Review article

Antimicrobial potential of kombucha against foodborne pathogens: a review

Jie Hou1, Rong Luo1, Hui Ni2, Ke Li3, Fedrick C Mgomi1, Luyao Fan1, Lei Yuan1, 2, 3*

1College of Food Science and Engineering, Yangzhou University, Yangzhou, Jiangsu, 225127, PR China;

2Fujian Provincial Key Laboratory of Food Microbiology and Enzyme Engineering, Xiamen 361021, China;

3Henan Key Laboratory of Cold Chain Food Quality and Safety Control, Zhengzhou University of Light Industry, Zhengzhou 450001, PR China


The survival of foodborne pathogens under stressful food processing conditions and in host’s gastrointestinal tract has been widely reported to cause the outbreak of human diseases. Generally, antibiotics have been used to eliminate the microbial flora of foodborne pathogens. However, the overuse of antibiotics has contributed to the emergence and spread of multi-drug-resistant foodborne pathogens. Kombucha is a beverage prepared by fermenting sugared tea or other substrates with a symbiotic culture of yeasts and bacteria, and has been proved to fight foodborne pathogens and affect gastrointestinal microbial flora to prevent foodborne illnesses. In this context, this review primarily focused on microbiological and chemical compositions of kombucha obtained by fermenting different substrates. It further discussed the antimicrobial activity of kombucha, as well as potential antimicrobial agents found in kombucha, and the limitations of kombucha in the food industry. In addition, the need for developing antimicrobial agents from kombucha has been discussed for potential applications. The information provided in this review indicates that kombucha could serve as an alternative approach to control pathogens in place of using antibiotics.

Key words: kombucha, foodborne pathogens, antimicrobial activity, food safety

*Corresponding author: Lei Yuan, College of Food Science and Engineering, Yangzhou University, 196 Huayang West Road, Yangzhou, Jiangsu 225127, China. Email: [email protected]

Received: 12 May 2021; Accepted: 23 July 2021; Published: 30 August 2021

DOI: 10.15586/qas.v13i3.920

© 2021 Codon Publications

This is an Open Access article distributed under the terms of the Creative Commons Attribution-NonCommercial-ShareAlike 4.0 International (CC BY-NC-SA 4.0). License (


Foodborne pathogens have gained much attention because of frequent outbreaks of serious safety issues. In general, foodborne illnesses are characterized by acute conditions such as hemorrhagic colitis and hemolytic uremic syndrome (caused by Escherichia coli O157:H7), listeriotic and abnormalities (caused by Listeria monocytogenes), bacterial gastroenteritis (caused by Campylobacter, Salmonella, and Vibrio parahaemolyticus), chronic arthritis (caused by Shigella and Salmonella), and even colon cancer (caused by Salmonella) (Buchanan et al., 2017; Li et al., 2019; Luo et al., 2018; Ssemanda et al., 2018). The World Health Organization (WHO) has reported around 600 million cases of foodborne diseases and 420,000 related deaths occurring annually due to pathogens (Shan et al., 2019).

Conventional physical treatment may efficiently deactivate foodborne pathogens but influence the functional and sensory properties of foods. Chemical agents, including antibiotics, also have been used widely to reduce microbiological contamination done by foodborne pathogens. However, misuse of antibiotics has contributed to the large spread of foodborne pathogens that have become resistant to multiple antibiotics (Caniça et al., 2019; Liao et al., 2020; Yuan et al., 2020). Consumption of contaminated food products could transfer resistant genes from antibiotic-resistant microorganisms to the human body through mobile genetic elements, leading to foodborne diseases, and hence enormous pressure on medical system (Das et al., 2017). This led researchers to develop novel methods to fight foodborne pathogens.

Recently, in order to reduce the heavy burden of foodborne diseases, natural antiseptic strategies have been increasingly developed to control foodborne pathogens (Ghazy et al., 2021; Kim et al., 2021; Savas et al., 2020; Zhu et al., 2021). Kombucha is known as anacidic and sweet refreshing beverage obtained from fermented sugared tea (Cardoso et al., 2020) or other substrates such as grape juice (Ayed et al., 2017), soursop (Tan et al., 2020), yarrow (Vitas et al., 2018), coffee (Bueno et al., 2021), and milk (Kruk et al., 2021). This fermentation is carried out by a characteristic consortium of yeasts, acetic acid bacteria, and often, but not always, lactic acid bacteria (Figure 1) (Coelho et al., 2020). The popularity of kombucha has spread widely due to its high nutritive and healthy values (Jayabalan et al., 2014; Leal et al., 2018).

Figure 1. Kombucha fermentation and its antimicrobial activities.

Although some studies have summarized the biological activities of kombucha (Coelho et al., 2020; Jayabalan et al., 2014; Leal et al., 2018), knowledge of its antimicrobial activity against foodborne pathogens and potential antimicrobial mechanism is still limited. In the context of above discussion, this review aimed to provide an overview of the current knowledge related to chemical and microbiological compositions of kombucha obtained by fermenting various substrates, the potential antimicrobial agents found in kombucha, and the limitations of kombucha in the food industry. In addition, need for developing antimicrobial agents from kombucha is discussed for potential applications. A comprehensive online literature review was conducted from January 2021 to July 2021 by using the following keywords on the web of science: kombucha and/or foodborne pathogens. In addition, for each selected publication, the reference section was examined to identify additional relevant publications. The information provided in this review indicates that kombucha could serve as an alternative approach to antibiotics to control pathogens.

Knowledge of Kombucha: Microbiological and Chemical Composition

Microbiological composition of kombucha

The difficulty in understanding kombucha fermentation is due to the high diversity of microorganisms and their complex interactions (May et al., 2019). Yeasts such as Zygosaccharomyces, Dekkera, Pichia, Hanseniaspora, Zygowilliopsis, Candida, and Saccharomyces, found in kombucha appear to be complicated, with the dominant species changing during the fermentation process (Coton et al., 2017; Villarreal-Soto et al., 2018). Yeasts have high fermentative ability to hydrolyze sucrose into fructose and glucose by action of invertase, followed by the generation of ethanol via glycolytic pathway. However, high concentration of alcohol is harmful to microorganisms through modifications to the structures, functions, and integrities of cellular membranes (Teoh et al., 2004). Ethanol can be reduced by aldehyde and alcohol dehydrogenase produced by acetic acid bacteria that oxidize it and produce acetic acid via the Krebs cycle (Gomes et al., 2018). This acetic acid conversion could be stopped when the desired taste of the beverage is achieved. The acidic conditions in kombucha protect the system from the invasion of pathogenic microorganisms (Coelho et al., 2020). The acetic acid bacteria contained in kombucha is mostly represented by Komagataeibacter, Acetobacter, Gluconacetobacter, and Gluconobacter (Arıkan et al., 2020; Tran et al., 2020). Genera of Komagataeibacter and Gluconacetobacter can also synthesize cellulose chains to form a surface biofilm by oxidation reactions with glucose dehydrogenase. This biofilm protects microorganisms from extreme environmental challenges such as ultraviolet (UV) radiation (Coelho et al., 2020). In addition, lactic acid bacteria (Lactobacillus, Lactococcus, Leuconostoc, and Oenococcus) have been reported to be frequently isolated and identified in kombucha from different areas (Marsh et al., 2014). It must be mentioned that the presence of lactic acid bacteria could improve the survival and biological functions of acetic acid bacteria (Nguyen et al., 2015; Pei et al., 2020). However, the microbial composition of kombucha samples, described in separate studies, is affected by various factors, including the incubation temperature and time, raw materials, source of sugar, and selected starter (Morales, 2020; Neffe-Skocińska et al., 2017). Hence, it is not easy to find out an exact consortium of kombucha for industry applications.

In most studies, the production of kombucha was carried out on a low scale, while vital fermentation parameters, such as oxygenation and sugar content distribution, are ignored (Arıkan et al., 2020; Cardoso et al., 2020; Kaewkod et al., 2019). To unravel a core microbial composition essential for controlling the fermentation of kombucha, Coton et al. (2017) conducted a study to explore microbial composition from kombucha fermentation on industrial level. Dominated yeasts are identified as Dekkera, Hanseniaspora, and Zygosaccharomyces, while Gluconobacter oxydans, Gluconacetobacter europaeus, Acetobacter peroxydans, and Gluconacetobacter saccharivorans are dominant acetic acid bacteria found in kombucha.

Chemical components of kombucha

High nutritive values of kombucha are partly attributed to its microbial community established during the fermentation of kombucha (Coelho et al., 2020). As shown in Table 1, the chemical analysis of kombucha has confirmed the presence of various compounds, such as organic acids (tartaric acid, malic acid, citric acid, acetic acid, quinic acid, oxalic acid, citric acid, succinic acid, D-glucuronic acid, and ascorbic acid), sugars (sucrose, glucose, and fructose), minerals (manganese, zinc, copper, cobalt, nickel, and lead), water-soluble vitamins (C, B1, B2, B6, and B12), and ethanol (Ivanišová et al., 2020; Kaewkod et al., 2019). However, the chemical composition of kombucha varies depending on the microbiological composition of the culture used for kombucha fermentation, types of substrates, and fermentation conditions (Kaewkod et al., 2019; Neffe-Skocińska et al., 2017).

Table 1. Chemical compositions of kombucha.

Chemical composition References
Organic acids: acetic acid (1.55 g/L), citric acid (0.05 g/L), and tartaric acid (0.23 g/L)
Sugars: sucrose (17.81 g/L), fructose (1.41 g/L), and glucose (9.35 g/L)
Alcohol: 0.4%
Minerals: manganese (1.57 mg/L), copper (0.14 mg/L), iron (0.31 mg/L), zinc (0.53 mg/L), lead (0.12 mg/L), cobalt (0.23 mg/L), and nickel (0.42 mg/L)
Ivanišová et al., 2020
Organic acids: quinic acid (0.46–0.47 g/L), oxalic acid (0.04–0.044 g/L), citric acid (0.03–0.086 g/L), malic acid (0.029–0.03 g/L), D-glucuronic acid (0.04–0.063 g/L), and acetic acid (1.42–1.65 g/L)
Sugars: sucrose (0.93–7.54 g/L), glucose (10.5–37.7 g/L), fructose (8.7–30.9 g/L)
Alcohol: (6.9–11.0 g/L)
Neffe-Skocin´ska et al., 2017
Organic acids: glucuronic acid (0.07–1.58 g/L), acetic acid (10.42–11.15 g/L), ascorbic acid (0.61–0.70 g/L), gluconic acid (41.42–70.11 g/L), and succinic acid (3.05 g/L) Kaewkod et al., 2019
Phenolic compounds: catechin (90.7 μg/mL), epicatechin (0.59 μg/mL), and rutin (7.02 μg/mL) Velic´anski et al., 2014
Phenolic compounds: catechin (8 μg/mL), rutin (30.19 μg/mL), caffeine (177.37 mg/L), and quercetin (1.22 mg/L) Barbosa et al., 2020
Phenolic compounds: epicatechin gallate (0.04 mg/mL), epicatechin (0.027 mg/mL), catechin (0.031 mg/mL), epigallocatechin (0.041 mg/mL), epigallocatechin gallate (0.135 mg/mL), and gallocatechin gallate (0.075 mg/mL) Zhao et al., 2018
Vitamins: B2 (8.3 mg/100 mL) and C (28.98 mg/L) Malbaša et al., 2011
Vitamins: B1 (74 mg/100 mL), B6 (52 mg/100 mL), and B12 (84 mg/100 mL) Bauer-Petrovska and Petrushevska-Tozi, 2000
Anionic minerals: fluoride (1.20–3.20 mg/g), chloride (0.96–3.13 mg/g), bromide (0.04 mg/g), iodide (0.44–1.04 mg/g), nitrate (0.18–0.34 mg/g), phosphate (0.04–0.08 mg/g), sulfate (1.02–4.20 mg/g) Kumar et al., 2008

Table 2. Antimicrobial activities of kombucha.

Antibacterial ability References
Listeria monocytogenes (22 mm), Staphylococcus epidermidis (22 mm), and Micrococcus luteus (21.5 mm) Battikh et al., 2013
Vibriocholera (20.667 mm), Shigella flexneri (20.44 mm), Salmonella typhimurium (19 mm), Escherichia coli (20.67 mm), and Staphylococcus aureus (18 mm) Bhattacharya et al., 2016
Candida krusei (15.81 mm), Candidaglabrata (16 mm), Candidaalbicans (12 mm), Candidatropicalis (14 mm), Haemophilus influenzae (10 mm), and Escherichia coli (4 mm) Ivanišová et al., 2020
Microsporum canis (>32 mm), Escherichia coli (16 mm), and Salmonella typhi (32 mm) Júnior et al., 2009
Escherichia coli (22.67 mm), Shigella dysenteriae (24.33 mm), Staphylococcus aureus (26 mm), and Bacillus cereus (26 mm) Valiyan et al., 2021
Staphylococcus aureus (2.6–4.5 mm) and Escherichia coli (1.0–4.5 mm) Lopes et al., 2021
Escherichi coli (15–15.33 mm), Pseudomonas aeruginosa (13.33 mm), Salmonella enterica serovar typhimurium (18–18.66 mm), Listeria monocytogene (10.33–12.66 mm), Enterococcus faecalis (12–12.33 mm), Micrococcus luteus (11–14 mm), Staphylococcus aureus (13.66–14 mm), and Staphylococcus epidermidis (10.66–14 mm) Deghrigue et al., 2013
Salmonella enteritidis (13.85 mm), Escherichi coli (13.67 mm), Proteus mirabilis (15 mm), Pseudomonas aeruginosa (14.4 mm), Erwinia carotovora (17.83 mm), Staphylococcus aureus (16 mm), and Bacillus cereus (14.33 mm) Četojević-Simin et al., 2012
Escherichia coli (2.7 mm), Pseudomonas aeruginosa (2.8 mm), Klebsiella pneumoniae (2.8 mm), Staphylococcus aureus (3 mm), Enterococcus faecalis (2.2 mm), Bacillus cereus (2.9 mm), Staphylococcus epidermidis (2.2 mm) Ayed et al., 2017
Escherichia coli (21–24.7 mm), Escherichia coli O157:H7 (20.3–24.3 mm), Shigella dysenteriae (19.3–21.7 mm), Salmonella Typhi (20–24.7 mm), Vibrio cholera (20–21 mm) Kaewkod et al., 2019
Escherichia coli (green tea kombucha MIC: 15.33 μL/mL; black tea kombucha MIC: 15 μL/mL), Pseudomonas aeruginosa (green tea kombucha MIC: 13.33 μL/mL; black tea kombucha MIC: 13.33 μL/mL), Salmonella enterica serovar typhimurium (green tea kombucha MIC: 18 μL/mL; black tea kombucha MIC: 18.66 μL/mL), and Listeria monocytogenes (green tea kombucha MIC: 12.66 μL/mL; black tea kombucha MIC: 10.33 μL/mL) Deghrigue et al., 2013
Escherichia coli (95.66–99.91%) and Staphylococcus aureus (97.7–100%) Tan et al., 2020
Escherichia coli (green tea kombucha MIC: 250 μL/mL; black tea kombucha MIC: >250 μL/mL), Staphylococcus aureus (green tea kombucha MIC: 250 μL/mL; black tea kombucha MIC: 250 μL/mL), Salmonella (green tea kombucha MIC: 250 μL/mL; black tea kombucha MIC: >250 μL/mL), and Listeria monocytogenes (green tea kombucha MIC: 250 μL/mL; black tea kombucha MIC: 250 μL/mL) Cardoso et al., 2020
Staphylococcus aureus (MIC: 78.12–312.5 μL/mL), Klebsiella pneumoniae (MIC: 19.53–312.5 μL/mL), Escherichia coli (MIC: 39.10–312.5 μL/mL), Bacillus subtilis (MIC: 9.77–156.25 μL/mL), Proteus vulgaris (MIC: 78.13–312.5 μL/mL) Vitas et al., 2018

Antimicrobial activities

Scientific studies have correlated antimicrobial activity of kombucha with acids production, polyphenols as well as the presence of lactic acid bacteria (Kaewkod et al., 2019; Morandi et al., 2020; Verrillo et al., 2021). However, the exact mechanism of action of kombucha on pathogenesis is still debatable.


Recent studies have indicated that the antimicrobial activity of kombucha against foodborne pathogens is mostly attributed to its low pH value, especially in the presence of acetic acid. It is known that acetic acid can inhibit a number of microorganisms by cytoplasmic acidification and accumulation of dissociated acid anion to toxic levels (Velićanski et al., 2014). For example, Steinkraus et al. (1996) showed that the inhibition effects of kombucha against E. coli, Helicobacter pylori, and S. aureus were attributed to the high concentration of acetic acid. The significance of organic acids found in kombucha was also pointed out by Kaewkod et al. (2019), who determined that kombucha tea showed antibacterial activity against E. coli, Salmonella Typhi, Vibriocholera, and Shigella dysenteriae, while neutralized kombucha did not reveal any antimicrobial activity against these microorganisms. Moreover, the antibacterial activity of heat-denatured kombucha indicates that this thermo-stable antimicrobial agent could be used in food products to control thermophilic spore-forming bacteria (Kaewkod et al., 2019). Tan et al. (2020) proved that acetic acid in kombucha could penetrate Gram-positive bacteria cells more easily than Gram-negative bacteria because of its lipophilic characteristics, and the highest proportion of microbial growth inhibition against E. coli and S. aureus was 99.83% and 100%, respectively. However, Sreeramulu et al. (2000) confirmed the presence of antimicrobial compounds in kombucha other than acetic acid and large proteins, as the antimicrobial activities against E. coli, Shigella sonnei, Salmonella typhimurium, S. enteritidis, and C. jejuni were shown even at neutral pH. Silva et al. (2021) also indicated that bioactivity could be more related to compounds present in kombucha than the total acidity produced during fermentation. Different varieties and quantities of substrates used in the production of kombucha result in beverages with different levels of organic acids and antimicrobial activity.


Besides organic acids, tea-derived phenolic compounds have been described as potential antimicrobial agents that regulate the composition of intestinal microbes to prevent foodborne diseases by destabilizing microbial cell surface and cytoplasmic membranes (Verrillo et al., 2021). A total of 127 phenolic compounds were identified in the green and black tea kombucha by ultra-high performance liquid chromatography-quadrupole time-of-flight mass spectrometry (UPLC-QTOF-MS), most of which were flavonoids (70.2%), followed by phenolic acids (18.3%), other polyphenols (8.4%), lignans (2.3%), and stilbenes (0.8%) (Cardoso et al., 2020). The high diversity of phenolic compounds in kombucha is due to their bio-transformation or degradation in tea varieties by enzymatic action or low pH during the fermentation of kombucha (Cardoso et al., 2020; Kaewkod et al., 2020).

Mizuta et al. (2020) proved the antibacterial activity of green tea kombucha fermented at different periods, even at low concentrations, against Alicycobacillus species, including Alicyclobacillus hesperidum, Alicyclobacillus herbarius, Alicyclobacillus cycloheptanicus, Alicyclobacillus acidiphilus, and Alicycobacillus acidoterrestris in orange juice. After exposure to the polyphenolic fractions of kombucha, the cell wall disruption and bacillus integrity deformation of Alicycobacillus species were observed by scanning electron microscopy (SEM). In addition, the increased antimicrobial activity of kombucha was due to the presence of metabolites produced by the kombucha consortium. In another study, the fermented sugared black tea exhibited potent bactericidal activity against enteric bacterial pathogens, including S. flexneri, E. coli, S. Typhimurium, and V. cholera. Isorhamnetin and catechin were proved as the main antimicrobial agents in the polyphenolic fraction of kombucha (Bhattacharya et al., 2016). It is hypothesized that the presence of hydroxylation at positions 5 and 7 of the A ring and position 3 of the C ring, and free hydroxyl group(s) in the B ring of flavonoids contributes to the antimicrobial activity of polyphenolic compounds. Moreover, in vivo studies have been carried out to confirm the promising antimicrobial activity. For example, Bhattacharya et al. (2020) suggested that these two polyphenolic fractions from kombucha can significantly inhibit the motility and gene expressions (motY and flaC) of V. cholera related to flagellar regulatory, and prevent bacterial colonization in intestinal epithelial cells at sub-inhibitory concentrations. On the other hand, phenolic compounds are reported to increase the abundance of probiotics, maintain intestinal homeostasis, and prevent the infections caused by foodborne pathogens (Faria et al., 2014).

Nevertheless, there are some studies that conflict the antibacterial activity of polyphenols. For example, Chou et al. (1999) reported that the tea fermentation process decreased its antibacterial activity with increasing fermentation time, which indicated that the antibacterial activity of tea-derived phenolic compounds could be partly destroyed by reduction in the concentration of catechins during enzymatic oxidation.

Lactic acid bacteria

Some studies have proved that the antimicrobial activity of kombucha is not exclusively due to organic acids and phenolic compounds but possibly because of lactic acid bacteria and their biologically active components such as bacteriocins (Alizadeh et al., 2020; Kadyan et al., 2021). The most important antimicrobial mechanism is the production of acid by lactic acid bacteria, as acidification may alter cell metabolism by damaging enzymes and the substructure and function of cell walls and membranes, interrupting nutrient absorption and inhibiting protein synthesis (Gao et al., 2019). In addition, bacteriocins are antimicrobial agents produced by diverse bacterial species that can alter the membrane by corrupting potassium ion and ATP and cause cell failure to balance intracellular pH (Simons et al., 2020). Therefore, the antimicrobial activity of bacteriocins against foodborne pathogens has gained increasing interest for their applications in the food industry.

Pediococcus pentosaceus and Pediococcus acidiliactici isolated from kombucha showed high antimicrobial activity against S. enterica Typhimurium, L. monocytogenes, Listeria ivanovii,B. cereus, Proteus hauseri, and S. aureus. Moreover, their antimicrobial activities were also proved against foodborne molds, including Penicillium expansum and Penicillium digitatum (Diguta et al., 2020). In another study, a novel bacteriocin produced by Lactobacillus plantarum isolated from the kombucha consortium was found to have antibacterial activity against Gram-negative (E. coli) and Gram-positive bacteria (Listeria innocua, Bacillus subtilis, L. monocytogenes, Clostridium butyricum, B. cereus, Bacillus megaterium, Micrococcus luteus, Brochothrix thermosphacta, and S. aureus) because of increasing cell membrane permeability and release of potassium ions (Pei et al., 2020).

However, the only concern is that the resistance genes may be transferred from pathogens to lactic acid bacteria, and the consumption of lactic acid bacteria carrying resistance genes may pass these genes to humans.

Interestingly, kombucha was reported to modulate gut microbiota in humans and its subsequent effect on health status. Jung et al. (2018) reported that genera of Allobaculum, Turicibacter, and Clostridium involved in the pathogenesis of non-alcoholic fatty liver disease were significantly decreased, whereas Lactobacillus increased after the consumption of kombucha, which could suppress accumulation of fat in the human liver. However, more clinical trials are required to evaluate the effects of kombucha on human microbiome.

It is worth mentioning that the antimicrobial activity of kombucha is not always positively affected by the fermentation process. Silva et al. (2021) proved that antimicrobial agents could be more related to the compounds present in tea infusion than to those produced during the fermentation process.

Toxicity and Limitation of Kombucha

Some pathogenic microorganisms, including Salmonella enterica, C. albicans, and Penicillium. spp, were isolated and identified from kombucha prepared without hygiene environments (Villarreal-Soto et al., 2020). Migration of ceramic containers used to store kombucha was reported to cause lead poisoning and gastrointestinal toxicity (Phan et al., 1998). Allergic reactions, nausea and dizziness, lactic acidosis, headache, and other potential toxicities have been reported in individuals (Jayabalan et al., 2014). Therefore, further studies involving more in vivo trials are required to validate the toxicity of ingredients found in kombucha.

Conclusion and Perspectives

Antibiotics are widely used to control infections caused by foodborne pathogens. However, the emergence of antibiotic-resistant bacteria has increased with the overuse of antibiotics. Therefore, towing to their health benefits and antimicrobial activities, natural products such as kombucha could be considered as effective, cheap, and easily available alternatives against foodborne pathogens without negative effects on the health of consumers. However, control of kombucha production is mostly empiric despite the increasing knowledge provided by scientific community, and its antimicrobial property is also not well understood. Besides the organic acids, phenolic compounds, and lactic acid bacteria, alcohol and aldehydes are other important antimicrobial agents found in kombucha but not considered. Utilization of omics technology is encouraged to further reveal the molecular-level antimicrobial mechanisms. Moreover, clinical trials are required to confirm the effects of kombucha consumption on gastrointestinal microbiota composition and health risks in humans.


This work is supported by the grants received from the China Postdoctoral Science Foundation (2021TQ0274), Fujian Provincial Key Laboratory of Food Microbiology and Enzyme Engineering, and Henan Key Laboratory of Cold Chain Food Quality and Safety Control (CCFQ2020).


Alizadeh, A.M., Hashempour-Baltork, F., Alizadeh-Sani, M., Maleki, M., Azizi-Lalabad, M. and Khosravi-Darani, K., 2020. Inhibition of Clostridium botulinum and its toxins by probiotic bacteria and their metabolites: an update review. Quality Assurance and Safety of Crops & Foods 12: 59–68. 10.15586/qas.v12iSP1.823

Arıkan, M., Mitchell, A.L., Finn, R.D. and Gurel, F., 2020. Microbial composition of kombucha determined using amplicon sequencing and shotgun metagenomics. Journal of Food Science 85: 455–464. 10.1111/1750-3841.14992

Ayed, L., Abid, S.B. and Hamdi, M., 2017. Development of a beverage from red grape juice fermented with the kombucha consortium. Annals of Microbiology 67: 111–121. 10.1007/s13213-016-1242-2

Barbosa, C.D., Baqueta, M.R., Santos, W.C.R., Gomes, D., Alvarenga, V.O., Albano, H., et al., 2020. Data fusion of UPLC data, NIR spectra and physicochemical parameters with chemometrics as an alternative to evaluating kombucha fermentation. LWT—Food Science and Technology 133: 109875. 10.1016/j.lwt.2020.109875

Battikh, H., Chaieb, K., Bakhrouf, A. and Ammar, E., 2013. Antibacterial and antifungal activities of black and green kombucha teas. Journal of Food Biochemistry 37: 231–236. 10.1111/j.1745-4514.2011.00629.x

Bauer-Petrovska, B. and Petrushevska-Tozi, L., 2000. Mineral and water soluble vitamin contents in the kombucha drink. International Journal of Food Science and Technology 35: 201–205. 10.1046/j.1365-2621.2000.00342.x

Bhattacharya, D., Bhattacharya, S., Patra, M.M., Chakravorty, S., Sarkar, S., Chakraborty, W., et al., 2016. Antibacterial activity of polyphenolic fraction of kombucha against enteric bacterial pathogens. Current Microbiology 73: 885–896. 10.1007/s00284-016-1136-3

Bhattacharya, D., Sinha, R., Mukherjee, P., Howlader, D.R., Nag, D., Sarkar, S., et al., 2020. Anti-virulence activity of polyphenolic fraction isolated from kombucha against Vibrio cholera. Microbial Pathogenesis 140: 103927. 10.1016/j.micpath.2019.103927

Buchanan, R.L., Gorris, L.G.M., Hayman, M.M., Jackson, T.C. and Whiting, R.C., 2017. A review of Listeria monocytogenes: an update on outbreaks, virulence, dose-response, ecology, and risk assessments. Food Control 75: 1–13. 10.1016/j.foodcont.2016.12.016

Bueno, F., Chouljenko, A. and Sathivel, S., 2021. Development of coffee kombucha containing Lactobacillus rhamnosus and Lactobacillus casei: Gastrointestinal simulations and DNA microbial analysis. LWT—Food Science and Technology 142: 110980. 10.1016/j.lwt.2021.110980

Caniça, M., Manageiro, V., Abriouel, H., Moran-Gilad, J. and Franz, C.M.A.P., 2019. Antibiotic resistance in foodborne bacteria. Trends in Food Science & Technology 84: 41–44. 10.1016/j.tifs.2018.08.001

Cardoso, R.R., Neto, R.O., dos Santos D’Almeida, C.T., do Nascimento, T.P., Pressete, C.G., Azevedo. L., et al., 2020. Kombuchas from green and black teas have different phenolic profile, which impacts their antioxidant capacities, antibacterial and antiproliferative activities. Food Research International 128: 108782. 10.1016/j.foodres.2019.108782

Četojević-Simin, D.D., Velićanski, A.S., Cvetković, D.D., Markov, S.L., Mrđanović, J.Z., Bogdanović, V.V., et al., 2012. Bioactivity of lemon balm kombucha. Food and Bioprocess Technology 5: 1756–1765. 10.1007/s11947-010-0458-6

Chou, C.C., Lin, L.L. and Chung, K.T., 1999. Antimicrobial activity of tea as affected by the degree of fermentation and manufacturing season. International Journal of Food Microbiology 48: 125–130. 10.1016/S0168-1605(99)00034-3

Coelho, R.M.D., de Almeida, A.L., do Amaral, R.Q.G., da Mota, R.N. and de Sousa, P.H.M., 2020. Kombucha: review. International Journal of Gastronomy and Food Science 22: 100272. 10.1016/j.ijgfs.2020.100272

Coton, M., Pawtowski, A., Taminiau, B., Burgaud, G., Deniel, F., Coulloumme-Labarthe, L., et al., 2017. Unraveling microbial ecology of industrial-scale kombucha fermentations by metabar coding and culture-based methods. FEMS Microbiology Ecology 93: fix048. 10.1093/femsec/fix048

Das, Q., Islam, M.R., Marcone, M.F., Warriner, K. and Diarra, M.S., 2017. Potential of berry extracts to control food borne pathogens. Food Control 73: 650–662. 10.1016/j.foodcont.2016.09.019

Deghrigue, M., Chriaa, J., Battikh, H., Abid, K. and Bakhrouf, A., 2013. Antiproliferative and antimicrobial activities of kombucha tea. African Journal of Microbiology Research 7: 3466–3470. 10.5897/AJMR12.1230

Diguta, C.F., Nitoi, G.D., Matei, F., Luta, G. and Cornea, C.P., 2020. The biotechnological potential of Pediococcus spp. isolated from kombucha microbial consortium. Foods 9: 1780. 10.3390/foods9121780

Faria, A., Fernandes, I., Norberto, S., Mateus, N. and Calhau, C., 2014. Interplay between anthocyanins and gut microbiota. Journal of Agricultural and Food Chemistry 62: 6898–6902. 10.1021/jf501808a

Gao, Z.H., Daliri, E.B.M., Wang, J., Liu, D.H., Chen, S.G., Ye, X.Q., et al., 2019. Inhibitory effect of lactic acid bacteria on food-borne pathogens: a review. Journal of Food Protection 82: 441–453. 10.4315/0362-028X.JFP-18-303

Ghazy, O.A., Fouad, M.T., Saleh, H.H., Kholif, A.E. and Morsy, T.A., 2021. Ultrasound-assisted preparation of anise extract nanoemulsion and its bioactivity against different pathogenic bacteria. Food Chemistry 341: 128259. 10.1016/j.foodchem.2020.128259

Gomes, R.J., Borges, M.F., Rosa, M.D.F., Castro-Gómez, R.J.H. and Spinosa, W.A., 2018. Acetic acid bacteria in the food industry: systematics, characteristics and applications. Food Technology & Biotechnology 5: 139–151. 10.17113/ftb.

Ivanišová, E., Meňhartová, K., Terentjeva, M., Harangozo, L., Kántor, A. and Kačániová M., 2020. The evaluation of chemical, antioxidant, antimicrobial and sensory properties of kombucha tea beverage. Journal of Food Science and Technology 57: 1840–1846. 10.1007/s13197-019-04217-3

Jayabalan, R., Malbasa, R.V., Loncar, E.S., Vitas, J.S. and Sathish Kumar, M., 2014. A review on kombucha tea—microbiology, composition, fermentation, beneficial effects, toxicity, and tea fungus. Comprehensive Reviews in Food Science and Food Safety 13: 538–550. 10.1111/1541–4337.12073

Jung, Y., Kim, I., Mannaa, M., Kim, J., Wang, S., Park, I., et al., 2018. Effect of kombucha on gut-microbiota in mouse having non-alcoholic fatty liver disease. Food Science and Biotechnology 28: 261–267. 10.1007/s10068-018-0433-y.

Júnior, R.J., Batista, R.A., Rodrigues, S.A., Filho, L.X. and Lima, Á.S., 2009. Antimicrobial activity of broth-fermented with kombucha colonies. Journal of Microbial & Biochemical Technology, 1: 72–78. 10.4172/1948-5948.1000014

Kadyan, S., Rashmi, H.M., Pradhan, D., Kumari, A., Chaudhari, A., Deshwal, G.K., et al., 2021. Effect of lactic acid bacteria and yeast fermentation on antimicrobial, antioxidative and metabolomic profile of naturally carbonated probiotic whey drink. LWT—Food Science and Technology 142: 111059. 10.1016/j.lwt.2021.111059

Kaewkod, T., Bovonsombut, S. and Tragoolpua, Y., 2019. Efficacy of kombucha obtained from green, oolong, and black teas on inhibition of pathogenic bacteria, antioxidation, and toxicity on colorectal cancer cell line. Microorganisms 7: 700. 10.3390/microorganisms7120700

Kim, T., Kim, J.H. and Oh, S.W., 2021. Grapefruit seed extract as a natural food antimicrobial: a review. Food and Bioprocess Technology 14: 626–633. 10.1007/s11947-021-02610-5

Kruk, M., Trzaskowska, M., Scibisz, I. and Pokorski, P., 2021. Application of the “SCOBY” and kombucha tea for the production of fermented milk drinks. Microorganisms 9: 123. 10.3390/microorganisms9010123

Kumar, S.D., Narayan, G. and Hassarajani, S., 2008. Determination of anionic minerals in black and kombucha tea using ion chromatography. Food Chemistry 111: 784–788. 10.1016/j.foodchem.2008.05.012

Leal, J.M., Suárez, L.V., Jayabalan, R., Oros, J.H. and Escalante-Aburto, A., 2018. A review on health benefits of kombucha nutritional compounds and metabolites. CyTA–Journal of Food 16: 390–399. 10.1080/19476337.2017.1410499

Li, L., Meng, H., Gu, D., Li, Y. and Jia. M., 2019. Molecular mechanisms of Vibrio parahaemolyticus pathogenesis. Microbiological Research 222: 43–51. 10.1016/j.micres.2019.03.003

Liao, X., Ma, Y., Daliri, E.B.M., Koseki, S., Wei, S., Liu, D., et al., 2020. Interplay of antibiotic resistance and food-associated stress tolerance in foodborne pathogens. Trends in Food Science & Technology 95: 97–106. 10.1016/j.tifs.2019.11.006

Lopes, D.R., Santos, L.O. and Prentice-Hernández, C., 2021. Antioxidant and antibacterial activity of a beverage obtained by fermentation of yerba-maté (Ilex paraguariensis) with symbiotic kombucha culture. Journal of Food Processing and Preservation 45: e15101. 10.1111/jfpp.15101

Luo, Y., Yi, W., Yao, Y., Zhu, N. and Qin. P., 2018. Characteristic diversity and antimicrobial resistance of Salmonella from gastroenteritis. Journal of Infection and Chemotherapy 24(4): 251–255. 10.1016/j.jiac.2017.11.003

Malbaša, R.V., Loncar, E.S., Vitas, J.S. and Canadanovic-Brunet, J.M., 2011. Influence of starter cultures on the antioxidant activity of kombucha beverage. Food Chemistry 127: 1727–1731. 10.1016/j.foodchem.2011.02.048

Marsh, A.J., O’Sullivan, O., Hill, C., Ross, R.P. and Cotter P.D., 2014. Sequence-based analysis of the bacterial and fungal compositions of multiple kombucha (tea fungus) samples. Food Microbiology 38: 171–178. 10.1016/

May, A., Narayanan, S., Alcock, J., Varsani, A., Maley, C. and Aktipis, A., 2019. Kombucha: a novel model system for cooperation and conflict in a complex multi-species microbial ecosystem. Peer J 7: e7565. 10.7717/peerj.7565

Mizuta, A.G., de Menezes, J.L., Dutra, T.V., Ferreira, T.V., Castro, J.C., da Silva, C.A.J., et al., 2020. Evaluation of antimicrobial activity of green tea kombucha at two fermentation time points against Alicyclobacillus spp. LWT—Food Science and Technology 130: 109641. 10.1016/j.lwt.2020.109641

Morales, D., 2020. Biological activities of kombucha beverages: the need of clinical evidence. Trends in Food Science & Technology 105: 323–333. 10.1016/j.tifs.2020.09.025

Morandi, S., Silvetti, T., Vezzini, V., Morozzo, E. and Brasca M., 2020. How we can improve the antimicrobial performances of lactic acid bacteria? A new strategy to control Listeria monocytogenes in Gorgonzola cheese. Food Microbiology 90: 103488. 10.1016/

Neffe-Skocińska, K., Sionek, B., Ścibisz, I. and Kołożyn-Krajewska, D., 2017. Acid contents and the effect of fermentation condition of kombucha tea beverages on physicochemical, microbiological and sensory properties. CyTA–Journal of Food 15: 601–607. 10.1080/19476337.2017.1321588

Nguyen, N.K., Dong, N.T.N., Nguye, H.T. and Le, P.H., 2015. Lactic acid bacteria: promising supplements for enhancing the biological activities of kombucha. Springer Plus 4: 91. 10.1186/s40064-015-0872-3

Pei, J., Jin, W., Abd El-Aty, A.M., Baranenko, D.A., Gou, X., Zhang H., et al., 2020. Isolation, purification, and structural identification of a new bacteriocin made by Lactobacillus plantarum found in conventional kombucha. Food Control 110: 106923. 10.1016/j.foodcont.2019.106923

Phan, T.G., Estell, J., Duggin, G., Beer, I., Smith, D. and Ferson, M.J., 1998. Lead poisoning from drinking Kombucha tea brewed in a ceramic pot. The Medical Journal of Australia 169: 644–646. 10.5694/j.1326-5377.1998.tb123448.x

Savas, E., Tavsanli, H., Catalkaya, G., Capanoglu, E. and Tamer, C.E., 2020. The antimicrobial and antioxidant properties of garagurt: traditional Cornelian cherry (Cornus mas) marmalade. Quality Assurance and Safety of Crops & Foods 12: 12–23. 10.15586/qas.v12i2.627

Shan, L., Wang, S., Wu, L. and Tsai, F.S., 2019. Cognitive biases of consumers’ risk perception of food-borne diseases in China: examining anchoring effect. International Journal of Environmental Research and Public Health 16: 2268. 10.3390/ijerph16132268

Steinkraus, K.H., Shapiro, K.B., Hotchkiss, J.H. and Mortlock, R.P., 1996. Investigations into the antibiotic activity of tea fungus/kombucha beverage. Acta Biotechnologica 16: 199–205. 10.1002/abio.370160219

Silva, K.A., Uekane, T.M., de Miranda, J.F., Ruiz, L.F., da Motta, J.C.B., Silva, C.B., et al., 2021. Kombucha beverage from non-conventional edible plant infusion and green tea: characterization, toxicity, antioxidant activities and antimicrobial properities. Biocatalysis and Agricultural Biotechnology 34: 102032. 10.1016/j.bcab.2021.102032

Simons, A., Alhanout, K. and Duval, R.E., 2020. Bacteriocins, antimicrobial peptides from bacterial origin: overview of their biology and their impact against multidrug-resistant bacteria. Microorganisms 8: 639. 10.3390/microorganisms8050639

Sreeramulu, G., Zhu, Y. and Knol, W., 2000. Kombucha fermentation and its antimicrobial activity. Journal of Agricultural and Food Chemistry 48: 2589–2594. 10.1021/jf991333m

Ssemanda, J.N., Reij, M.W., Bagabe, M.C., Muvunyi, C.M., Nyamusore, J., Joosten, H., et al., 2018. Estimates of the burden of illnesses related to foodborne pathogens as from the syndromic surveillance data of 2013 in Rwanda. Microbial Risk Analysis 9: 55–63. 10.1016/j.mran.2018.02.002

Tan, W.C., Muhialdin, B.J, and Hussin, A.S.M., 2020. Influence of storage conditions on the quality, metabolites, and biological activity of Soursop (Annona muricata. L.) kombucha. Frontiers in Microbiology 11: 603481. 10.3389/fmicb.2020.603481

Teoh, A.L., Heard, G. and Cox, J., 2004. Yeast ecology of kombucha fermentation. International Journal of Food Microbiology 95: 119–126. 10.1016/j.ijfoodmicro.2003.12.020

Tran, T., Grandvalet, C., Verdier, F., Martin, A., Alexandre, H. and Tourdot-Maréchal, R., 2020. Microbial dynamics between yeasts and acetic acid bacteria in kombucha: impacts on the chemical composition of the ceverage. Foods 9: 963. 10.3390/foods9070963

Valiyan, F., Koohsari, H. and Fadavi, A., 2021. Use of response surface methodology to investigate the effect of several fermentation conditions on the antibacterial activity of several kombucha beverages. Journal of Food Science and Technology 58: 1877–1891. 10.1007/s13197-020-04699-6

Velićanski, A., Cvetković, D.D., Markov, S.L., Šaponjac, V.T.T. and Vulić, J.J., 2014. Antioxidant and antibacterial activity of the beverage obtained by fermentation of sweetened lemon balm (Melissaofficinalis L.) tea with symbiotic consortium of bacteria and yeast. Food Technology and Biotechnology 52: 420–429. 10.17113/b.

Verrillo, M., Salzano, M., Cozzolino, V., Spaccini, R. and Piccolo, A., 2021. Bioactivity and antimicrobial properties of chemically characterized compost teas from different green composts. Waste Management 120: 98–107. 10.1016/j.wasman.2020.11.013

Villarreal-Soto, S.A., Beaufort, S., Bouajila, J., Souchard, J.P. and Taillandier, P., 2018. Understanding kombucha tea fermentation: a review. Journal of Food Science 83: 580–588. 10.1111/1750-3841.14068

Villarreal-Soto, S.A., Bouajila, J., Pace, M., Leech, J., Cotter, P.D., Souchard, J.P., Taillandier, P., et al., 2020. Metabolome-microbiome signatures in the fermented beverage, Kombucha. International Journal of Food Microbiology 333: 108778. 10.1016/j.ijfoodmicro.2020.108778

Vitas, J.S., Cvetanović, A.D., Mašković, P.Z., Švarc-Gajić, J.V. and Malbaša, R.V., 2018. Chemical composition and biological activity of novel types of kombucha beverages with yarrow. Journal of Functional Foods 44: 95–102. 10.1016/j.jff.2018.02.019

Yuan, L., Sadiq, F.A., Wang, N., Yang, Z.Q. and He, G., 2020. Recent advances in understanding the control of disinfectant-resistant biofilms by hurdle technology in the food industry. [Online]. Critical Reviews in Food Science and Nutrition. 10.1080/10408398.2020.1809345

Zhao, Z., Sui, Y., Wu, H., Zhou, C., Hu, X. and Zhang, J., 2018. Flavour chemical dynamics during fermentation of kombucha tea. Emirates Journal of Food and Agriculture 30: 732–741. 10.9755/ejfa.2018.v30.i9.1794

Zhu, Y., Li, C., Cui, H. and Lin L., 2021. Encapsulation strategies to enhance the antibacterial properties of essential oils in food system. Food Control 123: 107856. 10.1016/j.foodcont.2020.107856