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Thayne Montague*

Granier style thermal dissipation probes (TDP) have been used to estimate whole plant water loss on a variety of tree and vine species. However, studies using TDPs to investigate water loss of landscape tree species is rare. This research compared containerized tree water loss estimates of three landscape tree species using TDPs with containerized tree water loss estimates as measured by load cells. Over a three-year period, established, 5.0 cm caliper Bradford pear (Pyrus calleryana `Bradford'), English oak (Quercus robar), and sweetgum (Liquidambar styraciflua `Rotundiloba') trees in 75 L containers were placed on load cells, and water loss was measured for a 60-d period. One 3.0 cm TDP was placed into the north side of each trunk 30 cm above soil level. To reduce evaporation, container growing media was covered with plastic. Each night, plants were irrigated to soil field capacity and allowed to drain. To provide thermal insulation TDPs and tree trunks (up to 30 cm) were covered with aluminum foil coated bubble wrap. Hourly TDP water loss estimates for each species over a three-day period indicate TDP estimated water loss followed a similar trend as load cell estimated water loss. However, TDP estimates were generally less, especially during peak transpiration periods. In addition, mean, total daily water loss estimates for each species was less for TDP estimated water loss when compared to load cell estimated water loss. Although TDP estimated water loss has been verified for several plant species, these data suggest potential errors can arise when using TDPs to estimate water loss of select landscape tree species. Additional work is likely needed to confirm estimated sap flow using TDPs for many tree species.

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Guohai Xia and Lailiang Cheng

Four-year-old `Gala'/M.26 trees were grown under low (2.5 mm), medium (12.5 mm), or high (25 mm) N supply with balanced nutrients in sand culture and the cropload was adjusted to 5 fruit/cm2 trunk cross-sectional area at 10 mm king fruit. At about 100 days after bloom, exposed fruit on the south side of the canopy were chosen for monitoring chlorophyll fluorescence and fruit peel samples were taken for measuring xanthophyll cycle pigments, antioxidant enzymes, and metabolites. At noon, the efficiency of excitation transfer (Fv'/Fm') of the sun-exposed peel was higher in the low N treatment than in the medium or high N treatments. Photochemical quenching coefficient did not differ between fruits in different N treatments. The Photosystem II operating efficiency was higher in the peel of low N fruit compared with medium N or high N fruit. However, maximum quantum efficiency (Fv/Fm) of fruit peel after overnight dark adaptation was similar across the N treatments. The xanthophyll cycle pool size expressed on peel area basis was larger in the high N fruit than in the low N fruit. This corresponds well with the thermal dissipation capacity, as indicated by efficiency of excitation transfer. Over 95% of the xanthophyll cycle pool in the sun-exposed side was present in the form of zeaxanthin and antheraxanthin at noon regardless of N treatments. Activities of superoxide dismutase and all the antioxidant enzymes and metabolites in the ascorbate-glutathione cycle were higher in the high N fruit than in low N fruit. The results indicate that apple fruit with a good N status have a higher photoprotective capacity in terms of xanthophyll cycle-dependent thermal dissipation and detoxification of reactive oxygen species compared with low N fruit.

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Brandon R. Smith and Lailiang Cheng

The objective of this study was to quantify how photoprotective mechanisms in the leaves of `Concord' grapevines (Vitis labruscana Bailey) respond to a range of iron (Fe) supply. Own-rooted, 1-year-old container-grown vines were fertigated twice weekly for 11 weeks with a complete nutrient solution containing 1, 10, 20, 50, or 100 μm Fe from ferric ethylenediamine di (o-hydroxyphenylacetic) acid (Fe-EDDHA). Leaf total Fe content did not increase in response to Fe supply; however, “active” Fe (extracted with 2,2′-dipyridyl) and chlorophyll (Chl) increased on a leaf area basis as applied Fe increased. At the lowest active Fe level, leaf absorptance and the efficiency of excitation transfer (Fv′/Fm′) was lower, and nonphotochemical quenching (NPQ) was significantly greater. Photosystem II (PSII) quantum efficiency decreased curvilinearly, and the proportion of PSII reaction centers in the open state (qP) decreased linearly as active Fe content decreased. On a Chl basis, the xanthophyll cycle pool size [violaxanthin (V) + antheraxanthin (A) + zeaxanthin (Z)], lutein, and β-carotene increased curvilinearly as active Fe decreased, and neoxanthin (Neo) increased at the lowest Fe level. On a leaf area basis, as active Fe decreased, V+A+Z and β-carotene decreased curvilinearly, and lutein and Neo decreased linearly. At noon, conversion of V to A and Z increased as active Fe decreased. On a Chl basis, activities of antioxidant enzymes superoxide dismutase (SOD), monodehydroascorbate reductase (MDAR), and dehydroascorbate reductase (DHAR) increased curvilinearly, and glutathione reductase (GR) activity increased linearly as active Fe levels declined. Ascorbate peroxidase (APX) and catalase (CAT), on a Chl basis, were relatively constant. On a leaf area basis, a decrease in active Fe increased SOD and MDAR activity, whereas APX, CAT, DHAR and GR activity decreased. Antioxidant metabolites ascorbate (AsA), dehydroascorbate (DAsA), reduced glutathione (GSH) and oxidized glutathione (GSSG) also increased in response to Fe limitation when expressed on a Chl basis, whereas on a leaf area basis AsA and DAsA decreased and GSH increased curvilinearly. The GSH:GSSG ratio increased as active Fe declined, whereas the AsA:DAsA ratio did not change. In conclusion, both photoprotective mechanisms, xanthophyll cycle-dependent thermal dissipation and the ascorbate-glutathione antioxidant system, are enhanced in response to Fe deficiency to cope with excess absorbed light. In a low soil pH tolerant species such as V. labruscana, the foliar antioxidant system was upregulated in response to excess absorbed light from Fe deficiency-induced chlorosis, and there was no evidence of an increase in oxidative stress from high rates of applied Fe-EDDHA.

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Qin Shi, Yunlong Yin, Zhiquan Wang, Wencai Fan, and Jianfeng Hua

photochemistry to thermal dissipation ( Chaves et al., 2003 ). For DS-R plants, reduced SLA on days 5 and 8 was attributed to a strategy that allowed the minimization of light absorption. Meanwhile, decreasing of leaf Chl t accompanied by F v / F m and φPSII

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Lailiang Cheng and Fengwang Ma

Xanthophyll cycle conversion and the antioxidant system in the peel of apple fruit (Malus ×domestica Borkh. `Liberty') were monitored in the field over a diurnal course at about 3 months after full bloom. Compared with leaves, sun-exposed peel of apple fruit had much lower photosystem II operating efficiency at any given photon flux density (PFD) and a larger xanthophyll cycle pool size on a chlorophyll basis. Zeaxanthin (Z) level increased with rising PFD in the morning, reached the highest level during midday, and then decreased with falling PFD for the rest of the day. At noon, Z accounted for >90% of the xanthophyll cycle pool in the fruit peel compared with only 53% in leaves. Efficiency of excitation transfer to PSII reaction centers (F v′/F m′) was negatively related to the level of Z in fruit peel and leaves throughout the day. In fruit peel, the antioxidant enzymes in the ascorbate-glutathione cycle, ascorbate peroxidase (APX), monodehydroascorbate reductase (MDAR), dehydroascorbate reductase (DHAR) and glutathione reductase (GR) showed a diurnal pattern similar to that of incident PFD. In contrast, the activities of APX and GR in leaves did not change significantly during the day although activities of both MDAR and DHAR were higher in the afternoon than in the morning. In both fruit peel and leaves, superoxide dismutase activity did not change significantly during the day; catalase activity showed a diurnal pattern opposite to that of PFD with much lower activity in fruit peel than in leaves. The total ascorbate pool was much smaller in fruit peel than in leaves on an area basis, but the ratio of reduced ascorbate to oxidized ascorbate reached a maximum in the early afternoon in both fruit peel and leaves. The total glutathione pool, reduced glutathione and the ratio of reduced glutathione to oxidized glutathione in both fruit peel and leaves also peaked in the early afternoon. We conclude that the antioxidant system as well as the xanthophyll cycle responds to changing PFD over the course of a day to protect fruit peel from photooxidative damage.

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Brandon Smith* and Lailiang Cheng

One-year-old `Concord' grapevines (Vitis labrusca L.) were fertigated twice weekly for 11 weeks with a complete nutrient solution containing 1, 10, 20, 50 or 100 μmol iron (Fe) from ferric ethylenediamine di (o-hydroxyphenylacetic) acid (Fe-EDDHA). Leaf total Fe content did not increase in response to Fe supply, however both “active” Fe (extracted with 2, 2'-dipyridyl) and chlorophyll (Chl) content increased as applied Fe increased. At the lowest active Fe level, leaf absorptance and maximum PSII efficiency (Fv/Fm) were slightly decreased, and non-photochemical quenching was significantly greater. PSII quantum efficiency decreased curvilinearly as active Fe content decreased. On a Chl basis, the xanthophyll cycle pool size, lutein, and beta-carotene increased curvilinearly as active Fe decreased, and neoxanthin increased at the lowest Fe level. Activities of antioxidant enzymes superoxide dismutase, ascorbate peroxidase, monodehydroascorbate reductase, dehydroascorbate reductase, and glutathione reductase followed a similar trend and increased under Fe deficiency, when expressed on a Chl basis. Antioxidant metabolites also increased in response to Fe limitation. On a Chl basis, ascorbate (AsA), dehydroascorbate (DAsA), reduced glutathione (GSH) and oxidized glutathione (GSSG) content was greater at the lowest active Fe levels. We did not find a difference in the ratio of AsA to DAsA or GSH to GSSG. In conclusion, both photoprotective mechanisms, xanthophyll cyle-dependent thermal dissipation and the ascorbate-glutatione antioxidant system, are enhanced in response to iron deficiency to cope with excess absorbed light.

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Li-Song Chen and Lailiang Cheng*

To determine the cause of zonal chlorosis of `Honeycrisp' apple leaves, we compared CO2 assimilation, carbohydrate metabolism, xanthophyll cycle and the antioxidant system between chlorotic leaves and normal leaves. Chlorotic leaves accumulated higher levels of non-structural carbohydrates, particularly starch, sorbitol, sucrose, and fructose at both dusk and predawn, and no difference was found in total non-structural carbohydrates between predawn and dusk. CO2 assimilation and the key enzymes in the Calvin cycle, ribulose 1,5-bisphosphate carboxylase/oxygenase, NADP-glyceraldehyde-3-phosphate dehydrogenase, phosphoribulokinase, stromal fructose-1,6-bisphosphatase, and enzymes in starch and sorbitol synthesis, ADP-glucose pyrophosphorylase, cytosolic fructose-1,6-bisphosphatase, and aldose 6-phosphate reductase were significantly lower in chlorotic leaves than in normal leaves. However, sucrose phosphate synthase activity was higher in chlorotic leaves. Thermal dissipation of excitation energy was enhanced in chlorotic leaves under full sun, lowering the efficiency of excitation energy transfer to PSII reaction centers. This was accompanied by a corresponding increase in both xanthophyll cycle pool size (on a chlorophyll basis) and conversion of violaxanthin to antheraxanthin and zeaxanthin. The antioxidant system was up-regulated in chlorotic leaves in response to the increased generation of reactive oxygen species. These findings support the hypothesis that phloem loading and/or transport is partially or completely blocked in chlorotic leaves, and that excessive accumulation of non-structural carbohydrates may cause feedback suppression of CO2 assimilation via direct interference with chloroplast function and/or indirect repression of photosynthetic enzymes.

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Li-Song Chen and Lailiang Cheng

To determine the cause of a characteristic zonal chlorosis of `Honeycrisp' apple (Malus ×domestica Borkh.) leaves, we compared CO2 assimilation, carbohydrate metabolism, the xanthophyll cycle and the antioxidant system between chlorotic leaves and normal leaves. Chlorotic leaves accumulated higher levels of nonstructural carbohydrates, particularly starch, sorbitol, sucrose, and fructose at both dusk and predawn, and no difference was found in total nonstructural carbohydrates between predawn and dusk. This indicates that carbon export was inhibited in chlorotic leaves. CO2 assimilation and the key enzymes in the Calvin cycle, ribulose 1,5-bisphosphate carboxylase/oxygenase, NADP-glyceraldehyde-3-phosphate dehydrogenase, phosphoribulokinase, stromal fructose-1,6-bisphosphatase, and the key enzymes in starch and sorbitol synthesis, ADP-glucose pyrophosphorylase, cytosolic fructose-1,6-bisphosphatase, and aldose 6-phosphate reductase were significantly lower in chlorotic leaves than in normal leaves. However, sucrose phosphate synthase activity was higher in chlorotic leaves. In response to a reduced demand for photosynthetic electron transport, thermal dissipation of excitation energy (measured as nonphotochemical quenching of chlorophyll fluorescence) was enhanced in chlorotic leaves under full sun, lowering the efficiency of excitation energy transfer to PSII reaction centers. This was accompanied by a corresponding increase in both xanthophyll cycle pool size (on a chlorophyll basis) and conversion of violaxanthin to antheraxanthin and zeaxanthin. The antioxidant system, including superoxide dismutase and ascorbate peroxidase and the ascorbate pool and glutathione pool, was up-regulated in chlorotic leaves in response to the increased generation of reactive oxygen species via photoreduction of oxygen. These findings support the hypothesis that phloem loading and/or transport is partially or completely blocked in chlorotic leaves, and that excessive accumulation of nonstructural carbohydrates may cause feedback suppression of CO2 assimilation via direct interference with chloroplast function and/or indirect repression of photosynthetic enzymes.

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Shuyang Zhen and Marc W. van Iersel

-mediated thermal dissipation of excess absorbed light ( Demmig-Adams and Adams, 1992 ), and thus are often more susceptible to photoinhibition. Alternatively, a reduction in F v / F m could be the result of the sustention of high level of xanthophyll cycle

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Jinhong Yuan, Man Xu, Wei Duan, Peige Fan, and Shaohua Li

parameters is often used as an indicator of mechanisms for preventing overexcitation of RCs ( Ivanov and Edwards, 2000 ). To protect WRS leaves from damage by excess excitation energy, thermal dissipation of excitation energy in WRS leaves increased, as