Growth,morphology and gas exchange responses of two-year-old Quercus castaneifolia seedlings to flooding stress

Tolerance to flooding is crucial when thinking in promissory species for restoration of ecosystems prone to suffer soil water excess. In this study, we tested the flooding tolerance of two-years-old seedlings of Chestnut-leaved oak (Quercus castaneifolia C.A.Mey.) to determine whether it can be recommended for use in wetland restoration programs. Seedlings of Q. castaneifolia were subjected to three treatments: (1) control (C), (2) flooding for 60 days followed by a 42-day recovery period (F?+?R) and (3) continuous flooding for 102 days (F). Physiological performance, plant morphological changes and biomass accumulation were assessed. Results showed that, although net photosynthetic rates, stomatal conductance and transpiration decreased with prolonged flooding, when flood waters were removed, plants were able to recover their physiological activity (49–80% compared to controls). By contrast, when plants were continually flooded, their physiological activity decreased as well as the leaves experienced precocious senescence and wilting. Biomass responses paralleled physiological responses: leaf and root biomass were 42–49% higher under F?+?R treatment than under continuous flooding, and all plants under the F?+?R survived. Therefore, Q. castaneifolia appears as a promising species to be further studied when thinking re-vegetation of riverine areas and other temporarily flooded wetlands.     

Response of gas exchange to water stress in seedlings of woody angiosperms

Responses of net photosynthesis (A), leaf conductance to water vapor (g(wv)) and instantaneous water use efficiency (WUE) to decreasing leaf and soil water potentials (Psi(l), Psi(s)) were studied in three-month-old white oak (Quercus alba L.), post oak (Q. stellata Wangenh.), sugar maple (Acer saccharum Marsh.), and black walnut (Juglans nigra L.) seedlings. Quercus seedlings had the highest A and g(wv) when plants were well watered. As the soil was allowed to dry, both A and g(wv) decreased; however, trace amounts of A were observed at a Psi(l) as low as -2.9 MPa in Q. stellata and -2.6 MPa in Q. alba and A. saccharum. Photosynthesis was not measurable at Psi(l) lower than -2.2 MPa in J. nigra and water stress-induced leaflet senescence was observed in this species. Within each species, g(wv) showed a similar relationship to soil and leaf Psi, but the response to Psi(l) was shifted to more negative values by 1.2 to 1.6 MPa. As Psi(s) declined below -1 MPa, the difference between soil and leaf Psi diminished because of the suppression of transpiration. There was no indication that Psi(s) had a more direct influence on g(wv) than did Psi(l). Water use efficiency showed an initial increase as the soil dried, followed by a decline under severe water stress. Water use efficiency was highest in J. nigra, intermediate in Quercus species and lowest in A. saccharum. There was an evident relationship between gas exchange characteristics and natural distribution in these species, with the more xeric species showing higher A and g(wv) under both well-watered and water-stressed conditions. There was no trend toward increased efficiency of water use in the more xeric species.     

Root system of Quercus suber L. seedlings in response to herbaceous competition and different watering and fertilisation regimes

Different management practices are commonly applied to increase pasture yield of Mediterranean open woodlands, but the consequences of increasing competition for soil resources with these practices on tree recruitment are still unknown. In a greenhouse study, belowground competition of Quercus suber L. seedlings growing together with natural (Oak_NP) or improved pasture (Oak_IP) was evaluated, and their root systems compared with those of seedlings growing in bare soil (Oak_BS). Two watering levels and two regimes of P₂O₅ fertilisation were also tested. Because of competition, the Oak_IP seedlings had their fine root mass density, coarse root length, and shoot mass reduced by up to 40, 36, and 39%, respectively, when compared to Oak_NP seedlings. Oak_NP and Oak_BS seedlings showed similar average root density parameters and shoot mass values, indicating that Q. suber seedlings growing with natural pasture is a low competitive system. High availability of water and phosphorus did not mitigate the strength of competition between herbaceous plants and oak seedlings, and favoured the pasture to the detriment of the trees. Our findings suggest that P₂O₅ fertilisation and irrigation practices performed to improve herbaceous productivity will negatively influence recruitment of Q. suber seedlings.     

Chemical composition and variability of cork from Quercus suber L.

Summary The chemical composition of cork from Quercus suber L. was determined for virgin cork and for reproduction cork. Average chemical composition of virgin cork is: ash 0.7%, total extractives 15.3%, suberin 38.6%, lignin 21.7% and polysaccharides 18.2%. The carbohydrate composition shows that glucose represents 50.6% of all monosaccharides, xylose 35.0%, arabinose 7.0% and galactose and mannose, respectively, 3.6% and 3.4%. Reproduction cork has a composition similar to virgin cork. For the cases studied, reproduction cork had however less extractives and a higher amount of suberin. Significant differences were found for the chemical composition of virgin cork in relation to location of extractives and polysaccharide content. The between-tree variability was large and a variation within the tree was also recorded.The author is grateful for the help of collaborators throughout the chemical analysis of the many samples: Joaquina Ferreira, A. Velez Marques, M. Lurdes Ferreira, Rosa Amorim and Célia Constâncio.The work was financially supported by the Instituto de Ciência e Tecnologia dos Materiais (ICTM), Lisbon, Portugal     

Morphological and physiological acclimation of Quercus coccifera L. seedlings to water availability and growing medium

One of the main constraints of reforestation in the Mediterranean region is low summer water availability during the first years after out planting. Plant water availability depends on the precipitation regime, but also on the physical properties of the soil. Higher survival rates result when seedlings are soil acclimated. Our main goal was to describe the morpho-physiological responses of 6-, 10- and 18-month-old Quercus coccifera seedlings growing in a natural soil (terra rossa) or a standard nursery growing medium, and to assess in the nursery if seedlings growing in natural soil were more resistant to deficit irrigation. The high growth rate achieved after 10?C18 months by terra rossa-grown seedlings in contrast with those grown in the nursery substrate suggests that the former were acclimated to the soil. Higher photosynthetic rate (A), transpiration (E) and stomatal conductance (g_s) were observed in terra rossa seedlings, mainly during the first months. The higher carbon availability may account for the higher root nitrogen concentration in terra rossa-grown seedlings, which could favor their later field growth. Low-watered seedlings showed a certain degree of hardening, since after 18 months, they showed higher A, E, g_s and lower photoinhibition than well-watered seedlings, likely attributable to the sharp leaf-to-root biomass ratio reduction. Carbon isotope discrimination (??) values were similar to those of well-watered plants and indicated a non-stomatal component as the main factor controlling photosynthesis in these leaves. Eighteen-month-old low irrigated seedlings had the highest mortality. Overall, results suggest that nursery terra rossa-acclimated Q. coccifera seedlings with improved physiological status and hydraulic soil-root continuity would have a higher survival rate in the field.     

Development of cork oak (Quercus suber L.) seedlings in response to tree shelters and mulching in northwestern Tunisia

The need for reforestation in cork oak (Quercus suber L.) areas is challenged by difficulties. Principal among these is herbivory of young plants, vegetative competition, and slow growth rates of cork oak seedlings. We evaluated the early development of cork oak seedlings treated using tree shelters and mulching in northwestern Tunisia. We tested three tree shelter treatments (non-vented, vented, and control) to shield seedlings from animal damage and five mulch types to control competing vegetation (Italian Stone Pine, Lentisk, combination of Italian Stone Pine and Lentisk (organic mulches), gravel (inorganic mulch) and no mulch). At the end of the two-year experiment, sheltered seedlings were 89-99% taller than unshelteredseedlings and had higher numbers and lengths of shoot growth flushes. In contrast, both stem diameter growth and dry weight biomass (from samples extracted after two years) were significantly reduced inside tree shelters. Root-to-shoot ratio was not significantly different in sheltered vs. unsheltered seedlings, suggesting that tree shelters do not adversely affect this parameter. Mulching alone did not favour growth, but could be beneficial when combined with tree shelters. The combination of vented tree shelters and gravel mulch was the most effective treatment for promoting diameter, height and stem volume growth.     

Soil water stress: Its effects on phenology, physiology, and morphology of containerized Douglas-fir seedlings

Containerized Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) seedlings were subjected to six moisture-stress treatments (ranging from 7 to 65% soil water content by volume) for 12 weeks. At the end of this period, there were significant differences in phenological, physiological, and morphological responses among the seedlings in the various moisture-stress treatments. In general, seedlings grown under very high or very low soil moisture conditions were adversely affected, while those grown under moderate conditions (29 to 53% soil water content) exhibited optimum growth, bud development, and nutrient and starch reserves. The use of vector analysis was found to be helpful in data interpretation. The results indicate the importance of closely monitoring nursery moisture regimes in order to achieve the best seedling quality.     

Water relations and gas exchange of Acer saccharum seedlings in contrasting natural light and water regimes

Field measurements were made of leaf photosynthesis (A), stomatal conductance (g) and leaf water relations for sugar maple (Acer saccharum Marsh.) seedlings growing in a forest understory, small gap or large clearing habitat in southwestern Wisconsin, USA. Predawn water status, leaf gas exchange and plasticity in field and laboratory water relations characteristics were compared among contrasting light environments in a wet year (1987) and a dry year (1988) to evaluate possible interactions between light and water availability in these habitats. Leaf water potentials (Psi(leaf)) at predawn and midday were lower for clearing than gap or understory seedlings. Acclimation of tissue osmotic potentials to light environment was observed among habitats but did not occur within any of the habitats in response to prolonged drought. During a summer drought in 1988, decreases in daily maximum g (g(max)) and maximum A (A(max)) in clearing seedlings were correlated with predawn Psi(leaf), which reached a seasonal minimum of -2.0 MPa. Under well-watered conditions, diurnal fluctuations in Psi(leaf) of up to 2.0 MPa in clearing seedlings occurred along with large midday depressions of A and g. In a wet year, strong stomatal responses to leaf-to-air vapor pressure difference (VPD) in sunny habitats were observed over nine diurnal courses of gas exchange measurements on seedlings in a gap and a clearing. Increasing stomatal limitations to photosynthesis appeared to be responsible for the reduction in A at high VPD for clearing seedlings. In understory seedlings, however, low water-use efficiency and development of leaf water deficits in sunflecks was related to reduced stomatal limitations to photosynthesis relative to seedlings in sunny habitats. Predawn Psi(leaf) and VPD appear to be important factors limiting carbon assimilation in sugar maple seedlings in light-saturating irradiances, primarily through stomatal closure. The overall results are consistent with the idea that sugar maple seedlings exhibit "conservative" water use patterns and have low drought tolerance. Leaf water relations and patterns of water use should be considered in studies of acclimation and species photosynthetic performance in contrasting light environments.     

Interactive effects of nitrogen and water availabilities on gas exchange and whole-plant carbon allocation in poplar

Cuttings of balsam spire hybrid poplar (Populus trichocarpa var. Hastata Henry x Populus balsamifera var. Michauxii (Dode) Farwell) were grown in sand culture and irrigated every 2 (W) or 10 (w) days with a solution containing either 3.0 (N) or 0.5 (n) mol nitrogen m(-3) for 90 days. Trees in the WN (control) and wn treatments had stable leaf nitrogen concentrations averaging 19.4 and 8.4 mg g(-1), respectively, over the course of the experiment. Trees in the Wn and wN treatments had a similar leaf nitrogen concentration, which increased from 12.0 to 15.8 mg g(-1) during the experiment. By the final harvest, mean stomatal conductances of trees in the wN and wn treatments were less than those of trees in the Wn and WN treatments (1.8 versus 4.6 mm s(-1)). Compared to the WN treatment, biomass at the final harvest was reduced by 61, 72 and 75% in the Wn, wN and wn treatments, respectively. At the final harvest, WN trees had a mean total leaf area of 4750 +/- 380 cm(2) tree(-1) and carried 164 +/- 8 leaves tree(-1) with a specific leaf area of 181 +/- 16 cm(2) g(-1), whereas Wn trees had a smaller mean total leaf area (1310 +/- 30 cm(2) tree(-1)), because of the production of fewer leaves (41 +/- 6) with a smaller specific leaf area (154 +/- 2 cm(2) g(-1)). A greater proportion of biomass was allocated to roots in Wn trees than in WN trees, but component nitrogen concentrations adjusted such that there was no Wn treatment effect on nitrogen allocation. Compared with WN trees, rates of photosynthesis and respiration per unit weight of tissue of Wn trees decreased by 28 and 31%, respectively, but the rate of photosynthesis per unit leaf nitrogen remained unaltered. The wN and Wn trees had similar leaf nitrogen concentrations; however, compared with the Wn treatment, the wN treatment decreased mean total leaf area (750 +/- 50 cm(2) tree(-1)), number of leaves per tree (29 +/- 2) and specific leaf area (140 +/- 6 cm(2) g(-1)), but increased the allocation of biomass and nitrogen to roots. Net photosynthetic rate per unit leaf nitrogen was 45% lower in the wN treatment than in the other treatments. Rates of net photosynthesis and respiration per unit weight of tissue were 48 and 33% less, respectively, in wN trees than in Wn trees.     

Ectomycorrhizal inoculation with Pisolithus tinctorius increases the performance of Quercus suber L. (cork oak) nursery and field seedlings

Mediterranean ecosystems are characterized by large arid areas where the patchy distribution of trees offers little protection against harsh climate conditions for seedling establishment. Climate change is predicted to result in an increase in these arid regions, with pronounced effects on vegetation. Production of seedlings with developed ectomycorrhizas is a promising strategy for minimizing the initial transplant shock, thereby increasing plant survival and growth during the first, most critical years of a plantation. One important species in the Mediterranean basin is Quercus suber (cork oak), which occurs, together with other evergreen oak species, in an agro-silvo-pastoral system that represents an example of sustainable land use in Europe. In this study, a Pisolithus tinctorius isolate was used for ectomycorrhizal colonization of cork oak nursery seedlings, and the effects on aboveground plant growth and leaf structural and physiological parameters were investigated. Ectomycorrhizal development resulted in a significant increase in leaf area, dry weight, nitrogen content, and photosynthetic pigments, and mycorrhizal plants showed a higher photosynthetic capacity and water use efficiency. Nursery-inoculated plants established in the field showed increased survival and growth during the first year after transplant. These results indicate a potential for further enhancing the use of mycorrhizal inoculation as a cultivation practice in forest nurseries. Considering the difficulty of soil restoration under limiting environmental conditions, nursery inoculation with ectomycorrhizal fungi can be an important advantage for improving the quality of seedling stock and its performance after out-planting in the field, benefiting the regeneration of arid regions and the reintroduction of inocula of ectomycorrhizal fungi into these areas.     

Interactive effects of shade and irrigation on the performance of seedlings of three Mediterranean Quercus species

Shade and irrigation are frequently used to increase the success of Mediterranean Quercus spp. plantations. However, there is controversy about the combined effects of these treatments on plant performance. We assessed the effects of two irradiances (full sunlight and moderate shade) and two summer watering regimes (high (daily) and low (alternate days)) on leaf and whole-plant traits of 1-year-old seedlings of Quercus coccifera, Q. ilex subsp. ballota and Q. faginea grown outdoors for 8.5 months. Leaf traits included measures of morphology, nitrogen concentration, gas exchange and photochemical efficiency, and measures of whole-plant traits included biomass allocation patterns, growth phenology, across-summer leaf area change and relative growth rate (RGR). Moderate shade reduced leaf mass per area, increased photochemical efficiency, maximum carbon assimilation rate (Amax) and allocation to leaves, and prolonged the growing period in one or more of the species. Daily watering in summer increased Amax of Q. ilex and prolonged the growing period of Q. ilex and Q. faginea. Both treatments tended to increase RGR. The effect of shade was greater in the low-watering regime than in the high-watering regime for two of the 15 studied traits, with treatment effects being independent for the remaining 13 traits. Leaf nitrogen and the ability to maintain leaf area after the arid period, rather than biomass allocation traits, explained the variation in seedling RGR. Trait responsiveness to the treatments was low and similar among species and between study scales, being unexpectedly low in Q. faginea leaves. This may be because selective pressures on leaf plasticity act differently in deciduous and evergreen species. We conclude that moderate shade and daily summer watering enhance the performance of Mediterranean Quercus seedlings through species-specific mechanisms.     

Effects of cold storage and water stress on water relations and gas exchange of white spruce (Picea glauca) seedlings

To determine the effects of lifting time and storage on water-stress resistance of nursery-grown white spruce (Picea glauca (Moench) Voss) seedlings, we compared gas exchange, water relations and mortality of 3-year-old seedlings lifted in October 1991 and stored at -2 degrees C for 3 months with seedlings lifted in January 1992. The seedlings were placed in nutrient solution and subjected to -1.1 or -2.7 MPa water stress induced by polyethylene glycol 3350 for 9 days. Water stress, but not lifting time, had a significant effect on seedling net assimilation, symplastic volume and turgor loss point. In a second experiment, seedlings lifted in October 1991 were stored at -2 degrees C for 7 months and compared with seedlings lifted in May 1992. The seedlings were planted in pots, and their gas exchange and water relation parameters measured in response to gradual water stress. The results suggest that prolonged cold storage retards photosynthetic recovery of seedlings after planting. Higher rates of net assimilation in seedlings lifted in May were not directly related to their water status. Nonstomatal limitations were the primary factor influencing photosynthetic rate. We conclude that the inferior ability of cold-stored seedlings to tolerate water stress was due to poor osmotic adjustment and a lag in recovery of photosynthesis.     

Response of Quercus robur L. seedlings to north-south asymmetry of light within gaps in floodplain forests of Slovenia

To examine Quercus robur establishment and growth in low to intermediate light levels, we analysed regeneration in different microsites created by light asymmetry within gaps in two main floodplain forest regions of Slovenia: Dolinsko and Krakovo. Four years after a mast year in 1995, we installed systematic grids of 1×1 m plots in 11 gaps (0.03–0.40 ha) on wet and dry site variants. In 256 plots, seedling species, cover, density, height, height increment, browsing damage, tree architecture, understory vegetation species and cover, and direct and diffuse light were measured. The average seedling density in all gaps was 15/m², but the highest densities were found in gap positions with low diffuse light levels (10–20%). Competing understory vegetation was more abundant in positions with high diffuse light. In gaps on dry site variants a combination of low diffuse and high direct light was favourable for regeneration. The results indicated that Q. robur can successfully establish in gaps. Here, both light components were sources of within gap resource heterogeneity, therefore knowledge of light asymmetry can improve regeneration success.     

Morphological and photosynthetic responses of Quercus crispula seedlings to high-light conditions

We investigated morphological and photosynthetic responses of current-year seedlings of oak (Quercus crispula Blume) under high-light conditions. Quercus crispula seedlings were grown from seed in a relative photosynthetically active photon flux density (RPPFD) of 100, 10 or 2%. There was no difference in total dry mass between 100 and 10% RPPFD. At the end of the growing season, plants grown in 2% RPPFD had a lower total dry mass than those grown in 100 or 10% RPPFD. Seedlings grown in 100% RPPFD showed morphological acclimation, i.e., high root/shoot ratios and high leaf mass per area. De-epoxidation level in the xanthophyll cycle and activity of an antioxidant enzyme were highest in 100% RPPFD, but total chlorophyll concentration and photosynthetic rate were highest in 10% RPPFD. These results indicate that excess photons were generated in 100% RPPFD, leading to increased capacities for dissipation of received light energy through the xanthophyll cycle and for scavenging of reactive oxygen species through the water-water cycle. Nevertheless, a midday decrease in dark-adapted quantum yield of photosystem II (F(v)/F(m)) indicated that seedlings grown in 100% RPPFD were suffering from photoinhibition. We conclude that Q. crispula current-year seedlings have high morphological acclimation to high light but that photosynthetic efficiency cannot be maintained under high-light conditions even with a photoprotection system.     

Leaf water potential and gas exchange of eucalypt clonal seedlings to leaf solar protectant

This experiment was carried out in acclimatized greenhouses with seedlings of two hybrid clones of Eucalyptus urophylla×Eucalyptus grandis. A sunscreen protector consisting of 62.5% calcium carbonate was sprayed on the seedlings at weekly intervals. Water stress was induced by suspending irrigation until the soil reached 30% available water and water was then replaced so that it returned to field capacity. Gas exchange and leaf water status were measured after 50 days. The experiment was set up in a 4×2 factorial randomized block design in four distinct environments:(1) temperatures less than 21.2℃ and vapor pressure deficit of 0.15 kPa;(2) intermediate temperatures of 24.2℃ and vapor pressure deficit of 0.69 kPa;(3) high temperatures of 27.0℃ and high vapor pressure deficit of 1.4 kPa; and,(4) high temperature of 27.0℃ and vapor pressure deficit below 1.10 kPa. Two leaf sun protector treatments were used, with five replications each. High atmospheric demand acted as a stress factor for the seedlings during the initial growth phase.Applications of leaf sunscreen protector provided beneficial effects in maintaining optimum water status and gas exchanges of the plants under water stress.     

Water-use efficiency in cork oak (Quercus suber) is modified by the interaction of water and light availabilities

We studied the interaction of light and water on water-use efficiency in cork oak (Quercus suber L.) seedlings. One-year-old cork oak seedlings were grown in pots in a factorial experiment with four light treatments (68, 50, 15 and 5% of full sunlight) and two irrigation regimes: well watered (WW) and moderate drought stress (WS). Leaf predawn water potential, which was measured at the end of each of two cycles, did not differ among the light treatments. Water-use efficiency, assessed by carbon isotope composition (delta(13)C), tended to increase with increasing irradiance. The trend was similar in the WW and WS treatments, though with lower delta(13)C in all light treatments in the WW irrigation regime. Specific leaf area increased with decreasing irradiance, and was inversely correlated with delta(13)C. Thus, changes in delta(13)C could be explained in part by light-induced modifications in leaf morphology. The relationship between stomatal conductance to water vapor and net photosynthesis on a leaf area basis confirmed that seedlings in higher irradiances maintained a higher rate of carbon uptake at a particular stomatal conductance, implying that shaded seedlings have a lower water-use efficiency that is unrelated to water availability.     

Carbon sequestration of modern Quercus suber L. silvoarable agroforestry systems in Portugal: a YieldSAFE-based estimation

Modern alley cropping designs, with trees aligned in rows and adapted to operating farming machinery, have been suggested for Europe. This paper explores the potential for adoption of cork oak (Quercus suber L.) agroforestry in Portugal and estimates the potential carbon sequestration. Spatial modeling and Portuguese datasets were used to estimate target areas where cork oak could grow on farmland. Different implementation scenarios were then modeled for this area assuming a modern silvoarable agroforestry system (113 trees ha^?1 thinned at year 20 for establishing 50 trees ha^?1). The YieldSAFE process-based model was used to predict the biomass and carbon yield of cork oak under low and high soil water holding capacity levels. Approximately 353,000 ha are available in Portugal for new cork oak alley cropping. Assuming implementation rates between 10 % of the area with low soil water capacity (60 mm: 15 cm depth, coarse texture) and 70 % of the area with high soil water holding capacity (1,228 mm: 200 cm depth, very fine texture), then carbon sequestration could be 5 × 10⁶ and 123 × 10⁶ Mg CO₂ respectively. Due to higher yields on more productive land, scenarios of limited implementation in high productivity locations can sequester similar amounts of carbon as wide implementation on low productivity land, suggesting that a priori land classification assessments can improve the targeting of land and financial incentives for carbon sequestration.     

Effects of the interaction between drought and shade on water relations, gas exchange and morphological traits in cork oak (Quercus suber L.) seedlings

The combined effect of drought and light on different physiological and biochemical traits was assessed in cork oak (Quercus suber L.) seedlings grown under two levels of light availability and submitted to a long-standing drought. Watering was withdrawn after germination and seedlings were allowed to dry to a water content of ca. 50% of field capacity. At this point, water-stressed seedlings were grown under moderate drought and two light regimes: high light (HL—50%) and low light (LL—2%). Soil water in control plants was kept close to field capacity (90–100%) for both light environments. Water-relations parameters derived from P–V curves, gas exchange and water status at predawn (Ψ_pd) were evaluated at twice during the experiment. Nitrogen and chlorophyll contents were determined in the same leaves used for the gas exchange measurements. In addition, maximum rate of carboxylation (V_cmax) and electronic transport (J_max) were derived from A–C_i curves in well-watered seedlings.

The variation on moisture availability during the experiment was the same under both light environments. In control plants, Ψ_pd was over −0.3 MPa at the two harvests, while stressed seedlings decreased to −0.9 MPa, with no differences between light treatments. Water stress decreased osmotic potentials at full (Ψπ₁₀₀) and zero turgor (Ψπ₀). The regressions between both potentials and Ψ_pd showed a higher intercept in shade grown seedlings. This fact will point out the higher osmoregulation capacity in sun seedlings whatever water availability.

Nitrogen investment on a per leaf mass (N_mass), chlorophyll content (Chl_mass) and SLA tended to show a typical pattern of sun-shade acclimation. Thus, the three parameters increased with shade. Only for N_mass there was a significant effect of watering, since water stress increased N_mass.

LL plants showed a lower photosynthetic capacity in terms of maximum net photosynthesis at saturating light (A_max), which was related to a decrease in V_cmax and J_max. Both parameters varied with specific leaf area (SLA) in a similar way. The low-light environment brought about a higher nitrogen investment in chlorophyll, while under high-light environment the investment was higher in carboxylation (V_cmax) and electronic transport (F_max).

Stomatal conductance to water vapour (g_wv) and A_max were lower in low-light seedlings independently of watering. In addition, there was a trend to keep higher intrinsic water use efficiency (IWUE) under high light environment. The increase of IWUE under water stress was higher in HL seedlings. This was as consequence of the steeper decline in g_wv as Ψ_pd decreased. The decrease of A_max with Ψ_pd occurred in a similar way in LL and HL seedlings. Thus, the HL seedlings tended to sustain a higher ability to increase IWUE than LL seedlings when they were submitted to the same water stress.