Stability of individual anthocyanins from black carrots stored in light and darkness - Impact of acylation.

Black carrot anthocyanins have gained increasing attention as natural coloring agent, owing to their higher stability than anthocyanins from berries. The stability has been attributed to their higher degree of acylation. This study investigated the impact of acylation on the stability of individual anthocyanins during storage in light and darkness. We hypothesized that the acylated anthocyanins would be more stable than the non-acylated ones. The major five anthocyanins were fractioned by semi-preparative HPLC and stored at pH 4.5 in light and darkness to investigate how acylation affected the stability. The stability was evaluated by absorption spectroscopy and mass spectrometry (MS). Two of the anthocyanins were non-acylated; 3-xylosyl(glucosyl)galactoside and cyanidin 3-xylosylgalactoside, and three were acylated; cyanidin 3-xylosyl(sinapolyglucosyl)galacto-side, cyanidin 3-xylosyl(feruloylglu-cosyl)galactoside, and cyanidin 3-xylosyl(coumaroyl-glucosyl)galactoside. Both methods (spectroscopy and MS) showed a clear effect of acylation when stored in light, but surprisingly the two non-acylated anthocyanins, showed higher stability than the three acylated ones.

Effect of pulsed electric fields on carotenoid and phenolic bioaccessibility and their relationship with carrot structure.

Phenolic compounds (PC) and carotenoids from carrots are bound to dietary fibre or stored in vacuoles and chromoplasts, respectively. To exert their antioxidant effects these compounds must be released during digestion, which is hindered by such barriers. Pulsed electric fields (PEF) modify cell membrane permeability, thus enhancing their bioaccessibility. The effect of PEF on the carrot carotenoid and PC content and bioaccessibility was investigated. With this purpose, PEF-treated carrots (5 pulses of 3.5 kV cm-1) were stored for 24 h at 4 °C and microstructure was evaluated before subjecting them to in vitro digestion. PEF did not affect carotenoid content, whereas their bioaccessibility improved (11.9%). Likewise, PEF increased the content of some PC, e.g. coumaric acid (163.2%), probably caused by their better extractability. Conversely, caffeic acid derivatives decreased, which may be associated to greater contact with oxidative enzymes. Total PC bioaccessibility (20.8%) and some derivatives increased, e.g. caffeoylshikimic (68.9%), whereas some decreased (e.g. ferulic acid). Structural changes caused by PEF may improve bioaccessibility of carotenoids and PC by favouring their release and easy access to digestive enzymes. However, other antioxidants may be further degraded or entrapped during digestion. Therefore, PEF is an effective technology for obtaining carrots with enhanced carotenoids and phenolic bioaccessibility.

Ultrasonic pretreatment of carrot slices: Effects of sonication source on drying kinetics and product quality.

In this study effects of ultrasonic pretreatment with different sonication sources on drying behavior of carrot slices were evaluated. Fresh carrot slices were pretreated with ultrasonic probe at 65, 75 and 85 W or ultrasonic bath at 10, 20 and 30°C for 3, 5 and 10 min before air drying. Water gain % and solid loss % of pretreated samples were calculated and color values, ß-carotene content and rehydration ratios of dried samples were determined. Drying behavior of the pretreated samples was evaluated and the drying data were fitted to thin layer drying models. Constant drying rate period was not observed for the carrot slices; however two definite falling-rate periods having different slopes were obtained. Drying time was significantly reduced (up to 20%) depending on the type of pretreatment. The redness value, total color difference and Chroma values of pretreated and control samples were in the same group (p>0.05). ß-carotene content of ultrasonic bath pretreated samples were significantly higher than the samples pretreated with ultrasonic probe and the sample dried without any pretreatment as well (p<0.05). Also rehydration ratios of control samples were found lower than the ultrasound pretreated samples.

Thermal and Photochemical Stability of Anthocyanins from Black Carrot, Grape Juice, and Purple Sweet Potato in Model Beverages in the Presence of Ascorbic Acid.

Anthocyanins are natural dyes widely used in the food industry, but their chemical stability in beverages can be affected by the presence of additives. In the present paper, the interaction between anthocyanins and ascorbic acid (AA) is more particularly investigated. Ascorbic acid is an ubiquitous component in food products. In this study, the thermal stability at 43 °C and the photolysis stability in air and in an inert atmosphere (N2) of anthocyanins extracted from black carrot (BC), grape juice (GJ), and purple sweet potato (SP) were studied in the presence and absence of ascorbic acid (in citrate buffer at pH 3). Discriminating the main environmental factors (i.e., heat and light) affecting anthocyanin stability is a key point for better understanding the degradation pathways. The stability of the anthocyanins was followed by UV-vis spectrometry. Moreover, to understand the degradation mechanisms in both the presence and absence of ascorbic acid, various techniques such as fluorescence quenching, cyclic voltammetry, and electron-spin-resonance (ESR) spectroscopy were also used to furnish a full coherent picture of the chemical mechanisms associated with the anthocyanin degradation. In addition, molecular orbitals and bond-dissociation energies (BDE) were calculated to extend the investigation. Moreover, the effects of some supplementary stabilizers (chlorogenic acid, sinapic acid, tannic acid, fumaric acid, ß-carotene, isoquercitrin, myricitrin, green coffee bean extract, and rosemary extract) and sugars (sucrose, fructose, and glucose) on anthocyanins stability in the presence of ascorbic acid were examined.

The effect of high-power ultrasound on the quality of carrot juice.

The effect of high-power ultrasound treatment on enzymes' activity, physicochemical attributes (total soluble solids, pH, viscosity, turbidity, particle size distribution and colour) and carotenoids' content of carrot juice was investigated. The treatments were carried out at 20 kHz (0.95, 2.38, 3.80 W/ml power) in an ice bath for 2, 4, 6, 8, 10 min. The polyphenol oxidase and pectin methylesterase activity were decreased by 43.90 and 37.95% at 3.80 W/ml power and 10 min exposure time, respectively. With the increase of power and time, the effect of high-power ultrasound on the inactivation of enzymes was getting stronger. However, high-power ultrasound had no inactivation effect on peroxidase activity under all treatment conditions. The visual colour differences were not obvious after high-power ultrasound. The pH, total soluble solids and particle size distribution of carrot juice were not significantly affected (p > 0.05) under all treatment conditions, while turbidity was increased and carotenoids' content was decreased. The viscosity of carrot juice was decreased by 1.27% at 0.95 W/ml power and 8 min, while it was increased by 2.29% at 2.38 W/ml power and 8 min. The value of viscosity was negatively correlated with the activity of pectin methylesterase (Pearson's r = -0.481, p < 0.05). According to these results, we could conclude that the optimal treatment condition was 3.80 W/ml for 10 min. Overall, high-power ultrasound treatment inhibited browning, maintained taste and nutritional value and improved stability of carrot juice. Therefore, this technology could well be an option for processing of carrot juice and laid the theoretical foundation for the production of carrot juice and carrot compound beverage.

Effect of diflufenican on total carotenoid and phytoene production in carrot suspension-cultured cells.

MAIN CONCLUSION: Diflufenican increased 493-fold the level of phytoene. Diflufenican-induced inhibition of phytoene desaturase gene expression in carrot cells resulted in an increased production of phytoene. This work analyzes the effect of diflufenican, an inhibitor of phytoene desaturase, on the gene expression profiles of the biosynthetic pathway of carotenoids related with the production of these compounds in carrot cell cultures. The results showed that the presence of 10 µM diflufenican in the culture medium increased phytoene levels, which was 493-fold higher than in control cells after 7 days of treatment but did not alter cell growth in carrot cell cultures. The maximal production of phytoene was reached with 10 µM diflufenican after 7 days of incubation in the presence of light and with 30 g/L sucrose in the culture medium. Moreover, diflufenican decreased the expression of phytoene synthase and phytoene desaturase genes at all the times studied. This diflufenican-induced inhibition of phytoene desaturase gene expression in carrot cell cultures resulted in an increased production of phytoene. Our results provide new insights into the action of diflufenican in carrot cell cultures, which could represent an alternative more sustainable and environmentally friendly system to produce phytoene than those currently used.

Stability Analysis of Anthocyanins Using Alcoholic Extracts from Black Carrot (Daucus Carota ssp. Sativus Var. Atrorubens Alef.).

Anthocyanins are used for food coloring due their low toxicity and health benefits. They are extracted from different sources, but black carrot has higher anthocyanin content compared with common fruits and vegetables. Here, we study alcoholic anthocyanin extracts from black carrot to enhance their stability. The objective of our research is to determine if microencapsulation with tetraethyl orthosilicate (TEOS) is a feasible option for preventing black carrot anthocyanin degradation. Extraction solvents were solutions of (1) ethanol/acetic acid and (2) ethanol/citric acid. Samples were purified through a resin column and microencapsulated using TEOS. Fourier Transformed Infrared Spectroscopy (FTIR) spectra of samples were obtained, and degradation studies were performed under different conditions of UV radiation, pH and temperature. Antioxidant activity was evaluated with radical 2,2-diphenyl-1-picrylhydrazyl (DPPH) scavenging and electrochemical cupric reducing antioxidant capacity (CUPRAC). Color evaluation on food models were performed with CIE Lab at the beginning of experiments and after 25 days of storage. Results indicate that the more stable extracts against pH media changes are samples obtained with ethanol/acetic acid solution as extraction solvent. Extract purification through resin and TEOS microencapsulation had no significant effect on extract stability. In conclusion, although TEOS microencapsulation has proven to be effective for some dried materials from natural extracts in our previous research, we do not recommend its use for black carrot extracts considering our results in this particular case.

UVA, UVB and UVC Light Enhances the Biosynthesis of Phenolic Antioxidants in Fresh-Cut Carrot through a Synergistic Effect with Wounding.

Previously, we found that phenolic content and antioxidant capacity (AOX) in carrots increased with wounding intensity. It was also reported that UV radiation may trigger the phenylpropanoid metabolism in plant tissues. Here, we determined the combined effect of wounding intensity and UV radiation on phenolic compounds, AOX, and the phenylalanine ammonia-lyase (PAL) activity of carrots. Accordingly, phenolic content, AOX, and PAL activity increased in cut carrots with the duration of UVC radiation, whereas whole carrots showed no increase. Carrot pies showed a higher increase compared to slices and shreds. Phenolics, AOX, and PAL activity also increased in cut carrots exposed to UVA or UVB. The major phenolics were chlorogenic acid and its isomers, ferulic acid, and isocoumarin. The type of UV radiation affected phenolic profiles. Chlorogenic acid was induced by all UV radiations but mostly by UVB and UVC, ferulic acid was induced by all UV lights to comparable levels, while isocoumarin and 4,5-diCQA was induced mainly by UVB and UVC compared to UVA. In general, total phenolics correlated linearly with AOX for all treatments. A reactive oxygen species (ROS) mediated hypothetical mechanism explaining the synergistic effect of wounding and different UV radiation stresses on phenolics accumulation in plants is herein proposed.

Illuminating colors: regulation of carotenoid biosynthesis and accumulation by light.

Light stimulates the biosynthesis of carotenoids and regulates the development of plastid structures to accommodate these photoprotective pigments. Work with Arabidopsis revealed molecular factors coordinating carotenoid biosynthesis and storage with photosynthetic development during deetiolation, when underground seedlings emerge to the light. Some of these factors also adjust carotenoid biosynthesis in response to plant proximity (i.e., shade), a mechanism that was readapted in tomato to monitor fruit ripening progression. While light positively impacts carotenoid production and accumulation in most cases, total carotenoid levels decrease in roots of colored carrot cultivars when illuminated. The recent discovery that such cultivars might be photomorphogenic mutants provides an explanation for this striking phenotype.

Characterising genes associated with flowering time in carrot (Daucus carota L.) using transcriptome analysis.

Carrot is generally regarded as a biennial plant with an obligatory vernalization requirement. Early spring cultivation makes plants vulnerable to premature bolting, which results in a loss of commercial value. However, our knowledge of flowering time genes and flowering mechanisms in carrot remain limited. Bolting behavior of D. carota ssp. carota 'Songzi', a wild species sensitive to flower induction by vernalization and photoperiod, and orange cultivar 'Amsterdam forcing', and their offspring were investigated in different growing conditions. We performed RNA-seq to identify the flowering time genes, and digital gene expression (DGE) analysis to examine their expression levels. The circadian patterns of related genes were identified by qPCR. The results showed bolting behavior of carrot was influenced by low temperature, illumination intensity and photoperiod. A total of 45 flowering time-related unigenes were identified, which were classified into five categories including photoperiod, vernalization, autonomous and gibberellin pathway, and floral integrators. Homologs of LATE ELONGATED HYPOCOTYL (LHY) and CONSTANS-LIKE 2 (COL2) were more highly expressed under short day condition than under long day condition. Homologs of COL2, CONSTANS-LIKE 5 (COL5), SUPPRESSION OF OVEREXPRESSION OF CONSTANS 1 (SOC1), FLOWERING LOCUS C (FLC) and GIBBERELLIC ACID INSENSITIVE (GAI) were differentially expressed between 'Songzi' and 'Amsterdam forcing'. The homolog of COL2 (Dct43207) was repressed by light, but that of COL5 (Dct20940) was induced. A preliminary model of genetic network controlling flowering time was constructed by associating the results of DGE analysis with correlation coefficients between genes. This study provides useful information for further investigating the genetic mechanism of flowering in carrot.

Ionizing radiation from Chernobyl affects development of wild carrot plants.

Radioactivity released from disasters like Chernobyl and Fukushima is a global hazard and a threat to exposed biota. To minimize the deleterious effects of stressors organisms adopt various strategies. Plants, for example, may delay germination or stay dormant during stressful periods. However, an intense stress may halt germination or heavily affect various developmental stages and select for life history changes. Here, we test for the consequence of exposure to ionizing radiation on plant development. We conducted a common garden experiment in an uncontaminated greenhouse using 660 seeds originating from 33 wild carrots (Daucus carota) collected near the Chernobyl nuclear power plant. These maternal plants had been exposed to radiation levels that varied by three orders of magnitude. We found strong negative effects of elevated radiation on the timing and rates of seed germination. In addition, later stages of development and the timing of emergence of consecutive leaves were delayed by exposure to radiation. We hypothesize that low quality of resources stored in seeds, damaged DNA, or both, delayed development and halted germination of seeds from plants exposed to elevated levels of ionizing radiation. We propose that high levels of spatial heterogeneity in background radiation may hamper adaptive life history responses.

Possible involvement of a tetrahydrobiopterin in photoreception for UV-B-induced anthocyanin synthesis in carrot.

Our previous studies of action spectra for UV-B-induced anthocyanin accumulation in cultured carrot cells indicated that a reduced form of pterin, possibly tetrahydrobiopterin, contributes to UV-B photoreception. In this report, we provide additional evidence for the involvement of pterin in UV-B light sensing. UV-B-induced phenylalanine ammonia-lyase (PAL) activity was considerably suppressed by N-acetylserotonin (an inhibitor of tetrahydrobiopterin biosynthesis), and this suppression was partially recovered by adding biopterin or tetrahydrobiobiopterin. In addition, protein(s) specifically bound to biopterin were detected by radiolabeling experiments in N-acetylserotonin-treated cells. Furthermore, diphenyleneiodonium, a potent inhibitor of electron transfer, completely suppressed UV-B-induced PAL activity. These results suggest the occurrence of an unidentified UV-B photoreceptor (other than UVR8, the tryptophan-based UV-B sensor originally identified in Arabidopsis) with reduced pterin in carrot cells. After reexamining published action spectra, we suggest that anthocyanin synthesis is coordinately regulated by these two UV-B sensors.

Combating photooxidative stress in green hairy roots of Daucus carota cultivated under light irradiation.

The light-dependent generation of active oxygen species, which can disrupt normal metabolic process of plant, is termed as photo-oxidative stress. Plants are equipped with enzymatic and non-enzymatic antioxidative defence system to reduce the effect of such stress. Hairy root culture of Daucus carota when cultivated under continuous illumination (250 µmol m(-2)s(-1)) turned green. To know the reason behind that and photo-oxidative stress response in green hairy roots, activities of several antioxidant enzymes were measured. When compared with normal hairy roots, green hairy roots showed an enhanced superoxide dismutase (SOD) activity. Treatment with a SOD inhibitor diethyldithiocarbamate led to suppression of SOD activity in a concentration-dependent manner in green hairy roots. Interestingly, SOD-suppressed root showed three-fold enhanced caffeic acid glucoside accumulation in the soluble fraction as compared to untreated ones. While ascorbate peroxidase activity showed marginal increase in green hairy roots, a decrease in the activities of guaiacol peroxidase and catalase were observed. SDS-PAGE of crude protein profile from green hairy roots showed a distinct band, which was absent in normal hairy roots. MALDI-TOF-MS/MS analysis of the extracted protein confirmed it as the large subunit of RuBisCO. RT-PCR based expression analysis of betaine aldehyde dehydrogenase showed enhanced transcript levels in green hairy roots as compared to normal hairy roots, whereas reverse trends were observed with the transcripts accumulation for phenylalanine ammonia-lyase and chalcone synthase. These findings corroborate with the in vitro BADH activities in hairy roots, and thus indicate an important role of this stress enzyme in combating photo-oxidative stress in green hairy roots upon continuous light exposure.

Ultraviolet-B light treatment increases antioxidant capacity of carrot products.

BACKGROUND: The effect of ultraviolet-B (UV-B) light as a postharvest treatment to enhance the antioxidant content of carrots and fresh-cut carrot products was evaluated. Four levels of UV-B dose ranging from 1.3 to 12 kJ m⁻² were applied to whole, baby and various styles of cut carrots, and the changes in antioxidant capacity, total soluble phenolics and phenylalanine ammonia-lyase (PAL, EC 4.3.1.24) activity were measured after a 3 day incubation period at 15 °C and 45% relative humidity. RESULTS: Both cutting style and dose level were factors in determining carrot responses to UV-B treatment. Antioxidant capacity increased significantly (1.4-6.6-fold). Total soluble phenolic results correlated directly with those of antioxidant capacity (R² = 0.953), indicating that the enhancements achieved were due to an increase in phenolic content. High-performance liquid chromatography analysis revealed that 5-O-caffeoylquinic acid (5-CQA) was the primary phenolic responsible for this increase. Higher PAL activity was also observed in UV-B-treated samples, indicating that the increase in 5-CQA was a biological response to UV-B exposure. CONCLUSION: UV-B treatment has the potential to increase the nutritional value of carrots and offers an exciting opportunity to increase consumer accessibility to dietary choices that are rich in antioxidants.

Light-dependent changes in plastid differentiation influence carotenoid gene expression and accumulation in carrot roots.

Carrot is an important nutritional crop due to the high levels of pro-vitamin A carotenoids (ß-carotene and, to a lower extent, α-carotene) that accumulate in its storage root during secondary growth. In this work we show that in carrots, contrary to that reported for aerial organs of other plant species, light has a profound effect on root development by inhibiting root thickening, preventing the differentiation of chromoplasts and eventually repressing the expression of most genes required for the biosynthesis of ß-carotene and α-carotene and to a lesser extent genes for xanthophylls and apocarotenoids biosynthesis. We observed a correlation in the carotenoid profile and the patterns of gene expression during the development of root segments grown either in the light or in the dark, which suggests a transcriptional regulation for carotenoid synthesis during carrot root development. Furthermore, our work supports the conclusion that the differentiation of chromoplasts coincides with carotenoid accumulation during the later stages of development of underground storage roots.

Effect of UV-B light and different cutting styles on antioxidant enhancement of commercial fresh-cut carrot products.

Wounding stresses resulting from fresh-cut processing are known to enhance the antioxidant capacity (AC) of carrots by increasing the synthesis of phenolic compounds. Ultraviolet-B (UV-B) light exposure further promotes the formation of phenolic compounds. Changes in total soluble phenolics (TSP), 5-O-caffeoylquinic acid (5-CQA), total carotenoids, AC, and phenylalanine ammonia-lyase (PAL) activity of five commercial fresh-cut carrot products (baby carrots, carrot stixx, shredded carrots, crinkle cut coins, and oblong chips) were evaluated after exposure to UV-B dosage at 141.4 mJ/cm(2). Significant increases in TSP, AC and 5-CQA levels were observed for each sample following UV-B exposure. Increases in PAL activity were also observed in all carrot products, except crinkle cut coins. Total carotenoids of the carrot products were unchanged by UV-B exposure. Increases in AC levels corresponded directly with increases in the area/weight ratio (exposure area) of the fresh-cut carrot products.

Improved mechanical properties of retorted carrots by ultrasonic pre-treatments.

The use of ultrasound pre-processing treatment, compared to blanching, to enhance mechanical properties of non-starchy cell wall materials was investigated using carrot as an example. The mechanical properties of carrot tissues were measured by compression and tensile testing after the pre-processing treatment prior to and after retorting. Carrot samples ultrasound treated for 10 min at 60 °C provided a higher mechanical strength (P<0.05) to the cell wall structure than blanching for the same time period. With the addition of 0.5% CaCl(2) in the pre-treatment solution, both blanching and ultrasound treatment showed synergistic effect on enhancing the mechanical properties of retorted carrot pieces. At a relatively short treatment time (10 min at 60 °C) with the use of 0.5% CaCl(2), ultrasound treatment achieved similar enhancement to the mechanical strength of retorted carrots to blanching for a much longer time period (i.e. 40 min). The mechanism involved appears to be related to the stress responses present in all living plant matter. However, there is a need to clarify the relative importance of the potential stress mechanisms in order to get a better understanding of the processing conditions likely to be most effective. The amount of ultrasound treatment required is likely to involve low treatment intensities and there are indications from the structural characterisation and mechanical property analyses that the plant cell wall tissues were more elastic than that accomplished using low temperature long time blanching.

Effect of ultrasound and blanching pretreatments on polyacetylene and carotenoid content of hot air and freeze dried carrot discs.

The effect of ultrasound and blanching pretreatments on polyacetylene (falcarinol, falcarindiol and falcarindiol-3-acetate) and carotenoid compounds of hot air and freeze dried carrot discs was investigated. Ultrasound pretreatment followed by hot air drying (UPHD) at the highest amplitude and treatment time investigated resulted in higher retention of polyacetylenes and carotenoids in dried carrot discs than blanching followed by hot air drying. Freeze dried samples had a higher retention of polyacetylene and carotenoid compounds compared to hot air dried samples. Color parameters were strongly correlated with carotenoids (p<0.05). This study shows that ultrasound pretreatment is a potential alternative to conventional blanching treatment in the drying of carrots.

Response to UV-C radiation in topo I-deficient carrot cells with low ascorbate levels.

In animal cells, recent studies have emphasized the role played by DNA topoisomerase I (topo I) both as a cofactor of DNA repair complexes and/or as a damage sensor. All these functions are still unexplored in plant cells, where information concerning the relationships between DNA damage, PCD induction, and topo I are also limited. The main goal of this study was to investigate the possible responses activated in topo I-depleted plant cells under oxidative stress conditions which induce DNA damage. The carrot (Daucus carota L.) AT1-beta/22 cell line analysed in this study (characterized by an antisense-mediated reduction of top1beta gene expression of approximately 46% in association with a low ascorbate content) was more sensitive to UV-C radiation than the control line, showing consistent cell death and high levels of 8-oxo-dG accumulation. The topo I-depleted cells were also highly susceptible to the cross-linking agent mitomycin C. The death response was associated with a lack of oxidative burst and there were no changes in ascorbate metabolism in response to UV-C treatment. Electron and fluorescence microscopy suggested the presence of three forms of cell death in the UV-C-treated AT1-beta/22 population: necrosis, apoptotic-like PCD, and autophagy. Taken together, the data reported here support a reduced DNA repair capability in carrot topo I-deficient cells while the putative relationship between topo I-depletion and ascorbate impairment is also discussed.

Transgenic carrot plants accumulating ketocarotenoids show tolerance to UV and oxidative stresses.

Ketocarotenoids are strong antioxidant compounds which accumulate in salmon, shrimp, crustaceans and algae, but are rarely found naturally in higher plants. In this study, we engineered constitutive expression of an algal beta-carotene ketolase gene (bkt) in carrot plants to produce a number of ketocarotenoids from beta-carotene. These included astaxanthin, adonirubin, canthaxanthin, echinenone, adonixanthin and beta-cryptoxanthin. Leaves accumulated up to 56 microg/g total ketocarotenoids and contained higher beta-carotene levels but lower levels of alpha-carotene and lutein. The photosynthetic capacity of transgenic plants was not significantly altered by these changes. However, when high-expressing transgenic plants were exposed to UV-B irradiation, they grew significantly better than the wild-type controls. Similarly, leaf tissues exposed to various oxidative stresses including treatment with H(2)O(2) and methyl viologen showed less injury and retained higher levels of chlorophyll a+b. Total carotenoid extracts from transgenic leaves had higher antioxidant and free-radical scavenging activity in vitro compared to control leaves. Transgenic tissues also accumulated lower amounts of H(2)O(2) following exposure to oxidative stresses, suggesting that free radical and reactive oxygen species were quenched by the ketocarotenoids.