ALBERT

Notes: Leaf gas-exchange and chemical composition were investigated in seedlings of Quercus suber L. grown for 21 months either at elevated (700 μmol mol–1) or normal (350 μmol mol–1) ambient atmospheric CO2 concentrations, [CO2], in a sandy nutrient-poor soil with either ‘high’ N (0.3 mol N m–3 in the irrigation solution) or with ‘low’ N (0.05 mol N m–3) and with a constant suboptimal concentration of the other macro- and micronutrients. Although elevated [CO2] yielded the greatest total plant biomass in ‘high’ nitrogen treatment, it resulted in lower leaf nutrient concentrations in all cases, independent of the nutrient addition regime, and in greater nonstructural carbohydrate concentrations. By contrast, nitrogen treatment did not affect foliar N concentrations, but resulted in lower phosphorus concentrations, suggesting that under lower N, P use-efficiency in foliar biomass production was lower. Phosphorus deficiency was evident in all treatments, as photosynthesis became CO2 insensitive at intercellular CO2 concentrations larger than ≈ 300 μmol mol–1, and net assimilation rates measured at an ambient [CO2] of 350 μmol mol–1 or at 700 μmol mol–1 were not significantly different. Moreover, there was a positive correlation of foliar P with maximum Rubisco (Ribulose-1,5-bisphosphate carboxylase/oxygenase) carboxylase activity (Vcmax), which potentially limits photosynthesis at low [CO2], and the capacities of photosynthetic electron transport (Jmax) and phosphate utilization (Pmax), which are potentially limiting at high [CO2]. None of these potential limits was correlated with foliar nitrogen concentration, indicating that photosynthetic N use-efficiency was directly dependent on foliar P availability. Though the tendencies were towards lower capacities of potential limitations of photosynthesis in high [CO2] grown specimens, the effects were statistically insignificant, because of (i) large within-treatment variability related to foliar P, and (ii) small decreases in P/N ratio with increasing [CO2], resulting in balanced changes in other foliar compounds potentially limiting carbon acquisition. The results of the current study indicate that under P-deficiency, the down-regulation of excess biochemical capacities proceeds in a similar manner in leaves grown under normal and elevated [CO2], and also that foliar P/N ratios for optimum photosynthesis are likely to increase with increasing growth CO2 concentrations. Symbols: A, net assimilation rate (μmol m–2 s–1); Amax, light-saturated A (μmol m–2 s–1); α, initial quantum yield at saturating [CO2] and for an incident Q (mol mol–1); [CO2], atmospheric CO2 concentration (μmol mol–1); Ci, intercellular CO2 concentration (μmol mol–1); Ca, CO2 concentration in the gas-exchange cuvette (μmol mol–1); FB, fraction of leaf N in ‘photoenergetics’; FL, fraction of leaf N in light harvesting; FR, fraction of leaf N in Rubisco; Γ*, CO2 compensation concentration in the absence of Rd (μmol mol–1); Jmax*, capacity for photosynthetic electron transport; Jmc, capacity for photosynthetic electron transport per unit cytochrome f (mol e–[mol cyt f]–1 s–1); Kc, Michaelis-Menten constant for carboxylation (μmol mol–1); Ko, Michaelis-Menten constant for oxygenation (mmol mol–1); MA, leaf dry mass per area (g m–2); O, intercellular oxygen concentration (mmol mol–1); [Pi], concentration of inorganic phosphate (mM); Pmax*, capacity for phosphate utilization; Q, photosynthetically active quantum flux density (μmol m–2 s–1); Rd*, day respiration (CO2 evolution from nonphotorespiratory processes continuing in the light); Rubisco, ribulose-1,5-bisphosphate carboxylase/oxygenase; RUBP, ribulose-1,5-bisphosphate; Tl, leaf temperature (°C); UTPU*, rate of triose phosphate utilization; Vcmax*, maximum Rubisco carboxylase activity; Vcr, specific activity of Rubisco (μmol CO2[g Rubisco]–1 s–1]*given in either μmol m–2 s–1 or in μmol g–1 s–1 as described in the text.

Notes: We studied the effects of drought on leaf conductance (g) and on the concentration of abscisic acid (ABA) in the apoplastic sap of Lupinus albus L. leaves. Withholding watering for 5d resulted in complete stomatal closure and in severe leaf water deficit. Leaf water potential fully recovered immediately after rewatering, but the aftereffect of drought on stomata persisted for 2d. ABA and sucrose were quantified in pressurized leaf xylem extrudates. We assumed that the xylem sucrose concentration is negligible and hence that the presence of sucrose in leaf extrudates indicated that they were contaminated by phloem. To eliminate this interference, the concentration of ABA in leaf apoplast was estimated by extrapolation to zero sucrose concentration, using the regression between ABA and sucrose concentrations. The estimated apoplastic ABA concentration increased by 100-fold with soil drying and did not return to pre-stress values immediately following rewatering. g was closely related to the concentration of ABA in leaf apoplast. Furthermore, the feeding of exogenous ABA to leaves detached from well-watered plants brought about the same degree of depression in g as resulted from the drought-induced increase in ABA concentration. We therefore conclude that the observed changes in the concentration of ABA in leaf apoplast were quantitatively adequate to explain drought-induced stomatal closure and the delay in stomatal reopening following rewatering.

Notes: Wild-type and antisense rbcS tobacco (Nicotiana tabacum) plants were grown in a glasshouse in midsummer in Portugal with an irradiance of 1500–2000 μmol m−2s−1 and daytime temperatures of 30–35 °C. The Rubisco content of the transformants was lower by 35, 80 and over 90% than that of the wild-type. Gas exchange was measured over three separate days. There was a near-linear relation between Rubisco content and photosynthetic rate during the period of high irradiance, allowing a flux control coefficient of 0.83–0.89 to be estimated. The relation deviated slightly from linearity, because the internal CO2 concentration (c;) was higher in the transformants than in the wild-type (190 and 275 μmol mol−1 in plants with 35 and 80% less Rubisco, respectively, compared with 175 μmol mol−1 for wild-type), compensating to some extent for the decreased Rubisco content. This increase in ci occurred because the stomatal conductance (g) remained unaltered or was even higher in plants with decreased Rubisco, despite the lower rate of CO2 assimilation. As a consequence, water use efficiency declined. The decreased rate of photosynthesis was not accompanied by a stoichiometric decrease in apparent growth rate. These results are discussed in relation to earlier studies of the plant set in growth cabinets. It is concluded that tobacco can adjust over a wide range of growth conditions to avoid a onesided limitation by Rubisco, but that in extreme environmental conditions this capacity to adapt is exhausted.

Notes: The effects of long-term (4 year) CO2 enrichment (70 Pa versus 35 Pa) and nitrogen nutrition (8 mm versus 1 mm NO3–) on biomass accumulation and the development of photosynthetic capacity in leaves of cork oak (Quercus suber L., a Mediterranean evergreen tree) were studied. The evolution of photosynthetic parameters with leaf development was estimated by fitting the biochemical model of Farquhar et al. (Planta 149, 78–90, 1980) with modifications by Sharkey (Botanical Review 78, 71–75, 1985) to A–Ci response curves. CO2 enrichment had a small reduction effect on the development of the maximum CO2 fixation capacity by Rubisco (VCmax), and no effect over maximum electron transport capacity (Jmax), day-time respiration (Rd) and Triose-P utilization (TPU). However, there was a statistically significant effect of N fertilization and the interaction CO2 × N over the evolution of VCmax, Jmax and TPU. Relative stomatal limitation (estimated from A–Ci curves) was higher (+20%) for plants grown under ambient CO2 than for plants grown under elevated CO2. There was a significant effect of CO2 and N fertilization over total biomass accumulation as well as leaf area. Plants grown at elevated CO2 had 27% more biomass than plants grown at ambient CO2 when given high N. However, for plants grown under low N there was no significant effect of CO2 enrichment on biomass accumulation. Plants grown under low N also had significantly higher root : shoot ratios whereas there were no differences between CO2 treatments. The larger biomass accumulation of Q. suber under elevated CO2 is attributable to a higher availability of CO2 coupled to a larger leaf area, with no significant decrease in photosynthetic capacity under CO2 enrichment and elevated N fertilization. For low N fertilization, the effects of CO2 enrichment over leaf area and biomass accumulation are lost, suggesting that in native ecosystems with low N availability, the effects of CO2 enrichment may be insignificant.

Notes: Differences in maximum leaf conductance in grapevine plants growing in soils with contrasting water availabilities during mid-summer in Portugal could be accounted for by differences in the concentration of ABA in xylem sap. This conclusion is reinforced by the observation that the relationship between leaf conductance and endogenous ABA concentration can be mimicked by the application of exogenous ABA to leaves detached from irrigated plants. During the day, leaf conductance decreased after a morning peak, even when the leaves remained in a constant environment at a moderate temperature and leaf-to-air vapour pressure difference. This decline in leaf conductance was not a consequence of an increase in the xylem ABA concentration or the rate of delivery of this compound by the transpiratory stream. The afternoon depression in leaf conductance was associated with an apparent limitation in stomatal opening potential, which persisted even when detached leaves were fed with water and rehydrated. The reason for this inhibition has still to be identified.

Notes: Daily changes in photoprotective mechanisms were studied in sun leaves of Quercus suber L., Quercus ilex L., Olea europaea L. and Eucalyptus globulus Labill. trees during the summer in Portugal. Even though stomatal closure explained most of the diurnal variation in carbon assimilation along the summer, a decline in the photochemical yield of photosystem II (F′v/F′m) also occurred, as a result of an excess of intercepted solar radiation when carbon assimilation is limited by stomatal closure due to high vapour pressure deficits and/or soil water deficits. These changes were accompanied by the conversion of violaxanthin to antheraxanthin and zeaxanthin which were correlated with thermal dissipation of excess photon energy. In spite of a common general response, differences between species were observed -Olea europaea, which is a slow-growing tree, had the lowest net photosynthetic rates, the highest proportion of carotenoids in relation to chlorophyll and the highest rates of de-epoxidation of violaxanthin. This enabled a large thermal dissipation of the excess intercepted radiation but led to rather small values of light utilisation for photochemistry (ca 20%). In contrast, in E. globulus, a fast-growing tree, photosynthetic rates were the highest, thermal dissipation of absorbed radiation the lowest and maximal values of light utilisation for photochemistry reached ca 50%. The two Quercus species exhibited an intermediate response. A high degree of co-ordination is apparent between stomatal behaviour, photosynthetic capacity and photoprotection mechanisms.

Notes: Effects of flooding on young Populus deltoides, Salix nigra, Eucalyptus camaldulensis, E. globulus, Ulmus americana, Quercus rubra and Fraxinus pennsylvanica plants were studied. Flooding variously induced several sequential physiological disturbances, with stomatal closure among the earliest responses. Subsequent responses included inhibition of root growth, alterations in root and stem morphology, formation of adventitious roots, and leaf senescence. In amphistomatous species (Populus deltoides, Salix nigra, Eucalyptus camaldulensis) flooding rapidly induced stomatal closure on the adaxial leaf surface. It also significantly induced stomatal closure on the abaxial epidermis of Populus deltoides but not that of Salix nigra or Eucalyptus camaldulensis. In hypostomatous species (Eucalyptus globulus, Ulmus americana) flooding significantly induced stomatal closure on the abaxial surface within 3 days. Stomatal responses to flooding were not correlated with leaf water stress. In both long- and short-term experiments, flooding did not significantly increase plant water stress. These results deemphasized the importance of plant water stress in inducing plant responses to flooding. The importance of various hormones in inducing flooding symptoms is discussed.

Notes: Diurnal and seasonal changes in plant water potential, leaf diffusion resistance, and stem radial changes of Acer saccharum and Betula papyrifera trees were studied in northern Wisconsin during the 1974 and 1975 growing seasons. Water potential decreased during the day, following relatively high values in the morning, and increased in the late afternoon and evening. Diurnal patterns and actual values of water potential varied with species, soil water availability, and factors influencing transpiration (e.g., solar radiation, vapor pressure deficit, and transpiration flux density). When plant water deficits were not severe, leaf resistance of both species was rather stable during the day. During severe droughts, however, leaf resistance increased (stomata closed) during the day when light intensity was high. Leaf resistance at high light intensity was higher in Acer than in Betula. Stomatal closure with decreasing light intensity varied between species and among Acer trees. Tree stems of both species shrank during the day, as internal water deficits developed, and they expanded as trees rehydrated during the night. Stems of Acer shrank more than those of Betula. The amount of daily stem shrinkage increased as the season progressed if the trees were not under severe water deficits. During severe droughts the amount of diurnal stem shrinkage decreased. Shrinkage of stems lagged behind water potential changes by 1 to 2 h in Acer and less than 1 h in Betula. The relationship between stem radius and leaf water potential was not constant throughout the growing season.

Notes: Abstract. The effect of gradually-developing water-stress has been studied in Lupinus albus L., Helianthus annuus L., Vitis vinifera cv. Rosaki and Eucalyptus globulus Labill. Water was withheld and diurnal rhythms were investigated 4–8d later, when the predawn water deficit was more negative than in watered plants, and the stomata closed almost completely early during the photoperiod. The contribution of ‘stomatal’ and ‘non-stomatal’ components to the decrease of photosynthetic rate was investigated by (1) comparing the changes of the rate of photosynthesis in air with the changes of stomatal conductance and (2) measuring photosynthetic capacity in saturating irradiance and 15% CO2. Three species (lupin, eucalyptus and sunflower) showed larger changes of stomatal conductance than photosynthesis in air, and showed little or no decrease of photosynthetic capacity in saturating CO2. Photosynthesis in air also recovered fully overnight after watering the plants in the evening. In grapevines, stomatal conductance and photosynthesis in air changed in parallel, there was a marked decrease of photosynthetic capacity, and photosynthesis and stomatal conductance did not recover overnight after watering water-stressed plants. Relative water content remained above 90% in grapevine. We conclude that non-stomatal components do not play a significant role in lupins, sunflower or eucalyptus, but could in grapevine. The effect of water-stress on partitioning of photosynthate was investigated by measuring the amounts of sucrose and starch in leaves during a diurnal rhythm, and by measuring the partitioning of 14C-carbon dioxide between sucrose and starch. In all four species, starch was depleted in water-stressed leaves but sucrose was maintained at amounts similar to, or higher than, those in watered plants. Partitioning into sucrose was increased in lupins and eucalyptus, and remained unchanged in grapevine and sunflower. It is concluded that water-stressed leaves in all four species maintain high levels of soluble sugars in their leaves, despite having lower rates of field photosynthesis, decreased rates of export, and low amounts of starch in their leaves.

Notes: Abstract The transpiration, sap flow, stomatal conductance and water relations ofPinus pinaster were determined during spring and summer in a 64-year-old stand in Ribatejo (Portugal). The transpiration of the pine canopy was determined from sap flow or eddy covariance techniques. Canopy conductance values (g c) were estimated from inversion methods using eddy covariance or sap flow data, respectively, and from scaling-up methods using stomatal conductance values measured in the field and leaf area index (LAI) values. The transpiration was closely controlled by the stomatal conductance of pines (Ω was 0.05–0.15). For wet soil conditions, the various estimates ofg c showed reasonable agreement.g c peaked in the morning at 0.01 m×s-1, exhibited a midday depression and showed a secondary peak in late afternoon. This behaviour could be predicted simply on the basis of the stomatal sensitivity to air vapour pressure deficit. On a seasonal basis, monthly average values ofg c decreased from 4×10-3 m×s-1 in spring to 1.7×10-3 m·s-1 in late summer. Accordingly, the transpiration peaked at 3 mm×d-1 on wet soil in May. It decreased progressively during the summer drought to 0.8 mm×d-1 at the end of August. The minimal value of needle water potential was maintained at -1.9 MPa but predawn values decreased from -0.6 MPa in May to -0.9 MPa in July. It may have reached lower values in August. The amount of water stored in the trunk accounted for a 12% (10 kg×tree-1×day-1) of the daily transpiration in spring. The storage capacity of the canopy was within the same order of magnitude. The trunk storage increased to 25% (13 kg×tree-1×day-1) of the daily transpiration at the end of summer under drought conditions. The sap flow beneath the crown lagged accordingly behind transpiration with a time constant estimated between 26 min in spring and 40 min at the end of summer.