreasing Leaf Water Potential

Photosynthetic responses of intact leaves of the desert shrub Encelia farinosa were measured during a long term drought cycle in order to understand the responses of stomatal and nonstomatal components to water stress. Photosynthetic rate at high irradiance and leaf conductance to water vapor both decreased linearly with declining leaf water potential. The intercellular CO₂ concentration (c_i) remained fairly constant as a function of leaf water potential in plants subjected to a slow drought cycle of 25 days, but decreased in plants exposed to a 12-day drought cycle. With increasing water stress, the slope of the dependence of photosynthesis on c_i (carboxylation efficiency) decreased, the maximum photosynthetic rates at high c_i became saturated at lower values, and water use efficiency increased. Both the carboxylation efficiency and photosynthetic rates were positively correlated with leaf nitrogen content. Associated with lower leaf conductances, the calculated stomatal limitation to photosynthesis increased with water stress. However, because of simultaneous changes in the dependence of photosynthesis on c_i with water stress, increased leaf conductance alone in water-stressed leaves would not result in an increase in photosynthetic rates to prestressed levels. Both active osmotic adjustment and changes in specific leaf mass occurred during the drought cycle. In response to increased water stress, leaf specific mass increased. However, the increases in specific leaf mass were associated with the production of a reflective pubescence and there were no changes in specific mass of the photosynthetic tissues. The significance of these responses for carbon gain and water loss under arid conditions are discussed.     

The effect of light on stomatal control of gas exchange in Douglas fir (Pseudotsuga menziesii) saplings

Summary Attached twigs of young Pseudotsuga menziesii (Mirb.) Franco plants were subjected to variations in irradaince. Stomatal responsiveness to irradiance, measured in an open type gas exchange system, varied seasonally. During the autumn and winter, stomatal conductance was relatively unresponsive to changes in irradiance, but during the summer stomatal conductance decreased in response to reduced irradiance. The summer stomatal response to irradiance was such that a nearly constant ratio of stomatal conductance to net photosynthesis was maintained as irradiance was varied. This caused intercellular CO₂ concentration (c _i) and water use efficiency (net CO₂ uptake/transpiration) to also remain relatively constant. At constant irradiance, stomatal conductance was relatively insensitive to experimentally-induced changes in c _i. This, and the observation that c _i remained relatively constant as irradiance was varied, suggest that changes in c _i played a minor role in mediating the stomatal response to light.The ecological significance of the seasonal changes in stomatal response to light is discussed.     

Internal conductance to CO2 transfer of adult Fagus sylvatica: Variation between sun and shade leaves and due to free-air ozone fumigation

Two separate objectives were considered in this study. We examined (1) internal conductance to CO₂ (g_i) and photosynthetic limitations in sun and shade leaves of 60-year-old Fagus sylvatica, and (2) whether free-air ozone fumigation affects g_i and photosynthetic limitations. g_i and photosynthetic limitations were estimated in situ from simultaneous measurements of gas exchange and chlorophyll fluorescence on attached sun and shade leaves of F. sylvatica. Trees were exposed to ambient air (1× O₃) and air with twice the ambient ozone concentration (2× O₃) in a free-air ozone canopy fumigation system in southern Germany (Kranzberg Forest). g_i varied between 0.12 and 0.24 mol m⁻² s⁻¹ and decreased CO₂ concentrations from intercellular spaces (C_i) to chloroplastic (C_c) by approximately 55 μmol mol⁻¹. The maximum rate of carboxylation (V_cmax) was 22–39% lower when calculated on a C_i basis compared with a C_c basis. g_i was approximately twice as large in sun leaves compared to shade leaves. Relationships among net photosynthesis, stomatal conductance and g_i were very similar in sun and shade leaves. This proportional scaling meant that neither C_i nor C_c varied between sun and shade leaves. Rates of net photosynthesis and stomatal conductance were about 25% lower in the 2× O₃ treatment compared with 1× O₃, while V_cmax was unaffected. There was no evidence that g_i was affected by ozone.