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Cold-pressed oils are rich sources of bioactive substances, which may protect triacylglycerols from degradation during frying. Nevertheless, these substances may decompose under high temperature. This work considers the content of bioactive substances in blends and their changes during high-temperature heating. Blends of refined rapeseed oil with 5% or 25% in one of three cold-pressed oils (rapeseed, coriander and apricot) were heated at 170 or 200 °C in a thin layer on a pan. All non-heated blends and cold-pressed oils were tested for fatty acid profile, content and composition of phytosterols, tocochromanols, chlorophyll and radical scavenging activity (RSA) analyzed by 2,2-diphenyl-1-picrylhydrazyl (DPPH), and 2,2'-azino-bis(3-ethylbenzothiazoline-6-sulphonic acid) (ABTS) assays. Moreover, the stability of phytosterols, tocochromanols, DPPH and ABTS values was determined in heated blends. All tocochromanols were lost during the heating process, in particular, at 200 °C. However, there were some differences between homologues. α-Tocopherol and δ-tocopherol were the most thermolabile and the most stable, respectively. Phytosterols were characterized by very high stability at both temperatures. We observed relationships between ABTS and DPPH values and contents of total tocochromanols and α-tocopherol. The obtained results may be useful in designing a new type of fried food with improved health properties and it may be the basis for further research on this topic.

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Pretreatment of grape pomace seeds with a pulsed electric field (PEF) was applied to improve the extraction yield of cold-pressed grape seed oil. The effects of different PEF conditions, electric field intensities (12.5, 14.0 and 15.6 kV/cm), and durations (15 and 30 min) on the oil chemical composition were also studied. All PEF pretreatments significantly increased the oil yield, flow rate and concentration of total sterols (p < 0.05). In addition, similar trends were observed for total tocochromanols and phenolic compounds, except for PEF pretreatment under the mildest conditions (12.5 kV/cm, 15 min) (p < 0.05). Notably, the application of 15.6 kV/cm for 30 min resulted in the highest relative increase in oil yield and flow rate (29.6% and 56.5%, respectively) and in the concentrations of total tocochromanols, nonflavonoids, and flavonoids (22.1%, 60.2% and 81.5%, respectively). In addition, the highest relative increase in the concentration of total sterols (25.4%) was achieved by applying 12.5 kV/cm for 30 min. The fatty acid composition of the grape seed oil remained largely unaffected by the PEF pretreatments. These results show that PEF pretreatment effectively improves both the yield and the bioactive properties of cold-pressed grape seed oil.

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Reversed-phase separation of tocopherols (Ts) and tocotrienols (Ts) using C18 stationary phases results in the coelution of ß and γ positional isomers, leading to identification errors. This study investigates the potential of alternative stationary phase chemistries to effectively resolve tocochromanols, specifically focusing on the critical pair of ß and γ positional isomers. Initial screening of seven different stationary phases (C18, C18-PFP, C30, PFP, 5PYE, πNAP, and RP-Amide) was conducted. Linear solvent strength (LSS) studies were performed to assess the impact of the organic modifier (methanol) and temperature on the chromatographic performance parameters. Five columns were found to be suitable for the tocochromanol separation and two different chromatographical conditions per column were proposed. Elution order of tocochromanols was unique for 5PYE, πNAP and C30 columns in comparison to RP-Amide and PFP. Method development for the quantitative analysis of four tocopherol and four tocotrienol homologues was performed. The optimised method employed the RP-Amide (150 × 4.6 mm, 2.6 µm dp) superficially porous particle column, mobile phase of methanol:water of 92:8, v/v, with a flow rate of 1.0 mL/min, column oven temperature of 40 °C and fluorescence detection (λex 295 nm, λem 330 nm). The analysis run time was 10.5 min with 13.6 MPa back pressure. The method was validated and the obtained LOQs were found to be 1.30-3.13 µg/mL. The method developed was successfully applied for the determination of tocochromanols in twenty samples with unique tocochromanol profiles. Principal component analysis illustrated three distinct groups based on the tocochromanol profile.

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Postmenopausal women are susceptible to osteoporosis and osteoarthritis. Tocotrienol, a bone-protective nutraceutical, is reported to prevent osteoarthritis in male rats. However, its efficacy on joint health in oestrogen deficiency has not been validated. Besides, data on the use of emulsification systems in enhancing bioavailability and protective effects of tocotrienol are limited. Ovariectomised adult female Sprague-Dawley rats (3 months old) were treated with refined olive oil, emulsified (EPT, 100 mg/kg/day with 25% vitamin E content), non-emulsified palm tocotrienol (NEPT, 100 mg/kg/day with 50% vitamin E content) and calcium carbonate (1% w/v in drinking water) plus glucosamine sulphate (250 mg/kg/day) for 10 weeks. Osteoarthritis was induced with monosodium iodoacetate four weeks after ovariectomy. Baseline control was sacrificed upon receipt, while the sham group was not ovariectomised and treated with refined olive oil. EPT and NEPT prevented femoral metaphyseal and subchondral bone volume decline caused by ovariectomy. EPT decreased subchondral trabecular separation compared to the negative control. EPT preserved stiffness and Young's Modulus at the femoral mid-shaft of the rats. Circulating RANKL was reduced post-treatment in the EPT group. Joint width was reduced in all the treatment groups vs the negative control. The EPT group's grip strength was significantly improved over the negative control and NEPT group. EPT also preserved cartilage histology based on several Mankin's subscores. EPT performed as effectively as NEPT in preventing osteoporosis and osteoarthritis in ovariectomised rats despite containing less vitamin E content. This study justifies clinical trials for the use of EPT in postmenopausal women with both conditions.

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Bioactive lipids-major and minor-comprise an array of compounds belonging to different chemical categories. Among the minor bioactive lipids carotenoids, sterols and tocochromanols attract continuously the interest of food scientists, nutritionists and medical doctors for their importance in food processing, preservation and for their health properties. Provitamin A and non-provitamin A carotenoids are found in various food sources of plant and animal origin and are added to foods as colorants. Their interactions with other food ingredients are critical because of their role against reactive oxygen species. The role of cholesterol through the diet after decades of disputes is better justified whereas at the same time emphasis is given to the technological and health aspects of phytosterols, which became very efficiently part of the daily diet for many population groups. Last but not least the importance of vitamin E is in a continuous debate for over 100years whereas studies on tocotrienols are intensified as a result of a transient to palm oil product consumption globally. Chemistry, natural occurrence, absorption and metabolism, dietary intake and dietary recommendations, major health impacts and key technological issues are updated and discussed with the support of recent findings.

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Grape berries have been extensively studied in terms of antioxidant characterization, specifically in anthocyanin, total phenol, and tannin accumulation. However, very little is known about vitamin E composition and contents in this fruit. Aiming to examine the function of vitamin E during grape berries ripening, tocochromanol contents and composition were evaluated in berries and leaves of grapevines (Vitis vinifera L. cv. Merlot), from just before veraison to commercial harvest. We also determined the time-course evolution of tocochromanol accumulation in various fruit tissues, including the skin, pulp, and seeds, and measured the extent of primary and secondary lipid peroxidation, as well as fruit technological maturity parameters. Vitamin E accumulated at higher levels in leaves than in fruits, although the tissue-specific evaluation of tocochromanol contents revealed that berry skin is also rich in α-tocopherol whereas tocotrienols were present in seeds only. α-Tocopherol content decreased during ripening, more specifically in the skin, and it was accompanied by an increase in the extent of lipid peroxidation. Contents and variations in the levels of α-tocopherol, but not those of the other tocochromanols, were inversely related to changes in lipid peroxidation during fruit ripening, as indicated by tissue-specific variations in malondialdehyde contents. In conclusion, α-tocopherol is more abundant in leaves than fruit, yet it apears to exert a role in the modulation of the extent of lipid peroxidation in grape berries, more specifically in the skin, where α-tocopherol depletion and malondialdehyde accumulation may be related to an adequate progression of fruit ripening.

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The purpose of this study was to explore the effects of roasting linseeds on the pigment, lipid profile, bioactive components, and oxidative stability of the extracted oils. The linseed varieties Giza 11, Giza 12, Sakha 3, and Sakha 6 were roasted at 180 °C for 10 min, and the oils were extracted by cold pressing. The results showed that, after roasting, there was an increase in oil percentage and peroxide value, as well as small increases in p-anisidine and acid values. Roasting also caused an increase in chlorophyll content, while lutein and ß-carotene tend to slightly decrease, except in the Giza 11 variety. The total phenolics content was markedly enhanced after roasting. Omega-3 fatty acids were not affected by the roasting process. The total amounts of tocochromanol were found to decrease in the Giza 12 and Sakha 6 varieties after roasting. Plastochromanol-8 increased in all varieties after roasting. The phytosterol composition was minimally affected by roasting. Roasting enhanced the stability of the extracted oils, increasing the induction period and decreasing EC50 values. These results may thus help to discriminate between the different linseed varieties and serve to recommend the use of roasting to enhance the oxidative stability of extracted oil.

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Free and esterified (bound) tocopherols, tocotrienols and other tocochromanol-related compounds, often referred to "tocols", are lipophilic antioxidants of great importance for health. For instance, α-tocopherol is the only tocochromanol with vitamin E activity, while tocotrienols have a positive impact on health and are proposed in the prevention and therapy of so-called modern diseases. Tocopherols, tocotrienols and plastochromanol-8 are the most well-known tocochromanols; in turn, knowledge about tocodienols, tocomonoenols, and other rare tocochromanol-related compounds is limited due to several challenges in analytical chemistry and/or low concentration in plant material. The presence of free, esterified, and non-extractable tocochromanols in plant material as well as their biological function, which may be of great scientific, agricultural and medicinal importance, is also poorly studied. Due to the lack of modern protocols as well as equipment and tools, for instance, techniques suitable for the efficient and simultaneous chromatographical separation of major and minor tocochromanols, the topic requires attention and new solutions, and/or standardization, and proper terminology. This review discusses the advantages and disadvantages of different chromatographic techniques, tools and approaches used for the separation and detection of different tocochromanols in plant material and foodstuffs. Sources of tocochromanols and procedures for obtaining different tocochromanol analytical standards are also described. Finally, future challenges are discussed and perspective green techniques for tocochromanol determination are proposed along with best practice recommendations. The present manuscript aims to present key aspects and protocols related to tocochromanol determination, correct identification, and the interpretation of obtained results.

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Here we present comparative data on the inhibition of lipid peroxidation by a variety of tocochromanols in liposomes. We also show for the first time the potential neuroprotective role of all the vitamin E homologues investigated on the neuronally differentiated human neuroblastoma SH-SY5Y cell line. α-Tocopherol had nearly no effect in the inhibition of lipid peroxidation, while ß-, γ-, and δ-tocopherols inhibited the reaction completely when it was initiated in a lipid phase. Similar effects were observed for tocotrienol homologues. Moreover, in this respect plastochromanol-8 was as effective as ß-, γ-, and δ-tocochromanols. When the prenyllipids were investigated in a 1,1-diphenyl-2-picrylhydrazyl (DPPH) test and incorporated into different lipid carriers, the radical oxidation was most pronounced in liposomes, followed by mixed micelles and the micellar system. When the reaction of tocochromanols was examined in niosomes, the oxidation was most pronounced for α-tocopherol and plastochromanol-8, followed by α-tocotrienol. Next, using retinoic acid-differentiated SH-SY5Y cells, we tested the protective effects of the compounds investigated on hydrogen peroxide (H2O2)-induced cell damage. We showed that tocotrienols were more active than tocopherols in the oxidative stress model. Plastochromanol-8 had a strong inhibitory effect on H2O2-induced lactate dehydrogenase (LDH) release and H2O2-induced decrease in cell viability. The water-soluble α-tocopherol phosphate had neuroprotective effects at all the concentrations analyzed. The results clearly indicate that structural differences between vitamin E homologues reflect their different biological activity and indicate their potential application in pharmacological treatments for neurodegenerative diseases. In this respect, the application of optimal tocochromanol-carrying structures might be critical.

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Carotenoids may have different effects on cancer and its progression. The safety of carotenoid supplements was evaluated in vitro on human non-small cell lung cancer (NSCLC) adenocarcinoma A549 cells by the administration of three different oleoresins containing lycopene and other lipophilic phytochemicals, such as tocochromanols. The oleoresins, obtained by the supercritical CO2 green extraction technology from watermelon (Lyc W), gac(Lyc G) and tomato (Lyc T) and chlatrated in α-cyclodextrins, were tested in comparison to synthetic lycopene (Lyc S), by cell cycle, Annexin V-FITC/PI, clonogenic test, Mytosox, intracellular ROS, Western Blot for NF-kB and RT-PCR and ELISA for IL-8. The extracts administered at the same lycopene concentration (10 µM) showed conflicting behaviors: Lyc W, with the highest lycopene/tocochromanols ratio, significantly increased cell apoptosis, mitochondrial stress, intracellular ROS, NF-kB and IL-8 expression and significantly decreased cell proliferation, whereas Lyc G and Lyc T significantly increased only cell proliferation. Lyc S treatment was ineffective. The highest amount of lycopene in Lyc W was able to counteract and revert the cell survival effect of tocochromanols supporting the importance of evaluating the lycopene bio-availability and the real effect of antioxidant tocochromanols' supplementation which may not only have no anticancer benefits but may even increase cancer aggressivity.

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A greenhouse experiment was conducted to study the effects of the application of a plant-derived biostimulant (Bioup® TF) on fruit quality and composition of two clusters (cluster II and cluster VI) of the cherry tomato cultivars 'Eletta', 'Kaucana', and 'Top Stellina'. The biostimulant application promoted fruit yield by 12% (up to 1.3 kg m-2 in 'Kaucana') and increased the concentrations of important functional constituents like phytoene, γ-tocopherol and ß-tocopherol by up to 16, 25, and 23%, respectively. Fruits from late-ripe cluster VI showed higher fruit weights, D-fructose, and total sugar contents than those from early-ripe cluster II (by 15, 7 and 5%, respectively), but reduced concentrations of acyclic carotenoids (phytoene and lycopene) and tocochromanols (mainly γ-tocopherol, -44%). 'Top Stellina' showed the highest responsiveness to the biostimulant, as particularly (all-E)-ß-carotene, phytofluene, and γ-tocopherol concentrations increased, indicating a genotype-dependent effect of the treatment. However, fruits of all treated genotypes showed a contextual decrease in D-fructose and total sugars in response to the biostimulant (on average by 7 and 10%, respectively), indicating a metabolic load burdening the accumulation of lipophilic antioxidants in cherry tomatoes at the expense of their taste-related C pool.

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Polar compounds and polymers are regarded as the most reliable indicators of oil degradation during heating, and it is desirable to find methods to reduce these undesirable changes. The aim of this study was (1) to determine the effect of enrichment with black cumin cold-pressed oil (CP) or essential oil obtained from black cumin cold-pressed oil in an equivalent amount (ES) on limiting the polar compounds and polymers content in blends based on refined rapeseed oil during high-temperature heating in a thin layer; (2) to determine tocochromanol losses and their effect on the change content of the polar compounds and polymers. Four fortified oils were made from refined rapeseed oil and one of the four additives (10% CP, 20% CP, 0.1% ES, and 0.2% ES). All fortified oils and refined rapeseed oil as a control sample were heated at 170 and 200 °C on the pan in a thin layer and evaluated regarding loss of individual tocochromanol homologs by HPLC-FL, polar compounds content, oxidized triacylglycerols (TAG), and polymers content by HPSEC-ELSD. Additionally, the fatty acid profile in nonheated oil was investigated. Tocochromanol analysis showed loss in all the samples. At 170 °C polymers were not detected; no difference was noted for polar compounds and oxidized TAG content; only the 20% CP sample showed a higher level. At 200 °C the 10% CP sample exhibited a significant protective effect with the lowest content of polar compounds, oxidized TAG, and dimers.

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The upcoming climate change presents a great challenge for plant growth and development being extremes temperatures among the major environmental limitations to crop productivity. Understanding the repercussions of these extreme temperatures is of high importance to elaborate future strategies to confront crop damages. Tomato plants (Solanum lycopersicum L.) are one of the most cultivated crops and their fruits are consumed worldwide standing out for their organoleptic characteristics and nutritional value. Tomato plants are sensitive to temperatures below 12 °C and above 32 °C. In this study, Micro-Tom cultivar was used to evaluate the effects of extreme temperatures on the plant of tomato and the fruit productivity and quality from the stressed plants, either exposed to cold (4 °C for three nights per week) or heat (32 °C during the day, seven days per week) treatments. Total productivity and the percentage of ripe fruits per plant were evaluated together with foliar stress markers and the contents of photosynthetic pigments and tocochromanols. Fruit quality was also assessed determining lycopene contents, total soluble solids, total acidity and ascorbate contents. High temperatures altered multiple physiological parameters indicating a moderate stress, particularly decreasing fruit yield. As a response to this stress, plants enhanced their antioxidant contents both at leaf and fruit level. Low temperatures did not negatively affect the physiology of plants with similar yields as compared to controls, suggesting chilling acclimation. Both high and low temperatures, but most particularly the former, increased total soluble solids contents indicating that temperature control may be used as a strategy to modulate fruit quality.

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The objective of this study was to use accelerated-solvent-extraction to achieve antioxidant extracts from chia seeds oils, enriched in tocopherols and tocotrienols, namely tocochromanols. Nanotechnology applications have been also incorporated to develop an innovative formulation of chia seeds oil nanoemulsion that preserve its antioxidant potential after conditions of oxidative stress. Chia seeds oils proved to be a valuable source of tocochromanols, from 568.84 to 855.98 µg g-1, depending on the geographical provenance. Quantitative data obtained by LC-DAD-ESI-MS/MS showed outstanding levels of γ-Tocopherol, over 83%, followed far behind by Tocopherols-(α, ß, δ) and Tocotrienols-(α, ß, δ, γ)-tocotrienols. The characteristic tocochromanols fingerprint of chia seeds oils was positively correlated with the FRAP and DPPH antioxidant activity of the extracts (between 18.81 and 138.48 mg Trolox/g). Formulation of the Chia seeds oils as nanoemulsions did not compromised the antioxidant properties of fresh extracts. Interestingly, nanoemulsions retained about the 80% of the initial antioxidant capacity after UV-induced stress, where the non-emulsified oils displayed a remarkable reduction (50-60%) on its antioxidant capacity under the same conditions. These antioxidant chia seeds formulations can constitute a promising strategy to vectorizing vitamin E isomers, in order to be used for food fortification, natural additives and to increase the self-life of food products during packing.

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The objective of this research was to investigate the differences between glaucous and non-glaucous near-isogenic lines (NILs) of winter rye (Secale cereale L.) in terms of epicuticular wax layer properties (weight, composition, and crystal morphology), selected physiological and biochemical responses, yield components, above-ground biomass, and plant height under soil drought stress. An important aspect of this analysis was to examine the correlation between the above characteristics. Two different NIL pairs were tested, each consisting of a typical glaucous line and a non-glaucous line with a recessive mutation. The drought experiment was conducted twice (2015-2016). Our study showed that wax accumulation during drought was not correlated with higher leaf hydration and glaucousness. Environmental factors had a large impact on the response of the lines to drought in individual years, both in terms of physiological and biochemical reactions, and the composition of epicuticular leaf wax. The analysed pairs displayed significantly different responses to drought. Demonstration of the correlation between the components of rye leaf wax and the physiological and biochemical parameters of rye NILs is a significant achievement of this work. Interestingly, the study showed a correlation between the wax components and the content of photosynthetic pigments and tocopherols, whose biosynthesis, similarly to the biosynthesis of wax precursors, is mainly located in chloroplasts. This suggests a relationship between wax biosynthesis and plant response to various environmental conditions and drought stress.

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Plant prenyllipids, especially isoprenoid chromanols and quinols, are very efficient low-molecular-weight lipophilic antioxidants, protecting membranes and storage lipids from reactive oxygen species (ROS). ROS are byproducts of aerobic metabolism that can damage cell components, they are also known to play a role in signaling. Plants are particularly prone to oxidative damage because oxygenic photosynthesis results in O2 formation in their green tissues. In addition, the photosynthetic electron transfer chain is an important source of ROS. Therefore, chloroplasts are the main site of ROS generation in plant cells during the light reactions of photosynthesis, and plastidic antioxidants are crucial to prevent oxidative stress, which occurs when plants are exposed to various types of stress factors, both biotic and abiotic. The increase in antioxidant content during stress acclimation is a common phenomenon. In the present review, we describe the mechanisms of ROS (singlet oxygen, superoxide, hydrogen peroxide and hydroxyl radical) production in chloroplasts in general and during exposure to abiotic stress factors, such as high light, low temperature, drought and salinity. We highlight the dual role of their presence: negative (i.e., lipid peroxidation, pigment and protein oxidation) and positive (i.e., contribution in redox-based physiological processes). Then we provide a summary of current knowledge concerning plastidic prenyllipid antioxidants belonging to isoprenoid chromanols and quinols, as well as their structure, occurrence, biosynthesis and function both in ROS detoxification and signaling.

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The aim of this study was to analyse contents of phytosterols, carotenoids, tocochromanols and fatty acid composition in oil from nutlets of linden. Standard determination methods for these compounds were applied using liquid and gas chromatography. In the analysed oils from small and large-leaved lime the dominant acids included linoleic (53.1 and 54.1%), oleic (18.5 and 22.2%), palmitic (15.3 and 11.8%). Linden seed oil was characterised by very high contents of phytosterols (2-2.5 g/100 g). ß-Sitosterol was the dominant phytosterol (80-83%). Additionally squalene was detected at 806 and 607 mg/100 g, respectively, for Tilia cordata and T.platyphyllos. It was found that linden seed oil contains tocopherols (93%) and tocotrienols (7%). In terms of tocochromanol contents this oil is surpassed only by wheat germ oil. In terms of the α-T equivalent these oils contain 95 (Tilia cordata) and 50.6 mg/100 g vitamin E (Tilia platyphyllos).

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BACKGROUND: One of the requirements for proper nutrition and maintenance of good health is to supply the body, through diet, with an appropriately increased quantity of bioactive compounds. With this in mind, modern milling and baking industries keep introducing new types of products. The use of such additives as wholegrain flours and bran in baked products provided the basis for research in this paper. METHODS: The aim of the study was to conduct a qualitative and quantitative evaluation of marketable wheat, barley and oat grain, used as raw materials to produce dehulled kernels, ground grain, wholemeal flour and wheat flour type 550 (all-purpose or plain flour), as well as wheat bran. Additionally, analyses were performed to determine the chemical composition and contents of nutrients, selected bioactive compounds and antioxidant activity. RESULTS: The studied raw materials in commercial cereal differ in their chemical composition. Dehulling of wheat, barley and oat grains significantly contributed to the reduction of minerals, protein and total dietary fiber (TDF) contents, except for the amount of protein in dehulled wheat and oat grains. Oat bran, in contrast to other oat products, was characterized by the highest contents of minerals, protein, TDF, and the smallest amounts of saccharides and total starch. The lowest content of minerals was recorded in wheat flour type 550. Thermal processes affect the concentration of tocochromanols in the grain, with tocotrienols being more resistant to hydrothermal treatment than tocopherols. Grain dehulling also significantly decreased the total amount of tocochromanols in relation to the original grains. CONCLUSIONS: Wheat products differ in the contents of their individual components. They are characterized by high contents of tocochromanols, phenolic compounds and water-extractable arabinoxylans, with the exception of white refined wheat flour, which is mainly a source of saccharides. In the case of barley and oat products, the analysis showed no differences between these product groups. It was also shown that dehulling of barley and oat grain causes statistically significant differences in the contents of nutrients and natural antioxidants.

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Tocochromanols are a group of lipid-soluble antioxidants that include tocopherols, tocotrienols and plastochromanol-8. Here, we examined a putative differential accumulation of tocochromanols in photosynthetic and non-photosynthetic tissues (including leaves and whole fruits) of strawberry (Fragaria x ananassa cv. Albion) plants and evaluated their endogenous variations in response to a reiterated water deficit during a vegetative (non-productive) and a fruiting (productive) period. In addition, we evaluated the concentration of tocochromanols in achenes (true fruits) and flesh of strawberries (whole fruits) at the white and full-red stages both under optimal and stress conditions. Results showed that leaves mainly accumulated α-tocopherol, with plastochromanol-8 and γ-tocopherol being present at low amounts. In contrast, whole fruits did not accumulate plastochromanol-8, γ-tocopherol being the major tocochromanol in the achenes (true fruit) and α-tocopherol in the flesh. While α-tocopherol content in leaves increased up to seven-fold after 12 weeks of stress during the fruiting period, it kept unaltered during the vegetative period. Neither plastochromanol-8 nor γ-tocopherol contents increased in leaves of stressed plants. During the fruiting period, γ-tocopherol content increased in whole fruits of stressed plants (most of it being accumulated in the achenes). Among the compounds examined, the flesh of strawberries accumulated α-tocopherol only, both at the white and full-red stages. It is concluded that (i) α-tocopherol is the major tocochromanol in leaves, while γ-tocopherol is the major tocochromanol in achenes (ii) reiterated water deficit promotes the accumulation of α-tocopherol in leaves and γ-tocopherol in fruits, (iii) α-tocopherol not only accumulates in photosynthetic tissues (leaves and whole fruits at green stages), but also in non-photosynthetic tissues (flesh of whole fruits at the white and full-red stages), and (iv) achenes (true fruits) of strawberry plants are an extraordinary rich source of tocopherols.

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Tocochromanols (tocopherols, tocotrienols and plastochromanol-8), isoprenoid quinone (plastoquinone-9 and plastoquinol-9) and carotenoids (carotenes and xanthophylls), are lipid-soluble antioxidants in the chloroplasts, which play an important defensive role against photooxidative stress in plants. In this study, the interplay between the antioxidant activities of those compounds in excess light stress was analyzed in wild-type (WT) Arabidopsis thaliana and in a tocopherol cyclase mutant (vte1), a homogentisate phytyl transferase mutant (vte2) and a tocopherol cyclase overexpressor (VTE1oex). The results reveal a strategy of cooperation and replacement between α-tocopherol, plastochromanol-8, plastoquinone-9/plastoquinol-9 and zeaxanthin. In the first line of defense (non-radical mechanism), singlet oxygen is either physically or chemically quenched by α-tocopherol; however, when α-tocopherol is consumed, zeaxanthin and plastoquinone-9/plastoquinol-9 can provide alternative protection against singlet oxygen toxicity by functional replacement of α-tocopherol either by zeaxanthin for the physical quenching or by plastoquinone-9/plastoquinol-9 for the chemical quenching. When singlet oxygen escapes this first line of defense, it oxidizes lipids and forms lipid hydroperoxides, which are oxidized to lipid peroxyl radicals by ferric iron. In the second line of defense (radical mechanism), lipid peroxyl radicals are scavenged by α-tocopherol. After its consumption, plastochromanol-8 overtakes this function. We provide a comprehensive description of the reaction pathways underlying the non-radical and radical antioxidant activities of α-tocopherol, carotenoids, plastoquinone-9/plastoquinol-9 and plastochromanol-8. The interplay between the different plastid lipid-soluble antioxidants in the non-radical and the radical mechanism provides step by step insights into protection against photooxidative stress in higher plants.

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