Pomegranate (Punica granatum L.) is one of the oldest known edible fruits.

From Persia,
pomegranate native region, its cultivation spread into Asia, North Africa and Europe [1] [2]
(Duman et al., 2009; Chandra et al., 2010). The good adaptation to temperate climate favored its
wide diffusion throughout the Mediterranean area and the differentiation of several local
genotypes (Ferrara et al., 2011). Indeed, pomegranate species includes a very huge number of
domestic, wild and ornamental biotypes showing large variability in phenotypic traits such as
fruit size and yield, flowering season, seed-hardness, rind, and aril color. Chemical composition
of pomegranate fruits reflects the intra-species biodiversity, since large differences in sugars,
fatty acids and polyphenols were reported (Holland et al., 2009; Chandra et al., 2010; Teixeira da
Silva et al., 2013).

The rising consumers interest toward health-promoting foods is pressing the industries in re-
allocating resources on the development of novel functional foods (Peres et al., 2012; Valero-
Cases et al., 2017). This promotes the research on the optimization of new products with high
nutritional value to be used as carrier of functional compounds (Valero-Cases et al., 2017).

According to the scientific evidences related to its functional and health-promoting properties
(Viuda-Martos et al., 2010), an increasing demand for pomegranate (as fresh fruit or juice) has
been observed all over the world (Prospectiva, 2015).

The health-promoting properties of pomegranate are mainly related to the abundance of

ellagitannins and other polyphenolic compounds (Filannino et al., 2013), and to the correlated
antioxidant activity (Gil et al., 2000; Tzulker et al., 2007; Seeram et al., 2008). Indeed, it was
reported that the antioxidant activity of pomegranate is higher than red wine, green tea (Gil et al.,
2000), grape/cranberry, grapefruit, and orange juice (Roseblat and Aviram, 2006). Besides the
antioxidant activity, antiviral (Kotwal, 2007), anti-inflammatory (Giménez-Bastida et al., 2012),
and anti-atherosclerotic properties (Aviram et al., 2004) have been attributed to pomegranate
compounds. Moreover, the role of the unsaturated fatty acids (including isomers of conjugated
linoleic acid) contained in pomegranate seeds in preventing cardiovascular diseases, cancer,
asthma and in reducing cholesterol levels has recently been highlighted (Ferrara et al., 2011).

Overall, fruit juices are suitable for the development of functional foods because they are rich in
bioactive compounds and meets the consumer’s claims for healthy, tasty, and practical foods
(Fonteles and Rodrigues, 2018; Oliveira et al., 2018). Fermentation of plant material via lactic
acid bacteria has attracted industries consideration to produce fermented foods as alternative
source of probiotics (Soccol et al., 2010; Peres et al., 2012). Several studies demonstrated that
pomegranate juice fermentation (by lactic acid bacteria, yeasts or filamentous fungi) is a
promising tool for further improving its nutritional and functional profile (Trigueros et al.,
2014; Gumienna et al., 2016; Cano-Lamadrid et al., 2017). Fermentation with selected lactic acid
bacteria allowed to improve antioxidant activity, shelf-life and sensory properties of the
pomegranate juice (Filannino et al., 2013). Beside the metabolic conversion of the phenolic
compounds by lactic acid bacteria as an efficient mechanism for detoxification (Filannino et al.,
2018), the fermentation leads to an intense acidification with a correlated improvement in the
release of bioactive compounds. Hence, increasing their bioaccessibility and bioavailability (Hur
et al., 2014).
In this scenario, the present study proposes the exploitation of pomegranate juice and lactic acid
bacteria fermentation to obtain products with high antioxidant activity to be used in food and
beverage industry, to meet consumers requirements. Aiming at investigating the widest local
diversity, juices from twenty pomegranate accessions identified in Puglia region (Southeastern
Italy), were included in the study and compared to that obtained from Wonderful, the most
widespread pomegranate cultivar. Juices were supplemented with whey to promote fermentation,
which was carried out with a previously selected Lactobacillus plantarum strain as a starter. The
antioxidant activity of fermented juices was first evaluated through different in vitro assays and
then determined on murine fibroblasts cell cultures under artificially induced oxidative stress.
The phenolic profile and the sensory characteristics of the fermented juices showing the highest
antioxidant activity were also investigated.

Radical Scavenging Activity on 1,1-Diphenyl-2-Picrylhydrazyl (DPPH)

The scavenging activity of Pj and FPj on DPPH free radical was measured according to the
method of Shimada et al. (1992) with some modifications. Two milliliters of each Pj and FPj
were added to 2 ml of 0.1 mM DPPH dissolved in 95% ethanol. The mixture was shaken and
incubated at room temperature. The absorbance (517 nm) was measured after 10 min and used
for the calculation of the scavenged DPPH (Rizzello et al., 2010). The scavenging activity was
expressed as follows: DPPH scavenging activity (%) = [(blank absorbance - sample
absorbance) / blank absorbance] × 100. The whey solution used for pomegranate juice
supplementation was also analyzed (prior and after fermentation with L. plantarum PU1).
Butylated hydroxytoluene (BHT, 75 ppm) was used as the antioxidant reference (Rizzello et al.,
2010).

References

Alighourchi, H., Barzegar, M., and Abbasi, S. (2008). Anthocyanins characterization of 15

Iranian pomegranate (Punica granatum L.) varieties and their variation after cold storage and
pasteurization. Eur. Food Res. Technol. 227, 881–887. doi: 10.1007/s00217-007-0799-1

Ascacio-Valdés, J. A., Prado-Barragán, A., Buenrostro-Figueroa, J. J., Aguilera-Carbo, A.,

Rodríguez-Herrera, R., and Aguila, C. N. (2011). Ellagitannins: biosynthesis, biodegradation and
biological properties. J. Med. Plant Res. 5, 4696–4703.

Aviram, M., Rosenblat, M., Gaitini, D., Nitecki, S., Hoffman, A., Dornfeld, L., et al. (2004).
Pomegranate juice consumption for 3 years by patients with carotid artery stenosis reduces
common carotid intima-media thickness, blood pressure and LDL oxidation. Clin. Nutr. 23, 423–
433. doi: 10.1016/j.clnu.2003.10.002

Brighenti, V., Groothuis, S. F., Prencipe, F. P., Amir, R., Benvenuti, S., and Pellati, F. (2017).
Metabolite fingerprinting of Punica granatum L. (pomegranate) polyphenols by means of high-
performance liquid chromatography with diode array and electrospray ionization-mass
spectrometry detection. J. Chromatogr. 1480, 20–31. doi: 10.1016/j.chroma.2016.12.017

Bury, D., Jelen, P., and Kimura, K. (1998). Whey protein concentrate as a nutrient supplement
for lactic acid bacteria. Int. Dairy J. 8, 149–151. doi: 10.1016/S0958-6946(98)00033-8

Cano-Lamadrid, M., Trigueros, L., Wojdyło, A., Carbonell-Barrachina,ÁA., and Sendra, E.

(2017). Anthocyanins decay in pomegranate enriched fermented milks as a function of bacterial
strain and processing conditions. LWT-Food Sci. Technol. 80, 193–199. doi:
10.1016/j.lwt.2017.02.023

Carbonell-Capella, J. M., Buniowska, M., Barba, F. J., Esteve, M. J., and Frígola, A. (2014).
Analytical methods for determining bioavailability and bioaccessibility of bioactive compounds
from fruits and vegetables: a review. Compr. Rev. Food Sci. Food Saf. 13, 155–171. doi:
10.1111/1541-4337.12049

Cerdá, B., Espin, J. C., Parra, S., Martinez, P., and Tomas-Barberan, F. A. (2004). The potent in
vitro antioxidant ellagitannins from pomegranate juice are metabolized into bioavailable but poor
antioxidant hydroxy-6H-dibenzopyran-6-one derivatives by the colonic microflora of healthy
humans. Eur. J. Nutr. 43, 205–220.

Chandra, R., Babu, K. D., Jadhav, V. T., Jaime, A., and Silva, T. D. (2010). Origin, history and
domestication of pomegranate. Fruit. Veg. Cereal Sci. Biotech. 2, 1–6.

Chaturvedi, R. K., and Beal, M. F. (2013). Mitochondrial diseases of the brain. Free Radic. Biol.
Med. 63, 1–29. doi: 10.1016/j.freeradbiomed.2013.03.018

Coda, R., Rizzello, C. G., Pinto, D., and Gobbetti, M. (2012). Selected lactic acid bacteria
synthesize antioxidant peptides during sourdough fermentation of cereal flours. Appl. Environ.
Microbiol. 78, 1087–1096. doi: 10.1128/AEM.06837-11

Curiel, J. A., Pinto, D., Marzani, B., Filannino, P., Farris, G. A., Gobbetti, M., et al. (2015).
Lactic acid fermentation as a tool to enhance the antioxidant properties of Myrtus
communis berries. Microb. Cell Fact. 14:67. doi: 10.1186/s12934-015-0250-4

Curiel, J. A., Rodríguez, H., Acebrón, I., Mancheño, J. M., De Las Rivas, B., and Munoz, R.
(2009). Production and physicochemical properties of recombinant Lactobacillus
plantarum tannase. J. Agric. Food Chem. 57, 6224–6230. doi: 10.1021/jf901045s

Di Cagno, R., Filannino, P., and Gobbetti, M. (2017). Lactic acid fermentation drives the optimal
volatile flavor-aroma profile of pomegranate juice. Int. J. Food Microbiol. 248, 56–62. doi:
10.1016/j.ijfoodmicro.2017.02.014

Di Cagno, R., Minervini, G., Rizzello, C. G., Lovino, R., Servili, M., Taticchi, A., et al. (2011).
Exploitation of sweet cherry (Prunus avium L.) puree added of stem infusion through
fermentation by selected autochthonous lactic acid bacteria. Food Microbiol. 28, 900–909. doi:
10.1016/j.fm.2010.12.008
Duman, A. D., Ozgen, M., Dayisoylu, K. S., Erbil, N., and Durgac, C. (2009). Antimicrobial
activity of six pomegranate (Punica granatum L.) varieties and their relation to some of their
pomological and phytonutrient characteristics. Molecules 14, 1808–1817. doi:
10.3390/molecules14051808

Ferrara, G., Cavoski, I., Pacifico, A., Tedone, L., and Mondelli, D. (2011). Morpho-pomological
and chemical characterization of pomegranate (Punica granatum L.) genotypes in Apulia region.
Southeastern Italy. Sci. Hortic. 130, 599–606. doi: 10.1016/j.scienta.2011.08.016

Ferrara, G., Giancaspro, A., Mazzeo, A., Giove, S. L., Matarrese, A. M. S., Pacucci, C., et al.
(2014). Characterization of pomegranate (Punica granatum L.) genotypes collected in Puglia
region. Southeastern Italy. Sci. Hortic. 178, 70–78. doi: 10.1016/j.scienta.2014.08.007

Filannino, P., Azzi, L., Cavoski, I., Vincentini, O., Rizzello, C. G., Gobbetti, M., et al. (2013).
Exploitation of the health-promoting and sensory properties of organic pomegranate (Punica
granatum L.) juice through lactic acid fermentation. Int. J. Food Microbiol. 163, 184–192. doi:
10.1016/j.ijfoodmicro.2013.03.002

Filannino, P., Di Cagno, R., and Gobbetti, M. (2018). Metabolic and functional paths of lactic
acid bacteria in plant foods: get out of the labyrinth. Curr. Opin. Biotechnol. 49, 64–72. doi:
10.1016/j.copbio.2017.07.016

Fischer, U. A., Dettmann, J. S., Carle, R., and Kammerer, D. R. (2011). Impact of processing and
storage on the phenolic profiles and contents of pomegranate (Punica granatum L.) juices. Eur.
Food Res. Technol. 233:797. doi: 10.1007/s00217-011-1560-3

Fonteles, T. V., and Rodrigues, S. (2018). Prebiotic in fruit juice: processing challenges,
advances, and perspectives. Curr. Opin. Food Sci. 22, 55–61. doi: 10.1016/j.cofs.2018.02.001

Fracassetti, D., Costa, C., Moulay, L., and Tomás-Barberán, F. A. (2013). Ellagic acid
derivatives, ellagitannins, proanthocyanidins and other phenolics, vitamin C and antioxidant
capacity of two powder products from camu-camu fruit (Myrciaria dubia). Food Chem. 139,
578–588. doi: 10.1016/j.foodchem.2013.01.121

Fukuda, T., Ito, H., and Yoshida, T. (2003). Antioxidative polyphenols from walnuts (Juglans
regia L.). Phytochemistry 63, 795–801. doi: 10.1016/S0031-9422(03)00333-9

Gaschler, M. M., and Stockwell, B. R. (2017). Lipid peroxidation in cell death. Biochem.
Biophys. Res. Commun. 482, 419–425. doi: 10.1016/j.bbrc.2016.10.086

Giancaspro, A., Mazzeo, A., Giove, L. S., Zito, D., Marcotuli, I., Gallotta, A., et al. (2017).
Exploiting DNA-based molecular tools to assess genetic diversity in pomegranate (Punica
granatum L.) selections and cultivars. Fruits 72,292–305.

Gil, M. I., Tomás-Barberán, F. A., Hess-Pierce, B., Holcroft, D. M., and Kader, A. A. (2000).
Antioxidant activity of pomegranate juice and its relationship with phenolic composition and
processing. J. Agric. Food Chem. 48, 4581–4589. doi: 10.1021/jf000404a
Giménez-Bastida, J. A., Larrosa, M., Gonzalez-Sarrías, A., Tomas-Barberan, F., Espín, J. C., and
García-Conesa, M. T. (2012). Intestinal ellagitannin metabolites ameliorate cytokineinduced
inflammation and associated molecular markers in human colon fibroblasts. J. Agric. Food
Chem. 60, 8866–8876. doi: 10.1021/jf300290f

Gumienna, M., Szwengiel, A., and Gorna, B. (2016). Bioactive components of pomegranate fruit
and their transformation by fermentation processes. Eur. Food Res. Technol. 242, 631–640. doi:
10.1007/s00217-015-2582-z

Halliwell, B. (2012). Free radicals and antioxidants: updating a personal view. Nutr. Rev. 70,
257–265. doi: 10.1111/j.1753-4887.2012.00476.x

Hernádez, F., Melgarejo, P., Tomas-Barberan, F. A., and Artes, F. (1999). Evolution of juice
anthocyanins during ripening of new selected pomegranate (Punica granatum) clones. Eur.
Food Res. Technol. 210, 39–42. doi: 10.1007/s002170050529

Holland, D., Hatib, K., and Bar-Yàakov, I. (2009). Pomegranate: botany, horticulture,
breeding. Hortic. Rev. 35, 127–191. doi: 10.1002/9780470593776.ch2

Holst, B., and Williamson, G. (2008). Nutrients and phytochemicals: from bioavailability to
bioefficacy beyond antioxidants. Curr. Opin. Biotechnol. 19, 73–82. doi:
10.1016/j.copbio.2008.03.003

Hur, S. J., Lee, S. Y., Kim, Y. C., Choi, I., and Kim, G. B. (2014). Effect of fermentation on the
antioxidant activity in plant-based foods. Food Chem. 160, 346–356. doi:
10.1016/j.foodchem.2014.03.112

Iwamoto, K., Tsuruta, H., Nishitaini, Y., and Osawa, R. (2008). Identification and cloning of a
gene encoding tannase (tannin acylhydrolase) from Lactobacillus plantarum ATCC
14917T. Syst. Appl. Microbiol. 31, 269–277. doi: 10.1016/j.syapm.2008.05.004

Jiménez, N., Esteban-Torres, M., Mancheño, J. M., de las Rivas, B., and Muñoz, R. (2014).
Tannin degradation by a novel tannase enzyme present in some Lactobacillus
plantarum strains. Appl. Environ. Microbiol. 80, 2991–2997. doi: 10.1128/AEM.00324-14

Kapravelou, G., Martínez, R., Andrade, A. M., Chaves, C. L., López-Jurado, M., Aranda, P., et
al. (2015). Improvement of the antioxidant and hypolipidaemic effects of cowpea flours (Vigna
unguiculata) by fermentation: results of in vitro and in vivo experiments. J. Sci. Food Agric. 95,
1207–1216. doi: 10.1002/jsfa.6809

Khanbabaee, K., and van Ree, T. (2001). Tannins: classification and definition. Nat. Prod.
Rep. 18, 641–649. doi: 10.1039/B101061L

Kleerebezem, M., Boekhorst, J., van Kranenburg, R., Molenaar, D., Kuipers, O., Leer, R., et al.
(2003). Complete genome sequence of Lactobacillus plantarum WCFS1. Proc. Natl. Acad. Sci.
U.S.A. 100, 1990–1995. doi: 10.1073/pnas.0337704100
Kotwal, G. J. (2007). Genetic diversity-independent neutralization of pandemic viruses (e g.
HIV), potentially pandemic (e.g. H5N1 strain of influenza) and carcinogenic (e.g. HBV and
HCV) viruses and possible agents of bioterrorism (variola) by enveloped virus neutralizing
compounds (EVNCs). Vaccine 26, 3055–3058. doi: 10.1016/j.vaccine.2007.12.008

Mars, M., and Marrakchi, M. (1999). Diversity of pomegranate (Punica granatum L.)
germplasm in Tunisia. Genet. Resour. Crop Evol. 46, 461–467.

Martínez, J. J., Melgarejo, P., Hernández, F., Salazar, D. M., and Martínez, R. (2006). Seed
characterisation of five new pomegranate (Punica granatum L.) varieties. Sci. Hortic. 110, 241–
246. doi: 10.1016/j.scienta.2006.07.018

Mena, P., Calani, L., Dall’Asta, C., Galaverna, G., García-Viguera, C., Bruni, R., et al. (2012).
Rapid and comprehensive evaluation of (poly)phenolic compounds in pomegranate (Punica
granatum L.) juice by UHPLC-MSn. Molecules 17, 14821–14840. doi:
10.3390/molecules171214821

Mitsuta, H., Yasumoto, K., and Iwami, K. (1996). Antioxidative action of indole compounds
during the autoxidation of linoleic acid. Eiyo Shohkuryo 19, 210–214. doi:
10.4327/jsnfs1949.19.210

Mosmann, T. (1983). Toxicity determined in vitro by morphological alterations and neutral red
absorption. J. Immunol. Methods 65, 56–63. doi: 10.1016/0378-4274(85)90046-3

Mousavi, Z. E., Mousavi, S. M., Razavi, S. H., Emam-Djomeh, Z., and Kiani, H. (2011).
Fermentation of pomegranate juice by probiotic lactic acid bacteria. World J. Microbiol.
Biotechnol. 27, 123–128. doi: 10.1007/s11274-010-0436-1

Nath, A., Verasztó, B., Basak, S., Koris, A., Kovács, Z., and Vatai, G. (2016). Synthesis of
lactose-derived nutraceuticals from dairy waste whey-a review. Food Bioprocess Technol. 9, 16–
48. doi: 10.1007/s11947-015-1572-2

Oliveira, A., Amaro, A. L., and Pintado, M. (2018). Impact of food matrix components on
nutritional and functional properties of fruit-based products. Curr. Opin. Food Sci. 22, 153–159.
doi: 10.1016/j.cofs.2018.04.002

Osawa, T., and Namiki, M. (1985). Natural antioxidant isolated from Eucalyptus leaf waxes. J.
Agric. 33, 777–780. doi: 10.1021/jf00065a001

Peres, C. M., Peres, C., Hernández-Mendoza, A., and Malcata, F. X. (2012). Review on
fermented plant materials as carriers and sources of potentially probiotic lactic acid bacteria –
With an emphasis on table olives. Trends Food Sci. Technol. 26, 31–42. doi:
10.1016/j.tifs.2012.01.006

Petti, S., and Scully, C. (2009). Polyphenols, oral health and disease: a review. J. Dent. 3, 413–
423. doi: 10.1016/j.jdent.2009.02.003
Prazeres, A. R., Carvalho, F., and Rivas, J. (2012). Cheese whey management: a review. J.
Environ. Manage. 110, 48–68. doi: 10.1016/j.jenvman.2012.05.018

Prospectiva2020 (2015). World Pomegranate Market Supply, Demand and Forecast. Available
at: http://www.prospectiva2020.com/sites/default/files/report/files/re_-_pomegranate_-
_feb_2015.pdf (accessed February 2015).

Qian, Z. J., Jung, W. K., and Kim, S. K. (2008). Free radical scavenging activity of a novel
antioxidative peptide purified from hydrolysate of bullfrog skin. Rana
catesbeiana Shaw. Bioresour. Technol. 6, 1690–1698. doi: 10.1016/j.biortech.2007.04.005

Ragaee, S., Abdel-Aal, E. S. M., and Noaman, M. (2006). Antioxidant activity and nutrient
composition of selected cereals for food use. Food Chem. 98, 32–38. doi:
10.1016/j.foodchem.2005.04.039

Ren, B., Wu, M., Wang, Q., Peng, X., Wen, H., McKinstry, W. J., et al. (2013). Crystal structure
of tannase from Lactobacillus plantarum. J. Mol. Biol. 425, 2737–2751. doi:
10.1016/j.jmb.2013.04.032

Rizzello, C. G., Coda, R., Sánchez-Macías, D., Pinto, D., Marzani, B., Filanino, P., et al. (2013).
Lactic acid fermentation as a tool to enhance the functional features of Echinacea spp. Microb.
Cell Fact. 12:44. doi: 10.1186/1475-2859-12-44

Rizzello, C. G., Nionelli, L., Coda, R., De Angelis, M., and Gobbetti, M. (2010). Effect of
sourdough fermentation on stabilisation, chemical and nutritional characteristics of wheat
germ. Food Chem. 119, 1079–1089. doi: 10.1016/j.foodchem.2009.08.016

Rodrìguez, H., Curiel, J. A., Landete, J. M., de Las Rivas, B., de Felipe, F. L., Gòmez-Cordovés,
C., et al. (2009). Food phenolics and lactic acid bacteria. Int. J. Food Microbiol. 132, 79–90. doi:
10.1016/j.ijfoodmicro.2009.03.025

Roseblat, M., and Aviram, M. (2006). ““Antioxidative poperties of pomegranate: In,” in Vitro
Studies”, in Pomegranates: Ancient Roots to Modern Medicine, eds N. P. Seeram, D. Heber, R.
N. Schulman, and D. Heber (New York, NY: Taylor and Francis Group), 31–43.

Sarkosh, A., Zamani, Z., Fatahi, R., and Ranjbar, H. (2009). Evaluation of genetic diversity
among Iranian soft-seed pomegranate accessions by fruit characteristics and RAPD markers. Sci.
Hortic. 121, 313–319. doi: 10.1016/j.scienta.2009.02.024

Seeram, N. P., Aviram, M., Zhang, Y., Henning, S., Feng, L., Dreher, M., et al. (2008).
Comparison of antioxidant potency of commonly consumed polyphenol-rich beverages in the
United States. J. Agric. Food Chem. 56, 1415–1422. doi: 10.1021/jf073035s

Shahidi, F., and Zhong, Y. (2010). Lipid oxidation and improving the oxidative stability. Chem.
Soc. Rev. 39, 4067–4079. doi: 10.1039/B922183M
Shimada, K., Fujikawa, K., Yahara, K., and Nakamura, T. (1992). Antioxidative properties of
xanthan on the antioxidation of soybean oil in cyclodextrin emulsion. J. Agric. Food Chem. 40,
945–948. doi: 10.1021/jf00018a005

Simic, M. G., and Karel, M. (2013). Autoxidation in Food and Biological Systems. New York,
NY: Springer.

Singh, R. P., Sharad, S., and Kapur, S. (2004). Free radicals and oxidative stress in
neurodegenerative diseases: relevance of dietary antioxidants. J. Indian Acad. Clin. Med. 5, 218–
225.

Siso, G. M. L. (1996). The biotechnological utilization of cheese whey: a review. Bioresour.

Technol. 57, 1–11. doi: 10.1016/0960-8524(96)00036-3

Slinkard, K., and Singleton, V. L. (1997). Total phenol analysis: automation and comparison
with manual methods. Am. J. Enol. Vitic. 28, 49–55.

Soccol, C. R., de Souza-Vandenberghe, L. P., Spier, M. R., Pedroni-Mederios, A. B.,

Yamaguishi, C. T., De Dea Lindner, J., et al. (2010). The potential of probiotics: a review. Food
Technol. Biotech. 48, 413–434.

Tehranifar, A., Zarei, M., Nemati, Z., Esfandiyari, B., and Vazifeshenas, M. R. (2010).
Investigation of physic-chemical properties and antioxidant activity of twenty Iranian
pomegranate (Punica granatum L.) cultivars. Sci. Hortic. 126, 180–185. doi:
10.1016/j.scienta.2010.07.001

Teixeira da Silva, J. A., Rana, T. S., Narzary, D., Verma, N., Meshram, D. T., and Ranade, S. A.
(2013). Pomegranate biology and biotechnology: a review. Sci. Hortic. 160, 85–107. doi:
10.1016/j.scienta.2013.05.017

Tobi, S. E., Paul, N., and McMillan, T. J. (2000). Glutathione modulates the level of free radicals
produced in UVA-irradiated cells. ıJ. Photochem. B. 57, 102–112. doi: 10.1016/S1011-
1344(00)00084-1

Trigueros, L., Wojdyło, A., and Sendra, E. (2014). Antioxidant activity and proteinepolyphenol
interactions in a pomegranate (Punica granatum L.) yogurt. J. Agric. Food Chem. 62, 6417–
6425. doi: 10.1021/jf501503h

Tuberoso, C. I. G., Rosa, A., Bifulco, E., Melis, M. P., Atzeri, A., Pirisi, F. M., et al. (2010).
Chemical composition and antioxidant activities of Myrtus communis L. berries extracts. Food
Chem. 123, 1242–1251. doi: 10.1016/j.foodchem.2010.05.094

Tzulker, R., Glazer, I., Bar-Ilan, I., Holland, D., Aviram, M., and Amir, R. (2007). Antioxidant
activity, polyphenol content and related compounds in different fruit juices and homogenates
prepared from 29 different pomegranate accessions. Agric. Food Chem. 55, 9559–9570. doi:
10.1021/jf071413n
Valero-Cases, E., Nuncio-Jaìuregui, N., and Frutos, M. J. (2017). Influence of fermentation with
different lactic acid bacteria and in vitro digestion on the biotransformation of phenolic
compounds in fermented pomegranate juices. J. Agric. Food Chem. 65, 6488–6496. doi:
10.1021/acs.jafc.6b04854
Valko, M., Leibfritz, D., Moncol, J., Cronin, M. T. D., Mazur, M., and Telser, J. (2007). Free
radicals and antioxidants in normal physiological functions and human disease. Int. J. Biochem.
Cell Biol. 39, 44–84. doi: 10.1016/j.biocel.2006.07.001

Vaquero, I., Marcobal,Á., and Muñoz, R. (2004). Tannase activity by lactic acid bacteria isolated
from grape must and wine. Int. J. Food Microbiol. 96, 199–204. doi:
10.1016/j.ijfoodmicro.2004.04.004

Viuda-Martos, M., Fernandez-Lopez, J., and Perez-Alvarez, J. A. (2010). Pomegranate and its
many functional components as related to human health. Compr. Rev. Food Sci. Food Saf. 9,
635–654. doi: 10.1111/j.1541-4337.2010.00131.x

Yeh, C. T., and Yen, G. C. (2006). Modulation of hepatic phase II phenol sulfotransferase and
antioxidant status by phenolic acids in rats. J. Nutr. Biochem. 17, 561–569. doi:
10.1016/j.jnutbio.2005.10.008

Zhang, H., and Tsao, R. (2016). Dietary polyphenols, oxidative stress and antioxidant and anti-
inflammatory effects. Curr. Opin. Food Sci. 8, 33–42. doi: 10.1016/j.cofs.2016.02.002

[1] A. D. Duman, M. Ozgen, K. S. Dayisoylu, N. Erbil, and C. Durgac, "Antimicrobial

activity of six pomegranate (Punica granatum L.) varieties and their relation to some of
their pomological and phytonutrient characteristics," Molecules, vol. 14, no. 5, pp. 1808-
1817, 2009.
[2] R. Chandra, K. D. Babu, V. T. Jadhav, A. Jaime, and T. Silva, "Origin, history and
domestication of pomegranate," Fruit, Vegetable and Cereal Science and Biotechnology,
vol. 2, pp. 1-6, 2010.