Trends in Food Science & Technology 105 (2020) 323–333

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Trends in Food Science & Technology

journal homepage: www.elsevier.com/locate/tifs

Biological activities of kombucha beverages: The need of clinical evidence

Diego Morales a, b
a
AZTI, Food Research Division, Astondo Bidea 609, 48160, Derio, Spain
b
Department of Production and Characterization of Novel Foods, Institute of Food Science Research - CIAL (UAM+CSIC), C/ Nicolas Cabrera 9, Campus de
Cantoblanco, Universidad Autónoma de Madrid, 28049, Madrid, Spain

A R T I C L E I N F O A B S T R A C T

Keywords: Background: The growing consumption of kombucha beverages in the last decade has motivated an increase in
Kombucha the number of scientific studies investigating their biological activities. Their specific microbiological and
Fermentation chemical composition led to the generation of interesting bioactive fractions.
SCOBY
Scope and approach: Although many in vitro analyses have been described regarding bioactivities of kombucha
Antioxidant
Hypocholesterolemic
and kombucha ingredients, a significantly lower amount of in vivo tests have been published and, moreover,
Antihypertensive clinical trials are particularly missing. In this revision, several biological activities were reviewed through the
scientific literature (antioxidant, immune-modulatory, antiproliferative, hypocholesterolemic, antihypertensive,
hypoglycemic, antimicrobial), describing the utilized experimental model, the reported results and their po­
tential implications on human health. Furthermore, strengths and weaknesses of the current evidences around
kombucha bioactivities were pointed out.
Key findings and conclusions: It can be concluded that, despite of the interesting results obtained in vitro, further
research is required including more in vivo tests, bioaccessibility and bioavailability assessment and clinical
trials, as well as the isolation of bioactive compounds allowing the study of single molecules and their
functionality.

1. Introduction to kombucha beverages: fermentation,
microbiological and chemical composition, biological activities
The growing interest in functional foods has led to the development
of novel edible products with interesting biological activities.
Kombucha beverages should not be considered as novel foods since
reports indicated that they were originated about 200 B.C. ( Kapp &
Summer, 2019) and, although many people consume them thinking that
they are ‘functional food’, these products have not obtained approved
health claims from institutions such as EFSA and there is an obvious lack
of clinical trials testing the effects of kombucha or their ingredients on
human health.
Kombucha is the result of fermenting sugar (from different sources)
and traditionally tea but also other raw materials that are normally
vegetables (e.g. cereals or plant leaves) or animal products (e.g. milk)
and even mushrooms (Gamboa-Gomez et al., 2017; Lobo, Sagar, &
Shenoy, 2017; Malbasa, Vitas, Loncar, Grahovac, & Milanovic, 2014;
Sknepkek et al., 2018; Vitas, Malbasa, Grahovac, & Loncar, 2013).
This fermentation is carried out by a characteristic consortium of
yeasts and bacteria called SCOBY (Symbiotic Culture Of Bacteria and
Yeast) and it takes between 7 and 10 days to be developed (Kapp &
Summer 2019; Villarreal-Soto, Beaufort, Bouajila, Souchard, & Tail­
landier, 2018). The duration of the process could affect not only to the
sensorial features of the product but also to the stability and therefore
the biological activities of their components (Amarasinghe, Weer­
akkody, & Waisundara, 2018; Jayabalan, Subathradevi, Marimuthu,
Sathishkumar, & Swaminathan, 2008).
The SCOBY is integrated by a mixture of acidophilic yeasts and acetic
acid bacteria (AAB) that produce a microbial cellulose layer called “tea
fungus” that floats as a biofilm on the surface of the liquid medium.
Down the “tea fungus”, a sour liquid phase can be found, containing
acetic and gluconic acids, ethanol, amino acids, vitamins, phenolic
compounds, hydrolytic enzymes, etc. The coexistence of bacteria and
yeasts allows three types of fermentation (alcoholic, lactic, acetic), being
the first stages dominated by osmotolerant microorganisms and later by
acid-tolerant species (De Roos & de Vuyst, 2018; Villarreal-Soto et al.,
2018). This process involves yeasts and bacteria in different metabolic
activities using a great variety of pathways: yeasts hydrolyse sucrose
into glucose and fructose by invertase action and generate ethanol via
glycolysis (using preferably fructose as a substrate); AAB oxidize glucose

E-mail addresses: diegomoraleshdz@gmail.com, dmorales@azti.es.

https://doi.org/10.1016/j.tifs.2020.09.025
Received 12 August 2020; Received in revised form 21 August 2020; Accepted 25 September 2020
Available online 28 September 2020
0924-2244/© 2020 Elsevier Ltd. All rights reserved.
D. Morales Trends in Food Science & Technology 105 (2020) 323–333

to gluconic acid and glucuronic acid, use ethanol to generate acetic acid Table 1
and synthesize and polymerize the cellulose chains forming the fibrils Microbial species identified in kombucha beverages. AAB = Acetic acid bacteria,
network that was previously named as “tea fungus” where specific LAB = Lactic acid bacteria. (*)K. xylinum was previously named as Acetobacter
bacteria such as Komagataeibacter xilynus are particularly active (Jaya­ xylinum and Gluconacetobacter xylinus.
balan, Malbasa, Loncar, Vitas, & Sathishkumar, 2014; Villarreal-Soto Microorganism Genus Species Reference
et al., 2018). AAB Acetobacter Acetobacter aceti Jayabalan et al.
The microbial community of kombucha beverages depends on (2014)
fermentation conditions such as temperature, time, selected starter, etc. Acetobacter Coton et al.
and on the utilized raw materials and sugar sources. Thus, there is no lovaniensis (2017)
Acetobacter
unique or standardized kombucha’s microbiota but some of the identi­ okinawensis
fied species of AAB, lactic acid bacteria (LAB) and yeasts are included in Acetobacter Jayabalan et al.
Table 1. It should be mentioned that LAB can also be added to kombucha pasteurianus (2014)
preparations allowing to enhance antioxidant and antimicrobial capac­ Acetobacter Coton et al.
peroxydans (2017)
ities and increase glucuronic acid production (Nguyen, Dong, Nguyen, &
Acetobacter syzygii
Le, 2015). Among identified yeasts, the main genera are Candida, Acetobacter tropicalis
Debaryomyces, Dekkera, Eremothecium, Hanseniaspora, Kazachstania, Bacterium Bacterium gluconicum Jayabalan et al.
Kloeckera, Kluyveromyces, Lachancea, Merimbla, Meyerozyma, Pichia, (2014)
Saccharomyces, Saccharomycopsis, Schizosaccharomyces, Spor­ Gluconacetobacter Gluconacetobacter Coton et al.
europaeus (2017)
opachydermia, Starmera, Torulaspora, Zygowilliopsis, Zygosaccharomyces
Gluconacetobacter
and Zygotorulaspora; AAB genera reported are Acetobacter, Bacterium, hansenii
Gluconacetobacter, Gluconobacter, Komagataeibacter and Tanticharoenia; Gluconacetobacter Coton et al.
LAB genera detected were Lactobacillus and Oenococcus (Chakravorty intermedius (2017);
Nguyen,
et al., 2016; Coton et al., 2017; Dutta & Gachhui, 2007; Gaggia et al.,
Nguyen,
2019); Jayabalan et al., 2014; Kurtzman, Robnett, & Basehoar-Powers, Nguyen, and Le
2001; Nguyen, Nguyen, Nguyen, & Le, 2015; Villarreal-Soto et al., (2015)
2018; Yamada et al., 2012). Some species were firstly isolated in kom­ Gluconacetobacter Dutta and
bucha e.g. Zygosaccharomyces kombuchaensis and Gluconacetobacter kombuchae Gachhui (2007)
Gluconacetobacter Coton et al.
kombuchae (Coton et al., 2017; Kurtzman et al., 2001).
liquefaciens (2017)
Regarding the chemical composition of kombucha beverages, it also Gluconacetobacter
changes according to the factors that were mentioned for microbial di­ saccharivorans
versity. However, a general composition (Table 2) for kombucha can be Gluconacetobacter
oboediens
composed by: organic acids (as result of microbial activity e.g. acetic,
Gluconacetobacter
lactic, gluconic, glucuronic and oxalic acids), sugars (including a small rhaeticus
amount of non-hydrolysed sucrose but mainly glucose and fructose), Gluconacetobacter
proteins, ethanol, polyphenols (different molecular species depending xylinus
on the utilized raw material), vitamins (B1, B6, B12, C), minerals (Cu, Fe, Gluconobacter Glunobacter cerinus
Glunobacter entanii Gaggia et al.
Mn, Ni, Zn), anions (F− , Cl− , Br− , I− , NO−3 , HPO−4 SO-4) and specific
(2019)
bioactive compounds such as D-saccharic acid-1,4-lactone (DSL) (De Gluconobacter Coton et al.
Roos & de Vuyst, 2018; Greenwalt, Stainkraus, & Ledford, 2000; Jaya­ oxydans (2017);
balan et al., 2014; Villarreal-Soto et al., 2018; Watawana, Jayawardena, Jayabalan et al.
(2014)
Gunawardhana, & Waisundara, 2015).
Komagataeibacter Komagataeibacter Yamada et al.
In terms of the biological activities of kombucha ingredients, the xylinum (*) (2012)
scientific literature was reviewed in this article, paying special attention Tanticharoenia Tanticharoenia Coton et al.
to reported antioxidant, immune-modulatory, antitumoral, hypo­ sakaeratensis (2017)
cholesterolemic, antihypertensive and antimicrobial capacities, revising LAB Lactobacillus Lactobacillus nagelii Coton et al.
Lactobacillus (2017)
also the described experimental models, the identification (when
satsumensis
possible) of the bioactive molecules and the significance of the results on Oenococcus Oenococcus oeni
human health. Yeast Candida Candida boidinii Coton et al.
(2017)
Candida parapsilosis Chakravorty
2. Antioxidant activity
Candida stellimalicola et al. (2016)
Candida tropicalis
The term ‘oxidative stress’ is frequently used to describe the imbal­ Debaryomyces Debaryomyces
ance between free radicals production and antioxidant defence mecha­ hansenii
nisms and it leads to health disorders such as cancer, cardiovascular or Dekkera Dekkera anomala Coton et al.
(2017)
neurodegenerative diseases. The use of dietary antioxidants may help to Dekkera bruxellensis Coton et al.
correct these disturbances and to restore the balance and, therefore, (2017);
plants and mushrooms have been widely used as antioxidant sources Nguyen,
(Morales, Miguel, & Garces-Rimon, 2020; Morales, Piris, Nguyen, et al.
(2015)
Ruiz-Rodriguez, Prodanov, & Soler-Rivas, 2018).
Eremothecium Eremothecium ashbyii Chakravorty
Most of the works correlated this capacity to the phenolic com­ Eremothecium et al. (2016)
pounds content although some specific molecules can also play a key cymbalariae
role as antioxidants (Xu et al., 2017). Hanseniaspora Hanseniaspora meyeri
The capacity of kombucha and kombucha extracts to exert antioxi­ Hanseniaspora
uvarum
dant activities is the most widely studied capacity above the rest of
(continued on next page)
biological effects, leading to a significantly higher number of works.
Although most of authors usually correlated this activity to phenolics,

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Table 1 (continued ) Table 2

Microorganism Genus Species Reference
General chemical composition of kombucha beverages.
Compound Comments Reference
Hanseniaspora Coton et al.
valbyensis (2017) Proteins – Villarreal-Soto et al. (2018)
Hanseniaspora vineae Chakravorty Sugars There is remaining sucrose, Jayabalan et al. (2014),
Kazachstania Kazachstania exigua et al. (2016) but particularly glucose and Greenwalt, Steinkraus, and
Kazachstania telluris fructose Ledford (2000), Villarreal-Soto
Kloeckera Kloeckera apiculata Jayabalan et al. et al. (2018)
Kluyveromyces Kluyveromyces (2014) Polyphenols Several families can be found Villarreal-Soto et al. (2018)
africanum depending on the raw
Kuyveromyces Chakravorty material (tea or others)
marxianus et al. (2016) Organic acids Acetic, gluconic, glucuronic, De Roos and de Vuyst (2018),
Lachancea Lachancea fermentati lactic, oxalic Villarreal-Soto et al. (2018),
Lachancea kluyveri Watawana, Jayawardena,
Lachancea Gunawardhana, & Waisundara
thermotolerans (2015)
Merimbla Merimbla Vitamins B1, B6, B12, C Villarreal-Soto et al. (2018)
ingelheimense Minerals Cu, Fe, Mn, Ni, Zn
Meyerozyma Meyerozyma Anions F− , Cl− , Br− , I− , NO−3 ,
caribbica HPO−4 SO-4
Meyerozyma Ethanol –
guilliermondii D-saccharic Bioactive compound that is De Roos and de Vuyst (2018)
Pichia Pichia anomala Coton et al. acid-1,4- generated by
(2017) lactone Gluconacetobacter spp.
Pichia Coton et al.
membranifaciens (2017)
Pichia mexicana Chakravorty tested in vitro capability of tea-based kombucha beverages to scavenge
et al. (2016)
free radicals e.g. DPPH• and ABTS•+ and correlated higher radical
Saccharomyces Saccharomyces Chakravorty
cerevisiae et al. (2016), scavenging abilities to higher total phenolics content (Ahmed, Hikal, &
Coton et al. Abou-Taleb, 2020; Villarreal-Soto et al., 2019, 2020). Other works re­
(2017) ported that specific compounds played a key role as scavengers, such as
Saccharomycopsis Saccharomycopssis Chakravorty gluconic acid or thearubigins-derived compounds and different mole­
fibuligera et al. (2016)
Saccharomyces Saccharomyces Coton et al.
cules such as ascorbic acid or DSL that inhibits β-glucuronidase activity
uvarum (2017) reducing the production of oxidative, carcinogenic and toxic compounds
Schizosaccharomyces Schizosaccharomyces Villarreal-Soto (Bhattacharya, Gachhui, & Sil, 2011; Chu & Chen, 2006; Hrnjez et al.,
pombe et al. (2018) 2014). Apart from these assays, other authors evaluated the potential of
Sporopachydermia Sporopachydermia Chakravorty
tea kombucha to inhibit lipid peroxidation (Yang et al., 2009), the
lactativora et al. (2016)
Starmera Starmera amethionina ability of oak kombucha to reduce oxidative damage caused by
Starmera caribaea hydrogen peroxide to activated macrophages (THP-1) (Vazquez-Cabral
Torulaspora Torulaspora Coton et al. et al., 2017) and the recovery effect of oxidative damage on fibroblast
microellipsoides (2017) cell lines by using green, black and rooibos tea kombucha beverages
Zygowilliopsis Zygowlliopsis Chakravorty
californica et al. (2016)
(Gaggia et al., 2019).
Zygosaccharomyces Zygosaccharomyces Coton et al. In addition to tea leaves and oak, other substrates have been used
bailli (2017) obtaining kombucha beverages that exerted in vitro antioxidant activ­
Zygosaccharomyces Kurtzman et al. ities: red grape juice (Ayed, Abid, & Hamdi, 2017), African mustard
kombuchaensis (2001)
(Rahmani et al., 2019), milk, soy drink and soywhey with tea kombucha
Zygosaccharomyces Gaggia et al.
parabailli (2019) inoculum (Hrnjez et al., 2014; Tu, Tang, Azi, Hu, & Dong, 2019; Xia
Zygosaccharomyces Villarreal-Soto et al., 2019), yarrow (Vitas, Cvetanovic, Maskovic, Svarc-Gajic, &
rouxii et al. (2018) Malbasa, 2018), garlic (Pure & Pure, 2016a), banana peel (Pure & Pure,
Zygotorulaspora Zygotorulaspora Coton et al. 2016b) or reishi mushroom (Sknepkek et al., 2018), among a great
florentina (2017)
number of varied materials (Table 3).
Moreover, different sugar sources can be selected and this fact may
there are no works isolating specific phenols species and only some of affect the antioxidant activity of the resultant fermented beverage. For
them separated enriched fractions from the whole product. Due to this, instance, Muhialdin et al. (2019) observed that tea kombucha fermented
only statistical correlations between measured parameters (total with coconut palm sugar exhibited higher in vitro antioxidant activity
phenolic content and antioxidant activity) can be established but not than with white refined sugar or molasses sugar. These authors also
strict ‘single molecule-activity’ assumptions can be done. highlighted the importance of the fermentation time, indicating that
On the other hand, microbial fermentation seems to be an efficient prolonged procedures more than 14 days are not recommended since
process to improve the antioxidant activity since the bioconversion of they led to a decrease in antioxidant capacity. This reduction was also
phenols conjugated forms to free phenolics enhanced their potency. observed in green and black tea kombucha by Vohra, Fazry, Sairi, and
Thus, the microbial transformations of tea (or other materials) poly­ Babul-Arianah (2018) who described that jaggery led to higher radical
phenols release compounds resulting in higher activity (Ivanisova et al., scavenging activities than white sugar or honey.
2020; Torino et al., 2013). Tea leaves contain an interesting diversity of As it can be noticed in Table 3, most of the studies were carried out in
antioxidant phenolic compounds such as flavanols (catechin, epi­ vitro but some authors reported interesting results in animal models:
catechin, epigallocatechin, epicatechin gallate, epigallocatechin gallate, Yang et al. (2009) observed an increase in antioxidant enzyme activities
theaflavins, thearubigins) (Lorenzo & Munekata, 2016; Xu et al., 2017). (superoxide dismutase) and a decrease of malondialdehyde levels in
This fact, together with the fermentation ability to increase antioxidant mice after consumption of traditional tea kombucha (fermented by the
power and the final sensory characteristics, places tea as the main source mixed tea fungus) and these effects were significantly higher after
to develop kombucha functional beverages. Most of the related studies consumption of tea kombucha only fermented by Gluconacetobacter sp.

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Table 3
Biological activities of kombucha beverages evaluated in different experimental models.
Biological activity Raw material Bioactive compounds Experimental model Reference

Antioxidant Black tea (Camellia sinensis) Phenolic compounds Radical scavenging assays, Swiss Bhattacharya, Gacchui, & Sil
albino rats (2013)
Radical scavenging assays Villareal-Soto et al. (2019)
Phenolic compounds Chu and Chen (2006)
(thearubigins-derived
compounds)
Phenolic compounds Radical scavenging and lipid Battacharya et al. (2011)
(gluconic acid) and D- oxidation assays, murine
saccharic acid 1,4-lactone hepatocytes
Phenolic compounds Radical scavenging assays Muhialdin et al. (2019)
– Amarasinghe, Weerakkody,
and Waisundara (2017)
Phenolic compounds Reducing power assay Ivanisova et al. (2020)
Radical scavenging assays Villareal-Soto et al. (2020)
Black tea (Camellia sinensis), banana (Musa x Pure and Pure (2016a)
paradisiaca) peel, common nettle (Urtica
dioica)
Black tea (Camellia sinensis), rice (Oryza Ahmed et al. (2020)
sativa), barley (Hordeum vulgare)
Black tea (Camellia sinensis), winter savory – Radical scavenging and catalase Vitas et al. (2020)
(Satureja montana), peppermint (Mentha activity assays
piperita), stinging nettle (Urtica dioica), wild
thyme (Thymus serpyllum), elderberry
(Sambucus nigra), quince (Cydonia oblonga)
Green tea (Camellia sinensis) Sprague Dawley albino rats Ram et al. (2000)
Phenolic compounds Radical scavenging assays, Wistar Bellasoued et al. (2015)
rats
Phenolic compounds Radical scavenging assays Mizuta et al. (2020)
(catechins and phenolic
acids), ascorbic acid
Green and black tea (Camellia sinensis) – Vohra et al. (2018)
Phenolic compounds Cardoso et al. (2020)
Vitamin C and organic acids Malbasa, Loncar, Vitas, and
Canadanovic-Brunet (2011)
– Radical scavenging and lipid Jayabalan et al. (2008)
oxidation assays
Green and black tea (Camellia sinensis), Phenolic compounds L929 mouse fibroblasts Gaggia et al. (2019)
rooibos (Aspalathus linearis) (glucuronic acid)
Green tea (Camellia sinensis), cinnamon Phenolic compounds, organic Radical scavenging assays Shahbazi et al. (2018)
(Cinnamonum verum), cadamom (Elettaria acids
cardamomum), shirazi thyme (Zataria
multiflora Boiss)
Green, black and oolong tea (Camellia Phenolic compounds Kaewkod et al. (2019)
sinensis) (gluconic acid)
Tea (Camellia sinensis) – Sprague Dawley albino rats Dipti et al. (2003)
Antioxidant enzymes activity and Yang et al. (2009)
malondialdehyde determination
assays
Wistar rats Lobo et al. (2017)
Phenolic compounds Radical scavenging assays Watawana et al. (2016)
Gramza-Michalowska,
Kulczynski, Xindi, and
Gumienna (2016)
Oak (Quercus resinosa, Quercus arizonica, THP-1 cells Vazquez-Cabral et al. (2017)
Quercus convallata)
Oak (Quercus arizonica, Quercus convallata) Radical scavenging assays, Gamboa-Gomez et al. (2017)
C57BL/6 mice
Red grape (Vitis vinifera) juice Radical scavenging assays Ayed et al. (2017)
Yarrow (Achillea millefolium) Phenolic compounds, organic Vitas, Cvetanovic, Maskovic,
acids, ascorbic acid Svarc-Gajic, and Malbasa
(2018)
African mustard (Brassica tournefortii) Phenolic compounds Rahmani et al. (2019)
River red gum (Eucalyptus camaldulensis) and Gamboa-Gomez et al. (2016)
Mexican bay leaf (Litsea glaucescens)
Garlic (Allium sativum) Pure and Pure (2016a)
Bee polen, green tea (Camellia sinensis) – Utoiu et al. (2018)
Ganoderma lucidum extract – Radical scavenging and reducing Sknepkek et al. (2018)
power assays
Winery effluent, black tea (Camellia sinensis) Ascorbic and organic acids Radical scavenging assays Vukmanovic, Vitas, &
starter Malbasa (2020)
Soydrink (Glycine max), black tea (Camellia Phenolic compounds and Xia et al. (2019)
sinensis) kombucha inoculum vitamins
Soy whey (Glycine max), black tea (Camellia Phenolic compounds, Tu et al, (2019)
sinensis) kombucha inoculum isoflavone aglycones
(continued on next page)

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D. Morales Trends in Food Science & Technology 105 (2020) 323–333

Table 3 (continued )
Biological activity Raw material Bioactive compounds Experimental model Reference

Milk and peppermint (Mentha piperita) Ascorbic acid Vitas, Malbasa, Jokic, Loncar,
and Milanovic (2018)
Milk, green tea (Camellia sinensis) kombucha Vitamin C Hrnjez et al. (2014)
inoculum
Milk and wild thyme (Thymus serpyllum) Ascorbic acid, phenolic Malbasa et al. (2014)
compounds
Milk, stinging nettle (Urtica dioica), winter – Vitas et al. (2013)
savory (Satureja montana)
Immune-modulatory/ Black tea (Camellia sinensis) – 5-LOX inhibition assay Villareal-Soto et al. (2019)
Anti-inflammatory Villareal-Soto et al. (2020)
BALB/c mice Wang et al. (2016)
C57BL/6 mice Marzban et al. (2015)
Oak (Quercus resinosa, Quercus arizonica, Phenolic compounds THP-1 cells Vazquez-Cabral et al. (2017)
Quercus convallata) (quercetin glucuronide)
Antiproliferative/ Black tea (Camellia sinensis) Dimethyl 2-(2-hydroxy-2- A549, U2OS and 786-O cells Jayabalan et al. (2011)
Antitumoral methoxypropylidine)
malonate and vitexin
– HCT-116 cells Villareal-Soto et al. (2019)
Villareal-Soto et al. (2020)
Green and black tea (Camellia sinensis) Phenolic compounds A549, HCT8 and CACO-2 cells Cardoso et al. (2020)
(catechins and verbascoside)
Green, black and oolong tea (Camellia Phenolic compounds CACO-2 cells Kaewkod et al. (2019)
sinensis) (gluconic acid)
Tea (Camellia sinensis) – PC-3 cells Srihari, Arunkumar,
Arunakaran, and
Satyanarayana (2013)
Lemon balm tea (Melissa officinalis) HeLa, MCF-7 and HT-29 Cetojevic-Simin et al. (2012)
Yarrow (Achillea millefolium) Phenolic compounds, organic RD and HeLa cells Vitas, Cvetanovic, et al.
acids, ascorbic acid (2018)
African mustard (Brassica tournefortii) Phenolic compounds MCF-7 cells Rahmani et al. (2019)
Bee polen, green tea (Camellia sinensis) – CACO-2 and Hep-2 cells Utoiu et al. (2018)
Hypocholesterolemic Black tea (Camellia sinensis) – Wistar rats Aloulou et al. (2012)
Green tea (Camellia sinensis) Mice Yang et al. (2009)
Glucuronic acid Ducks Adriani et al. (2011)
Black tea (Camellia sinensis), snake fruit – Wistar rats Zubaidah, Afagni, Kalsum,
(Salacca zalacca) Srianta, and Blanc (2019)
Antihypertensive Black tea (Camellia sinensis), winter savory – ACE inhibition assay Vitas et al. (2020)
(Satureja montana), Peppermint (Mentha
piperita), stinging nettle (Urtica dioica), wild
thyme (Thymus serpyllum), elderberry
(Sambucus nigra), quince (Cydonia oblonga)
Tea (Camellia sinensis) Humans Hiremath et al. (2002)
River red gum (Eucalyptus camaldulensis) and Phenolic compounds ACE inhibition assay Gamboa-Gomez et al. (2016)
Mexican bay leaf (Litsea glaucescens)
Milk, green tea (Camellia sinensis) kombucha – Hrnjez et al. (2014)
inoculum
Milk Peptides released during Elkhtab et al. (2017)
fermentation
Antidiabetic/ Black tea (Camellia sinensis) – Wistar rats Srihari, Karthikesan,
Hypoglycemic Ashokkumar, and
Satyanarayana (2013)
Aloulou et al. (2012)
Black tea (Camellia sinensis), snake fruit Zubaidah, Afagni, et al.
(Salacca zalacca) (2019)
Tea (Camellia sinensis) Humans Hiremath et al. (2002)
Alpha-amylase and beta- Watawana et al. (2016)
glucosidase inhibition assays
Green and black tea (Camellia sinensis) Phenolic compounds Apha amylase inhibition assay Kallel et al. (2012)
Green, black and oolong tea (Camellia Alpha-amylase and beta- Watawana, Jayawardena, and
sinensis) glucosidase inhibition assays Waisundara (2018)
Oak (Quercus arizonica, Quercus convallata) Alpha-glucosidase and alpha- Gamboa-Gomez et al. (2017)
amilase inhibition assays, glucose
diffusion evaluation, C57BL/6
mice
Snake fruit (Salacca zalacca) – Wistar rats Zubaidah et al. (2018)
– Zubaidah, Ifadah, et al.
(2019)
Soydrink (Glycine max), black tea (Camellia Phenolic compounds Alpha-glucosidase and alpha- Xia et al. (2019)
sinensis) kombucha inoculum amilase inhibition assays
Antimicrobial/Antiviral Black tea (Camellia sinensis) Acetic acid Microbiological analyses Ivanisova et al. (2020)
– Vohra et al, (2018)
Phenolic compounds Microbiological analyses, New Bhattacharya et al. (2020)
Zealand White rabbits, BALB/c
suckling mice
Green tea (Camellia sinensis) Microbiological analyses Mizuta et al. (2020)
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Table 3 (continued )
Biological activity Raw material Bioactive compounds Experimental model Reference

Phenolic compounds
(catechins and phenolic
acids), ascorbic acid
Green and black tea (Camellia sinensis) – Battikh et al. (2012)
Phenolic compounds Cardoso et al. (2020)
(catechins and verbascoside)
Green, black and oolong tea (Camellia – Kaewkod et al. (2019)
sinensis)
Green tea (Camellia sinensis), licorice – Landrace swines Fu, Wu, Lv, He, and Jiang
(Glycyrrhiza glabra), Momordica grosvenori, (2015)
chrysanthemum (Dendrathema morifolium)
Green tea (Camellia sinensis), cinnamon Phenolic compounds, organic Microbiological analyses Shahbazi et al. (2018)
(Cinnamonum verum), cadamom (Elettaria acids
cardamomum), shirazi thyme (Zataria
multiflora Boiss)
Tea (Camellia sinensis) Bacteriocin Pei et al. (2020)
Lemon balm tea (Melissa officinalis) Acetic acid Cetojevic-Simin et al. (2012)
Red grape (Vitis vinifera) juice Phenolic compounds Ayed et al. (2017)
Yarrow (Achillea millefolium) Phenolic compounds, organic Vitas, Cvetanovic, et al.
acids, ascorbic acid (2018)
Garlic (Allium sativum) Phenolic compounds Pure and Pure (2016b)
Ganoderma lucidum extract – Sknepkek et al. (2018)
Soy whey (Glycine max), black tea (Camellia Phenolic compounds, Tu et al, (2019)
sinensis) kombucha inoculum isoflavone aglycones
Hepatoprotective Tea (Camellia sinensis) D-saccharic acid 1,4-lactone Mice Wang, Ji, et al. (2013)
Neuroprotective – Wistar rats Kabiri and Setorki (2016)
Anti- African mustard (Brassica tournefortii) Phenolic compounds Acetylcholinesterase inhibition Rahmani et al., (2019)
acetylcholinesterase assay
Detoxification Black tea (Camellia sinensis) Yeasts Hep 2 cells and brine shrimp Taheur et al. (2020)
Microbiota-modulator – C57BLKS db/db mice Jung et al. (2019)

A4 which shows strong ability to produce DSL. Lobo et al. (2017) sub­
jected Wistar rats to a pre-treatment with tea kombucha before inducing
them myocardial infarction. Their results suggested that the function­
alized product prevented membrane destabilization, scavenged
reactive-oxygen-species and lowered blood lipid levels. A hypercholes­
terolemic diet was also applied to this animal model and the adminis­
tration of green tea kombucha exerted a protective effect mediated by its
antioxidant activity (Bellassoued et al., 2015).
Previous studies also reported that orally-administered tea kombu­
cha decreased oxidative stress, DNA damage and lipid peroxidation in
Sprague-Dawley rats (Dipti et al., 2003 (Dipti et al., 2003; Ram et al.,
2000).
Up to now, no human trials have been carried out to validate the
observed antioxidant effects of kombucha beverages.
3. Immune-modulatory and anti-inflammatory activity
Inflammation is part of the non-specific immune response to infec­
tion, injury or irritation and this process can become continuous and
chronic in some disorders such as arthritis, allergy, cardiovascular dis­
ease and different types of cancer (Devi et al., 2015; Ferrero-Miliani,
Nielsen, Andersen, & Girardin, 2006; Wang, Huang, Lu, & Chang, 2013).
During the last decades, vegetal, animal and fungal edible matrices have
been explored to obtain bioactive fractions that show the ability not only
to inhibit inflammatory responses but also to induce changes in any
immune response included within the term ‘immune-modulation’ (Lee
& Paik, 2019; Morales et al., 2019a, 2020b; Tasneem, Liu, Li, Choudh­
ary, & Wang, 2019).
Fermented beverages such as kombucha have also been evaluated as
sources of anti-inflammatory or immune-modulatory ingredients but
just a few studies have been already published. Among them, Villar­
real-Soto et al. (2019, 2020) observed that black tea kombucha was able
to inhibit in vitro the 5-lipooxigenase activity, an enzyme involved in
fatty acids conversion to leucotrienes and implicated in inflammation.
Moreover, Vazquez-Cabral et al. (2017) applied oak kombucha to
hydrogen peroxide-activated macrophages (THP-1) and reported a
reduction in proinflammatory cytokines secretion (TNF-α and IL-6) and
NO production. Furthermore, two studies were carried out in animal
models: Wang et al. (2016) described the immune-stimulating effect of
black tea kombucha in immunosuppressive mice since it increased their
white blood cells counts, enhanced the phagocytic activity of peritoneal
macrophages and strengthened the activity of NK cells; Marzban et al.
(2015) demonstrated that black tea kombucha ameliorated experi­
mental autoimmune encephalomyelitis in mice as a model of multiple
sclerosis. Despite of these promising in vitro and in vivo results, no human
trials have been carried out yet testing the immunomodulatory activity
of tea kombucha or related beverages.
4. Antiproliferative and antitumoral activity
Cancer is the second leading cause of death worldwide after car­
diovascular diseases and giving rise to 1/6 of global deaths (Ahmadi &
Ebrahimzadeh, 2020). An enormous amount of resources has been
implemented to cancer treatments research and the disadvantages
related to anticancer drugs such as lack of specificity and side effects is
leading to a continuous and maintained search for novel and improved
antitumoral agents and, many times, these investigations target natural
and edible sources (Rayan, Raiyn, & Falah, 2017).
Kombucha preparations have also been tested as antitumoral and
antiproliferative substances in vitro and promising results have been
collected. Villarreal-Soto et al. (2019, 2020) pointed out that fermented
black tea kombucha fractions extracted with ethyl acetate and butanol
exerted a cytotoxic effect on HCT-116 (human colon carcinoma) cells
that was higher than those noticed for non-fermented tea fractions.
Jayabalan et al. (2011) also fractionated tea kombucha with different
solvents (apart from ethyl acetate and butanol, they also used chloro­
form and water) and these samples reduced cell invasion and cell
motility of A549 (human lung carcinoma), U2OS (human osteosarcoma)
and 786-O (human renal carcinoma) cell lines and they showed cyto­
toxic effects on U2OS and 786-O cells. After characterizing the fractions,
they postulated 2 compounds as the main responsible of this activity:
dimethyl 2-(2-hydroxy-2-methoxypropylidene) malonate and vitexin.
Phenolic compounds were also reported as relevant molecules in
antitumoral activities of kombucha beverages since Cardoso et al.

328
D. Morales
(2020) recorded different antiproliferative capacities of green tea and
black tea kombucha against A549, CACO-2 (colorectal carcinoma) and
HCT8 (ileocecal colorectal carcinoma) and explained these differences
because of the particular phenolic composition profile of each tea
kombucha.
Kombucha beverages that were not based on tea leaves were also
tested leading to different results. Kombucha made from yarrow in­
fusions exerted antiproliferative activity against RD (human rhabdo­
myosarcoma) and HeLa (human cervix carcinoma cells) lines (Vitas,
Cvetanovic, et al., 2018) and antitumoral capacity against CACO-2 cells
was observed for bee pollen fermented by SCOBY microorganisms
(Utoiu et al., 2018). African mustard leaves were also used to prepare
kombucha and its cytotoxicity against MCF-7 (breast cancer) cells was
reduced after fermentation (Rahmani et al., 2019). Moreover, lemon
balm (Melissa officinalis) tea kombucha had lower antitumoral capacity
than traditional kombucha (with Camellia sinensis tea) when applied to
different cell lines: HeLa, MCF-7 and HT-29 (colon carcinoma) (Ceto­
jevic-Simin et al., 2012).
These results must be validated with further research using animal
models that have not been utilized yet.
5. Hypocholesterolemic activity
As it was previously mentioned, cardiovascular diseases are the first
cause of death in the whole world and high serum levels of total
cholesterol and LDL-cholesterol are one of the major risk factors (Mo­
rales, Tejedor-Calvo, et al., 2019). Although severe hypercholesterole­
mia is normally treated with drugs that inhibit both cholesterol
absorption and endogenous synthesis, these therapeutic agents may be
replaced by functional foods containing lowering-cholesterol in­
gredients (Gil-Ramirez, Morales, & Soler-Rivas, 2018). Different animal
models fed kombucha products and their cholesterol and lipid levels
were measured. For instance, diabetes was induced to Wistar rats using
alloxan (specific toxin that destroys pancreatic β-cells) and the animals
presented increased pancreatic lipase activity that provoked higher lipid
absorption leading to higher levels of triglycerides and LDL-cholesterol.
When black tea kombucha was given to the rats, a significant reduction
of lipase activity as well as triglycerides and LDL-cholesterol levels was
noticed (Aloulou et al., 2012). This model was also subjected to diabetes
induction using streptozotozin and kombucha with black tea and snake
fruit administration was tested. These beverages, particularly snake fruit
kombucha, were effective enhancing the rats lipidic profile (reducing
total and LDL-cholesterol as well as triglycerides and increasing
HDL-cholesterol levels) (Zubaidah, Afgani, Kalsum, Srianta, & Blanc,
2019a). Moreover, tea kombucha was administered to ducks, decreasing
total and LDL-cholesterol but increasing HDL-cholesterol levels
(Adriani, Mayasari, Angga, & Kartasudjana, 2011).
Only another additional study was carried out in animals where tea
kombucha decreased total and LDL-cholesterol levels in mice serum
(Yang et al., 2009) and no human studies have been developed yet.
6. Antihypertensive activity
Hypertension is a chronic condition that causes a persistent elevation
of blood pressure in the arteries and it is considered as a relevant risk
factor for cardiovascular diseases. Since blood pressure is mainly regu­
lated by the renin-angiotensin system (RAS), the inhibition of RAS en­
zymes, particularly angiotensin-converting enzyme (ACE), has been
investigated as a therapeutic approach and the search for new molecules
able to act as potent inhibitors and to exert hypotensive effects has been
also carried out within the field of Food Science (Chaudhary, Sabikhi,
Hussain, & Kumar, 2020; Guo et al., 2019; Morales et al., 2018).
Kombucha and kombucha-based products have been tested as sour­
ces of compounds able to inhibit ACE activity. Among related works,
Vitas, Vukmanovic, Cakarevic, Popovic, and Malbasa (2020) recently
found that green tea kombucha showed higher inhibition than black tea
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Trends in Food Science & Technology 105 (2020) 323–333
kombucha. They also prepared kombucha from other sources noticing
higher inhibitory activities for beverages with elderberry or stinging
nettle and lower activities for those with winter savory, peppermint,
wild thyme or quince. Gamboa-Gomez et al. (2016) observed potent
ACE-inhibitory activities for kombucha beverages using river red gum or
Mexican bay leaves but they noticed that the fermentation process was
detrimental for this capacity that was higher in non-fermented infusions.
It is known that milk proteins contain relevant bioactive sequences
that could be released as active inhibitory peptides. With this aim,
Elkhtab, El-Alfy, Shenana, Mohamed, and Yousef (2017) utilized SCOBY
microorganisms for milk fermentation, measured ACE-inhibitory activ­
ity in vitro and identified bioactive peptides by mass spectrometry con­
firming their activity by synthesizing them. Several new peptides were
released during fermentation by kombucha consortium with low IC50
values (VAPFPEVFGK, LVYPFPGPLH, FVAPEPFVFGKEK). Moreover,
Hrnjez et al. (2014) fermented milk with kombucha inoculum too,
exerting higher ACE inhibitory activity than other products obtained by
probiotic and yoghurt starters.
As it can be seen in Table 3, no animal studies have been carried out
to assess antihypertensive activity. However, one human trial can be
found in the literature reporting blood pressure-lowering effects of
kombucha oral administration in individuals with mild hypertension for
3 months (Hiremath, Vaidehi, & Mushtari, 2002). However, these re­
sults were not explained in detail and the control group was missing.
Currently, there is one clinical trial that will be completed in 2021.
Blood pressure is being measured in subjects with high glucose levels
and, in this case, 4 groups have been defined since they are consuming
commercial kombucha, brewed kombucha, tea or tap water (Clin­
icaltrials.gov, 2019).
7. Antidiabetic/hypoglycemic activity
Diabetes is a complex metabolic disorder of hyperglycemia (90% of
diabetic adults are involved in diabetes mellitus type II) that may be
accompanied by cardiovascular diseases and illnesses related with
retina, kidney or nervous system (Bai et al., 2020). Current antidiabetic
therapies utilize drugs such as metformin or insulin but the use of hy­
poglycemic dietary ingredients can be an interesting alternative for
patients with moderate diabetes, avoiding drugs side effects and facili­
tating the administration (Harnedy et al., 2018).
Kombucha beverages have been tested in vitro regarding their ability
to inhibit enzymes related with the increase of blood glucose levels. For
that matter, Kallel, Desseaux, Hamdi, Stocker, and Ajandouz (2012)
evaluated the inhibitory activity of green and black tea kombuchas
against porcine pancreatic α-amylase, reporting that this capacity
increased with fermentation time and correlating it with phenolic
compounds content, particularly epigallocatechin gallate, gallocatechin
gallate and epicatechin gallate, that confirmed this inhibitory effect
when they were tested as separated pure compounds. In vitro inhibition
of α-amylase and β-glucosidase was also noticed by Watawana, Jaya­
wardena, Ranasinghe, and Waisundara (2016) and Watawana, Jaya­
wardena, & Waisundara et al. (2018) in green, black and oolong tea
kombuchas. Moreover, soy drink fermentation with kombucha SCOBY
was studied and the in vitro ability to inhibit α-amylase and α-glucosi­
dase activities was increased after the process (Xia et al., 2019). These
inhibitory capacities were observed for oak kombucha and the hypo­
glycemic effects were validated in vivo, leading to a decrease in fasting
glucose concentrations in obese mice with high saturated fat and fruc­
tose diets (Gamboa-Gomez et al., 2017). Srihari, Karthikesan, Ashok­
kumar, and Satyanarayana (2013) administered a lyophilized extract of
black tea kombucha to streptozotocin-induced diabetic Wistar rats and
the product reduced glycosylated haemoglobin and plasma insulin
levels. Snake fruit kombucha was also tested in this animal model and
led to a significant fasting plasma glucose reduction and a regeneration
of pancreatic β-cells that was higher than those recorded for black tea
kombucha (Zubaidah et al., 2018, 2019a and 2019b). Furthermore,
D. Morales
alloxan-induced diabetic rats fed black tea kombucha and the activity of
plasma and pancreatic α-amylase was significantly reduced (Aloulou
et al., 2012).
Hypoglycemic effect evaluation was included in the clinical trials
that were mentioned in the “Antihypertensive activity” section. In the
case of Hiremath et al. (2002) study, it was reported that kombucha
daily consumption (during 3 months) normalized blood sugar values in
non-insulin-dependent diabetes mellitus subjects (but without consid­
ering a control group, as it was previously commented). In the current
clinical trial (Clinicaltrials.gov, 2019), blood and fasting blood glucose
and insulin levels are being measured in subjects with high glucose
levels (in 4 different groups that were described in the previous section).
8. Antimicrobial/antiviral activity
The increasing appearance of microorganisms exhibiting resistance
to existing antimicrobials has promoted the search of novel molecules
able to inhibit the growth or to eliminate undesired microbes. Natural
antimicrobial compounds can be found in edible sources and they have
been tested because of their potential utility in food preservation and
their effects against pathogens (Sakkas & Papadopoulou, 2017). Thus,
novel strategies are crucial to overcome drug resistance: the use of
recently isolated compounds, the evaluation of molecules that were not
previously utilized against microbes and the new approaches to reen­
gineer natural products (around their effective natural scaffolds) to
potent antimicrobials (Rossiter, Fletcher, & Wuest, 2018).
Kombucha beverages have proved their antimicrobial capacity and,
although many studies correlated this activity with acetic acid produc­
tion during fermentation, there is scientific evidence informing that this
is not the only compound found in kombucha which is capable of
exerting antibacterial and antiviral effects. Thus, the presence of mole­
cules released from the source material (tea or others) is also relevant by
itself or carrying out synergistic actions with organic acids.
Consequently, the type of source material and ingredients could play
a key role in the antimicrobial activity of the final product. For instance,
green tea kombucha showed higher activity than black tea kombucha
against bacteria (Staphylococcus spp., Listeria monocytogenes, Micrococcus
luteus, Escherichia coli) and yeasts (Candida parapsilosis) (Battikh, Chaieb,
Bakhrouf, & Ammar, 2012; Cardoso et al., 2020). Cardoso et al. (2020)
explained this fact correlating the activity with higher abundance of
catechins among phenolic compounds in green tea that also contained
verbascoside as an exclusive compound. Green tea kombucha was also
reported as an effective inhibitor and bactericidal agent against Alicy­
clobacillus spp., known to cause spoilage in fruit juices (Mizuta et al.,
2020) and both green, black and oolong tea kombuchas displayed
antimicrobial activities against several enteric bacteria in correlation
with acetic acid content of the beverages (Kaewkod, Bovonsombut, &
Tragoolpua, 2019). This compound was also described as responsible of
the inhibitory effects of black tea kombucha against Candida spp. and
Haemophillum influenzae (Ivanisova et al., 2020) and the importance of
organic acids was pointed out by Ayed et al. (2017) that noticed that the
neutralization of red grape juice kombucha eliminated its bacteriostatic
power. The addition of other vegetal ingredients such as cinnamon could
also improve the content of organic acids (e.g. acetic and glucuronic)
and, therefore, antimicrobial activity of kombucha (Shahbazi, Gahruie,
Golmakani, Eskandari, & Movahedi, 2018).
Besides organic acids, other kombucha compounds were reported as
antimicrobial molecules. For instance, Tu et al. (2019) fermented soy
whey with black tea kombucha inoculum and obtained significant cor­
relations between flavonoid contents and antibacterial activity against
Staphylococcus aureus, Bacillus subtilis and E. coli. Moreover, Pei et al.
(2020) isolated a novel bacteriocin produced by the kombucha micro­
organism Lactobacillus plantarum, determining its aminoacidic sequence
(NIVWQLIGLPAQA) and confirming its bactericidal effects against
Gram + and Gram – bacteria but also its antifungal activity against
Aspergillus niger, Candida albicans and Saccharomyces cerevisiae.
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Trends in Food Science & Technology 105 (2020) 323–333
Kombucha products that were not based on tea leaves were also tested:
yarrow kombucha inhibited not only the bacterial growth but also
A. niger and C. albicans (Vitas, Cvetanovic, et al., 2018); lemon balm tea
exhibited antimicrobial activity against bacteria (Salmonella enteritidis,
E. coli) but no effects against yeasts (S. cerevisiae) and moulds (A. niger)
(Cetojevic-Simin et al., 2012); and garlic kombucha showed lower
antibacterial effects when compared to a fresh garlic extract (Pure &
Pure, 2016a).
Furthermore, some in vivo assays were carried out to confirm the
promising in vitro results. Bhattacharya et al. (2020) recently isolated a
polyphenolic fraction from black tea kombucha that inhibited Vibrio
cholerae swarming mobility, expression of flagellar regulatory genes,
bacterial protease secretion and mucin penetration in vitro. These results
were in concordance with posterior in vivo tests: the fraction inhibited
the fluid accumulation induced by V. cholerae in a rabbit ileal loop model
and the intestinal colonization in a sucking mice model. In addition, Fu,
Wu, Lv, He, & Jiang (2015) validated the antiviral effects against the
foot and mouth disease virus (that affects cloven-hoofed animals),
observing a protective effect in swines that were treated with a Chinese
herbal kombucha (prepared with green tea, licorice, Momordica gro­
vesnori and chrysanthemum) before intramuscular inoculation of the
virus.
Besides, another factor which is significantly relevant in functional
foods is the ability to differently modulate human gut microbiota and
the subsequent effect on human health status (Vamanu, 2018).
Regarding this, kombucha effects on microbiota composition must be
investigated but just a recent study has been already published using
mice having non-alcoholic fatty liver disease as experimental model.
The authors reported changes in host microbial populations (e.g. de­
creases of Allobacullum, Turibacter and Clostridium abundances) after tea
kombucha consumption and they suggested that these modifications
may help in the suppression of fat accumulation in the liver (Jung et al.,
2019). However, clinical trials are required to evaluate the effect of
kombucha beverages on human microbiome and their implications on
human health.
9. Other activities
Apart from the biological activities that were reviewed in the pre­
vious sections, other not so widely studied functions were found in the
literature.
Wang, Ji, et al. (2013) fermented tea kombucha with Gluconaceto­
bacter sp. A4, leading to a high DSL production and noticing a protective
effect against acetaminophen-induced liver injury in mice. Kabiri and
Setorki (2016) intraperitoneally injected tea kombucha to rats prior to
brain damage induction by ischemia and reperfusion and they reported a
neuroprotective effect of kombucha by oxidative damage inhibition and
neuronal cell death reduction. Rahmani et al. (2019) described in vitro
anti-acetylcholinesterase activity effects in African mustard leaves
kombucha possibly due to its phenolic profile, motivating further studies
to elucidate its neuroprotective capacity. Recently, Taheur et al. (2020)
described that kombucha yeasts were able to degrade aflatoxin B1, a
molecule that plays a key role in tea toxicity.
10. Current limitations, conclusions and future perspectives
This review illustrated the scientific evidences around the biological
activities of kombucha beverages. The current strengths are the prom­
ising results that were obtained in vitro proving antioxidant, immune-
modulatory, antiproliferative, hypocholesterolemic, antihypertensive,
hypoglycemic, antimicrobial, etc. abilities. However, a significantly
lower number of in vivo studies was carried out to confirm these effects.
Moreover, there is only one published clinical trial related with anti­
hypertensive and hypoglycemic effects but a clear description of the
study conditions and the control group is missing (Hiremath et al.,
2002). A more detailed human study is being developed and it will be
D. Morales
finished in 2021 (Clinicaltrials.gov, 2019). Undoubtedly, these studies
are mandatory to validate kombucha biological activities and required
to claim health declarations of these products as functional foods.
Other evidenced weakness is the absence of compound-activity
correlations. There is little information of isolated kombucha com­
pounds and their biological functions (just for DSL and specific peptides)
and the associations have been carried out calculating the content of
potentially bioactive compounds in enriched fractions (e.g. specific
phenolic compounds) and measuring particular activities. Thus, further
studies focused on purification of these molecules and bioactivities
evaluation are recommended.
In addition, bioaccessibility and bioavailability analyses of the
bioactive compounds must be carried out to obtain realistic approaches
of their potential effects after human consumption. Accordingly, many
studies reported effects that would suppose the consumption of non-
normal amounts of the product. Moreover, the ability of kombucha
beverages to modulate human microbiota must be investigated.
Most of the research works have been focussed on antioxidant ac­
tivity and phenolic compounds. However, a great diversity of bioactive
compounds can be found in kombucha beverages and the variety of raw
materials that can be fermented should motivate novel studies analysing
biological activities of, among others, released and produced peptides,
lipids, secondary metabolites, etc.
In conclusion, further research involving more in vivo tests, bio­
accessibility, bioavailability and microbiota-modulation studies, clinical
trials and isolation of identified and novel bioactive compounds are
required to validate the widely described in vitro biological effects of
kombucha beverages and ingredients.
Declaration of competing interest
None.
Acknowledgments
The author would like to thank EIT Food for the RIS Talent Fellow­
ship (EIT RIS Programmes).
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