Our experts continually monitor the health and wellness space, and we update our articles when new information becomes available. Polyphenols are beneficial plant compounds thought to offer various health benefits. This article reviews everything you need to know about…. Flavonoids, once known as vitamin P, are a large class of plant compounds found in deeply colored fruits, vegetables, cocoa, tea, and wine.

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Jul 12, Written By Kathryn Watson. Benlloch-González et al. HKT , NHX , and SOS genes play an essential role in the transport of ions such as Na, K, and Ca Shi et al. Besides regulating ion transporter genes, plants also respond to high temperature, salt, and drought stresses by regulating transcription factor networks such as heat stress transcription factors HSFs.

HSFs play an important part in the regulation of stress-responsive genes, such as HSP genes, in response to certain abiotic stresses including salt and heat stress Guo et al. HSP is found in bacteria, yeast, and multicellular plants and localized to different compartments of cell, which play a decisive roles in cell adaptation to heat stress Mishra and Grover, Previous reports show that HSFA1 transcription factor is upregulated in response to salt stress, osmotic pressure, and oxidative stress at the seedling stage Ogawa et al.

Over-expression of HSFA1 in glycine max has been shown to increase heat tolerance due to the induction of HsP70 and HsP22 genes Zhu et al. In contrast, a HSFA2-knockout mutant line was completely unable to mitigate heat tolerance; however, this mechanism is not specific to HSFA2 because other HSFs can also exert positive feedback on other HSFs Jacob et al.

The expression levels of these genes depend on the pattern of applied stress. In general, the expression pattern of a gene differs when stresses are combined or when the component stresses are applied individually Rizhsky et al. In response to long-term abiotic stress, plants have evolved to activate their antioxidant machinery, e.

It has been reported that flavonoid molecules enhance salt stress tolerance through the scavenging of free radicals, a process that reduces oxidative stress Chandran et al. Previous studies have shown that flavonoid biosynthesis is induced by salt stress and enhances plant quality Yan et al.

Quercetin is an important flavonoid that has effective antioxidant properties; it plays a key role in mitigating oxidative stress as well as enhancing salinity and high-light tolerance in Arabidopsis thaliana and Ligustrum vulgare Agati et al.

It has also been reported that the accumulation of kaempferol and quercetin is enhanced during combined heat and salt stress; however, this regulation does not occur with salt stress alone Martinez et al. Bian et al. Similarly, we previously reported that flavanol 3-hydroxylase is a key regulator of flavonoid biosynthesis that participates in the responses to several stresses including UV stress and salinity Liu et al.

In solanum lycopersicu m, during combined heat and salt stress, the accumulation of flavonoids is enhanced due to the upregulation of the F3H gene; however, F3H was downregulated with salt stress alone in solanum lycopersicu m plants, whereas F3H transcript levels increased in Reaumuria trigyna in response to salt stress Zhang et al.

As F3H has been shown to be involved in various stress responses due to the regulation of flavonoid biosynthesis, we investigated the physiological responses and phenotypic variation of rice F3H transgenic plants in response to combined salt and heat stress.

Overall, the aim of our study was to understand the response mechanisms of rice F3H transgenic and wild plants to combined salt and heat stress. In the current experiment, we used nagdong rice cultivar, we used F3H gene overexpressor line of nogdang cultivar OxF3H; Jan et al. The seeds were pretreated with fungicides overnight and then washed three times with double-distilled water.

After successful growth, the seedlings were exposed to light and kept in a greenhouse for further experimentation.

We selected three groups of nagdong rice cultivar to evaluate the effect of combined salt and heat stress on rice plants.

The control group, i. One treatment group, wild-type treated Wt-t , was exposed to stress conditions. The other treatment group contained F3H overexpressor OxF3H plants was exposed to stress condition; thus, the group was named OxF3H-treated OxF3H-t.

We used only one transgenic line as reported in our previous study for comparison with the wild-type control and treated groups. After 1 month, Wt-t and OxF3H-t plants were treated with mM NaCl in growth chamber maintaining 40°C temperature, whereas Wt-cont plants were kept under normal conditions.

RNA extraction, cDNA synthesis, and qRT-PCR were performed following the methods of Jan et al. The primer and accession number for each gene are listed in Table 1.

We used RNeasy Plant Mini Kits 50 from Qiagen for total RNA extraction, and qPCRBIO and qPCRBIO SYBR Green Kits from PCR Biosystems for cDNA synthesis and qRT-PCR, respectively. Actin was used as a reporter gene. We quantified both SA and ABA to determine the cross talk between the two in response to combined salt and heat stress.

Quantification and extraction of both SA and ABA were performed using previously published protocols Jan et al. For SA analysis, 0. The supernatant was then collected, and the MeOH of the supernatant was dried in a vacuum centrifuge. The top organic layer containing free SA was transferred to 4-ml tubes and dried with nitrogen gas.

The quantity of SA was calculated according to the peak value for authentic standards. For endogenous ABA analysis, 0. The filtrate was concentrated in a rotary evaporator.

The aqueous phase, brought to a pH value of approximately 3. The EtOAc extracts were then combined and evaporated. The dried residue was dissolved in a phosphate buffer pH 8.

The pH value of the phosphate buffer was adjusted to 3. The EtOAc extracts were combined again and evaporated. The hypersensitive response HR of Wt-t and OxF3H-t plants to combined salt and heat stress was compared with that of Wt-cont plants using Trypan blue histological staining and following the method of Koch and Slusarenko Cell death was quantified using the density of Trypan blue in each leaf.

Similarly, the tips and middle of the roots were collected for HR response analysis after 1 week of continuous exposure to combined salt and heat stress. Ethanol treatment decolorized the chlorophyll in the leaves; only a brown polymerization product, produced by DAB with H 2 O 2 , remained.

After boiling, the leaves were cooled at room temperature in ethanol before brown spots were detected. The supernatant was collected in a new 2-mL tube and used for further analysis. A sample, positive control, and blank reaction were used according to the scheme shown in Table 2.

Following the user manual, one vial of NADPH assay reagent was reconstituted in 1. Readings were taken six times. For malondialdehyde, the kit provided MDA lysis buffer, phosphotungstic acid, BHT X, TBA, and MDA standard 4.

The TBA solution was reconstituted by adding 7. A μL measure of 4. Reaction was run in three technical replicates, and data are calculated using the following formula:. We isolated kaempferol and quercetin from the samples using the protocol described by Neff and Chory with slight modifications.

All samples were analyzed in triplicate. Samples were collected after 1 week of continuous stress and crushed into fine powder in liquid nitrogen. From this powder, 0. The obtained solvent was quantified using inductively coupled plasma mass spectrometry 9ICP-MS; Optima DV, PerkinElmer, United States.

After 1 week of continuous exposure to stress conditions, the chlorophyll content was measured in randomly selected leaves using a soil plant analysis development chlorophyll meter SPAD.

All experiments were performed in triplicate, and the data from each replicate were pooled. A completely randomized design was used to compare the mean values of different treatments.

Data were graphically presented, and statistical analyses were performed using GraphPad Prism software version 5. In this study, we focused on flavanol 3-hydroxylase, which plays a key role in flavonoid biosynthesis and abiotic stress mitigation.

The phenotypes under control and stress conditions are shown in Figure 1. Although shoot length was reduced in the transgenic line when compared to Wt-control plants, the reduction was less than that in Wt-t plants, indicating that F3H overexpression is involved in plant growth and development.

Heat stress damages leaf tips, which adversely affects plant development and yield; it was previously reported that heat stress reduces leaf area, causes leaf tip burn and yellowing of the entire leaf, and finally leads to death Sarsu, Beside leaf tip burn, we also noted some necrotic symptoms on the middle area of the leaves of Wt-t and OxF3H-t plants, which were more severe in Wt-t plants Supplementary Figure S1.

Taken together, these results suggest that overexpression of F3H mitigates salt and heat stress and can enhance growth and development under stress conditions.

Figure 1. Evaluation of growth parameters of Wt-cont, Wt-t, and OxF3H-t plants. A Shoot length, B root length, C leaf width, and D leaf tip damage following stress exposure. Lower panels show photographic representations of each phenotype. These results demonstrate that overexpression of F3H could differentially regulate salt- and heat-related genes.

Figure 2. Overexpression of F3H significantly regulates salt- and heat-related genes during combined salt and heat stress. A—F Relative expression of F3H , HKT , NHX , SOS , HSF , and HSP , respectively, in Wt-control, Wt-t, and OxF3H-t plants.

The fold change of each gene was measured after 0-, 3-, 6-, , and h intervals of continuous stress exposure, with actin used as a reference gene. ROS are generated as because of common stresses such as wounds, pathogens, drought, salinity, and heat.

Beside promoting programmed cell death, high levels of ROS produce harmful effects due to lipid peroxidation, protein denaturation, and DNA damage Miller and Mittler, To evaluate H 2 O 2 generation and cell death, we performed DAB and Trypan blue staining DAB interacts with H 2 O 2 and generates a reddish-brown polymer, whereas Trypan blue staining shows dead cells; Hoang et al.

Cell death and H 2 O 2 accumulation in roots were determined after 1 week of exposure to stress conditions Figure 3. Similarly, root tips and the middle parts of the root also accumulated more blue stains compared with that in OxF3H-t plants Figure 3B ; thus, combined salt and heat stress seems to cause substantial levels of cell death in Wt-t plants relative to that in OxF3H-t plants, which may further affect physiological processes.

We also noted that the root tip of Wt-t plants, which showed the highest accumulation of blue staining, was more sensitive to the stress condition than that of the OxF3H-t plants.

These results suggest that F3H expression is associated with cell death and H 2 O 2 generation. Figure 3. Combined salt and heat stress causes severe oxidative stress in leaves and roots.

A,B Cell death in leaves and roots, respectively, according to Trypan blue staining. C H 2 O 2 generation in the leaves of Wt-cont, Wt-t, and OxF3H-t plants as detected using DAB staining. Salt and heat stress regulate osmotic imbalance, which can be adjusted by regulation of flavonoids and antioxidant machinery that ameliorate stress conditions by scavenging free radicals Kumar et al.

Peroxidation of membrane lipids is a biomarker that characterizes the level of oxidative damage in plants under stress conditions; it is usually measured as an increase in MDA content. We measured MDA in response to combined salt and heat stress was significantly increased in Wt-t plants compared with the levels in Wt-cont plants Figure 4B.

Thus, high levels of membrane lipid peroxidation apparently occurred in Wt-t plants, which led to oxidative damage in these plants. The accumulation patterns of GPx and MDA were antagonistically associated, i. Figure 4. Quantification of antioxidants and flavonoids in wild-type and transgenic plants in response to combined salt and heat stress.

ABA levels increased significantly in response to stress exposure in both Wt-t and OxF3H-t plants compared with those in Wt-cont plants Figure 5A ; however, ABA levels were consistently reduced as stress duration increased.

Accumulation of SA was lower in OxF3H-t plants relative to that in Wt-t plants, but SA levels increased as stress duration increased. Comparing the levels of SA and ABA in OxF3H-t plants, the hormones were regulated differentially under stress conditions.

Thus, in the F3H transgenic line, SA accumulation increased and ABA accumulation decreased in response to combined salt and heat stress. Figure 5.

Abscisic acid ABA and salicylic acid SA are differentially regulated in wild-type and transgenic plants during combined salt and heat stress. Contrastingly, OxF3H-t plants showed the highest level of chlorophyll content of the three plant groups Figure 6C , indicating that F3H can positively regulate photosynthesis during salt and heat stress.

Figure 6. The purpose of this work was to assess the activity of F3H in response to the combined stress of salt and heat, two of the most prominent and damaging abiotic stressors that coexist in arid and semiarid locations across the world.

Individually, salt and heat stressors have been extensively investigated, with significant advances made in understanding their related physiological and molecular processes. After validating stress tolerance in the laboratory, researchers can release a substantial number of transgenic crops into agricultural fields.

Here, we explored the likely function of flavonoid accumulation in response to combined salt and heat stress, as well as the physiological, biochemical, and molecular responses of plants. The shoot length, root length, leaf width, and leaf tip tolerance levels were all higher in OxF3H-t plants than in Wt-t plants Figure 1.

According to many experts, stress-related inhibition of cell elongation and cell division, irregular ion homeostasis, and osmotic and oxidative stress generated by salt stress could all contribute to decreased plant growth and biomass accumulation Rahman et al.

In addition to phenotypic variation in wild-type and transgenic lines under stress, we found that OxF3H-t and Wt-t plants had higher and lower photosynthesis levels, respectively, than those in control plants Figure 6C. According to previous studies, salt stress disrupts photosynthetic pigments in solanum lycopersicu m plants, which is associated with oxidative damage Rahman et al.

Our study indicates that when salt and heat stress are combined, Wt-t plants experience more oxidative stress and ROS accumulation than those observed in OxF3H-t plants, which results in greater chlorophyll degradation in Wt-t plants than in OxF3H-t plants. According to previous observations, oxidative stress stimulates the production of ROS, which enhance chlorophyllase activity, which in turn is responsible for the degradation of photosynthetic pigments Ahmad et al.

Under combined salt and heat combined stress, OxF3H-t plants produced more kaempferol and quercetin than were produced by Wt-t plants Figures 4C , D , suggesting that these compounds may be involved in protecting photosynthetic pigments from degradation. Our findings are consistent with those of Parvin et al.

We postulate that increased accumulation of kaempferol and quercetin Figures 4C , D detoxifies the damaging H 2 O 2 molecules produced under combined salt and heat stress due to the antioxidant nature of these compounds. Beside phenotypic variations, F3H overexpression causes transcriptional differences in salt- and heat-related genes in response to combined salt and heat stress.

F3H is a key gene in the flavonoid biosynthesis pathway, which is upregulated during abiotic stress. In a previous study, we discovered that F3H gene expression boosts kaempferol and quercetin accumulation in rice Jan et al. These findings suggest that flavonoids and genes involved in their biosynthesis may be implicated in the response to abiotic stressors such as UV radiation, dehydration, heat, and salinity.

Under combined salt and heat stress, we measured the expression levels of HKT , NHX , SOS , HSF , and HSP in both wild-type and transgenic plants Figure 2.

Compared with control plants, HKT , SOS , HSF , and HSP were significantly upregulated in OxF3H-t plants, whereas NHX was downregulated. Although these genes were expressed at higher levels in Wt-t plants than in control plants, the differences were not statistically significant. Individually, salt stress affects the expression of HKT , NHX , and SOS , but there is little information on how salt and heat stress interact to affect these genes Zhang et al.

Because HKT and SOS expression was significantly higher in F3H transgenic plants compared to wild-type and control plants, our findings indicate that F3H enhances HKT and SOS expression during heat and salt stress. However, based on our findings, we expected that the combination of salt and heat stress suppresses NHX expression in OxF3H-t plants compared its expression in control plants.

Similarly, in several previous studies, it has been reported that HSF and HSP play prominent roles in individual salt and heat stresses Nover et al. A previous study showed that heat and salt stress increased accumulation of kaempferol and quercetin via regulation of flavonoid biosynthesis-related genes Xu et al.

These findings lead us to conclude that F3H, HKT, NHX, SOS, HSF, HSP, kaempferol, and quercetin all work synergistically to increase the tolerance of plants to combined salt and heat stress.

High temperatures and salinity cause oxidative stress, which is mediated by ROS and results in the regulation of stress hormones including ABA, SA, JA, and ethylene. These stress hormones are important for stress mediation and the establishment of a stress—growth balance Yu et al.

When comparing OxF3H-t to Wt-t and control plants, we found that ABA accumulation was higher, while SA accumulation was lower Figure 5.

Thus, ABA and SA apparently exhibit antagonistic cross talk in response to combined salt and heat stress because SA levels continuously increased, whereas ABA levels decreased as the stress period lengthened. Furthermore, de Torres Zabala et al. Our findings demonstrate that although ABA levels are higher in OxF3H-t plants, ROS accumulation is lower in these plants relative to the levels detected in Wt-t plants Figure 3.

Crop J — Tiwari S, Singh P, Tiwari R, Meena KK, Yandigeri M, Singh DP, Arora DK Salt-tolerant rhizobacteria-mediated induced tolerance in wheat Triticum aestivum and chemical diversity in rhizosphere enhance plant growth. Biol Fertil Soils — Venkidasamy B, Rajendran V, Sathishkumar R Flavonoids antioxidants systems in higher plants and their response to stresses.

Springer International Publishing AG Antioxidants and Antioxidant Enzymes in Higher. Plan Theory — War AR, Paulraj MG, Hussain B, Buhroo AA, Ignacimuthu S, Sharma HC Effect of plant secondary metabolites on Helicoverpa armigera.

J Pest Sci — Wei J, Xu M, Zhang D, Mi H The role of carotenoid isomerase in maintenance of photosynthetic oxygen evolution in rice plant. Acta Biochim Biophys Sin — Wu J, Ji J, Wang G, Wu G, Diao J, Li Z, Chen X, Chen Y, Luo L Ecotopic expression of the Lyciumbarbarum b-carotene hydroxylase gene chyb enhances drought and salt stress resistance by increasing xanthophyll cycle pool in tobacco.

Plant Cell Tissue Organ Cult — Yan Q, Cui X, Lin S, Gan S, Xing H, Dou D GmCYP82A3, a soybean cytochrome P family gene involved in the jasmonic acid and ethylene signaling pathway, enhances plant resistance to biotic and abiotic stresses.

PLoS One e Yang J, Yen HE Early salt stress effects on the changes in chemical composition in leaves of ice plant and Arabidopsis - a fourier transform infrared spectroscopy study. Yu O, Jung W, Shi J, Croes RA, Fader GM, McGonigle B, Odell JT Production of the isoflavones genistein and daidzein in non-legume dicot and monocot tissues.

Plant Physiol 2 — Zahran HH Rhizobium -legume symbiosis and nitrogen fixation under severe conditions and in an arid climate. Micro Mol Biol Rev 63 4 — Zhan X, Shen Q, Chen J, Yang P, Wang X, Hong Y Rice sulfoquinovosyltransferase SQD2.

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Flavonoids, once known as vitamin P, are a large class of plant compounds found in deeply colored fruits, vegetables, cocoa, tea, and wine. While they're not typically able to prescribe, nutritionists can still benefits your overall health.

Let's look at benefits, limitations, and more. A new study found that healthy lifestyle choices — including being physically active, eating well, avoiding smoking and limiting alcohol consumption —…. Carb counting is complicated.

Take the quiz and test your knowledge! Together with her husband, Kansas City Chiefs MVP quarterback Patrick Mahomes, Brittany Mohomes shares how she parents two children with severe food…. While there are many FDA-approved emulsifiers, European associations have marked them as being of possible concern.

Let's look deeper:. Researchers have found that a daily multivitamin supplement was linked with slowed cognitive aging and improved memory. Dietitians can help you create a more balanced diet or a specialized one for a variety of conditions.

We look at their benefits and limitations. Liquid collagen supplements might be able to reduce some effects of aging, but research is ongoing and and there may be side effects. A Quiz for Teens Are You a Workaholic? How Well Do You Sleep? Health Conditions Discover Plan Connect. What Are Flavonoids?

Everything You Need to Know. Medically reviewed by Miho Hatanaka, RDN, L. Sources Function Health benefits Takeaway Flavonoids are compounds found in many plant products, including teas, citrus fruits, and vegetables. Which foods have flavonoids?

What do flavonoids do? What are the health benefits of flavonoids? How we reviewed this article: Sources. Healthline has strict sourcing guidelines and relies on peer-reviewed studies, academic research institutions, and medical associations.

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Jul 12, Written By Kathryn Watson. Oct 23, Medically Reviewed By Miho Hatanaka, RDN, L. Share this article. Read this next. Am J Bot — Shiu CT, Lee TM Ultraviolet-B-induced oxidative stress and responses of the ascorbate—glutathione cycle in a marine macroalga Ulva fasciata.

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J Agric Food Chem — Download references. Department of Plant, Soil and Environmental Science, University of Florence, Viale delle Idee 30, , Sesto Fiorentino, Firenze, Italy.

Consiglio Nazionale delle Ricerche, Istituto per la Protezione delle Piante, Via Madonna del Piano 10, I, Sesto Fiorentino, Firenze, Italy. You can also search for this author in PubMed Google Scholar. Correspondence to Massimiliano Tattini.

Reprints and permissions. Di Ferdinando, M. Flavonoids as Antioxidants in Plants Under Abiotic Stresses. In: Ahmad, P. eds Abiotic Stress Responses in Plants. Springer, New York, NY. Published : 17 October Publisher Name : Springer, New York, NY.

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Policies and ethics. Skip to main content. Abstract Flavonoids make a relevant contribution to the response mechanisms of higher plants to a plethora of abiotic stresses. Keywords Abiotic stresses Antioxidant enzymes Auxin movement Dihydroxy B-ring-substituted flavonoids Reactive oxygen species UV-radiation.

Buying options Chapter EUR eBook EUR Softcover Book EUR Hardcover Book EUR Tax calculation will be finalised at checkout Purchases are for personal use only Learn about institutional subscriptions. References Agati G, Tattini M Multiple functional roles of flavonoids in photoprotecion.

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Of all the products that we at Andaman Ag offer, CropBioLife , a flavonoid product made from the rind of oranges and imported from Australia, where it has a long history of application, is a product built for our times.

Plants easily synthesize CropBioLife to augment their own flavonoid production. Soil is the largest carbon store on Earth—holding more carbon than all plants and our atmosphere put together.

Plants and fungi struck a deal way back when. More than million years ago, plants began trading sugar carbon made from sunlight for some of the soil nutrients gathered by mycorrhizal fungi.

Plant phenols, which are a type of secondary metabolite found in plants, can play a role in mitigating heat stress. Heat stress and plants can occur when temperatures rise above the optimal range for their growth and development, leading to various physiological and biochemical changes.

First Name. Last Name. Sign Up Now. Home 5 News 5 Flavonoids Reduce Heat and Drought Stress. Flavonoids Reduce Heat and Drought Stress. Disclaimer My opinions and perspectives may differ from the information provided on the product label. Related Posts.

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Sign up now and get a FREE copy of the Best Diets for Cognitive Fitness. Stay on top of latest health news from Harvard Medical School. Recent Blog Articles. Flowers, chocolates, organ donation — are you in?

What is a tongue-tie? What parents need to know. Which migraine medications are most helpful? How well do you score on brain health? Shining light on night blindness.

Can watching sports be bad for your health? Beyond the usual suspects for healthy resolutions. April 14, These plant chemicals, found in most fruits and vegetables, may play an important role in cognitive fitness and dementia prevention.

Class in itself Flavonoids are phytochemicals commonly found in plant foods. Flavonoids and food There are six subclasses of flavonoids, which are abundant in most plant foods. Flavonoid subclass Common food sources Anthocyanidins berries red, blue, and purple , grapes, red wine Flavanols tea black, white, green, oolong , cocoa-based products, grapes, berries, apples, red wine Flavanones citrus fruit oranges, lemons, grapefruit Flavones parsley, thyme, celery, hot peppers Flavonols onions, peppers, broccoli, spinach, apples, berries, tea Polymers berries cranberries, blueberries, strawberries , wine, grapes, black tea Many factors in play Large human studies of flavonoids are still in their early phases, but initial findings show promise.

Color coordination So, to protect your brain from dementia, should you load up your plate with as many flavonoid-rich foods as possible? Share This Page Share this page to Facebook Share this page to Twitter Share this page via Email.

Print This Page Click to Print. Related Content. You might also be interested in…. A Guide to Cognitive Fitness In this Special Health Report, Harvard Medical School doctors share a six-step program that can yield important and lasting results. And if you're not a drinker, you can get almost all the same benefits from purple grape juice.

The pomegranate carries with it the mystique of ancient myth, but we moderns are beginning to realize that its health benefits are very real: pomegranate juice may have almost three times the antioxidant potency of an equal volume of green tea or red wine. Delicious, relaxing chamomile tea is home to the flavonoid called apigenin, one of a handful of flavonoids recently found to have mood-enhancing properties.

Currently the focus of intense study, they are thought to act on the same parts of the brain as common anti- anxiety drugs. In fact, certain synthetic flavonoids have been shown to have anxiolytic properties superior to diazepam.

Research is in its infancy, however. For now, take your apigenin with sugar and lemon. While many health benefits of flavonoids are not in dispute, there are a couple caveats to consider before sitting down to fill your face with flavonoids. First, scientists are only now beginning to understand the effects of flavonoids in the body.

As natural but real chemicals, flavonoids can interact with prescription drugs in a harmful way. The flavonoid naringenin found in grapefruit, for example, can interfere with the breakdown of certain drugs, magnifying their potency. It's best not to take any drugs with grapefruit juice unless the drug interaction profile of the medication is well known.

Second, taking flavonoid supplements is not the way to go. UC Davis' Dr. Mitchell cautions people not to think they can just take a supplement instead of consuming more fruits and vegetables.

Whole foods supply the added benefits of vitamins, minerals and fiber. Furthermore, the dosage furnished by supplements can vary widely but is likely to be much higher than what you'd receive from a healthy, balanced diet. Reseachers have yet to determine exactly what levels of flavonoids are optimally beneficial, or even whether flavonoids become harmful at very high doses.

As with all supplements, flavonoid supplements are not stringently regulated by the FDA. Mitchell, "and it's important not to view them as the latest fad cure-all.

If you're curious to learn about more whole food sources of flavonoids, the USDA has an online database of foods and their flavonoid content. You can find it at www. Flavonoids: Antioxidants Help the Mind Naturally occurring plant pigments, flavonoids are one of the reasons fruits and vegetables are so good for you.

By Erik Strand published July 8, - last reviewed on June 9, Share. We selected three groups of nagdong rice cultivar to evaluate the effect of combined salt and heat stress on rice plants.

The control group, i. One treatment group, wild-type treated Wt-t , was exposed to stress conditions. The other treatment group contained F3H overexpressor OxF3H plants was exposed to stress condition; thus, the group was named OxF3H-treated OxF3H-t. We used only one transgenic line as reported in our previous study for comparison with the wild-type control and treated groups.

After 1 month, Wt-t and OxF3H-t plants were treated with mM NaCl in growth chamber maintaining 40°C temperature, whereas Wt-cont plants were kept under normal conditions. RNA extraction, cDNA synthesis, and qRT-PCR were performed following the methods of Jan et al. The primer and accession number for each gene are listed in Table 1.

We used RNeasy Plant Mini Kits 50 from Qiagen for total RNA extraction, and qPCRBIO and qPCRBIO SYBR Green Kits from PCR Biosystems for cDNA synthesis and qRT-PCR, respectively. Actin was used as a reporter gene. We quantified both SA and ABA to determine the cross talk between the two in response to combined salt and heat stress.

Quantification and extraction of both SA and ABA were performed using previously published protocols Jan et al. For SA analysis, 0. The supernatant was then collected, and the MeOH of the supernatant was dried in a vacuum centrifuge.

The top organic layer containing free SA was transferred to 4-ml tubes and dried with nitrogen gas. The quantity of SA was calculated according to the peak value for authentic standards.

For endogenous ABA analysis, 0. The filtrate was concentrated in a rotary evaporator. The aqueous phase, brought to a pH value of approximately 3.

The EtOAc extracts were then combined and evaporated. The dried residue was dissolved in a phosphate buffer pH 8. The pH value of the phosphate buffer was adjusted to 3.

The EtOAc extracts were combined again and evaporated. The hypersensitive response HR of Wt-t and OxF3H-t plants to combined salt and heat stress was compared with that of Wt-cont plants using Trypan blue histological staining and following the method of Koch and Slusarenko Cell death was quantified using the density of Trypan blue in each leaf.

Similarly, the tips and middle of the roots were collected for HR response analysis after 1 week of continuous exposure to combined salt and heat stress. Ethanol treatment decolorized the chlorophyll in the leaves; only a brown polymerization product, produced by DAB with H 2 O 2 , remained.

After boiling, the leaves were cooled at room temperature in ethanol before brown spots were detected. The supernatant was collected in a new 2-mL tube and used for further analysis. A sample, positive control, and blank reaction were used according to the scheme shown in Table 2.

Following the user manual, one vial of NADPH assay reagent was reconstituted in 1. Readings were taken six times. For malondialdehyde, the kit provided MDA lysis buffer, phosphotungstic acid, BHT X, TBA, and MDA standard 4.

The TBA solution was reconstituted by adding 7. A μL measure of 4. Reaction was run in three technical replicates, and data are calculated using the following formula:.

We isolated kaempferol and quercetin from the samples using the protocol described by Neff and Chory with slight modifications. All samples were analyzed in triplicate. Samples were collected after 1 week of continuous stress and crushed into fine powder in liquid nitrogen.

From this powder, 0. The obtained solvent was quantified using inductively coupled plasma mass spectrometry 9ICP-MS; Optima DV, PerkinElmer, United States.

After 1 week of continuous exposure to stress conditions, the chlorophyll content was measured in randomly selected leaves using a soil plant analysis development chlorophyll meter SPAD.

All experiments were performed in triplicate, and the data from each replicate were pooled. A completely randomized design was used to compare the mean values of different treatments. Data were graphically presented, and statistical analyses were performed using GraphPad Prism software version 5.

In this study, we focused on flavanol 3-hydroxylase, which plays a key role in flavonoid biosynthesis and abiotic stress mitigation. The phenotypes under control and stress conditions are shown in Figure 1. Although shoot length was reduced in the transgenic line when compared to Wt-control plants, the reduction was less than that in Wt-t plants, indicating that F3H overexpression is involved in plant growth and development.

Heat stress damages leaf tips, which adversely affects plant development and yield; it was previously reported that heat stress reduces leaf area, causes leaf tip burn and yellowing of the entire leaf, and finally leads to death Sarsu, Beside leaf tip burn, we also noted some necrotic symptoms on the middle area of the leaves of Wt-t and OxF3H-t plants, which were more severe in Wt-t plants Supplementary Figure S1.

Taken together, these results suggest that overexpression of F3H mitigates salt and heat stress and can enhance growth and development under stress conditions. Figure 1. Evaluation of growth parameters of Wt-cont, Wt-t, and OxF3H-t plants.

A Shoot length, B root length, C leaf width, and D leaf tip damage following stress exposure. Lower panels show photographic representations of each phenotype. These results demonstrate that overexpression of F3H could differentially regulate salt- and heat-related genes.

Figure 2. Overexpression of F3H significantly regulates salt- and heat-related genes during combined salt and heat stress. A—F Relative expression of F3H , HKT , NHX , SOS , HSF , and HSP , respectively, in Wt-control, Wt-t, and OxF3H-t plants.

The fold change of each gene was measured after 0-, 3-, 6-, , and h intervals of continuous stress exposure, with actin used as a reference gene. ROS are generated as because of common stresses such as wounds, pathogens, drought, salinity, and heat.

Beside promoting programmed cell death, high levels of ROS produce harmful effects due to lipid peroxidation, protein denaturation, and DNA damage Miller and Mittler, To evaluate H 2 O 2 generation and cell death, we performed DAB and Trypan blue staining DAB interacts with H 2 O 2 and generates a reddish-brown polymer, whereas Trypan blue staining shows dead cells; Hoang et al.

Cell death and H 2 O 2 accumulation in roots were determined after 1 week of exposure to stress conditions Figure 3. Similarly, root tips and the middle parts of the root also accumulated more blue stains compared with that in OxF3H-t plants Figure 3B ; thus, combined salt and heat stress seems to cause substantial levels of cell death in Wt-t plants relative to that in OxF3H-t plants, which may further affect physiological processes.

We also noted that the root tip of Wt-t plants, which showed the highest accumulation of blue staining, was more sensitive to the stress condition than that of the OxF3H-t plants.

These results suggest that F3H expression is associated with cell death and H 2 O 2 generation. Figure 3. Combined salt and heat stress causes severe oxidative stress in leaves and roots. A,B Cell death in leaves and roots, respectively, according to Trypan blue staining. C H 2 O 2 generation in the leaves of Wt-cont, Wt-t, and OxF3H-t plants as detected using DAB staining.

Salt and heat stress regulate osmotic imbalance, which can be adjusted by regulation of flavonoids and antioxidant machinery that ameliorate stress conditions by scavenging free radicals Kumar et al.

Peroxidation of membrane lipids is a biomarker that characterizes the level of oxidative damage in plants under stress conditions; it is usually measured as an increase in MDA content. We measured MDA in response to combined salt and heat stress was significantly increased in Wt-t plants compared with the levels in Wt-cont plants Figure 4B.

Thus, high levels of membrane lipid peroxidation apparently occurred in Wt-t plants, which led to oxidative damage in these plants. The accumulation patterns of GPx and MDA were antagonistically associated, i. Figure 4. Quantification of antioxidants and flavonoids in wild-type and transgenic plants in response to combined salt and heat stress.

ABA levels increased significantly in response to stress exposure in both Wt-t and OxF3H-t plants compared with those in Wt-cont plants Figure 5A ; however, ABA levels were consistently reduced as stress duration increased.

Accumulation of SA was lower in OxF3H-t plants relative to that in Wt-t plants, but SA levels increased as stress duration increased. Comparing the levels of SA and ABA in OxF3H-t plants, the hormones were regulated differentially under stress conditions.

Thus, in the F3H transgenic line, SA accumulation increased and ABA accumulation decreased in response to combined salt and heat stress. Figure 5. Abscisic acid ABA and salicylic acid SA are differentially regulated in wild-type and transgenic plants during combined salt and heat stress.

Contrastingly, OxF3H-t plants showed the highest level of chlorophyll content of the three plant groups Figure 6C , indicating that F3H can positively regulate photosynthesis during salt and heat stress.

Figure 6. The purpose of this work was to assess the activity of F3H in response to the combined stress of salt and heat, two of the most prominent and damaging abiotic stressors that coexist in arid and semiarid locations across the world. Individually, salt and heat stressors have been extensively investigated, with significant advances made in understanding their related physiological and molecular processes.

After validating stress tolerance in the laboratory, researchers can release a substantial number of transgenic crops into agricultural fields.

Here, we explored the likely function of flavonoid accumulation in response to combined salt and heat stress, as well as the physiological, biochemical, and molecular responses of plants. The shoot length, root length, leaf width, and leaf tip tolerance levels were all higher in OxF3H-t plants than in Wt-t plants Figure 1.

According to many experts, stress-related inhibition of cell elongation and cell division, irregular ion homeostasis, and osmotic and oxidative stress generated by salt stress could all contribute to decreased plant growth and biomass accumulation Rahman et al. In addition to phenotypic variation in wild-type and transgenic lines under stress, we found that OxF3H-t and Wt-t plants had higher and lower photosynthesis levels, respectively, than those in control plants Figure 6C.

According to previous studies, salt stress disrupts photosynthetic pigments in solanum lycopersicu m plants, which is associated with oxidative damage Rahman et al. Our study indicates that when salt and heat stress are combined, Wt-t plants experience more oxidative stress and ROS accumulation than those observed in OxF3H-t plants, which results in greater chlorophyll degradation in Wt-t plants than in OxF3H-t plants.

According to previous observations, oxidative stress stimulates the production of ROS, which enhance chlorophyllase activity, which in turn is responsible for the degradation of photosynthetic pigments Ahmad et al.

Under combined salt and heat combined stress, OxF3H-t plants produced more kaempferol and quercetin than were produced by Wt-t plants Figures 4C , D , suggesting that these compounds may be involved in protecting photosynthetic pigments from degradation.

Our findings are consistent with those of Parvin et al. We postulate that increased accumulation of kaempferol and quercetin Figures 4C , D detoxifies the damaging H 2 O 2 molecules produced under combined salt and heat stress due to the antioxidant nature of these compounds.

Beside phenotypic variations, F3H overexpression causes transcriptional differences in salt- and heat-related genes in response to combined salt and heat stress.

F3H is a key gene in the flavonoid biosynthesis pathway, which is upregulated during abiotic stress. In a previous study, we discovered that F3H gene expression boosts kaempferol and quercetin accumulation in rice Jan et al. These findings suggest that flavonoids and genes involved in their biosynthesis may be implicated in the response to abiotic stressors such as UV radiation, dehydration, heat, and salinity.

Under combined salt and heat stress, we measured the expression levels of HKT , NHX , SOS , HSF , and HSP in both wild-type and transgenic plants Figure 2.

Compared with control plants, HKT , SOS , HSF , and HSP were significantly upregulated in OxF3H-t plants, whereas NHX was downregulated. Although these genes were expressed at higher levels in Wt-t plants than in control plants, the differences were not statistically significant.

Individually, salt stress affects the expression of HKT , NHX , and SOS , but there is little information on how salt and heat stress interact to affect these genes Zhang et al. Because HKT and SOS expression was significantly higher in F3H transgenic plants compared to wild-type and control plants, our findings indicate that F3H enhances HKT and SOS expression during heat and salt stress.

However, based on our findings, we expected that the combination of salt and heat stress suppresses NHX expression in OxF3H-t plants compared its expression in control plants.

Similarly, in several previous studies, it has been reported that HSF and HSP play prominent roles in individual salt and heat stresses Nover et al. A previous study showed that heat and salt stress increased accumulation of kaempferol and quercetin via regulation of flavonoid biosynthesis-related genes Xu et al.

These findings lead us to conclude that F3H, HKT, NHX, SOS, HSF, HSP, kaempferol, and quercetin all work synergistically to increase the tolerance of plants to combined salt and heat stress. High temperatures and salinity cause oxidative stress, which is mediated by ROS and results in the regulation of stress hormones including ABA, SA, JA, and ethylene.

These stress hormones are important for stress mediation and the establishment of a stress—growth balance Yu et al. When comparing OxF3H-t to Wt-t and control plants, we found that ABA accumulation was higher, while SA accumulation was lower Figure 5.

Thus, ABA and SA apparently exhibit antagonistic cross talk in response to combined salt and heat stress because SA levels continuously increased, whereas ABA levels decreased as the stress period lengthened.

Furthermore, de Torres Zabala et al. Our findings demonstrate that although ABA levels are higher in OxF3H-t plants, ROS accumulation is lower in these plants relative to the levels detected in Wt-t plants Figure 3.

This implies that OxF3H-t plants accumulate more ABA, which in turn lowers oxidative damage during combined salt and heat stress. Furthermore, we discovered that the transgenic line exhibited increased levels of kaempferol, quercetin, and ABA, which suggests that a link exists between flavonoid and hormone signaling in response to combined salt and heat stress.

Our findings are consistent with prior research showing that flavonoids were enhanced in ABA-, SA-, and JA-treated plants Thiruvengadam et al.

Moreover, based on the research of Hung and Kao , we may postulate that ABA and SA regulate the primary enzyme in the flavonoid production pathway, which results in increased levels of kaempferol and quercetin. Overall, our study demonstrates that wild-type and OsF3H transgenic rice plants respond differently to combined salt and heat stress in terms of their physiological, biochemical, and molecular responses.

During salt and heat stress, induced expression of F3H increases plant biomass and photosynthesis. Both salt and heat stress increase oxidative stress, which is mitigated by the high accumulation of kaempferol and quercetin, given that constant expression of F3H significantly enhances both of these flavonoid molecules, which are known to scavenge ROS.

Because of the antagonistic cross talk between ABA and SA in one transgenic line, we suggest that F3H is involved in the regulation of hormonal machinery in response to combined salt and heat stress. Overall, the overexpression of F3H seems to regulate the physiological, biochemical, and molecular machinery of rice plants during stress exposure involving salinity and heat.

RJ, S-HL, and K-MK designed the study. RJ, S-HL, and NK performed the experiments. MK, SAa, and S-HL performed the analyses. I-JL provided resources. RJ and S-HL wrote the manuscript. SAi produced the figures and revised the manuscript. All authors contributed to the article and approved the submitted version.

This work was supported by the National Research Foundation of Korea Grant funded by the Korean Government NRFM3E5E The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers.

Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher. Agati, G. The biosynthesis of flavonoids is enhanced similarly by UV radiation and root zone salinity in L. vulgare leaves.

The current world of global warming and a constantly changing environment have made life very stressful for living entities, which has driven the evolution of biochemical processes to cope with stressful environmental and ecological conditions.

As climate change continues, we anticipate more frequent occurrences of abiotic stresses such as drought, high temperature and salinity. However, plants are equipped with biosynthetic machinery that can supply thousands of bio-compounds that may be required for maintaining internal homeostasis or a relatively stable equilibrium.

Historically, plants adapt, and flavonoids are a catalyst to help in the long-term acclimation process. Flavonoids have the potential to assist plants in tolerating, resisting and escaping biotic and abiotic stresses generated by the external environment.

The function of flavonoids as signal compounds to communicate with rhizosphere microbes is indispensable, positively impacting the structure and function of the rhizosphere community.

We want to increase flavonoid production in plants in high-stress situations. Of all the products that we at Andaman Ag offer, CropBioLife , a flavonoid product made from the rind of oranges and imported from Australia, where it has a long history of application, is a product built for our times.

Plants easily synthesize CropBioLife to augment their own flavonoid production. Soil is the largest carbon store on Earth—holding more carbon than all plants and our atmosphere put together. Plants and fungi struck a deal way back when. More than million years ago, plants began trading sugar carbon made from sunlight for some of the soil nutrients gathered by mycorrhizal fungi.

Plant phenols, which are a type of secondary metabolite found in plants, can play a role in mitigating heat stress.

Heat stress and plants can occur when temperatures rise above the optimal range for their growth and development, leading to various physiological and biochemical changes. Discrimination at work is linked to high blood pressure. Icy fingers and toes: Poor circulation or Raynaud's phenomenon?

Science already touts the heart-health benefits of plant-based diets like the Mediterranean, MIND, and DASH diets. Now researchers have turned their attention to how plant-based eating also may prevent dementia, including Alzheimer's disease. And one class of nutrients in plants continues to stand out: flavonoids.

Tian-Shin Yeh, a researcher in the area of diet and cognitive function with Harvard's T. Chan School of Public Health. Flavonoids are phytochemicals commonly found in plant foods. They have six subclasses: anthocyanidins, flavanols, flavanones, flavones, flavonols, and polymers. A variety of fruits and vegetables contain flavonoids.

Some of the highest amounts are in berries, apples, citrus fruit oranges, lemons , grapes, spinach, legumes, kale, broccoli, soybeans, onions, tea, cocoa, and wine.

See "Flavonoids and food. Researchers believe flavonoids may help the brain in several ways. For instance, studies suggest they reduce cell-damaging free radicals and soothe inflammation.

Some early-stage animal studies have shown that flavonoids can block beta-amyloid plaque buildup in the brain, a trademark of Alzheimer's. Flavonoids also may enhance brain blood flow. There are six subclasses of flavonoids, which are abundant in most plant foods. tea black, white, green, oolong , cocoa-based products, grapes, berries, apples, red wine.

Large human studies of flavonoids are still in their early phases, but initial findings show promise. One of the most substantial to date was a recent study linking high flavonoid intake and a lower risk of Alzheimer's and related dementias conditions with symptoms similar to Alzheimer's.

The results were published online April 22, , by The American Journal of Clinical Nutrition. Using data from the Framingham Heart Study, researchers looked at the dietary habits of almost 3, people, average age 59, without any signs of dementia.

Over 20 years, people with the highest daily intake of flavonoids about milligrams had a lower risk of developing Alzheimer's or a related dementia than those who ate the smallest amounts about milligrams.

Still, the link between flavonoids and brain health might be a matter of coincidence. So, to protect your brain from dementia, should you load up your plate with as many flavonoid-rich foods as possible? Not really, according to Dr. Both salt and heat stress increase oxidative stress, which is mitigated by the high accumulation of kaempferol and quercetin, given that constant expression of F3H significantly enhances both of these flavonoid molecules, which are known to scavenge ROS.

Because of the antagonistic cross talk between ABA and SA in one transgenic line, we suggest that F3H is involved in the regulation of hormonal machinery in response to combined salt and heat stress. Overall, the overexpression of F3H seems to regulate the physiological, biochemical, and molecular machinery of rice plants during stress exposure involving salinity and heat.

RJ, S-HL, and K-MK designed the study. RJ, S-HL, and NK performed the experiments. MK, SAa, and S-HL performed the analyses. I-JL provided resources. RJ and S-HL wrote the manuscript.

SAi produced the figures and revised the manuscript. All authors contributed to the article and approved the submitted version. This work was supported by the National Research Foundation of Korea Grant funded by the Korean Government NRFM3E5E The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.

Agati, G. The biosynthesis of flavonoids is enhanced similarly by UV radiation and root zone salinity in L.

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