Anthocyanins as protectors of gut microbiota: mitigating the adverse effects of microplastic-induced disruption

Abstract

Anthocyanins, potent bioactive compounds found abundantly in berries, as well as in many other pigmented edible plants, have garnered significant attention for their health-promoting properties, particularly in relation to gut microbiota. This review focuses on the protective role of anthocyanins against gut microbiota disruption caused by microplastics, environmental pollutants that have triggered increased concerns in recent years for their impact on ecosystems and human health. By synthesizing current research, the mechanisms through which anthocyanins may exert their beneficial effects are explored, mitigating the negative health effects of microplastic ingestion. The paper also discusses the potential application of anthocyanin-rich functional foods and supplements as a strategy to preserve gut health in the face of rising environmental challenges.

Keywords

Microbiota / Microplastics / Anthocyanins

Author contributions

The authors confirm contribution to the paper as follows: draft manuscript preparation: Koistinen VM, Babu AF, Shad E, Zarei I; table and figures preparation: Koistinen VM, Babu AF. All authors reviewed the results and approved the final version of the manuscript.

Data availability

Data sharing not applicable to this article as no datasets were generated or analyzed during the current study.

Acknowledgments

This work was supported by the Ministry of Education and Culture, Finland through Seed Funding for the Finland-China Food and Health Network (FCFH) 2023, University of Eastern Finland. Koistinen VM was supported by the Lantmännen Research Foundation (Grant No. 2022H045). Babu AF received a working grant from the Finnish Cultural Foundation (Grant No. 00210224).

Conflict of interest

The authors declare that they have no conflict of interest.

References

[1] Mattioli R, Francioso A, Mosca L, Silva P. 2020. Anthocyanins: a comprehensive review of their chemical properties and health effects on cardiovascular and neurodegenerative diseases. Molecules 25:3809

[2] Hsu WY, Shipman PD, Thompson S. 2023. Molecular transformations and self-association in anthocyanin pigment patterns. Journal of Biosciences 49:4

[3] Ayvaz H, Cabaroglu T, Akyildiz A, Pala CU, Temizkan R, et al. 2022. Anthocyanins: metabolic digestion, bioavailability, therapeutic effects, current pharmaceutical/industrial use, and innovation potential. Antioxidants 12:48

[4] Sendri N, Bhandari P. 2024. Anthocyanins: a comprehensive review on biosynthesis, structural diversity, and industrial applications. Phytochemistry Reviews 23:1913-74

[5] Golovko TK. 2023. Plant anthocyanins: structure, biosynthesis regulation, functions, and ecology. Russian Journal of Plant Physiology 70:161

[6] Cappellini F, Marinelli A, Toccaceli M, Tonelli C, Petroni K. 2021. Anthocyanins: from mechanisms of regulation in plants to health benefits in foods. Frontiers in Plant Science 12:748049

[7] Alam MA, Islam P, Subhan N, Rahman MM, Khan F, et al. 2021. Potential health benefits of anthocyanins in oxidative stress related disorders. Phytochemistry Reviews 20:705-49

[8] EFSA Panel on Dietetic Products, Nutrition and Allergies. 2010. Scientific Opinion on the substantiation of health claims related to various food ( s )/food constituent ( s ) and protection of cells from premature ageing (ID 1668, 1917, 2515, 2527, 2530, 2575, 2580, 2591, 2620, 3178, 3179, 3180, 3181, 4329, 4415), antioxidant activity, antioxidant content and antioxidant properties (ID 857, 1306, 2515, 2527, 2530, 2575, 2580, 2591, 2629, 2728, 4327, 4365, 4380, 4390, 4394, 4455, 4464, 4507, 4694, 4705), protection of DNA, proteins and lipids from oxidative damage (ID 1196, 1211, 1216, 1306, 1312, 1440, 1441, 1666, 1668, 1692, 1900, 1914, 1948, 2023, 2158, 2517, 2522, 2527, 2575, 2591, 2620,2637,2639,2663,2860,3079,3276,3564,3818,4324,4329, 4351,4397,4416,4424,4507,4527,4528,4542,4611,4629,4659) and bioavailability of anthocyanins in black currants (ID 4220) pursuant to Article 13 (1) of Regulation (EC) No 1924/2006. EFSA Journal 8(10): 1752

[9] Chaiyasut C, Sivamaruthi B, Kesika P, Subasankari K. 2018. Beneficial effects of anthocyanins against diabetes mellitus associated conse-quences-A mini review. Asian Pacific Journal of Tropical Biomedicine 8:471-77

[10] Kapoor P, Tiwari A, Sharma S, Tiwari V, Sheoran B, et al. 2023. Effect of anthocyanins on gut health markers, Firmicutes-Bacteroidetes ratio and short-chain fatty acids: a systematic review via meta-analysis. Scientific Reports 13:1729

[11] Liang A, Leonard W, Beasley JT, Fang Z, Zhang P, et al. 2024. Antho-cyanins-gut microbiota-health axis: a review. Critical reviews in food science and nutrition 64:7563-88

[12] Verediano TA, Stampini Duarte Martino H, Dias Paes MC, Tako E. 2021. Effects of anthocyanin on intestinal health: a systematic review. Nutrients 13:1331

[13] Kumkum R, Aston-Mourney K, McNeill BA, Hernández D, Rivera LR. 2024. Bioavailability of Anthocyanins: whole Foods versus Extracts. Nutrients 16:1403

[14] Bhan C, Anita, Kumar N. 2024. Sources, impacts and distribution of microplastics in different environmental matrices: a review. Environmental Sustainability 7:171-80

[15] Jiang B, Kauffman AE, Li L, McFee W, Cai B, et al. 2020. Health impacts of environmental contamination of micro-and nanoplastics: a review. Environmental Health and Preventive Medicine 25:1-15

[16] Amobonye A, Bhagwat P, Raveendran S, Singh S, Pillai S. 2021. Environmental impacts of microplastics and nanoplastics: a current overview. Frontiers in Microbiology 12:768297

[17] Osman AI, Hosny M, Eltaweil AS, Omar S, Elgarahy AM, et al. 2023. Microplastic sources, formation, toxicity and remediation: a review. Environmental Chemistry Letters 21:2129-69

[18] Wright SL, Kelly FJ. 2017. Plastic and human health: a micro issue? Environmental Science & Technology 51:6634-47

[19] Pironti C, Ricciardi M, Motta O, Miele Y, Proto A, et al. 2021. Microplastics in the environment: intake through the food web, human exposure and toxicological effects. Toxics 9:224

[20] Andrady AL. 2011. Microplastics in the marine environment. Marine Pollution Bulletin 62:1596-605

[21] Eriksen M, Lebreton LCM, Carson HS, Thiel M, Moore CJ, et al. 2014. Plastic pollution in the world's oceans: more than 5 trillion plastic pieces weighing over 250,000 tons afloat at sea. PLoS one 9:e111913

[22] Yang H, Chen G, Wang J. 2021. Microplastics in the marine environment: Sources, fates, impacts and microbial degradation. Toxics 9:41

[23] Boucher J, Friot D. 2017. Primary microplastics in the oceans: a global evaluation of sources. Gland, Switzerland: The International Union for Conservation of Nature (IUCN). doi: 10.2305/IUCN.CH.2017.01.en

[24] Anderson PJ, Warrack S, Langen V, Challis JK, Hanson ML, et al. 2017. Microplastic contamination in lake Winnipeg, Canada. Environmental pollution 225:223-31

[25] Demarquoy J. 2024. Microplastics and microbiota: Unraveling the hidden environmental challenge. World Journal of Gastroenterology 30:2191

[26] Dawson AL, Kawaguchi S, King CK, Townsend KA, King R, et al. 2018. Turning microplastics into nanoplastics through digestive fragmentation by Antarctic krill. Nature communications 9:1001

[27] Ni J, Wu GD, Albenberg L, Tomov VT. 2017. Gut microbiota and IBD: causation or correlation? Nature Reviews Gastroenterology & Hepatology 14:573-84

[28] Tavelli R, Callens M, Grootaert C, Abdallah MF, Rajkovic A. 2022. Foodborne pathogens in the plastisphere: can microplastics in the food chain threaten microbial food safety? Trends in Food Science & Technology 129:1-10

[29] Chen W, Chu Q, Ye X, Sun Y, Liu Y, et al. 2021. Canidin-3-glucoside prevents nano-plastics induced toxicity via activating autophagy and promoting discharge. Environmental pollution 274:116524

[30] Chen W, Zhu R, Ye X, Sun Y, Tang Q, et al. 2022. Food-derived cyanidin-3-O-glucoside reverses microplastic toxicity via promoting discharge and modulating the gut microbiota in mice. Food & function 13:1447-58

[31] Zhang J, Liu W, Cui F, Kolehmainen M, Chen J, et al. 2025. Exploring the potential protective role of anthocyanins in mitigating micro/nano-plastic-induced reproductive toxicity: a steroid receptor perspective. Journal of Pharmaceutical Analysis 15:101148

[32] Faria A, Fernandes I, Norberto S, Mateus N, Calhau C. 2014. Interplay between anthocyanins and gut microbiota. Journal of Agricultural and Food Chemistry 62:6898-902

[33] Morais CA, de Rosso VV, Estadella D, Pisani LP. 2016. Anthocyanins as inflammatory modulators and the role of the gut microbiota. The Journal of nutritional biochemistry 33:1-7

[34] Wang H, Liu D, Ji Y, Liu Y, Xu L, et al. 2020. Dietary supplementation of black rice anthocyanin extract regulates cholesterol metabolism and improves gut microbiota dysbiosis in C57BL/6J mice fed a high-fat and cholesterol diet. Molecular nutrition & food research 64:1900876

[35] Ding R, Ma Y, Li T, Sun M, Sun Z, Duan J. 2023. The detrimental effects of micro-and nano-plastics on digestive system: An overview of oxidative stress-related adverse outcome pathway. Science of The Total Environment 878:163144

[36] Hu M, Palić D. 2020. Micro-and nano-plastics activation of oxidative and inflammatory adverse outcome pathways. Redox biology 37:101620

[37] Kadac-Czapska K, Ośko J, Knez E, Grembecka M. 2024. Microplastics and oxidative stress-current problems and prospects. Antioxidants 13:579

[38] Kunst C, Schmid S, Michalski M, Tümen D, Buttenschön J, et al. 2023. The influence of gut microbiota on oxidative stress and the immune system. Biomedicines 11:1388

[39] Caputi S, Diomede F, Lanuti P, Marconi GD, Di Carlo P, et al. 2022. Microplastics affect the inflammation pathway in human gingival fibroblasts: a study in the Adriatic Sea. International Journal of Environmental Research and Public Health 19:7782

[40] Huang W, Yan Z, Li D, Ma Y, Zhou J, et al. 2018. Antioxidant and antiinflammatory effects of blueberry anthocyanins on high glucoseinduced human retinal capillary endothelial cells. Oxidative Medicine and Cellular Longevity 2018:1862462

[41] Ma H, Johnson SL, Liu W, DaSilva NA, Meschwitz S, et al. 2018. Evaluation of polyphenol anthocyanin-enriched extracts of blackberry, black raspberry, blueberry, cranberry, red raspberry, and strawberry for free radical scavenging, reactive carbonyl species trapping, anti-glycation, anti- β-amyloid aggregation, and microglial neuroprotective effects. International Journal of Molecular Sciences 19:461

[42] Rehman SU, Ali Shah S, Ali T, Chung JI, Kim MO. 2017. Anthocyanins reversed D-galactose-induced oxidative stress and neuroinflammation mediated cognitive impairment in adult rats. Molecular Neurobiology 54:255-71

[43] Furuuchi R, Shimizu I, Yoshida Y, Hayashi Y, Ikegami R, et al. 2018. ?Boysenberry polyphenol inhibits endothelial dysfunction and improves vascular health. PloS one 13:e0202051

[44] Bicudo MOP, Ribani RH, Beta T. 2014. Anthocyanins, phenolic acids and antioxidant properties of juçara fruits (Euterpe edulis ) along the ontree ripening process. Plant Foods for Human Nutrition 69:142-47

[45] Ma Z, Du B, Li J, Yang Y, Zhu F. 2021. An insight into anti-inflammatory activities and inflammation related diseases of anthocyanins: a review of both in vivo and in vitro investigations. International Journal of Molecular Sciences 22:11076

[46] Duarte L, Chaves VC, dos Santos Nascimento MVP, Calvete E, Li M, et al. 2018. Molecular mechanism of action of Pelargonidin-3-O-glucoside, the main anthocyanin responsible for the anti-inflammatory effect of strawberry fruits. Food Chemistry 247:56-65

[47] Roth S, Spalinger MR, Gottier C, Biedermann L, Zeitz J, et al. 2016. Bilberry-derived anthocyanins modulate cytokine expression in the intestine of patients with ulcerative colitis. PLoS one 11:e0154817

[48] Le Phuong Nguyen T, Fenyvesi F, Remenyik J, Homoki JR, Gogolák P, et al. 2018. Protective effect of pure sour cherry anthocyanin extract on cytokine-induced inflammatory caco-2 monolayers. Nutrients 10:861

[49] Zhang Y, Meng Q, Yin J, Zhang Z, Bao H, et al. 2020. Anthocyanins attenuate neuroinflammation through the suppression of MLK3 activation in a mouse model of perioperative neurocognitive disorders. Brain Research 1726:146504

[50] Cui HX, Chen JH, Li JW, Cheng FR, Yuan K. 2018. Protection of anthocyanin from Myrica rubra against cerebral ischemia-reperfusion injury via modulation of the TLR4/NF- κ B and NLRP3 pathways. Molecules 23:1788

[51] Wu T, Tang Q, Yu Z, Gao Z, Hu H, et al. 2014. Inhibitory effects of sweet cherry anthocyanins on the obesity development in C57BL/6 mice. International Journal of Food Sciences and Nutrition 65:351-59

[52] Nazir N, Akbar A, Salar MZ, Ahmed MZ, Ishtiaq A. 2024. Pharmacological assessment of delphinidin in counteracting polystyrene microplastic induced renal dysfunction in rats. Journal of King Saud UniversityScience 36:103462

[53] Chen W, Zheng X, Yan F, Xu L, Ye X. 2024. Modulation of gut microbial metabolism by cyanidin-3-O-glucoside in mitigating polystyreneinduced colonic inflammation: insights from 16 S rRNa sequencing and metabolomics. Journal of agricultural and food chemistry 72:7140-54

[54] Wang M, Zhang Z, Sun H, He S, Liu S, et al. 2022. Research progress of anthocyanin prebiotic activity: A review. Phytomedicine 102:154145

[55] Doughari JH, Ndakidemi PA, Human IS, Benade S. 2012. Antioxidant, antimicrobial and antiverotoxic potentials of extracts of Curtisia dentata. Journal of Ethnopharmacology 141:1041-50

[56] Sun XH, Zhou TT, Wei CH, Lan WQ, Zhao Y, et al. 2018. Antibacterial effect and mechanism of anthocyanin rich Chinese wild blueberry extract on various foodborne pathogens. Food Control 94:155-61

[57] Ivanovski S, Haase HR, Bartold PM. 2001. Expression of bone matrix protein mRNAs by primary and cloned cultures of the regenerative phenotype of human periodontal fibroblasts. Journal of Dental Research 80:1665-71

[58] Lacombe A, Li RW, Klimis-Zacas D, Kristo AS, Tadepalli S, et al. 2013. Lowbush wild blueberries have the potential to modify gut microbiota and xenobiotic metabolism in the rat colon. PLoS One 8:e67497

[59] Sun H, Zhang P, Zhu Y, Lou Q, He S. 2018. Antioxidant and prebiotic activity of five peonidin-based anthocyanins extracted from purple sweet potato (Ipomoea batatas (L.) Lam.). Scientific Reports 8:5018

[60] Chen W, Tu P, Ye X, Tang Q, Yu T, et al. 2022. Cyanidin-3-O-glucoside impacts fecal discharge of polystyrene microplastics in mice: Potential role of microbiota-derived metabolites. Toxicology and Applied Pharmacology 453:116212

[61] Shipp J, Abdel-Aal ESM. 2010. Food applications and physiological effects of anthocyanins as functional food ingredients. The Open Food Science Journal 4:7-22

[62] EFSA Panel on Food Additives and Nutrient Sources added to Food. 2013. Scientific Opinion on the re-evaluation of anthocyanins (E 163) as a food additive. EFSA Journal 11:3145

[63] Lehto S, Buchweitz M, Klimm A, Straßburger R, Bechtold C, et al. 2017. Comparison of food colour regulations in the EU and the US: a review of current provisions. Food Additives & Contaminants: Part A 34:335-55

[64] Du L LüH, Chen Y, Yu X, Jian T, et al. 2023. Blueberry and blackberry anthocyanins ameliorate metabolic syndrome by modulating gut microbiota and short-chain fatty acids metabolism in high-fat diet-fed C57BL/6J mice. Journal of Agricultural and Food Chemistry 71:14649-65

[65] Della Lucia CM, Oliveira LA, Dias KA, Pereira SMS, da Conceição AR, et al. 2023. Scientific evidence for the beneficial effects of dietary blueberries on gut health: a systematic review. Molecular Nutrition & Food Research 67:2300096

[66] Stevenson D, Scalzo J. 2012. Anthocyanin composition and content of blueberries from around the world. Journal of Berry Research 2:179-89

[67] Wu Y, Han Y, Tao Y, Li D, Xie G, et al. 2020. In vitro gastrointestinal digestion and fecal fermentation reveal the effect of different encapsulation materials on the release, degradation and modulation of gut microbiota of blueberry anthocyanin extract. Food Research International 132:109098

[68] Lätti AK, Riihinen KR, Kainulainen PS. 2008. Analysis of anthocyanin variation in wild populations of bilberry (Vaccinium myrtillus L.) in Finland. Journal of Agricultural and Food Chemistry 56:190-96

[69] Li J, Wu T, Li N, Wang X, Chen G, et al. 2019. Bilberry anthocyanin extract promotes intestinal barrier function and inhibits digestive enzyme activity by regulating the gut microbiota in aging rats. Food & Function 10:333-43

[70] Wang L, Jiang G, Jing N, Liu X, Li Q, et al. 2020. Bilberry anthocyanin extracts enhance anti-PD-L 1 efficiency by modulating gut microbiota. Food & Function 11:3180-90

[71] Mauray A, Felgines C, Morand C, Mazur A, Scalbert A, et al. 2012. Bilberry anthocyanin-rich extract alters expression of genes related to atherosclerosis development in aorta of apo E-deficient mice. Nutrition, Metabolism and Cardiovascular Diseases 22:72-80

[72] Marques C, Fernandes I, Meireles M, Faria A, Spencer JPE, et al. 2018. Gut microbiota modulation accounts for the neuroprotective properties of anthocyanins. Scientific Reports 8:11341

[73] Pan P, Lam V, Salzman N, Huang YW, Yu J, et al. 2017. Black raspberries and their anthocyanin and fiber fractions alter the composition and diversity of gut microbiota in F-344 rats. Nutrition and Cancer 69:943-51

[74] Gu J, Thomas-Ahner JM, Riedl KM, Bailey MT, Vodovotz Y, et al. 2019. Dietary black raspberries impact the colonic microbiome and phytochemical metabolites in mice. Molecular nutrition & food research 63:1800636

[75] Zhang S, Xu M, Sun X, Liu X, Choueiry F, et al. 2022. Black raspberry extract shifted gut microbe diversity and their metabolic landscape in a human colonic model. Journal of Chromatography B 1188:123027

[76] Wang J, Yuan ZY, Wang XY, Zhu JX, Huang WF, et al. 2024. Antho-cyanins-rich cranberry extract attenuates DSS-induced IBD in an intestinal flora independent manner. Current Research in Food Science 9:100815

[77] Petersen C, Wankhade UD, Bharat D, Wong K, Mueller JE, et al. 2019. Dietary supplementation with strawberry induces marked changes in the composition and functional potential of the gut microbiome in diabetic mice. The Journal of Nutritional Biochemistry 66:63-69

[78] Sidor A, Gramza-Michałowska A. 2015. Advanced research on the antioxidant and health benefit of elderberry (Sambucus nigra) in food-a review. Journal of Functional Foods 18:941-58

[79] Pahlke G, Ahlberg K, Oertel A, Janson-Schaffer T, Grabher S, et al. 2021. Antioxidant effects of elderberry anthocyanins in human colon carcinoma cells: A study on structure-activity relationships. Molecular Nutrition & Food Research 65:2100229

[80] Koskela AKJ, Anttonen MJ, Soininen TH, Saviranta NMM, Auriola S, et al. 2010. Variation in the anthocyanin concentration of wild populations of crowberries (Empetrum nigrum L. subsp. hermaphroditum). Journal of Agricultural and Food Chemistry 58:12286-91

[81] Cremonini E, Mastaloudis A, Hester SN, Verstraeten SV, Anderson M, et al. 2017. Anthocyanins inhibit tumor necrosis alpha-induced loss of Caco-2 cell barrier integrity. Food & Function 8:2915-23

[82] Zhu Y, Zhang JY, Wei YL, Hao JY, Lei YQ, et al. 2020. The polyphenolrich extract from chokeberry (Aronia melanocarpa L.) modulates gut microbiota and improves lipid metabolism in diet-induced obese rats. Nutrition & metabolism 17:54

[83] Istas G, Wood E, Le Sayec M, Rawlings C, Yoon J, et al. 2019. Effects of aronia berry (poly)phenols on vascular function and gut microbiota: a double-blind randomized controlled trial in adult men. The American Journal of Clinical Nutrition 110:316-29

[84] Cao L, Lee SG, Melough MM, Sakaki JR, Maas KR, et al. 2020. Long-term blackcurrant supplementation modified gut microbiome profiles in mice in an age-dependent manner: an exploratory study. Nutrients 12:290

[85] de Moura Amália Soares dos Reis C, da Silva Vanderlei Aparecido de Lima LD, Cadorin Oldoni TL, Pereira C, Carpes ST. 2018. Optimization of phenolic compounds extraction with antioxidant activity from açaí blueberry and goji berry using response surface methodology. Emirates Journal of Food and Agriculture 30:180-89

[86] Mertens-Talcott SU, Rios J, Jilma-Stohlawetz P, Pacheco-Palencia LA, Meibohm B, et al. 2008. Pharmacokinetics of anthocyanins and antioxidant effects after the consumption of anthocyanin-rich acai juice and pulp (Euterpe oleracea Mart.) in human healthy volunteers. Journal of Agricultural and Food Chemistry 56:7796-802

[87] Song H, Shen X, Deng R, Zhang Y, Zheng X. 2021. Dietary anthocyaninrich extract of açai protects from diet-induced obesity, liver steatosis, and insulin resistance with modulation of gut microbiota in mice. Nutrition 86:111176

[88] Alqurashi RM, Alarifi SN, Walton GE, Costabile AF, Rowland IR, et al. 2017. In vitro approaches to assess the effects of açai (Euterpe oleracea) digestion on polyphenol availability and the subsequent impact on the faecal microbiota. Food Chemistry 234:190-98

[89] Zheng J, Ding C, Wang L, Li G, Shi J, et al. 2011. Anthocyanins composition and antioxidant activity of wild Lycium ruthenicum Murr. from Qinghai-Tibet Plateau. Food chemistry 126:859-65

[90] Yan Y, Peng Y, Tang J, Mi J, Lu L, et al. 2018. Effects of anthocyanins from the fruit of Lycium ruthenicum Murray on intestinal microbiota. Journal of Functional Foods 48:533-41

[91] Tian B, Zhao J, Zhang M, Chen Z, Ma Q, et al. 2021. Lycium ruthenicum anthocyanins attenuate high-fat diet-induced colonic barrier dysfunction and inflammation in mice by modulating the gut microbiota. Molecular Nutrition & Food Research 65:2000745

[92] Peng Y, Yan Y, Wan P, Dong W, Huang K, et al. 2020. Effects of longterm intake of anthocyanins from Lycium ruthenicum Murray on the organism health and gut microbiota in vivo. Food Research International 130:108952

[93] Peng Y, Yan Y, Wan P, Chen D, Ding Y, et al. 2019. Gut microbiota modulation and anti-inflammatory properties of anthocyanins from the fruits of Lycium ruthenicum Murray in dextran sodium sulfateinduced colitis in mice. Free Radical Biology and Medicine 136:96-108

[94] David L, Danciu V, Moldovan B, Filip A. 2019. Effects of in vitro gastrointestinal digestion on the antioxidant capacity and anthocyanin content of cornelian cherry fruit extract. Antioxidants 8:114

[95] Van Hul M, Geurts L, Plovier H, Druart C, Everard A, et al. 2018. Reduced obesity, diabetes, and steatosis upon cinnamon and grape pomace are associated with changes in gut microbiota and markers of gut barrier. American Journal of Physiology-Endocrinology and Metabolism 314:E334-E352

[96] Igwe EO, Charlton KE, Roodenrys S, Kent K, Fanning K, et al. 2017. Anthocyanin-rich plum juice reduces ambulatory blood pressure but not acute cognitive function in younger and older adults: a pilot crossover dose-timing study. Nutrition Research 47:28-43

[97] Bhaswant M, Brown L, Mathai ML. 2019. Queen Garnet plum juice and raspberry cordial in mildly hypertensive obese or overweight subjects: a randomized, double-blind study. Journal of functional foods 56:119-26

[98] Song H, Shen X, Deng R, Chu Q, Zheng X. 2022. Pomegranate peel anthocyanins prevent diet-induced obesity and insulin resistance in association with modulation of the gut microbiota in mice. European journal of nutrition 61:1837-47

[99] Zhu F, Yuan Z, Zhao X, Yin Y, Feng L. 2015. Composition and contents of anthocyanins in different pomegranate cultivars. Acta Horticulturae 1089:35-41

[100] Corrêa TAF, Tobaruela EdC, Capetini VC, Quintanilha BJ, Cortez RV, et al. 2023. Blood orange juice intake changes specific bacteria of gut microbiota associated with cardiometabolic biomarkers. Frontiers in Microbiology 14:1199383

[101] Lee HS. 2002. Characterization of major anthocyanins and the color of red-fleshed Budd Blood orange (Citrus sinensis). Journal of Agricultural and Food Chemistry 50:1243-46

[102] Zhang N, Jing P. 2023. Red cabbage anthocyanins attenuate cognitive impairment by attenuating neuroinflammation and regulating gut microbiota in aging mice. Journal of Agricultural and Food Chemistry 71:15064-72

[103] Żary-Sikorska E, Fotschki B, Fotschki J, Wiczkowski W, Juśkiewicz J. 2019. Preparations from purple carrots containing anthocyanins improved intestine microbial activity, serum lipid profile and antioxidant status in rats. Journal of Functional Foods 60:103442

[104] Kim HJ, Koo KA, Park WS, Kang DM, Kim HS, et al. 2020. Anti-obesity activity of anthocyanin and carotenoid extracts from color-fleshed sweet potatoes. Journal of Food Biochemistry 44:e13438

[105] Lee EJ, Yoo KS, Patil BS. 2011. Total carotenoid, anthocyanin, and sugar contents in sliced or whole purple (cv. Betasweet) and orange carrots during 4-week cold storage. Horticulture, Environment, and Biotechnology 52:402-07

[106] Condurache ( Lazăr) NN, Croitoru C, Enachi E, Bahrim GE, Stănciuc N, et al. 2021. Eggplant peels as a valuable source of anthocyanins: extraction, thermal stability and biological activities. Plants 10:577

[107] Liu D, Ji Y, Wang K, Guo Y, Wang H, et al. 2022. Purple sweet potato anthocyanin extract regulates redox state related to gut microbiota homeostasis in obese mice. Journal of Food Science 87:2133-46

[108] Mi W, Hu Z, Zhao S, Wang W, Lian W, et al. 2024. Purple sweet potato anthocyanins normalize the blood glucose concentration and restore the gut microbiota in mice with type 2 diabetes mellitus. Heliyon 10:e31784

[109] Steed LE, Truong VD. 2008. Anthocyanin content, antioxidant activity, and selected physical properties of flowable purple-fleshed sweetpotato purees. Journal of Food Science 73:S215-S221

[110] Fang JL, Luo Y, Jin SH, Yuan K, Guo Y. 2020. Ameliorative effect of anthocyanin on depression mice by increasing monoamine neurotransmitter and up-regulating BDNF expression. Journal of Functional Foods 66:103757

[111] Volden J, Bengtsson GB, Wicklund T. 2009. Glucosin olates, L-ascorbic acid, total phenols, anthocyanins, antioxidant capacities and colour in cauliflower (Brassica oleracea L. ssp. botrytis); effects of long-term freezer storage. Food Chemistry 112:967-76

[112] Marrelli M, Russo C, Statti G, Argentieri MP, Meleleo D, et al. 2022. Phytochemical and biological characterization of dry outer scales extract from Tropea red onion (Allium cepa L. var. Tropea)-A promising inhibitor of pancreatic lipase. Phytomedicine Plus 2:100235

[113] Sun M, Li D, Hua M, Miao X, Su Y, et al. 2022. Black bean husk and black rice anthocyanin extracts modulated gut microbiota and serum metabolites for improvement in type2 diabetic rats. Food & Function 13:7377-91

[114] Lee JH. 2010. Identification and quantification of anthocyanins from the grains of black rice (Oryza sativa L.) varieties. Food Science and Biotechnology 19:391-97

[115] Tian XZ, Li JX, Luo QY, Zhou D, Long QM, et al. 2021. Effects of purple corn anthocyanin on blood biochemical indexes, ruminal fluid fermentation, and rumen microbiota in goats. Frontiers in Veterinary Science 8:715710

[116] Xu H, Liu M, Liu H, Zhao B, Zheng M, et al. 2021. Anthocyanins from purple corn ameliorated obesity in high fat diet-induced obese mice through activating hepatic AMPK. Journal of Functional Foods 84:104582

[117] Lao F, Giusti MM. 2016. Quantification of purple corn (Zea mays L.) anthocyanins using spectrophotometric and HPLC approaches: method comparison and correlation. Food Analytical Methods 9:1367-80

[118] Takeoka GR, Dao LT, Full GH, Wong RY, Harden LA, et al. 1997. Characterization of black bean (Phaseolus vulgaris L.) anthocyanins. Journal of Agricultural and Food Chemistry 45:3395-400

[119] Le Roy CI, Wells PM, Si J, Raes J, Bell JT, et al. 2020. Red wine consumption associated with increased gut microbiota α-diversity in 3 independent cohorts. Gastroenterology 158:270-272.e2

[120] Boto-Ordóñez M, Urpi-Sarda M, Queipo-Ortuño MI, Tulipani S, Tinahones FJ, et al. 2014. High levels of Bifidobacteria are associated with increased levels of anthocyanin microbial metabolites: a randomized clinical trial. Food & Function 5:1932-38

[121] Cliff MA, King MC, Schlosser J. 2007. Anthocyanin, phenolic composition, colour measurement and sensory analysis of BC commercial red wines. Food research international 40:92-100

[122] Amer SA, Al-Khalaifah HS, Gouda A, Osman A, Goda NIA, et al. 2022. Potential effects of anthocyanin-rich Roselle (Hibiscus sabdariffa L.) extract on the growth, intestinal histomorphology, blood biochemical parameters, and the immune status of broiler chickens. Antioxidants 11:544

[123] Paraíso CM, Januário JGB, Mizuta AG, dos Santos SS, dos Santos Magon TF, et al. 2021. Comparative studies on chemical stability, antioxidant and antimicrobial activity from hot and cold hibiscus (Hibiscus sabdariffa L.) calyces tea infusions. Journal of Food Measurement and Characterization 15:3531-38