Canopy light heterogeneity drives leaf anatomical, eco-physiological, and photosynthetic changes in olive trees grown in a high-density plantation.

In the field, leaves may face very different light intensities within the tree canopy. Leaves usually respond with light-induced morphological and photosynthetic changes, in a phenomenon known as phenotypic plasticity. Canopy light distribution, leaf anatomy, gas exchange, chlorophyll fluorescence, and pigment composition were investigated in an olive (Olea europaea, cvs. Arbequina and Arbosana) orchard planted with a high-density system (1,250 trees ha(-1)). Sampling was made from three canopy zones: a lower canopy (<1 m), a central one (1-2 m), and an upper one (>2 m). Light interception decreased significantly in the lower canopy when compared to the central and top ones. Leaf angle increased and photosynthetic rates and non-photochemical quenching (NPQ) decreased significantly and progressively from the upper canopy to the central and the lower canopies. The largest leaf areas were found in the lower canopy, especially in the cultivar Arbequina. The palisade and spongy parenchyma were reduced in thickness in the lower canopy when compared to the upper one, in the former due to a decrease in the number of cell layers from three to two (clearly distinguishable in the light and fluorescence microscopy images). In both cultivars, the concentration of violaxanthin-cycle pigments and ß-carotene was higher in the upper than in the lower canopy. Furthermore, the de-epoxidized forms zeaxanthin and antheraxanthin increased significantly in those leaves from the upper canopy, in parallel to the NPQ increases. In conclusion, olive leaves react with morphological and photosynthetic changes to within-crown light gradients. These results strengthen the idea of olive trees as "modular organisms" that adjust the modules morphology and physiology in response to light intensity.

Effects of cadmium on cork oak (Quercus suber L.) plants grown in hydroponics.

Cork oak (Quercus suber L.) is an autochthonous tree species that is being used for reforestation in heavy-metal-contaminated areas in Spain. A hydroponics experiment was carried out to characterize the effects of Cd on several morphological and physiological parameters in this species, including shoot length, nutrient concentrations and allocation in different organs, leaf pigment concentrations, photosynthetic efficiency, root ferric chelate reductase (FCR) activity and organic acid concentrations in xylem sap. Four different Cd treatments were applied, adding Cd chelated with EDTA or as chloride salt at two different concentrations (10 and 50 µM Cd). After 1 month of Cd treatment, plant growth was significantly inhibited in all treatments. Results indicate that Cd accumulates in all organs 7- to 500-fold when compared with control plants. The highest Cd concentration was found in the 50 µM CdCl(2) treatment, which led to concentrations of ~30, 123 and 1153 µg Cd g(-1) dry weight in leaves, stems and roots, respectively. In the strongest Cd treatments the concentrations of P and Ca decreased in some plant parts, whereas the Mn leaf concentrations decreased with three of the four Cd treatments applied. The concentrations of chlorophyll and carotenoids on an area basis decreased, whereas the (zeaxanthin plus antheraxanthin)/(total violaxanthin cycle carotenoids) ratio and the non-photochemical quenching increased significantly in all Cd treatments. Cadmium treatments caused significant increases in the activity of the enzyme FCR in roots and in the concentrations of organic acids in xylem sap. Some of the physiological changes found support the fact that Cd induces a deficiency of Fe in cork oak, although the plant Fe concentrations were not reduced significantly. At higher concentrations the effects of Cd were more pronounced, and were more marked when Cd was in the free ion form than when present in the form of Cd-EDTA.

Changes in iron and organic acid concentrations in xylem sap and apoplastic fluid of iron-deficient Beta vulgaris plants in response to iron resupply.

In this study, the effects of Fe resupply on the composition of the xylem sap and apoplastic fluid of Fe-deficient sugar beet plants were investigated. Experiments were carried out in growth chambers with plants grown in hydroponics, and Fe resupply to Fe-deficient plants was carried out by adding 45muM Fe(III)-EDTA to the nutrient solution. In the short term (within 24h), Fe resupply caused marked changes in the xylem sap and apoplastic fluid composition and in leaf physiological parameters when de novo chlorophyll (Chl) synthesis was still beginning. Major changes included: (i) 10- and 5-fold increases in Fe concentrations in apoplastic fluid and xylem sap, respectively; (ii) marked decreases in the concentrations of organic acids in apoplastic fluid, but not in xylem sap and (iii) large decreases in the citrate/Fe ratios, both in apoplastic fluid and in xylem sap. Two to four days after Fe resupply, xylem sap and apoplastic fluid Fe and organic acid concentrations and pH reached values similar to those obtained in Fe-sufficient leaves. Leaf mesophyll ferric chelate-reductase (FC-R) activities and photosynthetic rates increased gradually during recovery from Fe deficiency.

Down co-regulation of light absorption, photochemistry, and carboxylation in Fe-deficient plants growing in different environments.

The regulation of photosynthesis through changes in light absorption, photochemistry, and carboxylation efficiency has been studied in plants grown in different environments. Iron deficiency was induced in sugar beet (Beta vulgaris L.) by growing plants hydroponically in controlled growth chambers in the absence of Fe in the nutrient solution. Pear (Pyrus communis L.) and peach (Prunus persica L. Batsch) trees were grown in field conditions on calcareous soils, in orchards with Fe deficiency-chlorosis. Gas exchange parameters were measured in situ with actual ambient conditions. Iron deficiency decreased photosynthetic and transpiration rates, instantaneous transpiration efficiencies and stomatal conductances, and increased sub-stomatal CO(2) concentrations in the three species investigated. Photosynthesis versus CO(2) sub-stomatal concentration response curves and chlorophyll fluorescence quenching analysis revealed a non-stomatal limitation of photosynthetic rates under Fe deficiency in the three species investigated. Light absorption, photosystem II, and Rubisco carboxylation efficiencies were down-regulated in response to Fe deficiency in a coordinated manner, optimizing the use of the remaining photosynthetic pigments, electron transport carriers, and Rubisco.

Fe Resupply to Fe-deficient Sugar Beet Plants Leads to Rapid Changes in the Violaxanthin Cycle and other Photosynthetic Characteristics without Significant de novo Chlorophyll Synthesis.

The effects of Fe resupply to Fe-deficient plants have been investigated in hydroponically-grown sugar beet. In the short-term (24 h) after Fe resupply, major changes were observed, although de novo chlorophyll (Chl) synthesis had not begun yet. Approximately 50% of the zeaxanthin was converted into violaxanthin, whereas the actual Photosystem II (PS II) efficiency increased by 69% and non-photochemical quenching (NPQ) and the amount of thermally dissipated energy decreased markedly (by 47% and 40%, respectively). At the same time, photosynthetic rate increased approximately by 50%. From one to two days after Fe resupply, there was a gradual increase in the leaf concentrations of Chl and other photosynthetic pigments, accompanied by a further conversion of zeaxanthin into violaxanthin, increases in actual PS II efficiency and photosynthetic rates and decreases in NPQ and the amount of thermally dissipated energy. At 72-96 h after Fe resupply, leaf pigment concentrations, photosynthetic rates and actual PS II efficiency had increased further, although both photosynthetic rate and leaf pigment concentrations were still lower than those found in Fe-sufficient leaves. Good correlations were observed between the amount of light thermally dissipated by the PS II antenna, NPQ and the antheraxanthin + zeaxanthin concentration after Fe resupply, confirming the photoprotective role of the xanthophyll cycle in Fe-deficient sugar beet leaves. Similar correlations were observed for lutein, suggesting a possible role of this pigment in photoprotection.

Effects of branch solid Fe sulphate implants on xylem sap composition in field-grown peach and pear: changes in Fe, organic anions and pH.

The effects of placing solid implants containing Fe sulfate in branches of Fe-deficient pear and peach trees on the composition of the xylem sap have been studied. Iron sulfate implants are commercially used in northeastern Spain to control iron chlorosis in fruit trees. Implants increased Fe concentrations and decreased organic acid concentrations in the xylem sap, whereas xylem sap pH was only moderately changed. The citrate to Fe ratios decreased markedly after implants, therefore improving the possibility that Fe could be reduced by the leaf plasma membrane enzyme reductase, known to be inhibited by high citrate/Fe ratios. In peach, the effects of the implants could be observed many months post treatment. In pear, some effects were still observed one year after the implants had taken place. Results obtained indicate that solid Fe sulfate implants were capable of significantly changing the chemical composition of the xylem sap in fruit trees.

Effects of Cd and Pb in sugar beet plants grown in nutrient solution: induced Fe deficiency and growth inhibition.

Effects of Cd and Pb toxicity were investigated in sugar beet (Beta vulgaris L.) grown in hydroponics under growth-chamber-controlled conditions. Chemical speciation calculations were used to estimate the chemical species in equilibrium. Cd, used as chloride salt or chelated to EDTA, decreased fresh and dry mass of both root and shoot, and increased root / shoot ratios. Plants developed few brownish roots with short laterals. Cd decreased N, P, Mg, K, Mn, Cu and Zn uptake, and facilitated Ca uptake. Leaves of plants treated with 10 or 50 µM Cd-EDTA and 10 µM CdCl2 developed symptoms of Fe deficiency. These symptoms included decreased leaf chlorophyll (Chl) and carotenoid concentrations, increased carotenoid / Chl and Chl a/b ratios, de-epoxidation of violaxanthin cycle pigments, and decreased photosynthetic rates and PSII efficiency. Plants treated with 50 µM CdCl2, however, had decreased growth but did not show marked leaf Fe-deficiency symptoms. All Cd treatments increased Fe(III)-chelate reductase activity in root tips, although Fe concentrations in shoots were similar to those found in control plants. Pb chelated with EDTA induced visual symptoms only at concentrations of 2 mM. Leaves of Pb-treated plants remained green and their edges were rolled inwards. Pb increased root fresh and dry mass with no changes in shoot mass, therefore increasing the root / shoot ratio. Changes in plant nutrient concentrations with Pb were only minor, although leaf Cu levels approached critical deficiency levels. No symptoms of Fe deficiency were apparent in leaves. Root tips of Pb-treated plants, however, had increased Fe(III)-chelate reductase activities.