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Received: 7 January 2020 Revised: 17 April 2020 Accepted: 21 April 2020

DOI: 10.1111/1541-4337.12574

COMPREHENSIVE REVIEWS IN FOOD SCIENCE AND FOOD SAFETY

Microbiological and technological parameters impacting the


chemical composition and sensory quality of kombucha

Thierry Tran1 Cosette Grandvalet1 François Verdier2 Antoine Martin2


Hervé Alexandre1 Raphaëlle Tourdot-Maréchal1

1 UMR Procédés Alimentaires et


Abstract
Microbiologiques, Université de Bourgogne
Franche-Comté/AgroSup Dijon, Équipe Vin Kombucha is a beverage made from sugared tea transformed by yeasts and acetic acid
Alimentation Micro-organismes Stress bacteria. Being originally homemade, it has become an industrially produced soft
(VAlMiS) Institut Universitaire de la Vigne et
du Vin Jules Guyot, 2 rue Claude Ladrey,
drink whose quality standards are poorly defined and whose production process is still
Dijon, 21000, France not fully controlled. Based on current knowledge in beverages, links between kom-
2 Biomère, 14 rue Audubon, Paris, 75012, bucha’s chemical composition and sensorial compounds are drawn. Macromolecules
France
create turbidity, whereas uncharacterized tea pigments derivatives participate in the
Correspondence color. Residual sugars bring sweetness and organic acids produced by acetic acid bac-
Thierry Tran, UMR Procédés Alimentaires et teria form its characteristic sour taste. Acetic acid is also part of its aroma profile,
Microbiologiques, Université de Bourgogne
although little data are available on the smell of kombucha. Carbon dioxide, poten-
Franche-Comté/AgroSup, Dijon, Équipe
Vin Alimentation Micro-organismes Stress tially polyphenols, and residual ethanol are involved in the mouthfeel. In this review,
(VAlMiS) Institut Universitaire de la Vigne after defining the key compounds that shape the characteristic sensory properties of
et du Vin Jules Guyot, rue Claude Ladrey, BP
27877, 21000, Dijon, France.
kombucha, the impact of different production parameters is discussed. Water compo-
Email: thierrytran01@gmail.com sition is determinant in the extraction of tea compounds along with the tea type and
infusion duration and temperature. The type and amount of sweeteners play a role in
Funding information
European Regional Development Fund;
the sweetness and influences the production kinetics. Similarly, the amount of inocu-
Biomère lum and its microbial composition have an effect on the production, but the role of the
vessels’ geometry and temperature are also essential parameters that can be used to
adjust the acidification phase’s duration. Despite the amount of research carried out,
further investigations of kombucha’s sensory characteristics are needed. Such research
could lead to a better definition of kombucha’s quality and to an improved control over
its production process.

KEYWORDS
fermentation, Kombucha, process, quality, sensory

1 I N T RO D U C T I O N Lončar, Vitas, & Sathishkumar, 2014; Villarreal-Soto, Beau-


fort, Bouajila, Souchard, & Taillandier, 2018). It is believed
Kombucha, also named “kombucha tea,” is a fermented bev- to originate from Asia but all the elements appearing in the
erage resulting from the activity of a microbial consortium scientific literature, especially about the Chinese origins of
including yeasts, acetic acid bacteria, and often (but not kombucha, are supported by unverified sources (Dufresne &
always) lactic acid bacteria in sugared tea liquor as liquid Farnworth, 2000; Troitino, 2017). Nevertheless, these sto-
medium (Dufresne & Farnworth, 2000; Jayabalan, Malbaša, ries continue to feed the mythology of kombucha and could

Compr Rev Food Sci Food Saf. 2020;1–21. wileyonlinelibrary.com/journal/crf3 © 2020 Institute of Food Technologists® 1
2 THE PARAMETERS OF KOMBUCHA’S QUALITY…

be used to support the marketing communication of indus- infusions of plants that are not tea (Camellia sinensis) or on
trialized products. Indeed, the main marketing arguments of other food matrices will not be discussed, nor will flavored
kombucha are its putative benefits for human health. Numer- kombuchas. Indeed, the use of aromatization ingredients such
ous in vivo and in vitro studies were carried out to establish as fruits extracts, herbs, or spices implies the addition of a
the existence of antioxidant (Bhattacharya, Gachhui, & Sil, variety of compounds. Among them, nitrogen sources, sugars,
2013; Gamboa-Gómez et al., 2016; Jayabalan, Subathradevi, or antimicrobial compounds can be the origin of significant
Marimuthu, Sathishkumar, & Swaminathan, 2008), antimi- modification in microbial dynamics and elaboration kinetics
crobial (Battikh, Bakhrouf, & Ammar, 2012; Sreeramulu, before and after bottling. The complexity of the effects that
Zhu, & Knol, 2000; Steinkraus, Shapiro, Hotchkiss, & Mort- could be induced by the addition of such ingredients makes
lock, 1996), and hepato-protective effects (Murugesan, 2009; inclusion of this topic injudicious and thus it is not addressed
Wang et al., 2014). Nevertheless, the existence of benefi- in the present review. For the review on kombucha made with
cial effects of kombucha for human health remains contro- plants other than tea, the reader can be directed to the review
versial because of the insufficient amount of decisive scien- of Emiljanowicz and Malinowska-Pańczyk (2019).
tific data (Ernst, 2003; Jayabalan et al., 2014; Martínez Leal, After defining kombucha and the way it is generally
Valenzuela Suárez, Jayabalan, Huerta Oros, & Escalante- produced, due to the lack of published sensory data on
Aburto, 2018) and because of multiple cases of diseases or kombucha beverage, the components of kombucha’s flavor
unexplained death following the overconsumption of kom- will be discussed by breaking down its chemical composition
bucha (Holbourn & Hurdman, 2017; Phan et al., 1998; Sung, and discussing the known relationship of these parameters
Kole, Jones, Christensen, & Gladstein, 2009). The active com- with sensory characteristic. Then, with all these elements
pounds of kombucha with potential benefits for human health in mind, different aspects of the production process will be
originate from tea polyphenols, in particular epigallocatechin presented in order to give the reader all the known levers that
gallate (Jayabalan, Marimuthu, & Swaminathan, 2007; Khan can be used to shape the product’s flavor and quality.
& Mukhtar, 2007), hydrolytic enzymes, vitamins (B1 , B2 ,
B6 , B12 , and C) (Bauer-Petrovska & Petrushevska-Tozi, 2000;
Kumar & Joshi, 2016), and organic acids such as gluconic 2 G ENERAL K NOWLEDG E A BOU T
acid, glucuronic acid (Nguyen, Dong, Le, & Nguyen, 2014; KOMBUCHA AND ITS PROCESS
Nguyen, Nguyen, Nguyen, & Le, 2015) or D-saccharic-1,4-
lactone acid (Wang, Gan, Tang, Wang, & Tan, 2010) produced The traditional way of making kombucha consists in the brew-
by the microorganisms. To preserve these potential benefits ing of black tea liquor to which sucrose is added. Most of the
and/or promote a clean label, some kombucha producers do scientific literature reports the use of infusion (initial temper-
not pasteurize nor filter their products. ature between 70 and 95 ◦ C) rather than decoction in order
During the last decades, kombucha transitioned from a to perform tea extraction (Ali & Shivanna, 2017; Dufresne
homemade fermented beverage to a commercialized soft & Farnworth, 2000; Jayabalan et al., 2014; Villarreal-Soto
drink produced industrially. A striking result is that the mar- et al., 2018). Thus, infusion will be used as the initial process
ket of kombucha is expected to exhibit a strong growth rate of in the context of the present review. After the liquid reaches
17.5% in the United States between 2019 and 2024 (Mordor room temperature, the infusion is inoculated with a kombucha
Intelligence, 2019). culture in the form of a pellicle fragment or as whole (tradi-
Despite a strong development of kombucha-producing tionally referred to as “tea fungus”) and/or broth (Dufresne
companies, the producers of this beverage suffer from a lack of & Farnworth, 2000; Greenwalt, Steinkraus, & Ledford, 2000;
technical knowledge similar to what exists for the production Jayabalan et al., 2008). In the course of kombucha production
of other beverages, such as wines or beers. This review aims a new cellulosic pellicle forms itself at the surface of the liq-
at giving the kombucha brewers, R&D staff, and researchers uid phase (Chen & Liu, 2000). Despite the lack of consensual
an innovative approach of kombucha as a commercialized fer- definition of a biofilm, the pellicle satisfies the definition of a
mented beverage. It will revolve around the identification and biofilm with aggregated and sessile cells without a solid sur-
control of the different dimensions, or components, of kom- face (Alhede et al., 2011). In the case of kombucha, the cellu-
bucha’s quality through the lens of the current available sci- losic pellicle is an air–liquid interface biofilm. Such structure
entific literature. The concept of quality developed on wine has already been reported by two studies focusing on the wine
was studied by Charters and Pettigrew (2007) and distin- matrix (David-Vaizant & Alexandre, 2018; Zara et al., 2005).
guished between extrinsic and intrinsic qualities. This review Kombucha pellicle will therefore be referred to as “biofilm”
will focus on the intrinsic quality of tea-based kombucha that in the context of the present review.
is bound to its structural features, in other words its sensory The different steps of kombucha elaboration are not stan-
properties. This then excludes all the aspects related to price, dardized. According to previous studies, sucrose concentra-
packaging, or marketing. Also, fermented products based on tion and tea amount can range from 50 to 100 g/L and
THE PARAMETERS OF KOMBUCHA’S QUALITY… 3

from 1.5 to 10 g/L, respectively, with steeping time between Other microbial families have been identified and
5 and 15 min (Blanc, 1996; Chen & Liu, 2000; Chu & involve lactic acid bacteria with namely the genera Lac-
Chen, 2006; De Filippis, Troise, Vitaglione, & Ercolini, 2018; tobacillus, Bifidobacterium, and Oenococcus (Chakra-
Goh et al., 2012a; Jayabalan et al., 2014; Kallel, Desseaux, vorty et al., 2016; Coton et al., 2017; Marsh et al.,
Hamdi, Stocker, & Ajandouz, 2012; Lončar, Djurić, Malbaša, 2014).
Kolarov, & Klašnja, 2006; Malbaša et al., 2006; Malbaša, The biological transformation driving the elaboration of
Lončar, & Djurić, 2008; Neffe-Skocińska, Sionek, Ścibisz, kombucha is a combination of microbial metabolic inter-
& Kołożyn-Krajewska, 2017; Reiss, 1994; Sievers, Lanini, relationships, which is not limited to a plain fermentation.
Weber, Schuler-Schmid, & Teuber, 1995). Namely, the order Consequently, this microbial process will be referred to as
and length of each step and the various amounts of tea, sugar, “elaboration” in the context of the present review. Kombucha
and inoculum can vary and be adapted depending on personal elaboration occurs generally following the process shown in
and empirical appreciations. Figure 1. At room temperature, the sucrose added in the tea
To trigger the transformation of sugared tea infusion in liquor is hydrolyzed by the yeasts into fructose and glucose,
kombucha, a microbial culture must be added to the sweetened which are subsequently converted into ethanol through alco-
tea medium. Jayabalan et al. (2014) mention, while describ- holic fermentation. This step is the basis of the symbiosis that
ing the inoculation process, that kombucha culture as tea fun- occurs in kombucha because acetic acid bacteria are unable to
gus has to be placed in the sugared tea broth. Nevertheless, metabolize sucrose. But yeasts provide them carbonated sub-
whether the process is home-made, produced at industrial strates that they can use. Since the system is in contact with
scale, or in research labs, several methods are reported. For air, the obligate aerobe acetic acid bacteria use the available
example, the inoculation step has been reported to be achieved glucose and ethanol to produce organic acids (Chen & Liu,
by the addition of the broth (Blanc, 1996; Jayabalan et al., 2000; Sievers et al., 1995), mainly acetic acid from the oxi-
2007; Loncar, Kanuric, Malbasa, Djuric, & Milanovic, 2014; dation of ethanol and gluconic acid obtained from glucose as
Malbaša et al., 2008), the addition of only the biofilm or part of their oxidative metabolism (De Ley, 1961; Lynch, Zan-
biofilm fragments (Jayabalan, Malini, Sathishkumar, Swami- nini, Wilkinson, Daenen, & Arendt, 2019). Although glucose
nathan, & Yun, 2010; Reiss, 1994; Sievers et al., 1995), or the is used by acetic acid bacteria, it is thought that a part can
addition of both (Chen & Liu, 2000; Goh et al., 2012a; Kallel remain available for yeasts as well as fructose as substrates
et al., 2012). for alcoholic fermentation (Sievers et al., 1995). The acidifi-
There is no single “culture” or microbial consortium for cation of the medium leads to the decrease of pH, which pre-
developing kombucha but instead a multitude of matrix- vents the development of pathogenic microorganisms under
dependent consortia whose origins are unknown. It appears the value of 3 (Leistner, 2000). It is worth noting that the
that the only constant element that defines a kombucha culture initial pH after inoculation is dependent on the total acid-
is the simultaneous presence of yeasts and acetic acid bacteria, ity of the inoculum. In parallel, the formation of a gelati-
lactic acid bacteria not being always present. nous mass is observed. This biofilm composed of cellulose
The acetic acid bacteria community is mostly repre- is produced by some acetic acid bacteria, more specifically
sented by the genera : Acetobacter (sp. okinawensis and Komagataeibacter xylinus (Zhang, Wang, Qi, Ren, & Qiang,
tropicalis), Gluconobacter (sp. oxydans), Gluconacetobacter 2018) (formerly named Acetobacter xylinum and Gluconace-
(sp. europaeus and saccharivorans), and Komagataeibacter tobacter xylinus [Yamada et al., 2012]). In a context of indus-
(sp. kombucha and xylinus) (Chakravorty et al., 2016; De Fil- trial production, the biofilm might be removed before bottling
ippis et al., 2018; Marsh, O’Sullivan, Hill, Ross, & Cotter, and smaller pieces can be eliminated by filtration. After bot-
2014; Reva et al., 2015). tling, further cellulose synthesis should be stopped because of
The yeast community is more variable and includes oxygen deprivation after consumption of residual oxygen by
genera such as Zygosaccharomyces (sp. lentus, bisporus microorganisms, namely acetic acid bacteria or yeast with res-
and bailii), Candida (sp. stellimalicola and tropicalis), piratory metabolism. Similarly, the acetic conversion can be
Lachancea (sp. thermotolerans and fermentati), Kloeck- stopped at any time according to the desired taste of the bev-
era/Hanseniaspora (sp. valbyensis), Torulaspora (sp. del- erage by putting the system in anaerobic conditions, which
brueckii), Rhodotorulaspora (sp. mucilagenosa), Pichia inhibits acetic acid bacteria and favors alcoholic fermentation
(sp. mexicana and occidentalis), Brettanomyces/Dekkera of yeasts, turning the residual sugars into ethanol and carbon
(sp. bruxellensis and anomala), Saccharomyces (sp. cere- dioxide. A sour and sparkling beverage is obtained. To our
visiae), Schizosaccharomyces (sp. pombe) and Saccharomy- knowledge, little investigation has been carried out on this
coides (Chakravorty et al., 2016; Coton et al., 2017; De secondary fermentation. Since different elaboration times are
Filippis et al., 2018; Markov, Cvetković, & Bukvić, 2005; used, chemical compositions reported in the literature are very
Marsh et al., 2014; Reva et al., 2015; Teoh, Heard, & Cox, diverse (Table 1 [Villarreal-Soto et al., 2018]). In order to
2004). provide a good acceptability of the product, the brewer will
4 THE PARAMETERS OF KOMBUCHA’S QUALITY…

F I G U R E 1 Kombucha metabolism and microbial interactions. (a) Kombucha is brewed by adding tea and table sugar to a small amount of
kombucha starter that contains yeast and acetic acid bacteria. These microbes begin to break down the sugar, leading to a metabolic cascade that ends
with a bubbly, acidic, and slightly alcoholic beverage. (b) During the process of elaboration, cooperative and competitive interactions occur among
microbes. The production of the public good invertase by yeast, the removal of waste products through metabolization of alcohol, and the generation
of the cellulose pellicle by bacteria are potentially cooperative functions. Antimicrobial metabolites, low pH, and the generation of a physical barrier
inhibit the growth of competitors (adapted from May et al., 2019)

TABLE 1 General chemical composition of Kombucha (Villarreal-Soto et al., 2018)


Fermentation
Compound Average composition Initial sucrose time (days) References
Organic acids Acetic acid 5.6 g/L 70 g/L 15 Blanc(1996)
Acetic acid 8.36 g/L 100 g/L 18 Jayabalan et al. (2007)
Acetic acid 11 g/L 100 g/L 30 Chen and Liu (2000)
Gluconic acid 39 g/L 100 g/L 60 Chen and Liu (2000)
Glucuronic acid 0.0160 g/L 70 g/L 21 Lončar et al. (2006)
Lactic acid 0.18 g/L 100 g/L 18 Jayabalan et al. (2007)
Vitamins Vitamin B1 0.74 mg/L 70 g/L 15 Bauer-Petrovska and
Vitamin B2 8 mg/100 mL 70 g/L 10 Petrushevska-Tozi (2000)
Vitamin B6 0.52 mg/L 70 g/L 15 Malbaša et al. (2011)
Vitamin B12 0.84 mg/L 70 g/L 15 Bauer-Petrovska and
Vitamin C 25 mg/L 70 g/L 10 Petrushevska-Tozi (2000)
Bauer-Petrovska and
Petrushevska-Tozi (2000)
Malbaša et al. (2011)
General Ethanol 5.5 g/L 100 g/L 20 Chen and Liu (2000)
composites Proteins 3 mg/mL 100 g/L 12 Jayabalan et al. (2007)
Tea polyphenols 7.8 mM (gallic acid 100 g/L 15 Chu and Chen (2006)
equivalent)
Minerals Cu, Fe, Mn, Ni, Zn 0.1 to 0.4 𝜇g/mL 70 g/L 15 Bauer-Petrovska and
Petrushevska-Tozi (2000)
Anions F− , Cl− , Br− , I− , NO3 − , 0.04 to 3.20 mg/g 100 g/L 7 Kumar and Joshi (2016)
HPO4 − , SO4 −

have to make sure that the acidity is not too high and ability; a lower pH would be too acidic (Lončar et al.,
the pH not too low in the case of long-lasting elaboration, 2006).
else the taste will not remain pleasant. The pH value of 3 The microbiological studies of kombucha elaboration show
appears to be a minimum threshold for kombucha accept- an increase in yeast and bacteria population during the
THE PARAMETERS OF KOMBUCHA’S QUALITY… 5

two first days following the inoculation of the sugared tea 3 DIS S ECTIO N O F TH E
liquor (Coton et al., 2017; De Filippis et al., 2018; Teoh CHEMICA L COMPOSITION OF
et al., 2004). Next, different variations in populations were KOMBUCHA IN RELAT IONSHIP TO
reported according to the consortium used (Chen & Liu, ITS POTENTIAL SENSORIAL
2000), the type of tea (Coton et al., 2017), or the tempera- I M PAC T
ture (De Filippis et al., 2018). The impact of these parame-
ters on the process and the final product will be developed in The perception of all food and beverages is conveyed by the
Section 4. consumer’s five senses: sight, hearing, touch, smell, and taste.
The microbial dynamics occurring in a given kombucha When drinking kombucha, the consumer experiences a mix of
consortium during the elaboration highlighted the domina- visual (aspect of the product), olfactive (aroma profile before
tion of one to three genera for the yeasts (Chakravorty et al., and after ingestion), taste (sweetness, sourness), and touch
2016; Reva et al., 2015; Teoh et al., 2004) and for bac- sensations (for example, via chemosensation of the tingling
teria (Chakravorty et al., 2016; Coton et al., 2017; Reva of carbon dioxide bubbles on the tongue) (Redondo, Gomez-
et al., 2015) over the important diversity of the other detected Martinez, & Marcos, 2014).
genera (with an abundance inferior to 1% per genus). One There are very few data available of descriptive sensory
predominant genus is often found per kingdom, for exam- analysis on tea kombucha beverages. The study of Neffe-
ple: Candida stellimalicola, (Chakravorty et al., 2016) or Skocińska et al. (2017) includes a sensory analysis of kom-
Dekkera anomala (Reva et al., 2015) for the yeasts and K. xyli- bucha beverages elaborated during 10 days, made of a mix
nus (Reva et al., 2015) for the bacteria. Some studies have of black and green tea (2 and 4 g/L, respectively) and
reported important variations of the proportion of the dom- 100 g/L sucrose at 20, 25, and 30 ◦ C. The sensory analy-
inant species over time, suggesting the occurrence of inter- sis was performed by a panel of 16 untrained people and
action between yeast species (Chakravorty et al., 2016; Teoh no statistical analysis was performed on the results obtained
et al., 2004) and between bacteria species (Coton et al., (Figure 2 [Neffe-Skocińska et al., 2017]). Therefore, the inter-
2017). pretation of these results should be handled with care. Never-
Currently, the control of kombucha production is mostly theless, the descriptors used are representative of the sensory
empiric despite the increasing amount of knowledge and characteristics of kombucha and help to define its main fea-
understanding provided by the scientific community. Yet, pro- tures. The descriptors with highest scores are: “color inten-
fessional producers of kombucha are confronted with several sity” and “clarity” for the visual attributes; “tea” and “cit-
difficulties for the control of its elaboration. First, there exists ric” for the olfactory attributes; and “tea,” “citric,” and “acid”
a wide variability of elaboration kinetics due to the complex for the flavor attributes. The descriptor “sweetness” was not
and hard-to-control microbial consortium, as opposed to a sin- assessed despite residual sugars ranging from 27 to 70 g/L.
gle culture fermentation (Villarreal-Soto et al., 2018). More- The descriptors “yeast flavor,” “acetic acid flavor,” “bitter fla-
over, the production of batches by successive inoculations vor,” “stinging flavor” have low scores. The low “stinging
(also called propagation) could lead to a modification or evo- flavor” score could be explained by a lack of secondary fer-
lution of the consortium in terms of composition, microbial mentation in closed vessels. Recent studies incorporated sen-
dynamics, or both. Finally, in the current context, refrigera- sory analysis but did not bring further detailed descriptive ele-
tion is not always sufficient to completely prevent microbial ments (Ivanišová et al., 2019; Shahbazi, Hashemi Gahruie,
activity after commercialization of unstabilized bottled kom- Golmakani, Eskandari, & Movahedi, 2018).
bucha (using thermal or filtration processes). Because of the The use of a more precise and standardized set of descrip-
possibility of yeast refermentation, the production of carbon tors could allow deeper investigations of the olfactive and gus-
dioxide resulting in bottle explosions is a real risk for kom- tative dimensions of kombucha. So, a more indirect approach
bucha producers. Moreover, the possible increase in alcohol is necessary to explore the sensory characteristics of kom-
content (Talebi, Frink, Patil, & Armstrong, 2017) could have bucha by relating the chemical constituents of kombucha to
consequences on regulatory levels. In fact, kombucha is clas- the known sensory properties of those compounds.
sified as nonalcoholic beverage only as long as its alcohol con-
tent does not exceed a threshold value. In United States, it is 3.1 The sight: the aspect of kombucha
set at 0.5% (Code of Federal Regulations, Title 27: Alcohol,
Tobacco and Firearms [Alcoholic content, 1993]), whereas Kombucha beverages can be either clear filtered or turbid
in the European Union, this limit is set at 1.2% (Regulation (as nonfiltered version). The turbidity of the latter is mainly
[Eu] No 1169/2011 Of The European Parliament And Of The due to the colloidal state of the aqueous beverage, defined as
Council [Official Journal of the European Union, 2011]). a suspension of particles. These particles are composed of
6 THE PARAMETERS OF KOMBUCHA’S QUALITY…

F I G U R E 2 Sensory profiles of the kombucha beverages after 10 days of elaboration process at 20, 25, and 30 ◦ C (K20, K25, and K30,
respectively). “o.” stands for “olfactive,” “int.” stands for “intensity,” and “f.” for “flavor” (Neffe-Skocińska et al., 2017)

microorganisms and large molecules or aggregates ranging that gives black tea its characteristic color (Harbowy et al.,
from 1 to 1000 nm. The presence of such bodies induces 1997). Nonetheless, theaflavin is not the main contributor
the scattering of the light. This phenomenon is known as the of black tea’s color. The second family named thearubigins,
Tyndall Effect (Petrucci, Herring, Madura, & Bissonnette, results from a higher degree of polymerization and acts as
2011). Although little investigation was carried on the col- the main pigment. The molecular structure of those polymeric
loids of kombucha, it can be speculated that they can result molecules is still not fully elucidated (Haslam, 2003). Chem-
from the aggregation of proteins (Jayabalan et al., 2007; Petro- ical structures of some of those compounds are displayed in
vić, Suturović, Lončar, & Malbaša, 1999), polyphenols, and Figure 3.
cellulose fibrils produced by acetic acid bacteria (Goh et al., Although little attention was given to the color and
2012b; Lin et al., 2013; Zhang et al., 2018). Colloids play a polyphenols of kombucha, two studies reported a significant
huge part of soft drinks’ intrinsic quality (Kappes & Schmidt, decrease in color intensity and increase in total phenolic con-
2007), but to our knowledge there are no data available men- tent during kombucha elaboration (Chakravorty et al., 2016;
tioning the mouthfeel of kombucha. Chu & Chen, 2006). The decrease of pH could be the cause
The color hue of kombucha is mainly due to the presence of of the change of color (United States Patent No. 4,552,776,
the pigment polyphenols extracted from the tea. The charac- 1985). It has also been hypothesized that the biological activ-
teristic color of black tea results from polyphenols oxidase- ity of the kombucha consortium may alter or even depolymer-
type enzymes, or the so-called “fermentation” of fresh tea ize the pigments extracted from the tea (Haslam, 2003) Thus,
leaves (Harris & Ellis, 1981). This process allows the oxi- a large part of kombucha pigments may be derivatives from
dation and polymerization of native polyphenols composed the tea polyphenols.
mainly of catechins (epicatechin, epigallocatechin, and their Another component of kombucha’s visual identity as a car-
gallic acid ester derivatives) into different classes of polymers bonated beverage is the aspect and abundance of the bub-
(Balentine, Wiseman, & Bouwens, 1997; Harbowy, Balen- bles that are dependent on the carbonation process that can
tine, Davies, & Cai, 1997). Two of those classes are pig- be obtained naturally after fermentation in closed vessel or
ments: one of them is theaflavin, a red-orange pigment dimer by injecting carbon dioxide artificially (forced carbonation)
THE PARAMETERS OF KOMBUCHA’S QUALITY… 7

F I G U R E 3 Chemical structure of some tea constituents (adapted from Dufresne and Farnworth [2000])

(Barker, Jefferson, & Judd, 2002; Descoins, Mathlouthi, Le olfactory experience (as for most food and beverages) (Acree
Moual, & Hennequin, 2006). Bubbles number is dependent & van Ruth, 2003), which is why this section relies mainly on
on nucleation sites, which mainly consist in particles (belong- speculations. Kombucha’s smell has been widely described
ing to the product itself or its vessel) (Lubetkin & Blackwell, as “cidery” (Dufresne & Farnworth, 2000; Greenwalt et al.,
1988; Wilt, 1986). The formation kinetics and size of bubbles 2000; Jayabalan et al., 2014). As many fermented bever-
is dependent on the interfacial tension between the gas and liq- ages, the odorant compounds originate from both the raw
uid phase (Jones, Evans, & Galvin, 1999), which means that material (the tea) and volatile metabolites produced by the
macromolecules such as polysaccharides and proteins play a microorganisms. Although black tea hosts numerous volatile
role in the visual aspect of a carbonated drink (Barker et al., molecules such as 3-hexenol (greenish), linalool (floral),
2002). geraniol (sweet, honey-like), 2-phenylethanol (honey-like),
damascenone (rose-like), or 2,5-dimethyl-4-hydroxy-3(2H)-
3.2 The smell: a vaporous idea of kombucha’s furanone (DMHF) (caramel-like) (Ho, Zheng, & Li, 2015;
aroma profile Robinson & Owuor, 2013; Teranishi, Wick, & Hornstein,
Little information is available about the volatile com- 1999), typical tea aroma never seems to be part of the
pounds of kombucha, their origins, and their relationship to characteristic aroma profile of kombucha. Instead, the
8 THE PARAMETERS OF KOMBUCHA’S QUALITY…

elaboration-related aromas dominate: the vinegary odor asso- compared to sucrose (although those values are modulated
ciated with acetic acid produced by acetic acid bacteria and by their concentrations [Stone & Oliver, 1969]). The second
the cidery odor associated with the activity of yeasts (Rosend, is other compounds contained in those ingredients, such as
Kuldjärv, Rosenvald, & Paalme, 2019; Wei, Wang, Zhang, those produced by the Maillard reactions in agave syrup or
Yuan, & Yue, 2019). In cider, higher alcohols are largely molasses (Willems & Low, 2012). These compounds can be
produced: amyl alcohols (banana, pear), butanol (balsamic), volatile impacting the aroma profile or nonvolatile such as
propanol (fermented, fruity), ethyl acetate (solvent, fruity minerals or pigments that can impact the visual aspect of the
like), and ethyl lactate (creamy, fruity). 2-phenylethanol, hex- beverage.
anol (green), octanol (citrus), and butanoic acid (cheesy) were The sweetness of the final product is thus dependent on the
also identified in a significant amount (Mangas, González, residual amount of sugars, which is conditioned by the initial
Rodríguez, & Blanco, 1996; Rosend et al., 2019; Valles, amount of sweetener added to the tea and the consumption
Bedrinana, Tascon, Simon, & Madrera, 2007; Williams & of this substrate by microorganisms during the elaboration.
Rosser, 1981). More details about the influence of the substrate and its initial
It is noteworthy that many yeast genera are common to quantity on elaboration are developed in Section 4.3.
both kombucha and cider, namely Candida, Hanseniaspora,
Pichia, Dekkera, and Saccharomyces (Morrissey, Davenport, 3.3.2 Sourness
Querol, & Dobson, 2004; Valles et al., 2007; Wei et al., 2019).
Organic acids of kombucha are mainly produced by acetic
Although it can be suggested that the production of volatile
acid bacteria, although the contribution of yeasts and lactic
metabolites follows similar pathways, there are currently too
acid bacteria should not be neglected. The major organic acids
few elements to make any further statement on this topic.
contributing to kombucha’s taste are acetic acid, gluconic
Thus, many questions remain open: what are the volatile com-
acid, and glucuronic acid, whereas the minor ones are lactic
pounds essential to kombucha aroma and what are their ori-
acid, malic acid, and succinic acid (Blanc, 1996; Chakravorty
gins? If a comparison with wine is conducted, the deglyco-
et al., 2016; De Filippis et al., 2018; Jayabalan et al., 2007;
sylation of some aroma precursors by yeast 𝛽-glucosidase
Malbaša et al., 2008; Neffe-Skocińska et al., 2017). The sen-
during fermentation might have very little impact on kom-
sory properties of these metabolites are detailed in Table 2.
bucha (Fia, Giovani, & Rosi, 2005; Hernandez, Espinosa,
It should be noted that organic acids also generate, on a
Fernandez-Gonzalez, & Briones, 2003). The enzymatic oxi-
lower level, a bitter and astringent taste (Rubico & McDaniel,
dation of polyphenols (see Section 3.1.) can lead to aroma
1992; Siebert, 1999). Yet, those chemical species are not the
release in black tea before kombucha elaboration (Ho et al.,
main origin of the bitter perception of kombucha.
2015; Zhou et al., 2017). On the contrary, the precursors are
present in green tea in their glycosylated form, which means
that the use of this raw material for kombucha elaboration 3.3.3 Bitterness
could unlock aroma potential through the enzymatic activity Bitterness in kombucha, if not masked by the sweetness, can
of yeasts. take origin from tea caffeine and polyphenols (Balentine et al.,
1997; Harbowy et al., 1997). The average perception taste
3.3 The taste: a complex combination of threshold for caffeine has been reported of 0.2 g/L (Paulus
sapid substances & Reisch, 1980) and Chakravorty et al. (2016) reported a caf-
feine content ranging above this value from 0.6 to 1 g/L in the
3.3.1 Sweetness course of black tea kombucha elaboration.
The traditional sweetener used for kombucha, which also acts Polyphenols are bitter and astringent secondary metabo-
as carbonated substrate for the microorganisms, is sucrose lites produced by plants, including tea (C. sinensis) (Harbowy
(Blanc, 1996; Dufresne & Farnworth, 2000; Jayabalan et al., et al., 1997; Lesschaeve & Noble, 2005). Before any treat-
2014; Villarreal-Soto et al., 2018). The use of sucrose con- ment, tea leaves are composed of 30 to 40% (w/w dry weight)
stitutes the basis of the symbiosis between yeast and bacteria, of polyphenols. After infusion in hot water, the proportion
the former breaking down sucrose via the activity of invertase in dry matter extracted is about the same (Balentine et al.,
into glucose and fructose that are usable by the latter (May 1997; Harbowy et al., 1997). The phenolic composition of a
et al., 2019). It is possible to bypass this necessary step for green tea infusion can be summed up (in weight percentage
sucrose hydrolysis by introducing sugars in the form of puri- of solid extract) as a majority of catechins (30 to 42%): epi-
fied or mixed ingredients such as glucose syrup, agave syrup, catechin, epigallocatechin, and their gallic acid esters (epicat-
or molasses (Malbaša et al., 2008; Reiss, 1994). These ingre- echin gallate and epigallocatechin gallate). Minor compounds
dients differ on two major aspects. The first is the difference (2%) are represented by flavonols (kaempferol, quercetin,
in relative sweetness: glucose and fructose possess a sweet- and myricetin) as aglycones and glycosides. Phenolic acids
ness intensity of 65 to 75% and 120% (w/w), respectively, such as gallic acid and theogallin are also found (2% in
THE PARAMETERS OF KOMBUCHA’S QUALITY… 9

TABLE 2 Chemical and sensorial properties of organic acids of kombucha (Da Conceicao Neta, Johanningsmeier, & McFeeters, 2007; Li &
Liu, 2015; Ramachandran, Fontanille, Pandey, & Larroche, 2006)
Molecular Number of carboxylic Taste perception
Acid weight (g/mol) pKa functions threshold (mg/L water) Sensory quality
Acetic 60 4.75 1 52.6 Tart and sour
Lactic 90 3.86 1 80.1 Acrid
Gluconic 196 3.86 2 Not determined Mild, soft, refreshing taste
Malic 134 3.40 and 5.11 2 7.3 Smooth tartness
Succinic 118 4.19 and 5.50 2 22 Tart, slightly bitter in
aqueous solutions
Citric 192 3.14, 4.77 and 6.39 3 4.3 Tart, delivers a “burst” of
tartness

total). Tea tannins are frequently mentioned, but tannins 3.4 The touch: the booze and the fizz
amounts are in very low concentrations (catechins being
wrongly included under the term “tannin”). As a matter of To our knowledge, despite the presence of astringent polyphe-
fact, proanthocyanidins (or condensed tannins) are catechins nols, kombucha is never described as astringent. This is prob-
polymers and are present in very low amounts in green tea ably due to the presence of sugars that inhibit the perception
and are mainly dimers (trimers are even rarer) (Fraser et al., of astringency, as reported by Lyman and Green (1990).
2012). Chemical structures of these compounds are detailed in The perception of ethanol and its influence on other percep-
Figure 3. tions have been intensively studied to elucidate its impact on
Flavan-3-ol, including catechins and proanthocyanidins, the quality of alcoholic beverages. The perception of ethanol
possess bitter and astringent sensory properties (Fontoin, occurs across gustatory, olfactory, and trigeminal (or irrita-
Saucier, Teissedre, & Glories, 2008; Kielhorn & Thorngate tion) systems (Cometto-Muñiz & Cain, 1990; Greenwalt et al.,
III, 1999; Lesschaeve & Noble, 2005; Peleg, Gacon, Schlich, 2000; Laska, Distel, & Hudson, 1997). Several studies have
& Noble, 1999) that may contribute to the mouthfeel of kom- reported that the olfaction and nasal irritation thresholds of
bucha. The fate of polyphenols during kombucha elabora- ethanol ranged around 0.01% (v/v) in water or below, with
tion still remains enigmatic. Jayabalan et al. (2007) reported the irritation (also referred to as “trigeminal”) threshold being
a general decrease of epicatechin, epigallocatechin, and their always higher (Cometto-Muñiz & Cain, 1990; Martin & Pang-
gallate derivatives during the first 9 days of elaboration and born, 1970; Mattes & DiMeglio, 2001). The taste threshold of
then an increase of the nongallated species until the 12th day, ethanol ranges around 1 to 2%. This means that ethanol, even
suggesting the hydrolysis of the ester bound. Based on the if not identifiable on the olfactory level, can impact the aro-
work of Zhu, Zhang, Tsang, Huang, and Chen (1997), an matic profile of kombucha. On the other hand, the taste of a
acidic hydrolysis is rather unlikely, which is why an enzy- regular kombucha, with alcohol content below 1%, should not
matic origin is speculated. However, the consequences of induce a perception of alcohol taste. At near threshold con-
this phenomenon on the taste and mouthfeel of kombucha centration of ethanol, Mattes and DiMeglio (2001) obtained a
remain unknown. A new study (Cardoso et al., 2020) using predominant description of ethanol taste as bitter. Therefore,
UPLC-MS (ultra-performance liquid chromatography – mass the crossing of this threshold could impact the perception of
spectrometry) reported a significant change of phenolic pro- kombucha for some consumers and potentially decrease the
files between black tea infusion and kombucha resulting global appeal of the product, as it can be that case after com-
from it. The diversity of compounds increased (27 new com- mercialization on the shelves of retailer (Talebi et al., 2017).
pounds), whereas the abundance of compounds decreased. Carbonation affects visual, taste, and trigeminal compo-
Nevertheless, the global profile did not change drastically nents. The characteristic oral perception of carbonated drinks
with flavonoids remaining the main phenolic compounds fol- is the irritation or trigeminal sensation of tingling (Dessirier,
lowed by phenolic acids. New compounds produced during Simons, Carstens, O’Mahony, & Carstens, 2000). Although
elaboration mainly belonged to the class of flavonoids. No sig- few investigations have been carried out, the results indicate
nificant change was observed for green tea and the kombucha an enhancement of sourness by carbonation (Cometto-Muniz,
made from it. This suggests that kombucha elaboration could Garcia-Medina, & Calvifio, 1987; Yau & McDaniel, 1992).
affect the phenolic profile, especially for black tea, but would In addition, carbonation has been shown to enhance the per-
not change it drastically. The initial phenolic profile obtained ception of cold and vice versa (Green, 1992). However, no
from infusion is therefore a defining step for the final product interaction seems to occur between sweetness and carbona-
regarding this class of compounds. tion (Odake, 2001).
10 THE PARAMETERS OF KOMBUCHA’S QUALITY…

3.5 Perceptual interactions - to bottling and stabilization steps if applicable.

As seen for ethanol and carbon dioxide, chemosensory stimuli These aspects are discussed in details in the next section of
do not work independently on perceptions but are involved in this review.
interactions even below their own thresholds (Dalton, Doolit-
tle, Nagata, & Breslin, 2000). It has been shown that the
visual aspect of beverages influenced significantly both olfac-
4 P RO CES S PARAM ETERS
tory and taste perception in noncarbonated aqueous solutions
IM PACTING TH E CO M P O S ITIO N
(DuBose & Cardello, 1980; Stillman, 1993). The amplifica-
AND QUALIT Y OF KOMBUCHA
tion of fruitiness by sourness (and to a lesser extent sweet-
ness) has been reported as a taste–smell interaction. On the
The production of kombucha has been mainly studied by
contrary, suppressive effects of sourness on sweetness have
focusing on the elaboration kinetics and the influence of dif-
also been determined as taste–taste interactions (Bonnans &
ferent conditions ranging from the choice of the substrate,
Noble, 1993; Nahon, Roozen, & De Graaf, 1996).
the temperature, the inoculum, and even the vessel geome-
Finally, temperature is a key parameter that impacts the
try (Jayabalan et al., 2014; Villarreal-Soto et al., 2018). This
physical chemistry of food and beverages and in particular
section proposes to expand this approach by adding elements
the volatility of molecules. Kombucha is usually consumed at
relative to the production of the tea liquor.
cold temperature (around 4 ◦ C), as it is commonly marketed
and perceived by the consumer as a soft drink. Consequently,
sensory evaluation of kombucha should then be assessed at 4.1 Parameters impacting the quality of the
the appropriate temperature. tea liquor
The initial composition of water plays a role in the final com-
3.6 What makes kombucha “refreshing”? position of the tea liquor: the medium in which microorgan-
isms of kombucha grow and exert their activities. Indeed, the
Even though the quality components of kombucha have not mineral and organic composition of water is variable from a
been defined, one of the most frequently used descriptor region to the other and associated with its hardness, depen-
for this beverage is the adjective “refreshing” (Dufresne & dent on the calcium and magnesium contents. It was demon-
Farnworth, 2000; Jayabalan et al., 2014; Reiss, 1994; strated that the calcium concentration in water is an important
Villarreal-Soto et al., 2018). The real temperature of the food parameter of mineral and organic compounds of tea leaves
product plays indeed a major role in the perception of “fresh- (Mossion, Potin-Gautier, Delerue, Le Hécho, & Behra, 2008;
ness” (Guinard, Souchard, Picot, Rogeaux, & Sieffermann, Spiro & Price, 1987; Spiro, Price, Miller, & Arami, 1987)
1998; Labbe, Gilbert, Antille, & Martin, 2009; Zellner & along with the time/temperature conditions of the infusion
Durlach, 2002) but it is not the only one. Several studies (Price & Spitzer, 1993; Price & Spitzer, 1994; Spiro, Jaganyi,
attempted to define the different components of “freshness.” & Broom, 1992).
Among them, low viscosity or thickness was judged more The minerals had different behaviors during the extraction
“refreshing” in liquids than gels (Labbe et al., 2009; McE- depending on the ions when using distilled water (Matsuura,
wan & Colwill, 1996; Zellner & Durlach, 2002). We can Hokura, Katsuki, Itoh, & Haraguchi, 2001; Ødegård & Lund,
cite also some aroma such as mint, citrus, or peach (Labbe 1997):
et al., 2009), acidity and low sweetness (Labbe et al., 2009;
McEwan & Colwill, 1996). These elements echo the sweet- - Sodium, potassium, and nickel are strongly
ness/sourness balance that has been described as being “basic extracted (>55%);
typical taste profile of all flavored soft drinks.” “Without this
- Magnesium, aluminum, manganese, and zinc are moder-
sweetener and acid balance, the beverage would taste totally
ately extracted (between 20 and 55%);
wishy-washy and unexciting” (Shachman, 2005).
Kombucha possesses all the characteristics of a refreshing - Calcium, iron, and copper are poorly extracted (<20%).
beverage: it is served cold, it is carbonated, sour with low
sweetness and viscosity, and exhibits fruity aroma. The iden- When mineral water is used, the ions with the lowest con-
tity of a defined product can be modulated by working on dif- centrations show variation of their extraction yield of 10%
ferent parameters of kombucha’s production process ranging maximum according to Mossion (2007) (approximately sev-
from: eral mg/L). The nature of the tea induces a strong variability
in extraction behaviors.
Calcium possesses a particular behavior, especially if the
- the initial formulation of the sweetened tea; infusion water possesses a significant amount of it as it is the
- the elaboration conditions; case for mineral water: an inversed flux was observed from
THE PARAMETERS OF KOMBUCHA’S QUALITY… 11

present in the liquid phase. In the biofilm, the domination of


Gluconobacter is effective in all modalities after 2 days of
elaboration to the detriment of O. oeni.

4.3 Impact of the carbohydrate substrate and


of its initial amount
The use of different carbohydrates as substrate has been inves-
tigated by Reiss (1994). Sucrose is hydrolyzed by yeasts into
glucose and fructose. In this study, glucose favors the pro-
duction of lactic acid and fructose the production of ethanol.
Moreover, maltose has been poorly consumed and lactose did
F I G U R E 4 Effect of water composition on dissolved organic not affect the yield of ethanol nor stimulated the production
carbon content extracted from tea leaves per gram of brewed leaves (mg of lactic acid. Acetic acid bacteria transform glucose in glu-
C/g) (Mossion et al., 2008) conic acid and fructose in acetic acid. Beside carbohydrates,
it was also reported that lactic acid enhanced the production
of biomass.
water to the tea leaves (Anderson, Hollins, & Bond, 1971;
The utilization of molasses with 50% sucrose content (wet
Mossion, 2007; Mossion et al., 2008). It is speculated that
weight) was studied at different rates (35, 50, and 70% of total
pectins present in the cell walls of the tea cells bind to Ca2+
batch volume) (Malbaša et al., 2008; Malbaša, Lončar, Djurić,
ions (Capel, Nicolai, Durand, Boulenguer, & Langendorff,
& Došenović, 2008). With 35 and 50 g/L of molasses, kinet-
2006; Spiro et al., 1987), inducing structure modifications that
ics of sucrose consumption, pH variation, and production of
inhibit the extraction of compounds (Figure 4 [Mossion et al.,
organic acids were similar; whereas with 70 g/L of molasses,
2008]).
the consumption of sucrose was much faster and the produc-
It has also been shown that the extraction of organic matter
tion of lactic acid was enhanced to the detriment of acetic acid
was enhanced by:
but with identical total acidity values. Moreover, the decrease
- the increase in temperature (ranging above 70 ◦ C) and infu- in pH was less intense, probably due to the buffering capacity
sion duration (between 1 min and 1 hr); of molasses.
The increase of the initial content in sucrose led to the
- a weaker mineral content;
increase of the production of cellulose until a limit concen-
- more specifically, a lower calcium content (corresponding tration (90 g/L) after which a decrease of cellulose produc-
to lower water hardness). tion could be observed. The increase of initial sucrose con-
centration between 70 and 110 g/L accelerates the decrease
In conclusion, temperature, calcium content of water, and
of pH following a dose effect. The growth of yeast and bacte-
the nature of the tea are the main parameters involved in the
rial populations was also stimulated by the increase of initial
making of the tea liquor. This matrix will affect the growth
sucrose concentration (Goh et al., 2012a). A similar trend was
and activities of the kombucha cultures’ microorganisms.
observed in the work of Blanc (1996) but to a lesser extent.

4.2 Impact of the nature of tea on microbial


dynamics 4.4 The inoculum
According to Kallel et al. (2012), a consumption of sucrose, As stated in Section 2, several methods are reported: using
glucose, fructose, and the production of organic acids and cel- a previous batch of kombucha, only the broth can be added
lulose are faster and more intense in black tea than in green (Blanc, 1996; Jayabalan et al., 2007; Loncar et al., 2014; Mal-
tea in identical elaboration conditions. This was not observed baša et al., 2008), only the biofilm or fragments (Jayabalan
in other studies, in which no impact on the physical chemistry et al., 2010; Reiss, 1994; Sievers et al., 1995), or both (Chen
could be observed (Jayabalan et al., 2007) or gave the opposite & Liu, 2000; Goh et al., 2012a; Kallel et al., 2012)
effect (Coton et al., 2017). On the contrary, at the microbiolog-
ical level, it was reported that the nature of tea did not impact 4.4.1 Nature of the inoculum
the dynamics of yeasts but the use of green tea allowed the Several studies have compared elaboration kinetics with dif-
development of Oenococcus oeni that was absent in the black ferent kombucha inocula. Chu and Chen (2006) observed dif-
tea modality (Figure 5 [Coton et al., 2017]). In the same study, ferent kinetics of antioxidant capacities and total phenolic
a higher bacterial biodiversity could be observed in green tea content increases over eight differently sourced kombucha
than in black tea, in which domination phenomenon is more cultures used as inocula.
12 THE PARAMETERS OF KOMBUCHA’S QUALITY…

F I G U R E 5 Relative abundance of bacterial species in green and black tea Kombucha elaboration based on 16S rDNA metabarcoding. The 28
most abundant OTUs (Operational Taxonomic Units) out of total 354 OTUs are presented on this figure (>1% relative abundance). Sampling was
performed at days 0, 2, 4, and 8 on both tea (two inner circles) and biofilm (two outer circles) samples. The two circles represent replicate samples for
each sample type (Coton et al., 2017).

The elaboration kinetics of a kombucha inoculum and two 4.4.2 Amount of inoculum used
controlled inocula prepared from kombucha isolates (a mix The addition of 15% inoculum (unknown chemical and micro-
of acetic acid bacteria and one yeast: Zygosaccharomyces sp. biological composition “from previous batch”) compared to
and Saccharomyces cerevisiae, respectively) were compared 10% demonstrated a consumption of sucrose, glucose, fruc-
(Malbaša, Lončar, Vitas, & Čanadanović-Brunet, 2011). Sim- tose and acidification faster at a given temperature (22 ◦ C or
ilar pH variations in black tea and green tea matrices were 30 ◦ C) on the first 10 days of elaboration (Figure 6 [Lončar
observed but the production of C and B vitamins showed sig- et al., 2006]). This suggests that increasing the quantity of
nificant differences with the original consortium producing inoculum accelerates the elaboration kinetics (Lončar et al.,
always the highest amount of vitamin B. The highest amount 2006).
of vitamin C was produced by the original consortium and the
controlled consortium including S. cerevisiae. Another study
using cocultures from isolated microorganisms (Gluconobac-
4.5 The temperature
ter intermedius coinoculated with Dekkera bruxellensis) was In the study of Lončar et al. (2006, 2014), temperature was
carried out in order to optimize the production of health ben- determined to be a parameter more impactful on elaboration
eficial glucuronic acid by playing with the relative proportion kinetics compared to the amount of added inoculum. When
of each strain with significant results (Nguyen et al., 2014). 22 ◦ C and 30 ◦ C elaboration temperatures were compared,
Currently, no link has been established between the presence the consumption of sugars and the acidification appeared to
of particular genera or species and general chemical profiles be faster at 30 ◦ C for a given quantity of added inoculum
of kombucha. (10% and 15%) on the first 10 days. Conclusion is that higher
THE PARAMETERS OF KOMBUCHA’S QUALITY… 13

F I G U R E 6 Sugar quantities as functions of time, temperature, and inoculum concentration (Lončar et al., 2006)

TABLE 3 Changes in total count of acetic acid bacteria, yeast, lactic acid bacteria, and pH values in Kombucha beverages during 10 days of
elaboration at 20, 25, and 30 ◦ C (adapted from Neffe-Skocińska et al. [2017])

Microbial species Temperature of Days of elaboration


(log CFU/mL)/pH elaboration (◦ C) 0 3 7 10
AAB 20 3.57 ± 0.2aA 4.60 ± 0.3aB 6.72 ± 0.1aC 7.39 ± 0.1bD
25 3.93 ± 0.1aA 4.90 ± 0.2aB 6.90 ± 0.2aC 7.61 ± 0.2bD
30 3.65 ± 0.2aA 5.15 ± 0.1aB 7.10 ± 0.2aC 6.77 ± 0.2aC
Yeast 20 4.02 ± 0.1 aA
5.86 ± 0.2 bB
7.00 ± 0.1 bC
7.83 ± 0.2cD
25 4.24 ± 0.1aA 7.00 ± 0.2cB 7.22 ± 0.1bB 7.43 ± 0.1bB
30 4.01 ± 0.2aA 4.73 ± 0.1aB 5.48 ± 0.1aC 36.40 ± 0.2aD
pH 20 3.08 ± 0.1aA 2.85 ± 0.1aA 2.88 ± 0.1aA 2.67 ± 0.1aA
25 3.07 ± 0.1 aB
2.80 ± 0.1 aA
2.79 ± 0.1 aA
2.77 ± 0.2aA
30 3.04 ± 0.1aB 2.81 ± 0.1aA 2.71 ± 0.1aA 2.63 ± 0.2aA
Data are expressed as mean ± standard deviation of n = 3 samples.
Means in the same column followed by different lowercase letters represent significant differences (P < 0.05).
Means in the same row followed by different uppercase letters represent significant differences (P < 0.05).
14 THE PARAMETERS OF KOMBUCHA’S QUALITY…

F I G U R E 7 Changes in pH value and total acidity of kombucha produced in flasks (a), cylinders (b), small reactors (c), and large reactors (d)
(adapted from Cvetković et al. [2008])

temperature also accelerates the elaboration kinetics. This rivorans population also enhanced the production of gluconic
is contradictory with the results of Neffe-Skocińska et al. and glucuronic acids.
(2017) who evaluated an optimal temperature of 25 ◦ C
because it could favor the yeasts’ activity whose metabolites 4.6 The geometry of the vessel
(monosaccharides and ethanol) are the only available sub-
strates for acetic acid bacteria at the beginning of the process According to Malbaša et al. (2006), the geometry of the elab-
(Table 3). Thus, the growth and metabolic activity of yeasts oration tank or vessel was a parameter more impactful than
in sucrose-based kombucha is the limiting step of the pro- the amount of added inoculum. In the same study, the param-
cess. Yet, no significant difference in sensory profile could be eter of geometric similarity for a cylindrical vessel was high-
determined between the kombucha produced at 20, 25, and lighted and defined as
30◦ C.
Another study (De Filippis et al., 2018) reported that a tem- ( )1∕3
𝐷𝑇 2 𝑉𝑇 2
perature of 30 ◦ C compared to 20 ◦ C increased the popula- =
𝐷𝑇 1 𝑉𝑇 1
tion of acetic acid bacteria. Each modality operated a selec-
tion of bacteria species: Gluconobacter saccharivorans was
in majority at 30 ◦ C to the detriment of G. xylinus, which was DTx is defined as the diameter of the cylinder x and VTx as
dominant at 20 ◦ C. At 30 ◦ C, the enhancement of G. saccha- the volume of the cylinder x. The elaboration kinetics seems to
THE PARAMETERS OF KOMBUCHA’S QUALITY… 15

be characterized by a relationship between the vessel’s diam- sumer (Monaco, 2019), it would need to take the following
eter and volume. elements into account:
Another parameter of the same type was investigated: the
specific interfacial surface (SIS) defined as - The rawness of the product, namely the preservation of
potential beneficial properties;
( −1 ) - The stability of the product over time;
Specific Interfacial Surface 𝑐𝑚
( ) - The accessibility of the product by offering a pleasant sen-
Surface of liquid 𝑐𝑚2 sory profile from appearance to taste.
= ( )
Volume of liquid 𝑐𝑚3
Therefore, decisions need to be taken about the microbial
and chemical stabilization of the product and the produc-
The accuracy of this model was tested with different tion process needs to be adapted accordingly. Consequently,
geometries on 56 vessels from 90 L reactors to 330 mL flasks interdependent parameters during the production process will
and could be validated (Cvetković, Markov, Djurić, Savić, impact the consumer perception of the final product. The
& Velićanski, 2008). This model was judged to be more knowledge of such complex systems is still lacking as pro-
efficient than the geometric similarity because the study ducers and scientists face many grey areas.
showed a higher fidelity of elaboration kinetics and durations The process of kombucha production was mainly studied
to the SIS mathematic model (Figure 7) (Cvetković et al., by the determination of the microbial composition and the
2008; Malbaša et al., 2006). The study shows that regard- understanding of the main technological roles attributed to
less of the shape of the vessel (flask, cylinder, or reactor) yeasts and bacteria through the consumption of carbohydrates
and the size, a larger SIS induced faster acidification kinet- and the production of organic acids. The impact of environ-
ics. The increase of SIS can be achieved by using a larger mental factors, such as temperature, the substrate content,
air/liquid interface and/or by reducing the volume of liquid. and the vessel geometry, was investigated. A characterization
Consequence results in better conditions for oxygen access of kombucha consortia composition involving a large num-
for acetic acid bacteria located in the biofilm or the liquid ber of samples at worldwide scale would help define kom-
phase. A smart use of the SIS parameter can help controlling bucha better and highlight signature genera, species, and co-
the speed of the acidification phase of kombucha elaboration. occurrences between microorganisms. The existence of an
As a conclusion, ways to increase the speed of the acidifica- effect of the geographical sourcing on the microbial composi-
tion phase of kombucha elaboration is to increase the amount tion could even lead to the concept of kombucha typicity. As
of inoculum (ranging between 10% and 15%), increase the a matter of fact, few studies have investigated the microbial
temperature up to around 30 ◦ C, and to maximize the SIS. dynamic and tried to explain the reasons why certain genera
The water composition, tea type, and the choice of substrates or species were dominant in given conditions during elabora-
interdependently influence composition of the initial matrix. tion (Coton et al., 2017; De Filippis et al., 2018; Teoh et al.,
Thus, these parameters need to be assessed on a case-by-case- 2004). Moreover, the intraspecific diversity was not investi-
basis depending on the targeted result. Eventually, the influ- gated to our knowledge. Indeed, the questions of the origin
ence of sugared tea liquor and microbial composition and the of kombucha consortia and their stability or evolution over
interaction of these both elements on the final product remain time, namely in relationship to the matrix composition (tea,
shrouded in mystery and there is a need for research to be car- substrate) and the process conditions, still remain unan-
ried out on these problematics. swered. Nevertheless, the available knowledge can already
allow the orientations of the microbial activity and the kinet-
ics of elaboration. Further control or prediction on kombucha
5 CONC LU S I O N A N D elaboration remains difficult when it comes to the organolep-
PE RS PECTIV E S tic profile.
Surprisingly, little data are available about the sensory of
Kombucha can be approached the same way as a carbonated kombucha per se and namely the compounds involved in its
soft drink: emphasis is put on its visual aspect, the aroma pro- aroma profile. To our knowledge, no research was carried
file, and the taste, in particular the sweetness/sourness bal- out to draw lines between the odorant volatile molecules and
ance. Yet, due to the infinity of combination of microbial the microbial composition. As a matter of fact, beside the
compositions, the multiple processes used at home and in the main metabolites such as organic acids, ethanol and the sub-
industry and the lack of specific quality standards in regu- stances of interest such as vitamins, gluconic, glucuronic, or
lation, the characterization of kombucha’s quality dimension D-saccharic-1,4-lactone acids, little is known of the metabolic
remains a challenge (Watson, 2019). If the authenticity of activity of the consortium. Without a doubt, microbial inter-
kombucha had to be defined by the expectations of the con- actions are occurring during the elaboration (Teoh et al.,
16 THE PARAMETERS OF KOMBUCHA’S QUALITY…

2004), first through the symbiosis between yeasts and acetic Science and Nutrition, 37(8), 693–704. https://doi.org/10.
acid bacteria and possibly through nutrient competition or/and 1080/10408399709527797
targeted chemical signals such as peptides or messenger Barker, G. S., Jefferson, B., & Judd, S. J. (2002). The control of bubble
size in carbonated beverages. Chemical Engineering Science, 57(4),
RNAs (Ivey, Massel, & Phister, 2013; Leroi & Pidoux, 1993;
565–573. https://doi.org/10.1016/S0009-2509(01)00391-8
Sieuwerts, Bron, & Smid, 2018; C. Wang, Mas, & Esteve-
Battikh, H., Bakhrouf, A., & Ammar, E. (2012). Antimicrobial effect of
Zarzoso, 2016). The utilization of “omics” techniques such as Kombucha analogues. LWT - Food Science and Technology, 47(1),
metabolomics or transcriptomics, which are nontargeted anal- 71–77. https://doi.org/10.1016/j.lwt.2011.12.033
yses of metabolites and gene expression, respectively, could Bauer-Petrovska, B., & Petrushevska-Tozi, L. (2000). Mineral and water
open doors in the study of microbial interactions in products soluble vitamin content in the Kombucha drink. International Jour-
as it is the case for wine (Liu et al., 2016). Kombucha is seen as nal of Food Science & Technology, 35(2), 201–205.
a promising model system for the study of microbial interac- Bhattacharya, S., Gachhui, R., & Sil, P. C. (2013). Effect of Kombucha,
a fermented black tea in attenuating oxidative stress mediated tissue
tion in symbiotic systems (May et al., 2019). The elucidation
damage in alloxan induced diabetic rats. Food and Chemical Toxicol-
of such interactions could give way to better control of elabo- ogy, 60, 328–340. https://doi.org/10.1016/j.fct.2013.07.051
ration processes in terms of timing and repeatability, and thus Blanc, P. J. (1996). Characterization of the tea fungus metabo-
spark significant interest in the dynamic kombucha commu- lites. Biotechnology Letters, 18(2), 139–142. https://doi.org/10.1007/
nity of producers and consumers. BF00128667
Bonnans, S., & Noble, A. C. (1993). Effect of sweetener type and
ACKNOW LEDGMENTS of sweetener and acid levels on temporal perception of sweet-
ness, sourness and fruitiness. Chemical Senses, 18(3), 273–283.
This work was supported by the Regional Council of Bour- https://doi.org/10.1093/chemse/18.3.273
gogne – Franche Comté, the “Fond Européen de DEveloppe- Capel, F., Nicolai, T., Durand, D., Boulenguer, P., & Langen-
ment Régional (FEDER). Authors declare no conflict of inter- dorff, V. (2006). Calcium and acid induced gelation of (ami-
est. We thank Jordi Ballester for his precious feedbacks on the dated) low methoxyl pectin. Food Hydrocolloids, 20(6), 901–907.
sensory aspects of the review. https://doi.org/10.1016/j.foodhyd.2005.09.004
Cardoso, R. R., Neto, R. O., dos Santos D’Almeida, C. T., do Nasci-
mento, T. P., Pressete, C. G., Azevedo, L., … de Barros, F. A. R.
AU THOR CONT RI B UT I O NS (2020). Kombuchas from green and black teas have different phe-
nolic profile, which impacts their antioxidant capacities, antibacte-
Thierry Tran took the lead of the writing of this review but all rial and antiproliferative activities. Food Research International, 128,
other authors provided critical and complementary elements 108782. https://doi.org/10.1016/j.foodres.2019.108782
to the manuscript. Chakravorty, S., Bhattacharya, S., Chatzinotas, A., Chakraborty,
W., Bhattacharya, D., & Gachhui, R. (2016). Kombucha tea
fermentation: Microbial and biochemical dynamics. Interna-
ORC ID tional Journal of Food Microbiology, 220, 63–72. https://doi.org/
Thierry Tran https://orcid.org/0000-0002-5666-3829 10.1016/j.ijfoodmicro.2015.12.015
Charters, S., & Pettigrew, S. (2007). The dimensions of wine qual-
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