Photosynthesis in response to sink-source activity and in relation to end products and activities of metabolic enzymes in peach trees

Fruits were removed (-fruit) or retained (+fruit) on limbs of 'Yanfengyihao' peach (Prunus persica (L.) Batch) trees. In addition to the -fruit and +fruit treatments, leaves on 1-year-old fruit-bearing shoots of 'Bayuecui' peach trees were covered with bags for varying periods of time. Net photosynthetic rate (P(n)), stomatal conductance and transpiration rate around midday significantly decreased in the -fruit treatment and significantly increased in bagged leaves after removal of the bags compared with values in the +fruit treatment. Leaf temperature (T(l)) was higher in the -fruit treatment than in the +fruit treatment, and P(n) decreased in the -fruit treatment, especially when T(l) exceeded 38 and 36 degrees C in 'Yanfengyihao' and 'Bayuecui', respectively. Fruit removal resulted in a significant accumulation of sorbitol and starch in leaves, but it did not decrease activities of enzymes involved in carbon metabolism in leaves of 'Yanfengyihao'. In contrast, activities of aldose-6-phosphate reductase and ADP-glucose pyrophosphorylase sometimes increased in leaves in the -fruit treatment. In 'Bayuecui', fruit removal significantly decreased maximal fluorescence and photochemical efficiency and increased minimal fluorescence around midday compared with values in the +fruit and bag treatments. The photosynthetic apparatus of peach may be damaged when T(l) exceeds a critical temperature under conditions of decreased sink demand. Decreased stomatal aperture and increased T(l) may regulate photosynthesis by affecting electron transport.     

Drought-induced oxidative stress in Canarian laurel forest tree species growing under controlled conditions

We studied photoprotection and antioxidative protection in the three major species of the Canarian laurel forest (Laurus azorica (Seub.) Franco, Persea indica (L.) K. Spreng and Myrica faya Aiton). Trees were exposed to drought under controlled conditions by withholding water until leaf relative water content (RWC) reached 50-55%. Drought reduced photosynthetic rate (P(N)) and was associated with decreased quantum yield of photosystem II (PSII) electron transport and increased non-photochemical quenching (NPQ) in L. azorica and M. faya, but did not increase NPQ in P. indica. Drought-treated trees of L. azorica had the highest de-epoxidation state (DPS) of the xanthophyll cycle and the highest zeaxanthin (Z) concentration, suggesting that this species had more effective photoprotective mechanisms than M. faya and P. indica. Moreover, beta-carotene remained unaltered in L. azorica trees during drought, suggesting that the chloroplasts of this species are better protected against oxidative stress than those of M. faya and P. indica. Increased antioxidation by ascorbate peroxidase, superoxide dismutase and glutathione reductase in L. azorica removed activated oxygen species (AOS) generated during drought treatment. Although M. faya was able to increase its energy dissipation rate by forming Z and thus increasing the DPS of the xanthophyll cycle, it did not respond to drought-induced oxidative stress with the result that beta-carotene degradation occurred. Persea indica did not activate an energy dissipation mechanism in response to drought treatment, hence formation of AOS was likely high in the drought-treated trees. In general, L. azorica was most resistant and P. indica most sensitive to photoinhibition and oxidative stress during drought.     

Girdling effects on fruit set and quantum yield efficiency of PSII in two Citrus cultivars

Girdling effects on fruitlet abscission, leaf chlorophyll, chlorophyll a fluorescence and carbohydrate concentration in various flowering and vegetative shoots were studied during natural fruit drop in two Citrus cultivars. Irrespective of shoot type, girdling delayed fruitlet abscission, but only fruitlets borne on leafy shoots had increased final fruit set. Chlorophyll a fluorescence analysis revealed differences in quantum yield efficiency of photosystem II of light adapted leaves (Phi(PSII)) among shoot types and in response to girdling. In young leaves of vegetative shoots, girdling decreased Phi(PSII), whereas Phi(PSII) increased from Day 30 after girdling in young leaves of leafy flowering shoots; however, Phi(PSII) did not change in mature leaves during fruit set in either control or girdled trees. Girdling altered leaf carbohydrate concentrations and the photosynthetic performance of the various shoot types. Our results indicate that, in Citrus, several carbohydrate-based regulatory mechanisms of photosynthesis coexist during carbohydrate accumulation brought about by girdling. It is concluded that the delay in fruitlet abscission and the increase in Phi(PSII )observed in girdled leafy flowering shoots are the mechanisms underlying the enhancement of fruit set after girdling.     

Increase of nitrogen to promote growth of poplar seedlings and enhance photosynthesis under NaCl stress

The solution culture method was used to study the effect of increasing nitrogen on the growth and pho-tosynthesis of poplar seedlings under 100 mmol L-1 NaCl stress. I Increase in nitrogen reduced stomatal limitation of leaves under NaCl stress, improved utilization of CO2 by mesophyll cells, enhanced photosynthetic carbon assimi-lation capacity, significantly alleviated saline damage of NaCl, and promoted the accumulation of aboveground and root biomass. I Increased nitrogen enhanced photochemical efficiency (ΦPSⅡ) and electron transport rates, relieved the reduction of maximum photochemical efficiency (Fv/Fm) under NaCl, and reduced the degree of photoinhibition caused by NaCl stress. Increased nitrogen applications reduced the proportion of energy dissipating in the form of ineffective heat energy and hence a greater proportion of light energy absorbed by leaves was allocated to photo-chemical reactions. Under treatment with increased nitro-gen, the synergistic effect of heat dissipation and the xanthophyll cycle in the leaves effectively protected pho-tosynthetic PSⅡ and enhanced light energy utilization of leaves under NaCl stress. The increased nitrogen promoted photosynthetic electron supply and transport ability under NaCl stress evident in enhanced functioning of the oxygen-evolving complex on the electron donor side of PS Ⅱ. It increased the ability of the receptor pool to accept electrons on the PSII electron acceptor side and improved the sta-bility of thylakoid membranes under NaCl stress. Therefore, increasing nitrogen applications under NaCl stress can promote poplar growth by improving the effi-ciency of light energy utilization.     

Physiological and biochemical leaf and tree responses to crop load in apple

Seven-year-old apple (Malus x domestica Borkh.) trees cv. 'Braeburn' on rootstock M.26 were flower-thinned to establish four crop loads, resulting in final mean fruit numbers per tree of 0, 100, 225 and 400. Mean fruit mass decreased by about 35% with each decrease in cropping density. Fruit from light-cropping trees had significantly advanced maturity as indicated by the harvest management criteria of background color and starch/iodine score, and other fruit quality characteristics such as soluble solids. Flesh firmness and dry matter also increased with decreasing crop load. Compared with fruiting trees, mean leaf photosynthetic rates of non-cropping trees were significantly lower (40%) between 75 days after full bloom (dafb) and fruit harvest, with a maximum reduction of almost 60% at 118 dafb. Photosynthetic activity decreased linearly with increasing concentration of leaf starch, but was positively and significantly related to stomatal conductance. Consequently, the accumulation of nonstructural carbohydrates in leaves of light-cropping or non-cropping trees may have led to end-product inhibition of photosynthesis. Increases in xanthophyll cycle carotenoids mediated non-radiative thermal energy dissipation in non-cropping trees, providing increased capacity for photoprotection but reducing photochemical efficiency.     

Hyperhydricity in apple: ultrastructural and physiological aspects

We studied the effects of hyperhydricity on subcellular ultrastructure and physiology of leaves during in vitro regeneration of apple plants. Morphological, anatomical and ultrastructural differences between healthy leaf tissues obtained from greenhouse-grown plants and healthy and hyperhydric leaves obtained from shoots raised from nodal shoot explants in a bioreactor were investigated by electron microscopy and confocal laser scanning microscopy. Compared with healthy leaves, hyperhydric leaves showed abnormal, often discontinuous development of the epidermis and cuticle. Stomata were malformed. The leaf lamina appeared thickened and was characterized by poor differentiation between the palisade and spongy mesophyll tissue. Hyperhydric leaves had a significantly lower chloroplast number per cell and chloroplasts showed reduced thylakoid stacking compared with healthy leaves. Hyperhydricity resulted in a general decrease in concentrations of reduced and oxidized pyridine nucleotides, reflecting a reduction in metabolic activity. The activities of antioxidant enzymes, such as superoxide dismutase, catalase, ascorbate peroxidase and glutathione reductase were higher in hyperhydric leaves than in healthy leaves, indicating that hyperhydricity was associated with oxidative stress. Chlorophyll fluorescence measurements provided evidence of oxidative damage to the photosynthetic machinery in hyperhydric leaves: photochemical efficiency of photosystem II, effective quantum efficiency and photochemical quenching were all lower in hyperhydric leaves compared with healthy leaves.     

Response of photosynthesis in second-generation Pinus radiata trees to long-term exposure to elevated carbon dioxide partial pressure

Second-generation Pinus radiata D. Don trees, propagated from cuttings of 4-year-old trees previously grown at ambient (36 Pa) and elevated (65 Pa) CO2 partial pressure (Ca) were grown under the same conditions in open-top chambers for a further year. As cuttings of the original trees, these second-generation trees were physiologically the same age as the first-generation trees with the only difference between the two being size. This allowed us to test the effects of tree size independently of age or duration of exposure. Total non-structural carbohydrate concentration, area-based nitrogen concentration, leaf mass per unit area and chlorophyll concentration measured in three foliage age cohorts were unaffected by either age or Ca. There were no signs of photosynthetic down-regulation in trees grown at elevated Ca. When measured at the growth Ca, photosynthetic rate in young needles during summer, autumn and spring was 34, 43 and 38% higher, respectively, in trees grown at elevated Ca than in trees grown at ambient Ca. In older needles, the corresponding photosythetic rate increases were 26, 47 and 49%. Water-use efficiency, determined by stable carbon isotope analysis, was 49% higher in foliage in the elevated Ca treatment than in foliage in the ambient Ca treatment. This increase was entirely due to photosynthetic enhancement, because stomatal conductance did not differ between treatments. We conclude that down-regulation of photosynthesis at elevated Ca is related to tree size rather than tree age or duration of exposure, and that enhanced photosynthetic rates can be maintained while sink strength is high enough to use the excess photosynthates.elevated CO2, needle age, photosynthetic down-regulation, photosynthetic enhancement, sink strength, water-use efficiency.     

Soluble sugars mediate sink feedback down-regulation of leaf photosynthesis in field-grown Coffea arabica

Source-sink relationships of field-grown plants of Coffea arabica L. cultivar 'Caturra' were manipulated to analyze the contribution of soluble sugars to sink feedback down-regulation of maximal leaf net CO2 assimilation rate (Amax). Total soluble sugar concentration (SSCm) and Amax were measured in the morning and afternoon on mature leaves of girdled branches bearing either high or low fruit loads. Leaf Amax was negatively correlated to SSCm, increased with fruit load and decreased during the day, indicating that limiting sink demand for carbohydrates caused SSCm to accumulate in the leaf tissue which results in down-regulation of Amax. To further analyze source-sink feedback on Amax, we compared Amax of mature, non-sink-limited coffee leaves fed with water or sucrose for 5, 10 or 30 min with that of non-fed control leaves. Sucrose-feeding reduced Amax compared with the control and water-feeding treatments, indicating that down-regulation of Amax is related to phloem sucrose concentration in coffee source leaves, independent of SSCm concentration in other leaf tissues. Although sucrose appeared to be more closely related to the mechanism underlying sink feedback down-regulation of Amax in coffee leaves than SSCm, Amax was closely related to SSCm by a nonlinear equation that may be useful for integrating sink limitations in coffee leaf photosynthetic models.     

Fruit load and branch ring-barking affect carbon allocation and photosynthesis of leaf and fruit of Coffea arabica in the field

Increasing fruit load (from no berries present to 25, 50 and 100% of the initial fruit load) significantly decreased branch growth on 5-year-old coffee (Coffea arabica L.) trees of the dwarf cultivar 'Costa Rica 95', during their third production cycle. Ring-barking the branches further reduced their growth. Berry dry mass at harvest was significantly reduced by increasing fruit load. Dry matter allocation to berries was four times that allocated to branch growth during the cycle. Branch dieback and berry drop were significantly higher at greater fruit loads. This illustrates the importance of berry sink strength and indicates that there is competition for carbohydrates between berries and shoots and also among berries. Leaf net photosynthesis (P(n)) increased with increasing fruit load. Furthermore, leaves of non-isolated branches bearing full fruit load achieved three times higher P(n) than leaves of isolated (ring-barked) branches without berries, indicating strong relief of leaf P(n) inhibition by carbohydrate demand from berries and other parts of the coffee tree when excess photoassimilates could be exported. Leaf P(n) was significantly higher in the morning than later during the day. This reduction in leaf P(n) is generally attributed to stomatal closure in response to high irradiance, temperature and vapor pressure deficit in the middle of the day; however, it could also be a feedback effect of reserves accumulating during the morning when climatic conditions for leaf P(n) were optimal, because increased leaf mass ratio was observed in leaves of ring-barked branches with low or no fruit loads. Rates of CO(2) emission by berries decreased and calculated photosynthetic rates of berries increased with increasing photosynthetic photon flux (PPF) especially at low PPFs (0 to 100 micromol m(-2) s(-1)). The photosynthetic contribution of berries at the bean-filling stage was estimated to be about 30% of their daily respiration costs and 12% of their total carbon requirements at PPF values commonly experienced in the field (200 to 500 micromol m(-2) s(-1)).     

The effect of defruiting at different stages of fruit development on leaf photosynthesis of "Golden Delicious" apple

Net photosynthetic rates (A) of leaves on 11-year-old, field-grown apple trees (Malus domestica Borkh. cv. Golden Delicious) were measured after removal of fruits at four different stages of development. Defruiting decreased A by 21, 42, 27 and 7% when fruits were growing at 311, 293, 229 and 113 mg(DW) day(-1), respectively. Photosynthesis was inhibited more in the afternoon than in the morning, but it was not affected during the first 8 h after fruit removal. Inhibition of A was positively correlated with crop sink strength, but it was not correlated with fruit relative growth rate or crop load. Defruiting decreased A at saturating irradiances (PPFD > 1000 micro mol m(-2) s(-1)), but did not modify the apparent quantum yield of single leaves. These results suggest that the overall effect of defruiting on carbon fixation is negligible in dense canopies, but it may be significant in sparse canopies and in single shoots.     

Influence of canopy light environment and nitrogen availability on leaf photosynthetic characteristics and photosynthetic nitrogen-use efficiency of field-grown nectarine trees

Relationships between CO(2) assimilation at light saturation (A(max)), nitrogen (N) content and weight per unit area (W(A)) were studied in leaves grown with contrasting irradiances (outer canopy versus inner canopy) and N supply rates in field-grown nectarine trees Prunus persica L. Batsch. cv. Fantasia. Both A(max) and N content per unit leaf area (N(A)) were linearly correlated to W(A), but leaves in the high-N treatment had higher N(A) and A(max) for the same value of W(A) than leaves in the low-N treatment. The curvilinear relationship between photosynthesis and total leaf N was independent of treatments, both when expressed per unit leaf area A(maxA) and N(A)) and per unit leaf weight (A(maxW) and N(W)), but the relationship was stronger when data were expressed on a leaf area basis. Both A(maxA) and N(A) were higher for outer canopy leaves than for inner canopy leaves and A(maxW) and N(W) were higher for leaves in the high-N treatment than for leaves in the low-N treatment. The relationship between A(max) and N resulted in a similar photosynthetic nitrogen-use efficiency at light saturation (A(max)NUE) for both N and light treatments. Photosynthetic nitrogen-use efficiency was similar among treatments throughout the whole light response curve of photosynthesis. Leaves developed in shade conditions did not show higher N-use efficiency at low irradiance. At any intercellular CO(2) partial pressure (C(i)), photosynthetic CO(2) response curves were higher for outer canopy leaves and, within each light treatment, were higher for the high-N treatments than for the low-N treatments. Consequently, most of the differences among treatments disappeared when photosynthesis was expressed per unit N. However, slightly higher assimilation rates per unit N were found for outer canopy leaves compared with inner canopy leaves, in both N treatments. Because higher daily irradiance within the canopies of the low-N trees more than compensated for the lower photosynthetic performances of these leaves compared to the leaves of high-N trees, daily carbon gain (and N-use efficiency on a daily assimilation basis) per leaf was higher for the low-N treatment than for the high-N treatment in both outer and inner canopy leaves.     

Transgenic poplar characterized by ectopic expression of a pine cytosolic glutamine synthetase gene exhibits enhanced tolerance to water stress

Physiological responses to water stress in hybrid poplar (INRA 7171-B4, Populus tremula L. x P. alba L.) lines transformed to overexpress a pine cytosolic glutamine synthetase (GS1) gene were compared with those of non-transgenic plants. Before, during and after a drought treatment, net photosynthetic rates (Anet) were higher in transgenic than in non-transgenic plants. Stomatal conductance (gs) was higher in transgenic than in non-transgenic plants before, but not after exposure to drought. Before drought treatment, a sudden reduction in photosynthetic photon flux caused a greater burst of CO2 efflux in transgenic than non-transgenic plants, indicating greater photorespiratory activity. Drought caused greater reductions in photochemical quenching, photosystem II (PSII) antennae transfer efficiency (Fv'/Fm') and light-adapted PSII yield (PhiPSII) in non-transgenic than in transgenic plants, especially at low irradiances. Antennae-based thermal dissipation was higher in transgenic plants than in non-transgenic plants both during the imposition of drought and 1 or 3 days after the relief of drought. Under severe water stress and subsequently, transgenic plants maintained a higher expression of glutamine synthetase, glutamate synthase and Rubisco and higher concentrations of chlorophyll and glycine than non-transgenic plants. These findings indicate that overexpression of pine cytosolic GS1 enhanced sustained photosynthetic electron transport capacity during severe stomatal limitation. The data also suggest that ectopic expression of cytosolic GS increases photorespiratory activity, and that this serves as a protective sink for electrons from photosynthetic reaction centers.     

Effects of fruit thinning on ascorbate–glutathione cycle metabolism in black currants (<Emphasis Type="Italic">Ribes nigrum</Emphasis> L.)

To elucidate mechanisms regulating ascorbic acid (AsA) biosynthesis and accumulation in the fruit and leaves of black currants, AsA and the activities of key enzymes in the ascorbate–glutathione (AsA–GSH) cycle were measured from fruit set to fruit ripening during fruit thinning treatments of three common commercial black currant cultivars that differed in their AsA levels: ‘Risager’ (low), ‘Brodtrop’ (medium) and ‘Adelinia’ (high). Treatments were 50% fruit reduction (50% of total fruit set) by hand, control was no thinning. Fruit thinning treatment significantly increased AsA content in fruit of all three cultivars from weeks 2 to 8, significantly decreased AsA content in leaves from weeks 3 to 8. Dehydroascorbate reductase and monodehydroascorbate reductase activities in fruit and leaves had a similar pattern, increasing during week 2, rose until they peaked in week 4. Ascorbate peroxidase activity in fruit in the thinning treatment was slightly lower than in the control. Fruit thinning was shown to be a good model to test AsA biosynthesis regulation and accumulation in black currants. The results from our study provided strong evidence that AsA–GSH cycle involved in AsA synthesis and accumulation in fruit.     

Carbon and nitrogen assimilation in red oaks (Quercus rubra L.) subject to defoliation and nitrogen stress

To examine how rates of net photosynthesis and N uptake of red oak seedlings respond to defoliation under contrasting conditions of N availability, nitrogen-deficient plants were grown in sand culture and subjected to partial defoliation and increased N availability under low light conditions. Both photosynthesis and N uptake rates were measured regularly before and after the treatments. Defoliation resulted in elevated rates of net photosynthesis in both low-N and high-N trees, but the high-N trees were able to maintain the high photosynthetic rates for a longer period of time. Nitrogen availability did not affect the photosynthetic rate of the undefoliated plants. Nitrogen uptake was not affected by the defoliation treatment, but was increased by increasing N availability in both the defoliated and undefoliated plants. Nitrogen uptake rates increased less than would be expected on the basis of N availability alone, but the uptake rates were apparently not limited by carbon supply in the short term. Suboptimal concentrations of N in plant tissues resulted in a strong sink for N even in the absence of refoliation.     

Tree- and needle-age-dependent variations in antioxidants and photoprotective pigments in Norway spruce needles at the alpine timberline

To cope with environmental stress, plants are equipped with antioxidative (e.g., ascorbate, glutathione and alpha-tocopherol) and photoprotective (e.g., xanthophyll cycle pigments) defense systems. We investigated the defense capacities of three tree age classes (mature, sapling and seedling) of Norway spruce (Picea abies (L.) Karst.) at a field site near the timberline. Biochemical data were expressed on both a needle dry mass and a surface area basis. Compared with current-year needles, previous-year needles contained higher mass- and area-based concentrations of chlorophylls and alpha-tocopherol, and a larger xanthophyll cycle pool that was in a more epoxidized state. Total glutathione concentration was lower, the glutathione pool was more reduced and the ascorbate pool was more oxidized in previous-year needles than in current-year needles. Needle concentrations of glutathione and alpha-tocopherol increased and chlorophyll concentration decreased with increasing tree age when expressed on a surface area basis. On a dry mass basis, these trends were reversed or nonexistent. The ascorbate pool was more reduced and the glutathione pool was more oxidized in needles of mature trees than in needles of saplings and seedlings. The proportion of protective xanthophyll cycle pigments decreased and the de-epoxidation state increased with increasing tree age. We conclude that tree age and the basis of expression of antioxidant concentration--surface area or dry mass--are important in scaling from seedlings to large trees.     

A recommended rate for application of Chaetomium globosum ND35 fungus fertilizer on poplar plantations in China

Previous studies showed that Chaetomium globosum ND35 fungus fertilizer can improve the microbial community structure and enzyme activities of replanted soil.However, it remains unclear whether can improve the physiological and ecological characteristics of plants under successive rotation.In this study, we investigated the photosynthetic, physiological, and biochemical indexes including photosynthetic parameters, chlorophyll fluorescence, and chlorophyll content of 1-year-old poplar seedlings under seven different doses(range from 0 to1.67 g kg~(-1)) of C.globosum ND35 fungus fertilizer to study the effects of fungus fertilizer on photosynthesis of Poplar.Our results showed that:(1) With increasing application of fungus fertilizer in replanted soil, chlorophyll content of poplar leaves(Chl) increased, while physiological indexes such as electron transport rate(ETR),net photosynthetic rate(P_n), quantum efficiency(U),nitrate reductase(NR) activity and root vigor initially increased and then declined.Meanwhile, heat dissipation that depended on the xanthophyll cycle declined and nonphotochemical quenching(NPQ) initially increased and then decreased.When the dose of C.globosum ND35 fungus fertilizer was 0.67 g kg~(-1)(T3) and 1.00 g kg~(-1)(T4), excess light energy of photosynthetic apparatus was reduced, and photosynthetic apparatus distributed more light energy to the direction of photochemical reactions,which improved the efficiency of energy use.Plant height and biomass of leaves, stems, and roots were maximum at T3.We conclude that applying appropriate amounts of C.globosum ND35 fungus fertilizer can improve root physiological activity and capacity for use of light by poplar leaves.This can improve the operating states of the photosynthetic apparatus and lead to increased photosynthetic efficiency of poplar leaves and accumulation of dry matter.This suggests a strategy to alleviate the successive rotation obstacle of soil nutrient depletion.     

Modeling photosynthesis in olive leaves under drought conditions

We quantified parameters for a model of leaf-level photosynthesis for olive, and tested the model against an independent dataset. Specific temperature-dependence parameters of the model for olive leaves were measured, as well as the relationship of the model parameters with area-based leaf nitrogen (N) content. The effect of soil water deficit on leaf photosynthesis was examined by applying two irrigation treatments to 29-year-old trees growing in a plantation: drip irrigation sufficient to meet the crop water requirements (I) and dry-farming (D). In both treatments, leaves had a higher photosynthetic capacity in April than in August. In August, photosynthetic capacity was lower in D trees than in I trees. Leaf photosynthetic capacity was linearly and positively related to leaf N content on an area basis (N(a)) and to leaf mass per unit area (LMA), and the regression slope varied with irrigation treatment. The seasonal reduction in N(a) was used in the model to predict photosynthesis under drought conditions. Olive leaves showed a clear limitation of photosynthesis by triose phosphate utilization (TPU) even at 40 degrees C, and the data suggest that olive invests fewer resources in TPU than other species. The seasonal decrease in photosynthetic capacity moderated the stomatal limitation to carbon dioxide (CO(2)) fixation as soil water deficit increased. Further, it enabled leaves to operate close to the transition point between photosynthetic limitation due to RuBP carboxylation capacity and that due to RuBP regeneration capacity, and resulted in a near constant value of internal CO(2) concentration from April to August. Under well watered conditions, N-use efficiency of the olive leaves was enhanced at the expense of reduced water-use efficiency.     

Multivariate patterns of biochemical responses of Pinus ponderosa trees at field plots in the San Bernardino Mountains, southern California

Most environmental stress conditions promote the production of potentially toxic active oxygen species in plant cells. Plants respond with changes in their antioxidant and photoprotective systems. Antioxidants and pigments have been widely used to measure these responses. Because trees are exposed to multiple man-made and natural stresses, their responses are not reflected by changes in single stress markers, but by complex biochemical changes. To evaluate such response patterns, explorative multivariate statistics have been used. In the present study, 12 biochemical variables (chloroplast pigments, state of the xanthophyll cycle, alpha-tocopherol, ascorbate and dehydroascorbate, glutathione and oxidized glutathione) were measured in previous-year needles of field-grown Pinus ponderosa Dougl. ex Laws. The trees were sampled in two consecutive years in the San Bernardino Mountains in southern California, where a pollution gradient is overlaid by gradients in natural stresses (drought, altitude). To explore irradiance effects, needle samples were taken directly in the field (sun exposed) and from detached, dark-adapted branches. A principal component analysis on this data set (n = 80) resulted in four components (Components 1-4) that explained 67% of the variance in the original data. Component 1 was positively loaded by concentrations of alpha-tocopherol, total ascorbate and xanthophyll cycle pools, as well as by the proportion of de-epoxides in the xanthophyll cycle. It was negatively loaded by the proportion of dehydroascorbate in the ascorbate pool. Component 2 was negatively loaded by chlorophyll concentrations, and positively loaded by the ratios of lutein and beta-carotene to chlorophyll and by the de-epoxidation state of the xanthophyll cycle. Component 3 was negatively loaded by GSH concentrations and positively loaded by the proportions of GSSG and tocopherol concentrations. Component 4 was positively loaded by neoxanthin and negatively loaded by beta-carotene. The four components could be assigned to the concerted action of the biochemical protection system: high scores on Component 1 represent highly activated antioxidative defense, changes in pigment composition are represented in Components 2 and 4, and the glutathione system, which is important for antioxidant regeneration, is represented in Component 2. Although Component 1 scores were generally higher (indicating activation of antioxidant defense) in light-adapted needles relative to dark-adapted needles, they were also site dependent with increased scores at sites with less pollution, but higher natural stress impacts. High scores of Components 2 and 3 at the highest elevation site, which was only moderately polluted, indicated an increase in photoprotection by pigments and activation of the glutathione system. Significant differences between light- and dark-adapted needles in Components 2 and 3 were only found at the site with the highest pollution. Use of accumulated variables (components) instead of single biochemical variables enabled recognition of response patterns at particular sites and a better comparison with results of other studies is expected. Typical response patterns could be assigned to particular environmental stress combinations, providing a means of assessing potential biological risks within individual forest stands.     

Acclimation of shade-developed leaves on saplings exposed to late-season canopy gaps

We hypothesized that photoinhibition of shade-developed leaves of deciduous hardwood saplings would limit their ability to acclimate photosynthetically to increased irradiance, and we predicted that shade-tolerant sugar maple (Acer saccharum Marsh.) would be more susceptible to photoinhibition than intermediately shade-tolerant red oak (Quercus rubra L.). After four weeks in a canopy gap, photosynthetic rates of shade-developed leaves of both species had increased in response to the increase in irradiance, although final acclimation was more complete in red oak. However, photoinhibition occurred in both species, as indicated by short-term reductions in maximum rates of net photosynthesis and the quantum yield of oxygen evolution, and longer-term reductions in the efficiency of excitation energy capture by open photosystem II (PSII) reaction centers (dark-adapted F(v)/F(m)) and the quantum yield of PSII in the light (phi(PSII)). The magnitude and duration of this decrease were greater in sugar maple than in red oak, suggesting greater susceptibility to photoinhibition in sugar maple. Photoinhibition may have resulted from photodamage, but it may also have involved sustained rates of photoprotective energy dissipation (especially in red oak). Photosynthetic acclimation also appeared to be linked to an ability to increase leaf nitrogen content. Limited photosynthetic acclimation in shade-developed sugar maple leaves may reflect a trade-off between shade-tolerance and rapid acclimation to a canopy gap.     

Leaf morphological and physiological adjustments to the spectrally selective shade imposed by anthocyanins in Prunus cerasifera

Prunus domestica L. has green leaves, whereas Prunus cerasifera Ehrh. var. atropurpurea has red leaves due to the presence of mesophyll anthocyanins. We compared morphological and photosynthetic characteristics of leaves of these species, which were sampled from shoots grafted in pairs on P. domestica rootstocks, each pair comprising one shoot of each species. Two hypotheses were tested: (1) anthocyanins protect red leaves against photoinhibition; and (2) red leaves display shade characteristics because of light attenuation by anthocyanins. Parameters were measured seasonally, during a period of increasing water stress, which caused a similar drop in shoot water potential in each species. As judged by predawn measurements of maximum PSII yield, chronic photoinhibition did not develop in either species and, despite the anthocyanic screen, the red leaves of P. cerasifera displayed lower light-adapted PSII yields and higher non-photochemical quenching than the green leaves of P. domestica. Thus, it appears that, in this system, anthocyanins afford little photoprotection. As predicted by the shade acclimation hypothesis, red leaves were thinner and had a lower stomatal frequency, area- based CO2 assimilation rate, apparent carboxylation efficiency and chlorophyll a:b ratio than green leaves. However, red leaves were similar to green leaves in conductivity to water vapor diffusion, dry-mass-based chlorophyll concentrations and carotenoid:chlorophyll ratios. The data for red leaves indicate adaptations to a green-depleted, red-enriched shade, rather than a neutral or canopy-like shade. Thus, green light attenuation by anthocyanins may impose a limitation on leaf thickness. Moreover, the selective depletion of light at wavelengths that are preferentially absorbed by PSII and chlorophyll b may lead to adjustments in chlorophyll pigment ratios to compensate for the uneven spectral distribution of internal light. The apparent photosynthetic cost associated with lost photons and reduced leaf thickness, and the absence of a photoprotective advantage, suggest that there are other, yet to be identified, benefits for permanently anthocyanic leaves of P. cerasifera.