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-photochemical quenching increased in PRD-treated hot pepper compared with full irrigation ( Shao et al., 2010 ). The dissipation of xanthophyll pigments in the photosynthetic structure prevented the damage of excess light to PSII ( Niyogi, 2000 ; Ort, 2001 ). Through the

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Chlorophylls and xanthophylls were monitored in broccoli (Brassica oleracea L. var. italica Plen.) florets stored in air, air + 10 ppm ethylene, or 10% CO2 + 1% O2 controlled atmosphere (CA) at 15 °C. Chlorophylls a and b, as measured with high-performance liquid chromatography, decreased in florets held in air. The decrease was accelerated by ethylene treatment and suppressed in CA. Chlorophyllide a and pheophorbide a were present in fresh broccoli florets, but the levels decreased significantly in all treatments during storage. The oxidized product of chlorophyll a, 132-hydroxychlorophyll a, did not accumulate. Xanthophylls decreased, but new pigments, suggested to be esterified xanthophylls, formed with yellowing in stored florets. The chlorophyll degradative pathway in broccoli florets was not altered by ethylene or CA and differed from that reported for parsley (Petroselium crisum Nym.) and spinach (Spinacia oleracea L.) leaves.

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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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Abstract

Eleven of 39 varieties and lines resulted in good canned slice color in both unusually warm and unusually cool growing seasons for carrots. A number of lines showed seasonal interaction in color, with better color in the cool season, while others resulted in consistently fair or poor color in both growing seasons. No single pigment was highly correlated with color across the range of environmental and genetic diversity encountered in the study. Beta-carotene was the only single component showing a significant positive correlation with color. The highest multiple relationship with color considered beta-carotene, other carotenes except alpha carotene, and xanthophylls. Within a season this multiple correlation accounted for 47 to 50% of the color variance (R of .710 for spring and .686 for fall grown carrots).

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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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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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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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About 80 days after full bloom, well-exposed fruit on the south part of the canopy of mature Liberty/M.9 apple trees were randomly assigned to one of the following two treatments. Some fruit were turned about 180 degrees to expose the original shaded side to full sun whereas the rest served as untreated controls. On day 0, 1, 2, 4, 7, and 10 after treatment, fruit peel samples were taken from the original shaded side of the treated fruit and both the sun-exposed side and the shaded side of the control fruit at midday to determine photosynthetic pigments and enzymatic and non-enzymatic antioxidants. Maximum photosystem II efficiency of the original shaded side decreased sharply after 1 day exposure to full sun, and then gradually recovered to a similar value of the sun-exposed side of the control fruit by day 10. The shaded side of the control fruit had much lower xanthophyll cycle pool size and conversion and antioxidant enzymes and soluble antioxidants of the ascorbate-glutathione cycle than the sun-exposed side. In response to full sun exposure, xanthophyll cycle pool size of the original shaded side increased, reaching a similar value of the sun-exposed side by day 10. Ascorbate peroxidase, monodehydroascorbate reductase, dehydroascorbate reductase, and glutathione reductase and total pool size and reduction state of both ascorbate and glutathione of the original shaded side all increased to the corresponding values found in the sun-exposed side of the control fruit over a 10-day period. It is concluded that both xanthophyll cycle and the ascorbate-glutathione cycle in the original shaded side are up-regulated in response to fullsun exposure to minimize photo-oxidative damage and contributes to its re-acclimation to full sun.

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, 2002 ). Diets deficient in vitamin A cause night blindness in humans ( Rao and Rao, 2007 ). Human macula pigments are a mixture of carotenoids, lutein, and other xanthophylls. These compounds prevent free radical production in the retina, protecting the

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Abstract

Carotenoid synthesis was studied in tissue cultures derived from different carrot phenotypes of red, dark orange, orange, light orange, yellow and white root colors. Cultures derived from a single genetic source or even from a single root often differed in color. Yellow, pink, orange-red and red cultures were derived from red roots; yellow and orange cultures from dark orange and light orange roots; and yellow cultures from the roots of the other genetic sources. The total carotenoid content in tissue cultures partly reflected that of the parent root, but was generally lower than in the intact root. Cultures derived from white roots were exceptions to this; the carotenoid contents in these cultures sometimes exceeded the contents of the parent roots. Carotenes predominated in cultures and roots of the red and orange sources; xanthophylls were the main components in cultures and roots of the yellow and white sources. Individual cultures from one root differed in quantities of carotenes, but the main carotene was the same as in the root, lycopene for the red sources and B-carotene for the orange sources.

The effects of kinetin, 2,4-dichlorophenoxyacetic acid (2,4-D) and gibberellic acid3 (GA3) on growth and pigmentation were tested in 13 selected cultures. Kinetin inhibited and 2,4-D enhanced carotenoid synthesis in several of the cultures. GA had no effect on pigment formation. Kinetin and 2,4-D affected the relative quantities of carotenoid components in one orange-red culture, R 1-1, but little in the orange-red culture R 1-2. Low total carotenoid content was associated with a relatively high xanthophyll content and a low lycopene to B-carotene ratio.

Variation in culture pigmentation due to mutations was not confirmed. Plants regenerated from tissue cultures which had been derived from a single root had the same phenotype even though the colors and carotenoid contents of the callus cultures differed widely. If plantlet regeneration is limited only to cells with a diploid chromosome number, aneuploidy, which is very common in callus cultures, may have contributed to the observed pigmentation differences in the cultures.

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