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https://doi.org/10.17113/ftb.61.01.23.7811

Uloga bakterija octenog vrenja u proizvodnji hrane i pića

Natália Norika Yassunaka Hata orcid id orcid.org/0000-0003-2319-1156 ; Departamento de Ciência e Tecnologia de Alimentos/Centro de Ciências Agrárias/Universidade Estadual de Londrina - UEL, Rodovia Celso Garcia Cid, km 380, 86057-970, Londrina, PR, Brazil
Monica Surek orcid id orcid.org/0000-0003-4048-3046 ; Departamento de Análises Clínicas/Universidade Federal do Paraná – UFPR, Campus III – Sede Botânico, 80210-170, Curitiba, PR, Brazil
Daniele Sartori orcid id orcid.org/0000-0002-0465-9932 ; Departamento de Bioquímica e Biotecnologia/Centro de Ciências Exatas/Universidade Estadual de Londrina - UEL, Rodovia Celso Garcia Cid, km 380, 86057-970, Londrina, PR, Brazil
Rodrigo Vassoler Serrato orcid id orcid.org/0000-0001-8868-6825 ; Departamento de Bioquímica e Biologia Molecular/Universidade Federal do Paraná – UFPR, Centro Politécnico, 81531-990, Curitiba, PR, Brazil
Wilma Aparecida Spinosa orcid id orcid.org/0000-0001-9532-0135 ; Departamento de Ciência e Tecnologia de Alimentos/Centro de Ciências Agrárias/Universidade Estadual de Londrina - UEL, Rodovia Celso Garcia Cid, km 380, 86057-970, Londrina, PR, Brazil


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Sažetak

Bakterije octenog vrenja su široko rasprostranjeni mikroorganizmi u prirodi. Iako uzrokuju kvarenje hrane, od velikog su industrijskog značaja, no njihova funkcionalnost još uvijek nije dovoljno istražena. U procesu octeno-kiselog vrenja pretvaraju etanol, šećere i poliole u različite organske kiseline, aldehide i ketone. Ti metaboliti nastaju nakon niza biokemijskih reakcija u različitim fermentiranim proizvodima i pićima, kao što su ocat, kombucha, vodeni kefir, lambic pivo i kakao. Osim toga, metabolizam bakterija octenog vrenja može se iskoristiti za dobivanje proizvoda od industrijskog značaja, kao što su prekursori glukonske i askorbinske kiseline. Razvoj novih voćnih napitaka sa zdravim i funkcionalnim svojstvima dobivenih octeno-kiselim vrenjem predstavlja zanimljiv segment razvoja prehrambene industrije, jer takvi proizvodi mogu zadovoljiti potrebe velikog broja potrošača. Egzopolisaharidi koje proizvode bakterije octenog vrenja, poput levana i bakterijske celuloze, imaju jedinstvena svojstva, ali potrebno ih je proizvesti u većoj mjeri da bi se mogla proširiti njihova primjena u proizvodnji hrane i pića. U ovom revijalnom prikazu su istaknuti značaj i primjena bakterija octenog vrenja u proizvodnji raznovrsne hrane, njihova uloga u razvoju novih napitaka, te mogućnosti primjene levana i bakterijske celuloze.

Ključne riječi

octena kiselina; hrana; napitak; octeno-kiselo vrenje

Hrčak ID:

301283

URI

https://hrcak.srce.hr/301283

Datum izdavanja:

25.4.2023.

Podaci na drugim jezicima: engleski

Posjeta: 1.734 *




INTRODUCTION

Acetic acid bacteria (AAB) are mesophilic, Gram-negative bacteria that belong to the Acetobacteraceae family. They can be single, in pairs or chains and have an ellipsoidal to elongated shape (rods). Their width varies from 0.4 to 1.0 µm, and their length ranges from 0.8 to 4.5 µm. AAB do not sporulate (1).

They show positive catalase and negative oxidase reactions, and have a strictly aerobic metabolism with oxygen as the terminal electron acceptor (1). According to Laureys et al. (2), AAB grow well between pH=5.0 and 6.5, but can also grow at pH=3.0-4.0 and even lower. The optimum temperature for growth is between 25 and 30 °C, and typically, no growth occurs above 34 °C (2,3). However, thermotolerant strains can continue to grow at 37 °C, and some strains can even grow at temperatures as high as 42 °C (4,5).

To date, twenty genera have been described in the family Acetobacteraceae, among which the ones with the highest number of species are Acetobacter, Gluconobacter, Asaia, Komagataeibacter and Gluconacetobacter (6,7). The group can oxidize various types of sugars, sugar alcohols and alcohols to their respective aldehydes, ketones and corresponding organic acids through an incomplete oxidation process called oxidative fermentation, from which they obtain their energy (3). Acetobacter and Komagataeibacter spp., for example, are specialized in converting ethanol to acetic acid via two successive oxidative steps and are thus common in alcoholic and acidic environments, such as the vinegar industry (8,9). They also have a complete set of citric acid cycle (CAC) enzymes, which are required for further oxidation of organic acids to CO2 and H2O (3). In contrast, Gluconobacter spp. occur preferentially in sugar-rich niches and are particularly proficient in the oxidation of sugars and sugar alcohols (10,11). Due to a lack of CAC enzymes, Gluconobacter species are unable to oxidize acetate to CO2 and H2O, but they are useful in the biotechnological synthesis of precursor compounds of vitamin C (l-sorbose), gluconic acid, and its derivatives, dihydroxyacetone and miglitol (12,13).

The bacterium Gluconacetobacter diazotropicus is the most well-known member of the genus Gluconacetobacter, which plays an important role as nitrogen-fixing bacteria in plants. In addition, G. diazotropicus produces indole-3-acetic acid and gibberellins A1 and A3, which are important hormones that control plant growth (14,15).

The genus Asaia, on the other hand, has been linked to beverage spoilage and has recently been found to be a symbiotic microorganism in malaria-carrying mosquitos (16). In addition, the role of AAB in the production of exopolysaccharides (EPS) is highlighted, the most valuable being bacterial cellulose and levan, produced mainly by the species of Komagataeibacter, Kozakia, Gluconacetobacter, Neoasaia and Gluconobacter (3,8). AAB are widely distributed in alcoholic, sugar-rich and acidic environments (14). They are often seen only as spoilage agents in wine, where acidity is undesirable, but their role in the production of fermented foods such as vinegar, kombucha, water kefir, lambic and cocoa, also in the bioconversion of specific products, such as ascorbic acid, as well as the applicability and functionality of levan and bacterial cellulose (Fig. 1) is still quite limited. Therefore, the objective of this review is to demonstrate the many advantages of AAB in this area, also encouraging their research in other application areas.

Fig. 1 Acetic acid bacteria are involved in the production of various foods, beverages, chemicals and exopolysaccharides: a) vinegar types from different raw materials, b) kombucha, c) water kefir, d) lambic, e) cocoa, f) organic acids (gluconic and ascorbic), g) new fruit drinks, and h) exopolysaccharides (bacterial cellulose and levan). Parts of the figure designed by iStock
FTB-61-85-f1

AAB IN FOOD AND BEVERAGE FERMENTATIONS

Vinegar

Vinegar production has been around for over 10 000 years (3). Despite not being considered a ’food’ and not having a high nutritional value, vinegar is consumed by people of all social classes all over the world, and it differs in terms of the used raw materials, manufacturing technologies and its wide range of applications (Fig. 1a) (3,17).

The definition and standards of vinegar identity and quality have some local differences, but in general, food regulatory agencies consider vinegar to be the result of a double fermentation (first alcoholic, then acetic) of sugar-rich substrates (18). In Brazil, the MAPA (Ministry of Agriculture, Livestock and Supply) defines a product as fermented by acetic acid if the minimum volatile acidity is 4% (g/100 mL, expressed in acetic acid) obtained from acetic fermentation of fruit, cereal, other vegetables, honey, vegetable mixture or hydroalcoholic mixture already fermented in alcoholic fermentation. The acetic acid fermented product can be called a vinegar of plus the name of the used substrate (19).

The substrate used in the processing of vinegar is mostly of plant origin, including fruits (e.g. grapes, apple, mango, etc.), cereals, onion and cider, except for honey vinegar and whey vinegar (18,20). The chemical composition of the raw material has a strong influence on the selection of microorganisms and determines the dominant species involved in the acetification process (21).Table 1 shows the main AAB species involved in vinegar production (1,7,22-60).

Table 1 Acetic acid bacteria (AAB) associated with fermented foods and beverages and with the synthesis of chemical compounds and polysaccharides
Food/Beverage /Chemical compound/PolysaccharideAAB species associated with the fermentationReference
Vinegar
Fruit vinegar (apple cider, orange, red and white wine, persimmon, white and red grape, apricot and blueberry)Acetobacter pasteurianus, A. aceti, A. estunensis, A. pomorum, A. syzygii, Komagataeibacter europaeus, Novacetimonas hansenii (formerly K. hansenii), K. kakiaceti, K. oboediens, K. melaceti, K. melomenusus, N. pomaceti (formerly K. pomaceti), K. rhaeticus, K. saccharivorans, Gluconacetobacter entanii, N. maltaceti (formerly K. maltaceti), K. nataicola, K. intermedius, K. xylinus and Gluconobacter oxydans1,7,22-27
Cereal vinegar (rice grain and rice hull, wheat bran and glutinous rice)K. europaeus, K. kakiaceti and K. medellinensis27-29
Cheese whey vinegarA. aceti and A. pasteurianus30,31
Kombucha
A. papayae, A. indonesiensis, A. lovaniensis, A. okinawensis, A. peroxydans, A. syzgii, A. tropicalis, K. takamatsuzukensis, K. oboediens, K. eurapaeus, K. saccharivorans, K. intermedius, K. xylinus, K. rhaeticus, Novacetimonas hansenii (formerly K. hansenii), Gluconacetobacter liquefaciens, G. entanii, Gluconobacter cerinus, G. oxydans and Tanticharoemia sakaeratensis32–37
Water kefir
A. indonesiensis, A. fabarum, A. orientalis, A. tropicalis, A. okinawensis, A. lovaniensis, A. lovaniensis, K. intermedius, K. saccharivorans, Novacetimonas hansenii (formerly K. hansenii), G. cerinus, G. japonicus and G. liquefaciens38-40
Lambic
A. orientalis, G. cerevisiae, G. wancherniae, A. pasteurianus, A. aceti, A. lovaniensis, A. lambici and A. pomorum41–43
Cocoa
A. pasteurianus, A. syzygii, A. tropicalis, A. ghanensis, A. indonesiensis, A. okinawensis, Novacetimonas hansenii (formerly K. hansenii), G. oxydans, G. frateurii, G. diazotrophicus and Granulibacter bethesdensis44-48
Gluconic acid, dihydroxyacetone, vitamin C precursors, miglitol
G. oxydans8,12,49,50
New beverages from AAB
Acetobacter sp. and G. japonicus51-53
Levan
G. albidus, G. cerinus, G, oxydans, G. frateurii, Kozakia baliensis, Neoasaia chiangmaiensis, Tanticharoenia sakaeratensis, Novacetimonas hansenii (formerly K. hansenii), K. xylinus, A. pasteurianus and G. diazotrophicus54-58
Bacterial cellulose
Novacetimonas hansenii (formerly K. hansenii), K. nataicola, K. rhaeticus, K. swingsii, K. maltaceti and K. xylinus24,59,60

Generally, vinegar is made through two fermentation processes: alcoholic fermentation and acetic acid fermentation (61). Under anaerobic conditions, yeasts (typically strains of Saccharomyces cerevisiae) convert fermentable sugars to ethanol, whereas in acetic acid fermentation, AAB convert ethanol to acetic acid by the activity of two membrane-bound enzymes located on the outer surface of the cytoplasmic membrane (periplasmic side) (3,18). First, alcohol dehydrogenase (ADH) oxidizes ethanol to acetaldehyde, which is then oxidized to acetic acid by aldehyde dehydrogenase (ALDH) (61). On the other hand, some rice and cereal vinegar made from starchy raw materials include a distinct saccharification step prior to alcoholic fermentation (62).

Three main methods are used in industrial vinegar production: (i) the slow, traditional Orleans or French (surface acetification carried out in wooden barrels), (ii) the fast generator (production under forced aeration with wood chips or other inert material), and (iii) the rapid submerged (modern or industrial; batch acetification with forced aeration and agitation) (18,63). In the production of traditional vinegar, the acetification is typically made using a culture from a previous batch known as ’seed-vinegar’ or ’mother of vinegar’, whereas selected cultures are added to ensure large-scale production, higher yield, better safety, process stability, shorter fermentation time and product losses, as well as avoiding undesirable characteristics caused by uncontrolled fermentation (26,64). The use of specific AAB starters, however, is still far from being a common practice (26).

The microbiota that leads to vinegar production is complex and includes several genera of AAB; however, Acetobacter and Komagataeibacter species have a strong capacity to produce acetic acid, and both genera also exhibit high resistance to high ethanol and acetic acid concentrations, which are essential characteristics required for industrial vinegar production (3,65). Thermotolerant AAB have also been introduced for the production of a variety of valuable products, including vinegar (50). They are advantageous for industrial vinegar fermentation because they allow for stable fermentation with lower cooling costs, particularly in tropical countries (66,67).

Vinegar has a wide range of applications. Different types of vinegar are frequently used as preservatives, flavouring agents, and as ingredients in mayonnaise, salad dressings, mustard and other condiments (17,20,68). Its use as a routine medicine for humans and animals dates back to remote antiquity; in addition, it can be used as a cleaning agent and in some countries even as a healthy drink (3,17).

Although vinegar is traditionally used as a flavouring and food preservative, recent scientific studies have reported that regular consumption of vinegar can promote beneficial physiological health effects (3,65,69). Among the therapeutic properties of vinegar are its antibacterial activity, regulation of blood pressure and glycaemia, antioxidant activity, prevention of cardiovascular diseases, and prevention of obesity (8,70). In addition to acetic acid, some types of vinegar contain several bioactive compounds, such as polyphenols, that contribute to their taste, smell, and specific functions. Considering that different types of vinegar can be produced using different raw materials, processes, and species of AAB, understanding the relationship between the compounds present and the functionality of vinegar is of great importance (20,70).

Kombucha

Kombucha is a popular drink usually consumed in Asia (35) (Fig. 1b). It is a nonalcoholic refreshing beverage, with accentuated acidity and a specific flavour (71). Traditionally, it is made by fermenting sweetened tea, but other plant (e.g. cereals or leaves) or animal (e.g. milk) raw materials, as well as mushrooms, can also be used (72). Fermentation lasts approx. 7 to 10 days and occurs quickly after adding the cellulosic layer called tea fungus or SCOBY (symbiotic colony of bacteria and yeast) to the sweetened tea (73). The microbial composition of kombucha varies significantly from batch to batch depending on its origin, substrate and fermentation conditions (74). It predominantly comprises AAB, which include Komagataeibacter and Gluconobacter, while Acetobacter species are less abundant (71). Komagataeibacter xylinus is considered one of the most important species involved in the fermentation of kombucha due to its superior capacity for cellulose synthesis (37).Table 1 shows the main AAB species involved in kombucha fermentation (3237).

In addition to AAB, a wide range of yeast species can be found in the kombucha, including species of the genera Zygosaccharomyces, Candida, Kloeckera/Hanseniaspora, Torulaspora, Pichia, Brettanomyces/Dekkera, Saccharomyces, Lachancea, Saccharomycoides, Schizosaccharomyces and Kluyveromyces (34,75). Lactic acid bacteria (LAB), such as Lactobacillus, Lactococcus and Bifidobacterium species, can occur (75); however, they do not seem to be a crucial part of the kombucha microbial ecosystem since they are not always present (2).

In this symbiotic relationship, the yeast community hydrolyses the sucrose present in the medium to glucose and fructose, and produces ethanol from glucose. AAB use glucose, fructose and ethanol to produce gluconic acid, glucuronic acid, acetic acid, d-saccharic acid-1,4-lactone and bacterial cellulose (37,71). The produced acetic acid further stimulates the production of ethanol by the yeasts, and this ethanol is then converted to acetic acid by AAB. The continuous accumulation of ethanol and acetic acid in the medium prevents contamination by various pathogenic microorganisms (73).

The composition of kombucha also includes vitamins, phenolic compounds, amino acids and some hydrolytic enzymes (35,75). The chemical profile of kombucha may be responsible for its multiple health benefits when associated with regular consumption of the drink (35). Recently, tea has attracted the attention of researchers and consumers due to its in vitro biological activities, such as antimicrobial, antioxidant and anti-inflammatory activities, and anticarcinogenic potential. However, more clinical investigations and in vivo evaluations should be carried out to confirm the health benefits of the beverage (75,76).

Water kefir

Water kefir is a sparkling, refreshing, low-alcohol drink with acidic and fruity flavours (71) (Fig. 1c). It is obtained by the spontaneous fermentation of sugar solution, dried fruits, and water kefir grains (insoluble dextran). These translucent grains, whose shape resembles that of a cauliflower, contain microorganisms that act as an inoculum for the fermentation process (77).

The microbial composition of the grains consists of yeasts, LAB and AAB, with the bacterial community presenting a higher diversity. Some of the main microorganisms of water kefir are LAB, such as Lentilactobacillus hilgardii, Liquorilactobacillus nagelii, Lacticaseibacillus paracasei and Bifidobacterium aquikefiri, and yeasts such as Saccharomyces cerevisiae and Dekkera bruxellensis (39,40). During the first 24 h of fermentation, yeasts (mainly S. cerevisiae) convert sucrose to alcohol. Furthermore, yeasts hydrolyze sucrose by the action of invertase, promoting an increase in glucose and fructose, which are then metabolized by LAB and AAB (3,39). In the advanced stages, ethanol concentrations decrease due to oxidation of ethanol to acetic acid by AAB (e.g. Gluconobacter, Komagataeibacter and Acetobacter) (40).Table 1 shows the main AAB species involved in water kefir fermentation (3840).

At the end of fermentation, the main products are ethanol, lactic acid, acetic acid and other metabolites, such as mannitol, glycerol, esters, aldehydes and other organic acids (78,79). Kefir drinks, such as kombucha, have been linked to numerous health benefits. The beverage is well known for having potentially ’probiotic’ and antimicrobial properties against a wide range of pathogenic bacteria (78,80). Furthermore, studies have shown that water kefir has antihyperlipidemic properties (81), antioxidant (39,82) and anticarcinogenic activities, hepatoprotective, anti-inflammatory and gastroprotective effects, among others (79). Due to the numerous positive effects of kefir, several substrates have been investigated for the adaptation of its grains (39,8284). This has enabled the emergence of new functional drinks with characteristics similar to those of the traditional brown sugar kefir (78).

Lambic

Lambic, originally from Belgium, is probably one of the oldest beer styles brewed to date (85,86) (Fig. 1d). It is a refreshing, alcoholic, acidic beer with fruity notes and only a few residual carbohydrates. The spontaneous fermentation occurs in the presence of water, barley malt, unmalted wheat and aged dry hops, and maturation takes up to three years in wooden barrels (71).

An in-depth examination of the dominance of microbial profile of lambic fermentation during three years revealed four distinct phases (21).Table 1 shows the main AAB species involved in lambic fermentation (4143). The first phase (from the start up to 1 month of fermentation) begins with the members of the Enterobacteriaceae family, which are inhibited by ethanol accumulation produced by wild (oxidative) yeast, acidification by enterobacteria and AAB (A. orientalis) and glucose reduction by microbial growth in general (71,87). Gluconobacter species such as G. cerevisiae have also been isolated during this phase, probably as a result of the combination of a monosaccharide-rich environment and low ethanol concentrations (42).

The second phase of ethanol fermentation, referred to as the most important, extends until the fourth month, with yeasts (Saccharomyces cerevisiae, S. bayanus/pastorianus and S. uvarum) being the main representatives for the conversion of carbohydrates into ethanol and carbon dioxide (88). After 4 to 10 months of fermentation, the acidification phase (third phase) takes place. This phase is characterized by the predominance of the LAB and AAB species, which together produce lactic and acetic acids, resulting in a pH drop below 3.5 (41,87). During this fermentation stage, the most frequently isolated LAB species are homofermentative Pediococcus damnosus and heterofermentative Lactobacillus brevis, whereas the most frequently reported AAB species are A. pasteurianus and A. lambici. LAB species produce both lactic and acetic acid from saccharides, while AAB species oxidize ethanol to acetic acid and produce acetoin from the lactic acid produced by LAB species (41,89).

After the acidification phase, the final or maturation phase begins and can last for several years (89). LAB, AAB and primarily Brettanomyces yeast species are present during this phase. The LAB and AAB species that proliferate during this phase are typically the same species that are present in the previous phase (41). B. bruxellensis and other species belonging to the genus Brettanomyces play a key role in the final flavour formation of lambic (90) since they synthesize precursor compounds responsible for the characteristic Brett flavour (volatile phenolic compounds 4-vinylguaiacol and 4-vinylphenol) of lambic and several ethyl esters, such as ethyl acetate and ethyl lactate. Together with Brettanomyces, AAB may also participate in ethyl acetate formation (43).

AAB are abundant during major periods of the first fermentation year of traditional lambic production, producing much larger concentrations of acetic acid and acetoin (from in the liquid/air interphase of the casks). The formation of acetic acid by AAB and the subsequent formation of ethyl acetate are desired compounds for complex lambic; however, excessive AAB development must be controlled to avoid an unfavourable flavour profile (42,87).

Cocoa

Cocoa bean (Theobroma cacao L.) is the main raw material for the manufacture of chocolate (91) (Fig. 1e). Taken from freshly harvested cocoa pods, raw cocoa beans are subjected to a complex fermentation that involves physical and biochemical transformations in all bean structures and lasts 3–7 days depending on various factors including the seed genetic origin, agro-ecological conditions and the used method (92,93). This process is primarily regulated by yeast, LAB, AAB and Bacillaceae (particularly Bacillus) that use the cocoa bean pulp as a growing substrate. Several molecules are released during microbial fermentation of cocoa, giving chocolate its distinctive aromatic profile, reducing bitterness and astringency, and finally killing the embryo to prevent its germination (94).

During the fermentation of cocoa beans, three main stages can be identified (94). Yeasts are the most prevalent microorganisms in the first 24 h of fermentation, converting pulp sugars to ethanol and carbon dioxide via alcoholic fermentation (anaerobiosis) (45). They are involved in the production of pectinolytic enzymes, which degrade pectin and allow oxygen to enter the cocoa pulp (93). Furthermore, yeasts produce a large number of aroma compound precursors that significantly contribute to the development of the chocolate aroma profile (47). Alternatively, simultaneously with yeast, fructophilic LAB species use fructose as an energy source, with or without citric acid conversion, and heterofermentative LAB species grow by converting glucose to lactic acid, acetic acid, ethanol, carbon dioxide and/or mannitol (95).

The second stage is distinguished by an increase in the lactic acid concentrations as a result of an increase in the LAB populations and a reduction in yeasts (94). Lactic acid produced in seeds is important to activate endogenous enzymes and contribute to the generation of chocolate flavour and aroma (45).

During the third phase, the high ethanol concentrations (metabolized by yeast) and oxygen ingress because of pulp liquefaction provide conditions for the growth of AAB (44).Table 1 shows the main AAB species involved in cocoa bean fermentation (4448). Ethanol, which is the main energy source, is converted into acetic acid by AAB, particularly Acetobacter species, while lactic acid produced by LAB serves as the primary carbon source (95). Lactic acid is primarily oxidized into acetoin and, to a lesser extent, acetic acid due to low pyruvate decarboxylase activity in Acetobacter (95). The entry of acetic acid and ethanol, along with the temperature increase (approx. 50 °C), causes the death of the seed embryo and induces a series of endogenous reactions that produce flavour, aroma and colour precursors of the chocolate raw material (47,96). As a result, the counts of yeast, LAB and AAB decrease, favouring the growth of Bacillus spores in the later stages of cocoa bean fermentation (94,95).

Among the different genera of AAB, Acetobacter is the most common during cocoa fermentation (45). It includes A. ghanensis and A. senegalensis, which predominate mainly at the beginning of fermentation, whereas A. pasteurianus and occasionally Acetobacter lovaniensis, Acetobacter syzygii or Acetobacter tropicalis prevail during the last phase, when the concentration of ethanol is high (45,95). A. pasteurianus prevails during the fermentation of cocoa due to its ability to oxidize ethanol, lactic acid and mannitol, as well as its tolerance to acidity and heat (95). On the other hand, Gluconobacter species can be prevalent at the beginning of the fermentation due to their preference for sugar metabolism. However, their growth is undesirable, as this can result in the production of gluconic acid from glucose and off-flavours, impacting the final quality of cocoa beans (45).

OTHER METABOLITES PRODUCED BY AAB

AAB can participate as biocatalysts in the industrial manufacturing of a wide range of compounds, in addition to being employed commercially in the manufacturing of vinegar and other fermented foods (8,14) (Fig. 1f). Gluconobacter strains, in particular G. oxydans, can carry out oxidative fermentation of sugars and sugar alcohols, resulting in the formation of l-sorbose, ketogluconic acids, dihydroxyacetone (DHA) and cyclic ketones, among other compounds (21,50). The oxidative fermentation of l-sorbose from d-sorbitol is the most classic example observed during the production of vitamin C by Gluconobacter. Other precursor intermediates, such as 2-keto-d-gluconic acid (2KGA) from gluconic acid, 2,5-diketo-d-gluconic acid (25DKGA) and 5-keto-d-gluconic acid (5KGA), are also present in the synthesis route (14,21). The 5KGA has potential applications in the synthesis of tartaric and xylaric acids, in addition to being a precursor for the manufacture of aromatic compounds such as 4-hydroxy-5-methyl-2,3-dihydrofuranone-3, a valuable product used in the food industry (97). The microbial production of DHA from glycerol has been explored in the pharmaceutical industry, and it can be used as a tanning agent and as an intermediate for the synthesis of various chemicals and surfactants (14,98). Gluconobacter species can also be applied in the biotransformation of miglitol precursors, a drug used for the treatment of type II diabetes; in the production of gluconic acid, considered a multifunctional acid in the food, feed, beverage, textile, pharmaceutical and construction industries; and in the manufacture of shikimic acid, a key intermediate for numerous antibiotics (8,12,49,50) (Table 1).

DEVELOPMENT OF NEW PRODUCTS FROM AAB

Currently, consumers have shown a growing interest in foods that, in addition to satisfying their hunger, can also prevent nutrition-related diseases and improve mental health (99). According to the Food and Agriculture Organization of the United Nations (FAO), fruits constitute an important part of a healthy diet. In addition to being a source of dietary fibre, vitamins, minerals and beneficial phytochemicals, fruits may help lower the risk factors for diseases such as overweight/obesity, chronic inflammation, high blood pressure and high cholesterol (100).

Fruit-based fermentation has improved the nutritional and functional quality of beverages. In addition, rising consumer demand for lactose-free products with low fat content and few additives makes this type of fermentation a promising tool for meeting the needs of obese people with cardiovascular diseases, allergies, intolerances, vegans and vegetarians (101103).

The development of fruit drinks containing probiotic bacteria stands out among current research (104106). These microorganisms improve mineral bioavailability, digestibility and organoleptic properties such as colour, flavour and aroma, in addition to providing a functional beverage (103). However, the acidic environment, as well as the presence of anti-nutritional and inhibitory factors in fruits, make maintaining bacterial viability and stability during processing and storage a major challenge (102). Given this, the fermentation of fruit drinks with AAB becomes a viable alternative, since they can oxidize a wide range of substrates and are typically found in sugar-rich and highly acidic environments (Table 1).

Fruit vinegar drinks, for instance, are gaining popularity in North America (Fig. 1g). By definition, the product must be made from at least one type of fruit and contain at least 300 g of fruit juice for each litre of fermented product. These beverages have been categorized according to the volume fraction of acetic acid as drinks with low acidity (φ<3%) and high acidity (φ=5-7%) (107). Furthermore, the used fermentation method and the volume fraction of acetic acid may affect the content of total sugars and soluble solids, titratable acidity and density (107). Regarding its benefits, in vivo animal studies have shown that tomato vinegar drinks can prevent visceral obesity and insulin resistance (51), while pomegranate vinegar drinks have been shown to reduce visceral adipose tissue in humans (108). Other research suggests that fruit vinegar drinks such as cranberry, blueberry and tomato could be used to treat hypertension and hypercholesterolaemia (107). However, according to Chang et al. (109), continuous consumption of vinegar drinks should be avoided to prevent gastrointestinal injuries.

Another recent approach has been focused on gluconic acid fermentation. Although works related to this type of fermentation are still scarce, their results are very promising. The gluconic acid fermentation of strawberry drink by G. japonicus converts glucose into gluconic acid while keeping the fructose naturally present in the fruit as a sweetener. This allows diabetics to consume a drink that keeps phenolic compounds (nonanthocyanins) and antioxidant activity practically unchanged (110). Furthermore, its composition remains stable for 15 days at room temperature (27−30 °C) and up to 30 days when refrigerated (4 °C) (53). In another study, Hornedo-Ortega et al. (111) compared the antioxidant activity and anthocyanin composition of alcoholic and gluconic acid-fermented strawberry drinks. The authors concluded that the gluconic acid fermentation of strawberry beverages by G. japonicus is a novel process that preserves the anthocyanin composition and shows higher antioxidant activity values than alcoholic fermentation (111). Ordóñez et al. (112) also confirmed the safety of strawberry drink fermented with gluconic acidby demonstrating that none of the eight studied biogenic amines was detected. In addition to the bioactive compounds found in fruits, gluconic acid and its derivatives have been shown to have prebiotic properties. This acid promotes the growth of Lactobacillus sp. and Bifidobacterium adolescentis in the human colon and alters the metabolic profile of the intestines (113). Gluconic acid and its derivatives are approved for use in food and are commonly used to preserve and/or improve the sensory properties of dairy products and soft drinks (113). The presence of higher proportions of gluconic acid in kombucha, for example, improved the taste of the drink in a study conducted by Li et al. (114). Gluconic acid contributes to the pleasantly sour taste, while the formed acetic acid produces an acidic and astringent off-flavour (114).

EXOPOLYSACCHARIDES PRODUCED BY AAB

Microbial polysaccharides are produced by a wide range of bacteria, presenting extreme diversity in terms of chemical structure and composition (115). AAB, for example, can produce large amounts of EPS, including both homopolysaccharides such as levan and bacterial cellulose and heteropolysaccharides such as acetan or xylinan and gluconacetan (116118). Due to the importance of levan and bacterial cellulose (Fig. 1h) in research and industrial applications, some of their characteristics and main applications in the food industry are discussed in this paper.

Levan

Levan is a polymer composed of d-fructofuranosyl residues joined by β-(2,6) bonds in the main chain and β-(2,1) bonds in the side chain, in addition to having a d-glucopyranosyl residue at the end of the main chain (119).Fig. 2 shows the structure of the levan. Levan is commonly biosynthesized by a restricted number of plants in the nature, but it can also be produced by several microorganisms, including Archaea, fungi and a wide range of bacteria (58,120). Among the AAB, species of the genera Acetobacter, Gluconobacter, Gluconacetobacter, Kozakia, Komagataeibacter, Tanticharoenia and Neoasaia are also capable of producing it (5458) (Table 1).

Fig. 2 Chemical structure of levan
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Levan is synthesized and polimerized in the extracellular matrix by the action of the enzyme levansucrase, whose main function is to transfer fructose residues from sucrose by transfructosylation reactions (121). The enzyme has high specificity for sucrose and lower activity for fructose, mannose, raffinose, mannitol, etc. On the other hand, inhibition is observed in the presence of glucose and other sugars that have a configuration similar to glucose, such as lactose, galactose and maltose, as well as other sugar alcohols (121,122).

In general, the structure of levans produced by different organisms is similar but differs in terms of molecular mass, degrees of polymerization (DP) and branching (122,123). Plant levans have a lower molecular mass and DP than bacterial levans. Plant levans have a molecular mass of 2000 to 33000 Da and a DP<100, whereas bacterial levans have multiple branches (2 to 12%) and a molecular mass of 2 to 100 million Da with DP>100 (121,123,124).

In comparison with polysaccharides formed by pyranose rings, the structural characteristic of levan, in the form of furanose rings, plays an important role in the conformation of molecules in a solution, providing additional flexibility. Furthermore, the semiflexible chain of the rings interacts intramolecularly and intermolecularly, resulting in a densely packed spherical structure and aqueous solutions of low viscosity (room temperature) at concentrations where other polysaccharides would form pastes or gels (121,122,124126).

Xu et al. (127) observed that aqueous solutions of levan (Brenneria sp. EniD312) exhibit Newtonian fluid behaviour at low content (3%; m/V) and non-Newtonian fluid (pseudoplastic fluid) behaviour at high contents (6, 9 and 12%; m/V) when studying the rheological properties of levan. Levan solutions derived from Zymomonas mobilis and Erwinia herbicola showed similar results (128). At amounts between 1 and 8%, however, the behaviour of Bacillus subtilis levan solutions was completely Newtonian (128). According to Xu et al. (127), levan could be a good additive in the food industry, since its non-Newtonian behaviour is interesting for the manufacture of dairy products, syrups and salad dressings.

Levan solutions also exhibit an atypical behaviour when compared to other polysaccharides, in which gel formation is not observed (129,130). However, Jakob et al. (131), when establishing the structure/function relationship of isolated AAB levans, suggested that in a solution, increasing their molecular mass reinforces intramolecular interactions to achieve a more compact structure characteristic of a ’microgel’ with hydrocolloid properties. The authors also emphasise that levans produced by AAB may thus offer new possibilities for applications in food.

Unlike many other polymers, levan does not swell in water, but it has a high solubility in hot water, a variable solubility in cold water and is insoluble in most organic solvents, with the exception of dimethyl sulfoxide (DMSO). The high solubility of levan in water is mainly attributed to its β-(2,6) bond rather than to the β-(2,1) bond, and the ramifications could only be a supporting factor (121,122,126). Levans are nonreducing agents that are not hydrolyzed by yeast invertases or amylases, but they are susceptible to acid hydrolysis (122). They decompose at approx. 225 °C, and their glass transition temperature is 141 °C (126). Another important property of levan is its adhesive strength. Although sugars are characterized by stickiness, the adhesive strength of levan is significantly higher than that of other natural polymers. The branches contribute to its cohesive strength, and the ability to form adhesive bonds with a wide range of substrates is given to its large number of hydroxyl groups. Levan is commonly referred to as a ’green’ adhesive because it is water-removable and has high-value applications in several areas (8,121,125).

In the food area, several studies have explored the effects of levan-type fructooligosaccharides (L-FOS) and levan as prebiotics on probiotic bacteria and the complex gut microbiota; however, there is no conclusive evidence from human trials (132136) (Fig. 3). In animal models, for instance, levan supplementation increased Lactobacillus and Bifidobacteria viability, while inhibiting Escherichia coli and Clostridium perfringens (136,137). Adamberg et al. (132), using the human faecal microbiota, reported that levan alters the composition of the faecal microbiota and the profile of metabolites, making it a potential candidate for prebiotics. Using metagenomic sequencing to assess the prebiotic activity of levan, Cheng et al. (138) also verified alterations in the intestinal microbiota of mice and the stimulation of the production of short-chain fatty acids.

Fig. 3 Levan applications in food area. Parts of the figure designed by Freepik
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In bakery, EPS are known to improve the rheological properties of dough and the texture, nutritional value, shelf life and machinability of wheat, rye and gluten-free bread (56). Jakob et al. (139) evaluated the functional effects of different AAB levans (G. frateurii TMW 2767, G. cerinus DSM 9533, N. chiangmaiensis NBRC 101099 and K. baliensis DSM 14400) added to wheat-based bread. The addition of w(EPS)=1 or 2% to the flour resulted in an increase in the volume, a noticeable softening of the fresh bread and a delay in the hardening of the bread after a week of storage. Although LAB and yeasts are common microorganisms in the sourdough (54), Hermann et al. (56) reported that AAB, such as N. chiangmaiensis NBRC 101099 and K. baliensis DSM 14400, can grow on a variety of flour types (wheat, whole wheat, spelled and rye) and produce large amounts of levan. Later, gluten-free bread types were made with buckwheat and molasses dough fermented by G. albidus TMW 2.1191 and K. baliensis NBRC 16680, and their volume, crumb hardness and sensory characteristics were evaluated (54). Bread made from the sourdough had better sensory and quality characteristics, such as higher specific volume and lower crumb hardness. However, the authors pointed out that strong acidification during fermentation could become a challenge in a large-scale production (54).

Other advantages of applying levan in foods include its use as a fat substitute. Fructans have fat-like properties that improve the flavour and spreadability of dairy products. Furthermore, high-molecular-mass levans are rarely detected by taste sensors, and odour detection is almost imperceptible due to their low volatility (120). Their low viscosity and high solubility make them an excellent substitute for gum arabic, as the latter also has excellent stabilizing and emulsifying properties for food applications (122). In the food packaging industry, levan as a component in edible starch films increases their barrier and mechanical properties in addition to being a cost-effective alternative (119). Gan et al. (140) also developed levan/pullulan/chitosan edible films enriched with ε-polylysine and applied them to strawberry. As a result, they demonstrated that films could help preserve the postharvest strawberry quality by minimizing water loss, inhibiting microbial development and decreasing the respiration rate during storage.

Although this study is not focused on biomedical or cosmetic applications of levan, this EPS has shown several bioactive properties, including antitumour, antimicrobial, anti-inflammatory, hypocholesterolaemic, antidiabetic and immunostimulating activities (120,121). As a result, in addition to being a useful EPS for food production, its consumption alone or in food can provide several health benefits to the host.

Bacterial cellulose

Bacterial cellulose is a linear glucan composed of several glucose monomers linked by β-(1–4) bonds (8) (Fig. 4). This biopolymer can be synthesized by various microorganisms, such as algae and fungi, as well as by various bacteria, including Achromobacter, Alcaligenes, Aerobacter, Agrobacterium, Azotobacter, Gluconacetobacter, Pseudomonas, Rhizobium, Sarcina, Dickeya and Rhodobacter (141,142). Among these bacteria, Komagataeibacter species (AAB) (Table 1) (24,59,60) are frequently utilized in research and commercial production, and are employed as model strains because of their high productivity and ability to metabolize a wide range of carbon/nitrogen sources (60).

Fig. 4 Chemical structure of bacterial cellulose
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When cultivated under controlled conditions, Komagataeibacter produces highly porous bacterial cellulose structures in the form of pellicules (static culture) or as fibrous suspensions, pellets, spheres or irregular masses (agitated culture) (142,143). The synthesis of bacterial cellulose from glucose involves several individual enzymes, catalytic complexes and regulatory proteins. Briefly, β-glucan chains are formed first, followed by the assembly and crystallization of cellulose chains. In this final stage, the cellulose chains are released from the cell and self-assemble into fibrils (144). When compared to plant cellulose, bacterial celulose has a great number of unique physicochemical and mechanical properties, including higher crystallinity, degree of polymerization, water absorbing and holding capacity, tensile strength and biological adaptability (60). Moreover, plant-derived cellulose usually consists of hemicellulose and lignin, necessitating harsh chemical treatments to remove these impurities (145). Bacterial cellulose generated by microbial fermentation, on the other hand, has a greater purity and requires less energy and chemical processing for purification (145). As a result, bacterial cellulose has been used in a variety of food-related applications (Fig. 5) since it is a dietary fibre that has been approved as generally recognized as safe (GRAS) food by the US Food and Drug Administration (FDA) (146).

Fig. 5 Different applications of bacterial cellulose (BC) in the food industry. Parts of the figure designed by Freepik and iStock
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The ’nata de coco’, a traditional food consumed in the Philippines and other Southeast Asian countries, is one of the most well-known industrial applications of bacterial cellulose. In its manufacturing process, bacterial cellulose is synthesized by fermenting coconut water and then cleaning, washing, chopping and immersing it in sugar syrup to serve as a dessert (142,145,147). A variety of nata-like products have been developed to meet consumer demands. To change the colour and flavour of the dessert, different fruit juices, syrups and other ingredients have been employed. Other items containing nata de coco, such as fruit-flavoured pudding, drinks and jellies, have been marketed all over the world.

In other food systems, bacterial celulose has also shown promise as a stabilizer, noncaloric bulking agent and texture modifier. After heat sterilization, the use of an aqueous suspension of bacterial cellulose in liquids such as chocolate drinks prevents cocoa precipitation and stabilizes the dispersion (148). In pasty condiments, bacterial cellulose reduces the stickiness and controls the syneresis during storage (148). Furthermore, it promotes firmness in solid foods such as tofu, while it changes the texture of kamaboko by increasing hardness and fraturability (148). The addition of bacterial cellulose to meat products such as hamburger and sausage can also reduce fat content without compromising tenderness and juiciness, as well as produce stable emulsions (148).

Following the example of earlier products that were lower in calories, Guo et al. (149) demonstrated that adding bacterial cellulose/soy protein isolate blends to ice cream as a cream substitute can result in ice creams with low calories, melt resistance and good texture properties. Surimi products (150), cheese (151), meatballs (152), pork Frankfurters (153) and mayonnaise (154) are among other applications of bacterial cellulose as a fat replacer. These findings suggest that bacterial cellulose could be widely used as a food additive in processed foods to improve their quality and shelf life while also lowering the calories in the final products.

Bacterial cellulose has also been used as a vegetarian meat preparation when combined with Monascus extract obtained from a naturally red pigmented mould (155). The product has a natural meat flavour and is resistant to colour and morphological changes. Furthermore, because of its nonanimal origin, this ingredient may be a suitable substitute for animal-based products for some dietary restrictions (60,142).

It has attracted interest in research related to the immobilization of enzymes, cells and probiotics for use in food. Because of its superior characteristics compared to plant cellulose, bacterial cellulose provides stability to enzymes against temperature and pH variations (156). When compared to free laccase, laccase immobilized on magnetically modified bacterial cellulose showed superior thermal stability at 70 °C and maintained 65% of its initial activity after 8 cycles of use (157). Similarly, Chen et al. (158) obtained a retention of 69% of the original activity after seven recyclings when immobilizing fungal laccase on natural bacterial cellulose.

The bacterial cellulose has also been explored as a carrier for cell immobilization, primarily for yeasts in the winemaking process. This approach reduces inoculum preparation costs, as the yeast can be recovered and separated at the end of the fermentation process (145). BC has been shown to protect wine yeast from adverse conditions such as high osmotic pressure and low pH (159). As a result, the growth of immobilized yeast was better than that of free yeast (159). Furthermore, the metabolic activities of immobilized yeast in BC were reported to be higher than those of free yeast (160). Immobilized yeast in BC was also shown to have no negative impact on the sensory quality of the final product during repeated batch fermentation (161) and can increase the amount of alcohol produced compared to free cells (162).

Studies have proven that bacterial cellulose is an effective matrix for the immobilization of probiotic bacteria (163). In this context, Fijałkowski et al. (164) showed that as an immobilization support it improves probiotic viability, protecting against adverse conditions of the gastrointestinal tract (164). Furthermore, the authors established that the immobilization efficiency depends on the cellulose form, its synthesis and immobilization methods (164). Similarly, Phromthep and Leenanon (165) demonstrated that the bacterial cellulose produced from fruit juice residues and coconut milk improved the survival of immobilized Lactobacillus plantarum compared to free cells. Under prolonged incubation, Żywicka et al. (166) used the bacterial cellulose pellicle as a support for immobilization during prolonged incubation and reported that the cell viability of L. delbrueckii was affected after 72 h. On the other hand, the viability of L. acidophilus 016 immobilized in bacterial cellulose nanofibres was found to be 71% for up to 24 days when stored at ambient temperature (35 °C) (163). These findings show a potential because one of the requirements for a microorganism to be administered for therapeutic purposes is that it remains viable in the food that will be consumed (164).

More recently, bacterial cellulose has been reported as a matrix for probiotic films (167,168). The films can be used as coatings or wrapped over a variety of foods, providing consumer health benefits, as well as potentially inhibiting the growth of spoilage bacteria and fungi on food surfaces, thus extending the shelf life of the product (167,168). Similarly, the development of bacterial cellulose-based films and probiotic-derived bioactive metabolites (so-called postbiotics) has also gained attention for antimicrobial food packaging (169,170). For meat applications, the rapid release of postbiotics from the bacterial cellulose-based films into food is ideal for food with a finite shelf life, as it can effectively control foodborne pathogens such as Listeria monocytogenes while also extending the shelf life without affecting the sensorial attributes of the meat (169,170). In addition, several studies have performed ex situ and in situ modifications of bacterial cellulose to improve its properties for use in food packaging. However, to assess its potential usefulness as active packaging, more research is needed to investigate the mechanical properties, permeability, interactions and release rates in semisolid and solid food model media (171).

Other recent applications of bacterial cellulose include its use in dough leavening and baking trials to improve the rheological and sensory properties of gluten-free bakery products (59) and as a food-grade emulsion stabilizer (172174), whose function can be extended to cosmetics and medical emulsions. In addition, bacterial cellulose can serve as an excellent matrix for the development of pH-sensitive indicators containing anthocyanins from different fruits, vegetables and flowers. Anthocyanins can display different colour spectra under acidic and basic conditions. Since the colour spectrum of each indicator has a direct link to pH changes in the food product, these pH-sensitive indicators can be used to track pH changes and monitor the freshness and spoilage of foods (fish, fruits and shrimp) and beverages (175-179).

CONCLUSIONS

Acetic acid bacteria (AAB) are well known for causing wine spoilage. However, their importance and functionality in food applications were demonstrated in this review. Through oxidative fermentation, AAB can produce a variety of metabolites. Their role in various biochemical processes during food fermentation, such as vinegar, kombucha, water kefir, cocoa and lambic, results in unique sensory and biochemical characteristics, as well as health benefits. Although the commercial production of vinegar-based drinks is well established, research on fruit drinks fermented by AAB is still relatively new and scarce. Beverage production from fruits through gluconic acid fermentation is very promising since several fruits can be used (including nonstandard fruits, for example) and can be more adaptable to AAB metabolism than to the growth of LAB. Furthermore, due to the formation of gluconic acid and the presence of phenolic compounds in the fruits, the functional drink could meet the demand of lactose-intolerant consumers, those allergic to milk proteins, and those seeking a vegetarian, vegan and healthy diet. Due to its bioactive properties, levan could also be explored in beverage development. Levan could be produced in situ during the gluconic acid fermentation of fruit juices since several species of Gluconobacter can produce levan from sucrose and oxidize glucose to gluconic acid. Bacterial cellulose has numerous applications in food, with several products already marketed worldwide. However, similar to levan, its main challenge lies in the large-scale production and in reducing production costs to expand its applications in this area.

ACKNOWLEDGEMENTS

This work was supported by the National Council of Technological and Scientific Development (CNPq) (142379/2017-4).

Notes

[1] Conflicts of interest CONFLICT OF INTEREST

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

REFERENCES

1 

Sengun IY, Kilic G, Charoenyingcharoen P, Yukphan P, Yamada Y. Investigation of the microbiota associated with traditionally produced fruit vinegars with focus on acetic acid bacteria and lactic acid bacteria. Food Biosci. 2022;47:101636. https://doi.org/10.1016/j.fbio.2022.101636

2 

Laureys D, Britton SJ, De Clippeleer J. Kombucha tea fermentation: a review. J Am Soc Brew Chem. 2020;78(3):165–74. https://doi.org/10.1080/03610470.2020.1734150

3 

Lynch KM, Zannini E, Wilkinson S, Daenen L, Arendt EK. Physiology of acetic acid bacteria and their role in vinegar and fermented beverages. Compr Rev Food Sci Food Saf. 2019;18(3):587–625. https://doi.org/10.1111/1541-4337.12440 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/33336918

4 

Es-Sbata I, Lakhlifi T, Yatim M, El-Abid H, Belhaj A, Hafidi M, et al. Screening and molecular characterization of new thermo- and ethanol-tolerant Acetobacter malorum strains isolated from two biomes Moroccan cactus fruits. Biotechnol Appl Biochem. 2021;68(3):476–85. https://doi.org/10.1002/bab.1941 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/32410247

5 

Matsushita K, Azuma Y, Kosaka T, Yakushi T, Hoshida H, Akada R, et al. Genomic analyses of thermotolerant microorganisms used for high-temperature fermentations. Biosci Biotechnol Biochem. 2016;80(4):655–68. https://doi.org/10.1080/09168451.2015.1104235 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/26566045

6 

Qiu X, Zhang Y, Hong H. Classification of acetic acid bacteria and their acid resistant mechanism. AMB Express. 2021;11:29. https://doi.org/10.1186/s13568-021-01189-6 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/33595734

7 

Brandão PR, Crespo MTB, Nascimento FX. Phylogenomic and comparative analyses support the reclassification of several Komagataeibacter species as novel members of the Novacetimonas gen. nov. and bring new insights into the evolution of cellulose synthase genes. Int J Syst Evol Microbiol. 2022;72(2):005252. https://doi.org/10.1099/ijsem.0.005252 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/35175916

8 

Gomes RJ, Borges M de F, Rosa M de F, Castro-Gómez RJH, Spinosa WA. Acetic acid bacteria in the food industry: Systematics, characteristics and applications. Food Technol Biotechnol. 2018;56(2):139–51. https://doi.org/10.17113/ftb.56.02.18.5593 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/30228790

9 

Peng W, Meng D, Yue T, Wang Z, Gao Z. Effect of the apple cultivar on cloudy apple juice fermented by a mixture of Lactobacillus acidophilus, Lactobacillus plantarum, and Lactobacillus fermentum. Food Chem. 2021;340:127922. https://doi.org/10.1016/j.foodchem.2020.127922 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/32889211

10 

Jakob F, Gebrande C, Bichler RM, Vogel RF. Insights into the pH-dependent, extracellular sucrose utilization and concomitant levan formation by Gluconobacter albidus TMW 2.1191. Antonie van Leeuwenhoek. 2020;113:863–73. https://doi.org/10.1007/s10482-020-01397-3 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/32130597

11 

Moghadami F, Fooladi J, Hosseini R. Introducing a thermotolerant Gluconobacter japonicus strain, potentially useful for coenzyme Q10 production. Folia Microbiol (Praha). 2019;64:471–9. https://doi.org/10.1007/s12223-018-0666-4 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/30680590

12 

da Silva GAR, Oliveira SS de S, Lima SF, do Nascimento RP, Baptista AR de S, Fiaux SB. The industrial versatility of Gluconobacter oxydans: Current applications and future perspectives. World J Microbiol Biotechnol. 2022;38:134. https://doi.org/10.1007/s11274-022-03310-8 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/35688964

13 

La China S, Zanichelli G, De Vero L, Gullo M. Oxidative fermentations and exopolysaccharides production by acetic acid bacteria: a mini review. Biotechnol Lett. 2018;40:1289–302. https://doi.org/10.1007/s10529-018-2591-7 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/29987707

14 

Mamlouk D, Gullo M. Acetic acid bacteria: physiology and carbon sources oxidation. Indian J Microbiol. 2013;53:377–84. https://doi.org/10.1007/s12088-013-0414-z PubMed: http://www.ncbi.nlm.nih.gov/pubmed/24426139

15 

Sebring RL, Duiker SW, Berghage RD, Regan JM, Lambert JD, Bryant RB. Gluconacetobacter diazotrophicus inoculation of two lettuce cultivars affects leaf and root growth under hydroponic conditions. Appl Sci (Basel). 2022;12(3):1585. https://doi.org/10.3390/app12031585

16 

Bassene H, Niang EHA, Fenollar F, Doucoure S, Faye O, Raoult D, et al. Role of plants in the transmission of Asaia sp., which potentially inhibit the Plasmodium sporogenic cycle in Anopheles mosquitoes. Sci Rep. 2020;10:7144. https://doi.org/10.1038/s41598-020-64163-5 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/32346047

17 

Spinosa WA, dos Santos Júnior V, Galvan D, Fiorio JL, Gomez RJHC. Vinegar rice (Oryza sativa L.) produced by a submerged fermentation process from alcoholic fermented rice. Food Sci Technol. 2015;35(1):196–201. https://doi.org/10.1590/1678-457X.6605

18 

Ho CW, Lazim AM, Fazry S, Zaki UKHH, Lim SJ. Varieties, production, composition and health benefits of vinegars: A review. Food Chem. 2017;221:1621–30. https://doi.org/10.1016/j.foodchem.2016.10.128 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/27979138

19 

Ministry of Agriculture, Livestock and Supply (MAPA). Instrução Normativa n.° 6 de abril de 2012. Estabelece os padrões de identidade e qualidade e a classificação dos fermentados acéticos. Diário Oficial da União República Federativa do Brasil, Brasília, DF, Brasil; 2012 (in Portuguese).

20 

Li S, Li P, Feng F, Luo LX. Microbial diversity and their roles in the vinegar fermentation process. Appl Microbiol Biotechnol. 2015;99:4997–5024. https://doi.org/10.1007/s00253-015-6659-1 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/25971198

21 

Matsushita K, Toyama H, Tonouchi N, Okamoto-Kainuma A, editors. Acetic acid bacteria. Ecology and physiology. Tokyo, Japan: Springer Japan; 2016. https://doi.org/10.1007/978-4-431-55933-7 https://doi.org/10.1007/978-4-431-55933-7

22 

Marič L, Cleenwerck I, Accetto T, Vandamme P, Trček J. Description of Komagataeibacter melaceti sp. nov. and Komagataeibacter melomenusus sp. nov. isolated from apple cider vinegar. Microorganisms. 2020;8(8):1178. https://doi.org/10.3390/microorganisms8081178 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/32756518

23 

Cepec E, Trček J. Antimicrobial resistance of acetobacter and Komagataeibacter species originating from vinegars. Int J Environ Res Public Health. 2022;19(1):463. https://doi.org/10.3390/ijerph19010463 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/35010733

24 

Greser AB, Avcioglu NH. Optimization and physicochemical characterization of bacterial cellulose by Komagataeibacter nataicola and Komagataeibacter maltaceti strains isolated from grape, thorn apple and apple vinegars. Arch Microbiol. 2022;204:465. https://doi.org/10.1007/s00203-022-03083-6 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/35802199

25 

Buyukduman E, Kirtil HE, Metin B. Molecular Identification and technological properties of acetic acid bacteria isolated from malatya apricot and home-made fruit vinegars. Microbiol Biotechnol Lett. 2022;50(1):81–8. https://doi.org/10.48022/mbl.2109.09017

26 

Milanović V, Osimani A, Garofalo C, De Filippis F, Ercolini D, Cardinali F, et al. Profiling white wine seed vinegar bacterial diversity through viable counting, metagenomic sequencing and PCR-DGGE. Int J Food Microbiol. 2018;286:66–74. https://doi.org/10.1016/j.ijfoodmicro.2018.07.022 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/30048915

27 

Kim DH, Chon JW, Kim H, Seo KH. Development of a novel selective medium for the isolation and enumeration of acetic acid bacteria from various foods. Food Control. 2019;106:106717. https://doi.org/10.1016/j.foodcont.2019.106717

28 

Peng MY, Zhang XJ, Huang T, Zhong XZ, Chai LJ, Lu ZM, et al. Komagataeibacter europaeus improves community stability and function in solid-state cereal vinegar fermentation ecosystem: Non-abundant species plays important role. Food Res Int. 2021;150(Part B):110815. https://doi.org/10.1016/j.foodres.2021.110815 https://doi.org/10.1016/j.foodres.2021.110815

29 

Jiang Y, Lv X, Zhang C, Zheng Y, Zheng B, Duan X, et al. Microbial dynamics and flavor formation during the traditional brewing of Monascus vinegar. Food Res Int. 2019;125:108531. https://doi.org/10.1016/j.foodres.2019.108531 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/31554138

30 

Parrondo J, Herrero M, García LA, Díaz MA. Note - Production of vinegar from whey. J Inst Brew. 2003;109(4):356–8. https://doi.org/10.1002/j.2050-0416.2003.tb00610.x

31 

Lustrato G, Salimei E, Alfano G, Belli C, Fantuz F, Grazia L, et al. Cheese whey recycling in traditional dairy food chain: Effects of vinegar from whey in dairy cow nutrition. Acetic Acid Bact. 2013;2 s1:e8. https://doi.org/10.4081/aab.2013.s1.e8

32 

Santana de Carvalho D, Trovatti Uetanabaro AP, Kato RB, Aburjaile FF, Jaiswal AK, Profeta R, et al. The space-exposed kombucha microbial community member Komagataeibacter oboediens showed only minor changes in its genome after reactivation on earth. Front Microbiol. 2022;13:782175. https://doi.org/10.3389/fmicb.2022.782175 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/35369445

33 

Tran T, Grandvalet C, Winckler P, Verdier F, Martin A, Alexandre H, et al. Shedding light on the formation and structure of kombucha biofilm using two-photon fluorescence microscopy. Front Microbiol. 2021;12:725379. https://doi.org/10.3389/fmicb.2021.725379 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/34421883

34 

Coton M, Pawtowski A, Taminiau B, Burgaud G, Deniel F, Coulloumme-Labarthe L, et al. Unraveling microbial ecology of industrial-scale Kombucha fermentations by metabarcoding and culture-based methods. FEMS Microbiol Ecol. 2017;93(5):fix048. https://doi.org/10.1093/femsec/fix048 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/28430940

35 

De Filippis F, Troise AD, Vitaglione P, Ercolini D. Different temperatures select distinctive acetic acid bacteria species and promotes organic acids production during Kombucha tea fermentation. Food Microbiol. 2018;73:11–6. https://doi.org/10.1016/j.fm.2018.01.008 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/29526195

36 

Gaggìa F, Baffoni L, Galiano M, Nielsen DS, Jakobsen RR, Castro-Mejía JL, et al. Kombucha beverage from green, black and rooibos teas: A comparative study looking at microbiology, chemistry and antioxidant activity. Nutrients. 2019;11(1):1. https://doi.org/10.3390/nu11010001 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/30577416

37 

La China S, De Vero L, Anguluri K, Brugnoli M, Mamlouk D, Gullo M. Kombucha tea as a reservoir of cellulose producing bacteria: Assessing diversity among Komagataeibacter isolates. Appl Sci (Basel). 2021;11(4):1595. https://doi.org/10.3390/app11041595

38 

Patel S, Tan J, Zhang SJ, Priour S, Lima A, Ngom-bru C, et al. A temporal view of the water kefir microbiota and flavour attributes. 2022;80:103084. https://doi.org/10.1016/j.ifset.2022.103084 https://doi.org/10.1016/j.ifset.2022.103084

39 

Bueno RS, Ressutte JB, Hata NNY, Henrique-Bana FC, Guergoletto KB, de Oliveira AG, et al. Quality and shelf life assessment of a new beverage produced from water kefir grains and red pitaya. Lebensm Wiss Technol. 2021;140:110770. https://doi.org/10.1016/j.lwt.2020.110770

40 

Pendón MD, Bengoa AA, Iraporda C, Medrano M, Garrote GL, Abraham AG. Water kefir: Factors affecting grain growth and health-promoting properties of the fermented beverage. J Appl Microbiol. 2022;133(1):162–80. https://doi.org/10.1111/jam.15385 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/34822204

41 

Bongaerts D, De Roos J, De Vuysta L. Technological and environmental features determine the uniqueness of the lambic beer microbiota and production process. Appl Environ Microbiol. 2021;87(18):e00612-21. https://doi.org/10.1128/AEM.00612-21 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/34232060

42 

De Roos J, Verce M, Aerts M, Vandamme P, De Vuyst L. Temporal and spatial distribution of the acetic acid bacterium communities throughout the wooden casks used for the fermentation and maturation of lambic beer underlines their functional role. Appl Environ Microbiol. 2018;84(7):e02846–17. https://doi.org/10.1128/AEM.02846-17 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/29352086

43 

De Roos J, Verce M, Weckx S, De Vuyst L. Temporal shotgun metagenomics revealed the potential metabolic capabilities of specific microorganisms during lambic beer production. Front Microbiol. 2020;11:1692. https://doi.org/10.3389/fmicb.2020.01692 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/32765478

44 

Verce M, Schoonejans J, Hernandez Aguirre C, Molina-Bravo R, De Vuyst L, Weckx S. A Combined metagenomics and metatranscriptomics approach to unravel Costa Rican cocoa box fermentation processes reveals yet unreported microbial species and functionalities. Front Microbiol. 2021;12:641185. https://doi.org/10.3389/fmicb.2021.641185 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/33664725

45 

Viesser JA, de Melo Pereira GV, de Carvalho Neto DP, Favero GR, de Carvalho JC, Goés-Neto A, et al. Global cocoa fermentation microbiome: revealing new taxa and microbial functions by next generation sequencing technologies. World J Microbiol Biotechnol. 2021;37:118. https://doi.org/10.1007/s11274-021-03079-2 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/34131809

46 

Serra JL, Moura FG. Pereira GV d. M, Soccol CR, Rogez H, Darnet S. Determination of the microbial community in Amazonian cocoa bean fermentation by Illumina-based metagenomic sequencing. Lebensm Wiss Technol. 2019;106:229–39. https://doi.org/10.1016/j.lwt.2019.02.038

47 

Agyirifo DS, Wamalwa M, Otwe EP, Galyuon I, Runo S, Takrama J, et al. Metagenomics analysis of cocoa bean fermentation microbiome identifying species diversity and putative functional capabilities. Heliyon. 2019;5(7):e02170. https://doi.org/10.1016/j.heliyon.2019.e02170 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/31388591

48 

Ouattara HG, Niamké SL. Mapping the functional and strain diversity of the main microbiota involved in cocoa fermentation from Cote d’Ivoire. Food Microbiol. 2021;98:103767. https://doi.org/10.1016/j.fm.2021.103767 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/33875203

49 

Gao L, Wu X, Zhu C, Jin Z, Wang W, Xia X. Metabolic engineering to improve the biomanufacturing efficiency of acetic acid bacteria: advances and prospects. Crit Rev Biotechnol. 2020;40(4):522–38. https://doi.org/10.1080/07388551.2020.1743231 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/32212873

50 

Saichana N, Matsushita K, Adachi O, Frébort I, Frebortova J. Acetic acid bacteria: A group of bacteria with versatile biotechnological applications. Biotechnol Adv. 2015;33(6):1260–71. https://doi.org/10.1016/j.biotechadv.2014.12.001 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/25485864

51 

Seo KI, Lee J, Choi RY, Lee HI, Lee JH, Jeong YK, et al. Anti-obesity and anti-insulin resistance effects of tomato vinegar beverage in diet-induced obese mice. Food Funct. 2014;5(7):1579–86. https://doi.org/10.1039/c4fo00135d PubMed: http://www.ncbi.nlm.nih.gov/pubmed/24867606

52 

Hornedo-Ortega R, Krisa S, Carmen García-Parrilla M, Richard T. Effects of gluconic and alcoholic fermentation on anthocyanin composition and antioxidant activity of beverages made from strawberry. Lebensm Wiss Technol. 2016;69:382–9. https://doi.org/10.1016/j.lwt.2016.01.070

53 

Álvarez-Fernández MA, Hornedo-Ortega R, Cerezo AB, Troncoso AM, García-Parrilla MC. Determination of nonanthocyanin phenolic compounds using high-resolution mass spectrometry (UHPLC-Orbitrap-MS/MS) and impact of storage conditions in a beverage made from strawberry by fermentation. J Agric Food Chem. 2016;64(6):1367–76. https://doi.org/10.1021/acs.jafc.5b05617 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/26803927

54 

Ua-Arak T, Jakob F, Vogel RF. Influence of levan-producing acetic acid bacteria on buckwheat-sourdough breads. Food Microbiol. 2017;65:95–104. https://doi.org/10.1016/j.fm.2017.02.002 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/28400025

55 

Aramsangtienchai P, Kongmon T, Pechroj S, Srisook K. Enhanced production and immunomodulatory activity of levan from the acetic acid bacterium, Tanticharoenia sakaeratensis. Int J Biol Macromol. 2020;163:574–81. https://doi.org/10.1016/j.ijbiomac.2020.07.001 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/32629048

56 

Hermann M, Petermeier H, Vogel RF. Development of novel sourdoughs with in situ formed exopolysaccharides from acetic acid bacteria. Eur Food Res Technol. 2015;241:185–97. https://doi.org/10.1007/s00217-015-2444-8

57 

Idogawa N, Amamoto R, Murata K, Kawai S. Phosphate enhances levan production in the endophytic bacterium Gluconacetobacter diazotrophicus Pal5. Bioengineered. 2014;5(3):173–9. https://doi.org/10.4161/bioe.28792 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/24717418

58 

Anguluri K, La China S, Brugnoli M, De Vero L, Pulvirenti A, Cassanelli S, et al. Candidate acetic acid bacteria strains for levan production. Polymers (Basel). 2022;14(10):2000. https://doi.org/10.3390/polym14102000 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/35631879

59 

Vigentini I, Fabrizio V, Dellacà F, Rossi S, Azario I, Mondin C, et al. Set-up of bacterial cellulose production from the genus Komagataeibacter and its use in a gluten-free bakery product as a case study. Front Microbiol. 2019;10:1953. https://doi.org/10.3389/fmicb.2019.01953 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/31551945

60 

Lin D, Liu Z, Shen R, Chen S, Yang X. Bacterial cellulose in food industry: Current research and future prospects. Int J Biol Macromol. 2020;158:1007–19. https://doi.org/10.1016/j.ijbiomac.2020.04.230 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/32387361

61 

Cantadori E, Brugnoli M, Centola M, Effredi E, Colonello A, Gullo M. Date Fruits as raw material for vinegar and non-alcoholic fermented beverages. Foods. 2022;11(13):1972. https://doi.org/10.3390/foods11131972 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/35804787

62 

Zhang Q, Fu C, Zhao C, Yang S, Zheng Y, Xia M, et al. Monitoring microbial succession and metabolic activity during manual and mechanical solid-state fermentation of Chinese cereal vinegar. Lebensm Wiss Technol. 2020;133:109868. https://doi.org/10.1016/j.lwt.2020.109868

63 

Al-Dalali S, Zheng F, Xu B, Abughoush M, Li L, Sun B. Processing technologies and flavor analysis of Chinese cereal vinegar: A comprehensive review. Food Anal Methods. 2023;16:1–28. https://doi.org/10.1007/s12161-022-02328-w

64 

Plioni I, Bekatorou A, Terpou A, Mallouchos A, Plessas S, Koutinas AA, et al. Vinegar production from corinthian currants finishing side-stream: Development and comparison of methods based on immobilized acetic acid bacteria. Foods. 2021;10(12):3133. https://doi.org/10.3390/foods10123133 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/34945684

65 

Xia T, Zhang B, Duan W, Zhang J, Wang M. Nutrients and bioactive components from vinegar: A fermented and functional food. J Funct Foods. 2020;64:103681. https://doi.org/10.1016/j.jff.2019.103681

66 

Matsumoto N, Osumi N, Matsutani M, Phathanathavorn T, Kataoka N, Theeragool G, et al. Thermal adaptation of acetic acid bacteria for practical high-temperature vinegar fermentation. Biosci Biotechnol Biochem. 2021;85(5):1243–51. https://doi.org/10.1093/bbb/zbab009 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/33686416

67 

Taweecheep P, Naloka K, Matsutani M, Yakushi T, Matsushita K, Theeragool G. In vitro thermal and ethanol adaptations to improve vinegar fermentation at high temperature of Komagataeibacter oboediens MSKU 3. Appl Biochem Biotechnol. 2019;189:144–59. https://doi.org/10.1007/s12010-019-03003-3 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/30957194

68 

Budak NH, Aykin E, Seydim AC, Greene AK, Guzel-Seydim ZB. Functional properties of vinegar. J Food Sci. 2014;79(5):R757–64. https://doi.org/10.1111/1750-3841.12434 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/24811350

69 

Ousaaid D, Mechchate H, Laaroussi H, Hano C, Bakour M, El Ghouizi A, et al. Fruits vinegar: Quality characteristics, phytochemistry, and functionality. Molecules. 2022;27(1):222. https://doi.org/10.3390/molecules27010222 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/35011451

70 

Samad A, Azlan A, Ismail A. Therapeutic effects of vinegar: A review. Curr Opin Food Sci. 2016;8:56–61. https://doi.org/10.1016/j.cofs.2016.03.001

71 

De Roos J, De Vuyst L. Acetic acid bacteria in fermented foods and beverages. Curr Opin Biotechnol. 2018;49:115–9. https://doi.org/10.1016/j.copbio.2017.08.007 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/28863341

72 

Morales D. Biological activities of kombucha beverages: The need of clinical evidence. Trends Food Sci Technol. 2020;105:323–33. https://doi.org/10.1016/j.tifs.2020.09.025

73 

Laavanya D, Shirkole S, Balasubramanian P. Current challenges, applications and future perspectives of SCOBY cellulose of Kombucha fermentation. J Clean Prod. 2021;295:126454. https://doi.org/10.1016/j.jclepro.2021.126454

74 

Nyhan LM, Lynch KM, Sahin AW, Arendt EK. Advances in kombucha tea fermentation: A review. Appl Microbiol. 2022;2(1):73–103. https://doi.org/10.3390/applmicrobiol2010005

75 

Villarreal-Soto SA, Beaufort S, Bouajila J, Souchard JP, Taillandier P. Understanding kombucha tea fermentation: A review. J Food Sci. 2018;83(3):580–8. https://doi.org/10.1111/1750-3841.14068 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/29508944

76 

Kapp JM, Sumner W. Kombucha: a systematic review of the empirical evidence of human health benefit. Ann Epidemiol. 2019;30:66–70. https://doi.org/10.1016/j.annepidem.2018.11.001 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/30527803

77 

Lynch KM, Wilkinson S, Daenen L, Arendt EK. An update on water kefir: Microbiology, composition and production. Int J Food Microbiol. 2021;345:109128. https://doi.org/10.1016/j.ijfoodmicro.2021.109128 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/33751986

78 

Fiorda FA, de Melo Pereira GV, Thomaz-Soccol V, Rakshit SK, Pagnoncelli MGB, Vandenberghe LP de S, et al. Microbiological, biochemical, and functional aspects of sugary kefir fermentation - A review. Food Microbiol. 2017;66:86–95. https://doi.org/10.1016/j.fm.2017.04.004 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/28576377

79 

Guzel-Seydim ZB, Gökırmaklı Ç, Greene AK. A comparison of milk kefir and water kefir: Physical, chemical, microbiological and functional properties. Trends Food Sci Technol. 2021;113:42–53. https://doi.org/10.1016/j.tifs.2021.04.041

80 

Waldherr FW, Doll VM, Meißner D, Vogel RF. Identification and characterization of a glucan-producing enzyme from Lactobacillus hilgardii TMW 1.828 involved in granule formation of water kefir. Food Microbiol. 2010;27(5):672–8. https://doi.org/10.1016/j.fm.2010.03.013 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/20510787

81 

Rocha-Gomes A, Escobar A, Soares JS, Alves A, Silva D, Riul TR. Chemical composition and hypocholesterolemic effect of milk kefir and water kefir in Wistar rats. Rev Nutr. 2018;31(2):137–45. https://doi.org/10.1590/1678-98652018000200001

82 

Ozcelik F, Akan E, Kinik O. Use of Cornelian cherry, hawthorn, red plum, roseship and pomegranate juices in the production of water kefir beverages. Food Biosci. 2021;42:101219. https://doi.org/10.1016/j.fbio.2021.101219

83 

Darvishzadeh P, Orsat V, Martinez JL. Process optimization for development of a novel water kefir drink with high antioxidant activity and potential probiotic properties from Russian olive fruit (Elaeagnus angustifolia). Food Bioprocess Technol. 2021;14:248–61. https://doi.org/10.1007/s11947-020-02563-1

84 

Destro TM, Prates D da F, Watanabe LS, Garcia S, Biz G, Spinosa WA. Organic brown sugar and jaboticaba pulp influence on water kefir fermentation. Cienc Agrotec. 2019;43:e005619. https://doi.org/10.1590/1413-7054201943005619

85 

Spitaels F, Wieme AD, Janssens M, Aerts M, Van Landschoot A, De Vuyst L, et al. The microbial diversity of an industrially produced lambic beer shares members of a traditionally produced one and reveals a core microbiota for lambic beer fermentation. Food Microbiol. 2015;49:23–32. https://doi.org/10.1016/j.fm.2015.01.008 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/25846912

86 

Thompson Witrick K, Duncan SE, Hurley KE, O’Keefe SF. Acid and volatiles of commercially available lambic beers. Beverages. 2017;3(4):51. https://doi.org/10.3390/beverages3040051

87 

De Roos J, Vandamme P, De Vuyst L. Wort substrate consumption and metabolite production during lambic beer fermentation and maturation explain the successive growth of specific bacterial and yeast species. Front Microbiol. 2018;9:2763. https://doi.org/10.3389/fmicb.2018.02763 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/30510547

88 

Spitaels F, Wieme AD, Janssens M, Aerts M, Daniel HM, Van Landschoot A, et al. The microbial diversity of traditional spontaneously fermented lambic beer. PLoS One. 2014;9(4):e95384. https://doi.org/10.1371/journal.pone.0095384 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/24748344

89 

De Roos J, De Vuyst L. Microbial acidification, alcoholization, and aroma production during spontaneous lambic beer production. J Sci Food Agric. 2019;99(1):25–38. https://doi.org/10.1002/jsfa.9291 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/30246252

90 

Serra Colomer M, Funch B, Forster J. The raise of Brettanomyces yeast species for beer production. Curr Opin Biotechnol. 2019;56:30–5. https://doi.org/10.1016/j.copbio.2018.07.009 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/30173102

91 

Ho VTT, Fleet GH, Zhao J. Unraveling the contribution of lactic acid bacteria and acetic acid bacteria to cocoa fermentation using inoculated organisms. Int J Food Microbiol. 2018;279:43–56. https://doi.org/10.1016/j.ijfoodmicro.2018.04.040 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/29727857

92 

Pacheco-Montealegre ME, Dávila-Mora LL, Botero-Rute LM, Reyes A, Caro-Quintero A. Fine resolution analysis of microbial communities provides insights into the variability of cocoa bean fermentation. Front Microbiol. 2020;11:650. https://doi.org/10.3389/fmicb.2020.00650 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/32351482

93 

Papalexandratou Z, Kaasik K, Kauffmann LV, Skorstengaard A, Bouillon G, Espensen JL, et al. Linking cocoa varietals and microbial diversity of Nicaraguan fine cocoa bean fermentations and their impact on final cocoa quality appreciation. Int J Food Microbiol. 2019;304:106–18. https://doi.org/10.1016/j.ijfoodmicro.2019.05.012 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/31176963

94 

Figueroa-Hernández C, Mota-Gutierrez J, Ferrocino I, Hernández-Estrada ZJ, González-Ríos O, Cocolin L, et al. The challenges and perspectives of the selection of starter cultures for fermented cocoa beans. Int J Food Microbiol. 2019;301:41–50. https://doi.org/10.1016/j.ijfoodmicro.2019.05.002 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/31085407

95 

De Vuyst L, Leroy F. Functional role of yeasts, lactic acid bacteria and acetic acid bacteria in cocoa fermentation processes. FEMS Microbiol Rev. 2020;44(4):432–53. https://doi.org/10.1093/femsre/fuaa014 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/32420601

96 

Almeida OGG, Pinto UM, Matos CB, Frazilio DA, Braga VF, von Zeska-Kress MR, et al. Does Quorum Sensing play a role in microbial shifts along spontaneous fermentation of cocoa beans? An in silico perspective. Food Res Int. 2020;131:109034. https://doi.org/10.1016/j.foodres.2020.109034 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/32247478

97 

Salusjärvi T, Povelainen M, Hvorslev N, Eneyskaya EV, Kulminskaya AA, Shabalin KA, et al. Cloning of a gluconate/polyol dehydrogenase gene from Gluconobacter suboxydans IFO 12528, characterization of the enzyme and its use for the production of 5-ketogluconate in a recombinant Escherichia coli strain. Appl Microbiol Biotechnol. 2004;65:306–14. https://doi.org/10.1007/s00253-004-1594-6 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/15060755

98 

Sengun IY, Karabiyikli S. Importance of acetic acid bacteria in food industry. Food Control. 2011;22(5):647–56. https://doi.org/10.1016/j.foodcont.2010.11.008

99 

Donno D, Turrini F. Plant foods and underutilized fruits as source of functional food ingredients: Chemical composition, quality traits, and biological properties. Foods. 2020;9(10):1974. https://doi.org/10.3390/foods9101474 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/33076549

100 

Fruit and vegetables – your dietary essentials. The international year of fruits and vegetables, 2021, background paper. Rome, Italy: Food and Agriculture Organization of the United Nations (FAO); 2020. Available from: https://doi.org/10.4060/cb2395en https://doi.org/10.4060/cb2395en

101 

Ayed L, M’Hir S, Hamdi M. Microbiological, Biochemical, and functional aspects of fermented vegetable and fruit beverages. J Chem. 2020;2020:5790432. https://doi.org/10.1155/2020/5790432

102 

Min M, Bunt CR, Mason SL, Hussain MA. Nondairy probiotic food products: An emerging group of functional foods. Crit Rev Food Sci Nutr. 2019;59(16):2626–41. https://doi.org/10.1080/10408398.2018.1462760 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/29630845

103 

Marrero SC, Martínez-Rodríguez A, Pérez SEM, Moya SP. New trends and applications in fermented beverages. In: Grumezescu AM, Holban AM, editors. Fermented beverages: Volume 5: The science of beverages. Duxford, UK: Elsevier Inc.; 2019. pp. 531–66. https://doi.org/10.1016/B978-0-12-815271-3.00002-6 https://doi.org/10.1016/B978-0-12-815271-3.00002-6

104 

Kaprasob R, Kerdchoechuen O, Laohakunjit N, Somboonpanyakul P. B vitamins and prebiotic fructooligosaccharides of cashew apple fermented with probiotic strains Lactobacillus spp., Leuconostoc mesenteroides and Bifidobacterium longum. Process Biochem. 2018;70:9–19. https://doi.org/10.1016/j.procbio.2018.04.009

105 

Yang X, Zhou J, Fan L, Qin Z, Chen Q, Zhao L. Antioxidant properties of a vegetable–fruit beverage fermented with two Lactobacillus plantarum strains. Food Sci Biotechnol. 2018;27(6):1719–26. https://doi.org/10.1007/s10068-018-0411-4 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/30483436

106 

Mantzourani I, Kazakos S, Terpou A, Mallouchos A, Kimbaris A, Alexopoulos A, et al. Assessment of volatile compounds evolution, antioxidant activity, and total phenolics content during cold storage of pomegranate beverage fermented by Lactobacillus paracasei K5. Fermentation (Basel). 2018;4(4):95. https://doi.org/10.3390/fermentation4040095

107 

Nandasiri R, Rupasinghe V. Inhibition of low density lipoprotein oxidation and angiotensin converting enzyme in vitro by functional fruit vinegar beverages. J Food Process Beverages. 2013;1(1):4.

108 

Park JE, Kim JY, Kim J, Kim YJ, Kim MJ, Kwon SW, et al. Pomegranate vinegar beverage reduces visceral fat accumulation in association with AMPK activation in overweight women: A double-blind, randomized, and placebo-controlled trial. J Funct Foods. 2014;8:274–81. https://doi.org/10.1016/j.jff.2014.03.028

109 

Chang J, Han SE, Paik SS, Kim YJ. Corrosive esophageal injury due to a commercial vinegar beverage in an adolescent. Clin Endosc. 2020;53(3):366–9. https://doi.org/10.5946/ce.2019.066 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/31405264

110 

Álvarez-Fernández MA, Hornedo-Ortega R, Cerezo AB, Troncoso AM, García-Parrilla MC. Effects of the strawberry (Fragaria ananassa) purée elaboration process on nonanthocyanin phenolic composition and antioxidant activity. Food Chem. 2014;164:104–12. https://doi.org/10.1016/j.foodchem.2014.04.116 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/24996312

111 

Hornedo-Ortega R, Álvarez-Fernández MA, Cerezo AB, Troncoso AM, García-Parrilla MC. Influence of storage conditions on the anthocyanin profile and color of an innovative beverage elaborated by gluconic fermentation of strawberry. J Funct Foods. 2016;23:198–209. https://doi.org/10.1016/j.jff.2016.02.014

112 

Ordóñez JL, Sainz F, Callejón RM, Troncoso AM, Torija MJ, García-Parrilla MC. Impact of gluconic fermentation of strawberry using acetic acid bacteria on amino acids and biogenic amines profile. Food Chem. 2015;178:221–8. https://doi.org/10.1016/j.foodchem.2015.01.085 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/25704705

113 

Cañete-Rodríguez AM, Santos-Dueñas IM, Jiménez-Hornero JE, Ehrenreich A, Liebl W, García-García I. Gluconic acid: Properties, production methods and applications—An excellent opportunity for agro-industrial byproducts and waste biovalorization. Process Biochem. 2016;51(12):1891–903. https://doi.org/10.1016/j.procbio.2016.08.028

114 

Li R, Xu Y, Chen J, Wang F, Zou C, Yin J. Enhancing the proportion of gluconic acid with a microbial community reconstruction method to improve the taste quality of Kombucha. Lebensm Wiss Technol. 2022;155:112937. https://doi.org/10.1016/j.lwt.2021.112937

115 

Roca C, Alves VD, Freitas F, Reis MAM. Exopolysaccharides enriched in rare sugars: Bacterial sources, production, and applications. Front Microbiol. 2015;6:288. https://doi.org/10.3389/fmicb.2015.00288 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/25914689

116 

Brandt JU, Born FL, Jakob F, Vogel RF. Environmentally triggered genomic plasticity and capsular polysaccharide formation are involved in increased ethanol and acetic acid tolerance in Kozakia baliensis NBRC 16680. BMC Microbiol. 2017;17:172. https://doi.org/10.1186/s12866-017-1070-y PubMed: http://www.ncbi.nlm.nih.gov/pubmed/28797225

117 

Kornmann H, Valentinotti S, Duboc P, Marison I, Von Stockar U. Monitoring and control of Gluconacetobacter xylinus fed-batch cultures using in situ mid-IR spectroscopy. J Biotechnol. 2004;113(1–3):231–45. https://doi.org/10.1016/j.jbiotec.2004.03.029 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/15380658

118 

Semino CE, Dankert MA. In vitro biosynthesis of acetan using electroporated Acetobacter xylinum cells as enzyme preparations. J Gen Microbiol. 1993;139(11):2745–56. https://doi.org/10.1099/00221287-139-11-2745 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/8277256

119 

Mantovan J, Bersaneti GT, Faria-Tischer PCS, Celligoi MAPC, Mali S. Use of microbial levan in edible films based on cassava starch. Food Packag Shelf Life. 2018;18:31–6. https://doi.org/10.1016/j.fpsl.2018.08.003

120 

Öner ET, Hernández L, Combie J. Review of Levan polysaccharide: From a century of past experiences to future prospects. Biotechnol Adv. 2016;34(5):827–44. https://doi.org/10.1016/j.biotechadv.2016.05.002 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/27178733

121 

Srikanth R, Reddy CHSSS, Siddartha G, Ramaiah MJ, Uppuluri KB. Review on production, characterization and applications of microbial levan. Carbohydr Polym. 2015;120:102–14. https://doi.org/10.1016/j.carbpol.2014.12.003 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/25662693

122 

Han W. Youn. Microbial Levan. In: Neidleman SL, Laskin AI, editors. Advances in applied microbiology. San Diego, CA, USA: Academic Press; 1990. pp. 171-94. https://doi.org/10.1016/S0065-2164(08)70244-2 https://doi.org/10.1016/S0065-2164(08)70244-2

123 

Mutanda T, Mokoena MP, Olaniran AO, Wilhelmi BS, Whiteley CG. Microbial enzymatic production and applications of short-chain fructooligosaccharides and inulooligosaccharides: Recent advances and current perspectives. J Ind Microbiol Biotechnol. 2014;41(6):893–906. https://doi.org/10.1007/s10295-014-1452-1 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/24793124

124 

Stojković B, Sretenovic S, Dogsa I, Poberaj I, Stopar D. Viscoelastic properties of levan-DNA mixtures important in microbial biofilm formation as determined by micro- and macrorheology. Biophys J. 2015;108(3):758–65. https://doi.org/10.1016/j.bpj.2014.10.072 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/25650942

125 

Combie J. Properties of levan and potential medical uses. In: Marchessault RH, Ravenelle F, Zhu XX, editors. Polysaccharides for drug delivery and pharmaceutical applications (ACS Symposium), vol. 934. Washington, DC, USA: ACS Symposium Series; 2006. pp. 263-9. https://doi.org/10.1021/bk-2006-0934.ch013 https://doi.org/10.1021/bk-2006-0934.ch013

126 

Manandhar S, Vidhate S, D’Souza N. Water soluble levan polysaccharide biopolymer electrospun fibers. Carbohydr Polym. 2009;78(4):794–8. https://doi.org/10.1016/j.carbpol.2009.06.023

127 

Xu W, Liu Q, Bai Y, Yu S, Zhang T, Jiang B, et al. Physicochemical properties of a high molecular weight levan from Brenneria sp. EniD312. Int J Biol Macromol. 2018;109:810–8. https://doi.org/10.1016/j.ijbiomac.2017.11.056 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/29133100

128 

Benigar E, Dogsa I, Stopar D, Jamnik A, Cigić IK, Tomšič M. Structure and dynamics of a polysaccharide matrix: Aqueous solutions of bacterial levan. Langmuir. 2014;30(14):4172–82. https://doi.org/10.1021/la500830j PubMed: http://www.ncbi.nlm.nih.gov/pubmed/24654746

129 

Benigar E, Tomšič M, Sretenovic S, Stopar D, Jamnik A, Dogsa I. Evaluating SAXS results on aqueous solutions of various bacterial Levan utilizing the string-of-beads model. Acta Chim Slov. 2015;62(3):509–17. https://doi.org/10.17344/acsi.2015.1437 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/26454583

130 

Majee SB, Avlani D, Biswas GR. Rheological behavior and pharmaceutical applications of bacterial exopolysaccharides. J Appl Pharm Sci. 2017;7(9):224–32. https://doi.org/10.7324/JAPS.2017.70931

131 

Jakob F, Pfaff A, Novoa-carballal R, Rübsam H, Becker T, Vogel RF. Structural analysis of fructans produced by acetic acid bacteria reveals a relation to hydrocolloid function. Carbohydr Polym. 2013;92(2):1234–42. https://doi.org/10.1016/j.carbpol.2012.10.054 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/23399151

132 

Adamberg K, Tomson K, Talve T, Pudova K, Puurand M, Visnapuu T, et al. Levan enhances associated growth of Bacteroides, Escherichia, Streptococcus and Faecalibacterium in fecal microbiota. PLoS ONE. 2015;10(12):e0144042. https://doi.org/10.1371/journal.pone.0144042 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/26629816

133 

Adamberg S, Tomson K, Vija H, Puurand M, Kabanova N, Visnapuu T, et al. Degradation of fructans and production of propionic acid by bacteroides thetaiotaomicron are enhanced by the shortage of amino acids. Front Nutr. 2014;1:21. https://doi.org/10.3389/fnut.2014.00021 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/25988123

134 

Porras-Domínguez JR, Ávila-Fernández Á, Rodríguez-Alegría ME, Miranda-Molina A, Escalante A, González-Cervantes R, et al. Levan-type FOS production using a Bacillus licheniformis endolevanase. Process Biochem. 2014;49(5):783–90. https://doi.org/10.1016/j.procbio.2014.02.005

135 

Li J, Kim IH. Effects of levan-type fructan supplementation on growth performance, digestibility, blood profile, fecal microbiota, and immune responses after lipopolysaccharide challenge in growing pigs. J Anim Sci. 2013;91(11):5336–43. https://doi.org/10.2527/jas.2013-6665 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/24045486

136 

Zhao PY, Wang JP, Kim IH. Effect of dietary levan fructan supplementation on growth performance, meat quality, relative organ weight, cecal microflora, and excreta noxious gas emission in broilers. J Anim Sci. 2013;91(11):5287–93. https://doi.org/10.2527/jas.2012-5464 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/24045474

137 

Zhao PY, Wang JP, Kim IH. Evaluation of dietary fructan supplementation on growth performance, nutrient digestibility, meat quality, fecal microbial flora, and fecal noxious gas emission in finishing pigs. J Anim Sci. 2013;91(11):5280–6. https://doi.org/10.2527/jas.2012-5393 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/24045483

138 

Cheng R, Cheng L, Zhao Y, Wang L, Wang S, Zhang J. Biosynthesis and prebiotic activity of a linear levan from a new Paenibacillus isolate. Appl Microbiol Biotechnol. 2021;105:769–87. https://doi.org/10.1007/s00253-020-11088-8 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/33404835

139 

Jakob F, Steger S, Vogel RF. Influence of novel fructans produced by selected acetic acid bacteria on the volume and texture of wheat breads. Eur Food Res Technol. 2012;234:493–9. https://doi.org/10.1007/s00217-011-1658-7

140 

Gan L, Jiang G, Yang Y, Zheng B, Zhang S, Li X, et al. Development and characterization of levan/pullulan/chitosan edible films enriched with ε-polylysine for active food packaging. Food Chem. 2022;388:132989. https://doi.org/10.1016/j.foodchem.2022.132989 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/35447595

141 

Poddar MK, Dikshit PK. Recent development in bacterial cellulose production and synthesis of cellulose based conductive polymer nanocomposites. Nano Select. 2021;2(9):1605–28. https://doi.org/10.1002/nano.202100044

142 

Ullah H, Santos HA, Khan T. Applications of bacterial cellulose in food, cosmetics and drug delivery. Cellulose. 2016;23:2291–314. https://doi.org/10.1007/s10570-016-0986-y

143 

Andriani D, Apriyana AY, Karina M. The optimization of bacterial cellulose production and its applications: A review. Cellulose. 2020;27:6747–66. https://doi.org/10.1007/s10570-020-03273-9

144 

Azeredo HMC, Barud H, Farinas CS, Vasconcellos VM, Claro AM. Bacterial cellulose as a raw material for food and food packaging applications. Front Sustain Food Syst. 2019;3:7. https://doi.org/10.3389/fsufs.2019.00007

145 

Marestoni LD, Barud H da S, Gomes RJ, Catarino RPF, Hata NNY, Ressutte JB, et al. Commercial and potential applications of bacterial cellulose in Brazil: Ten years review. Polímeros. 2021;30(4): https://doi.org/10.1590/0104-1428.09420

146 

Shi Z, Zhang Y, Phillips GO, Yang G. Utilization of bacterial cellulose in food. Food Hydrocoll. 2014;35:539–45. https://doi.org/10.1016/j.foodhyd.2013.07.012

147 

Iguchi M, Yamanaka S, Budhiono A. Bacterial cellulose - a masterpiece of nature’s arts. J Mater Sci. 2000;35:261–70. https://doi.org/10.1023/A:1004775229149

148 

Okiyama A, Motoki M, Yamanaka S. Bacterial cellulose IV. Application to processed foods. Top Catal. 1993;6(6):503–11. https://doi.org/10.1016/S0268-005X(09)80074-X

149 

Guo Y, Zhang X, Hao W, Xie Y, Chen L, Li Z, et al. Nanobacterial cellulose/soy protein isolate complex gel as fat substitutes in ice cream model. Carbohydr Polym. 2018;198:620–30. https://doi.org/10.1016/j.carbpol.2018.06.078 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/30093042

150 

Lin S, Chen L, Chen H. Physical characteristics of surimi and bacterial cellulose composite gel. 2009;34(4):1363–79. https://doi.org/10.1111/j.1745-4530.2009.00533.x https://doi.org/10.1111/j.1745-4530.2009.00533.x

151 

Karahan AG, Kart A, Akoǧlu A, Çakmakçi ML. Physicochemical properties of low-fat soft cheese Turkish Beyaz made with bacterial cellulose as fat mimetic. Int J Dairy Technol. 2011;64(4):502–8. https://doi.org/10.1111/j.1471-0307.2011.00718.x

152 

Lin KW, Lin HY. Quality characteristics of chinese-style meatball containing bacterial cellulose (nata). J Food Sci. 2004;69(3:SNQ107-SNQ111. https://doi.org/10.1111/j.1365-2621.2004.tb13378.x https://doi.org/10.1111/j.1365-2621.2004.tb13378.x

153 

Yu SY, Lin KW. Influence of bacterial cellulose (nata) on the physicochemical and sensory properties of Frankfurter. J Food Sci. 2014;79(6):C1117–22. https://doi.org/10.1111/1750-3841.12494 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/24888552

154 

Akoğlu A, Cakir I, Karahan AG, Cakmakci ML. Effects of bacterial cellulose as a fat replacer on some properties of fat-reduced mayonnaise. Rom Biotechnol Lett. 2017;23(3):13674–80.

155 

Purwadaria T, Gunawan L, Gunawan AW. The production of nata colored by Monascus purpureus J1 pigments as functional food. Microbiol Indones. 2010;4(1):2. https://doi.org/10.5454/mi.4.1.2

156 

Wu SC, Lia YK. Application of bacterial cellulose pellets in enzyme immobilization. J Mol Catal, B Enzym. 2008;54(3–4):103–8. https://doi.org/10.1016/j.molcatb.2007.12.021

157 

Drozd R, Rakoczy R, Wasak A, Junka A, Fijałkowski K. The application of magnetically modified bacterial cellulose for immobilization of laccase. Int J Biol Macromol. 2018;108:462–70. https://doi.org/10.1016/j.ijbiomac.2017.12.031 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/29223754

158 

Chen L, Zou M, Hong FF. Evaluation of fungal laccase immobilized on natural nanostructured bacterial cellulose. Front Microbiol. 2015;6:1245. https://doi.org/10.3389/fmicb.2015.01245 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/26617585

159 

Ton NMN, Nguyen MD, Pham TTH, Le VVM. Influence of initial pH and sulfur dioxide content in must on wine fermentation by immobilized yeast in bacterial cellulose. Int Food Res J. 2010;17:743–9.

160 

Nguyen DN, Ton NMN, Le VVM. Optimization of Saccharomyces cerevisiae immobilization in bacterial cellulose by ‘adsorption- incubation’ method. Int Food Res J. 2009;16:59–64.

161 

Ton NMN, Le VVM. Application of immobilized yeast in bacterial cellulose to the repeated batch fermentation in wine-making. Int Food Res J. 2011;18(3):983–7.

162 

Żywicka A, Peitler D, Rakoczy R, Junka AF, Fijałkowski K. Wet and dry forms of bacterial cellulose synthetized by different strains of Gluconacetobacter xylinus as carriers for yeast immobilization. Appl Biochem Biotechnol. 2016;180:805–16. https://doi.org/10.1007/s12010-016-2134-4 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/27188971

163 

Jayani T, Sanjeev B, Marimuthu S, Uthandi S. Bacterial Cellulose Nano Fiber (BCNF) as carrier support for the immobilization of probiotic, Lactobacillus acidophilus 016. Carbohydr Polym. 2020;250:116965. https://doi.org/10.1016/j.carbpol.2020.116965 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/33049863

164 

Fijałkowski K, Peitler D, Rakoczy R, Zywicka A. Survival of probiotic lactic acid bacteria immobilized in different forms of bacterial cellulose in simulated gastric juices and bile salt solution. Lebensm Wiss Technol. 2016;68:322–8. https://doi.org/10.1016/j.lwt.2015.12.038

165 

Phromthep K, Leenanon B. Survivability of immobilized Lactobacillus plantarum cells within bacterial cellulose in mamao juice. Int Food Res J. 2017;24(3):939–49.

166 

Żywicka A, Wenelska K, Junka A, Chodaczek G, Szymczyk P, Fijałkowski K. Immobilization pattern of morphologically different microorganisms on bacterial cellulose membranes. World J Microbiol Biotechnol. 2019;35:11. https://doi.org/10.1007/s11274-018-2584-7 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/30604023

167 

Oliveira-Alcântara AV, Abreu AAS, Gonçalves C, Fuciños P, Cerqueira MA, Gama FMP, et al. Bacterial cellulose/cashew gum films as probiotic carriers. Lebensm Wiss Technol. 2020;130:109699. https://doi.org/10.1016/j.lwt.2020.109699

168 

Motalebi Moghanjougi Z, Rezazadeh Bari M, Alizadeh Khaledabad M, Almasi H, Amiri S. Biopreservation of white brined cheese (Feta) by using probiotic bacteria immobilized in bacterial cellulose: Optimization by response surface method and characterization. Lebensm Wiss Technol. 2020;117:108603. https://doi.org/10.1016/j.lwt.2019.108603

169 

Rasouli Y, Moradi M, Tajik H, Molaei R. Fabrication of anti-Listeria film based on bacterial cellulose and Lactobacillus sakei-derived bioactive metabolites; application in meat packaging. Food Biosci. 2021;42:101218. https://doi.org/10.1016/j.fbio.2021.101218

170 

Shafipour Yordshahi A, Moradi M, Tajik H, Molaei R. Design and preparation of antimicrobial meat wrapping nanopaper with bacterial cellulose and postbiotics of lactic acid bacteria. Int J Food Microbiol. 2020;321:108561. https://doi.org/10.1016/j.ijfoodmicro.2020.108561 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/32078868

171 

Cazón P, Vázquez M. Bacterial cellulose as a biodegradable food packaging material: A review. Food Hydrocoll. 2021;113:106530. https://doi.org/10.1016/j.foodhyd.2020.106530

172 

Zhai X, Lin D, Liu D, Yang X. Emulsions stabilized by nanofibers from bacterial cellulose: New potential food-grade Pickering emulsions. Food Res Int. 2018;103:12–20. https://doi.org/10.1016/j.foodres.2017.10.030 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/29389597

173 

Paximada P, Koutinas AA, Scholten E, Mandala IG. Effect of bacterial cellulose addition on physical properties of WPI emulsions. Comparison with common thickeners. Food Hydrocoll. 2016;54(Part B):245–54. https://doi.org/10.1016/j.foodhyd.2015.10.014 https://doi.org/10.1016/j.foodhyd.2015.10.014

174 

Yan H, Chen X, Song H, Li J, Feng Y, Shi Z, et al. Synthesis of bacterial cellulose and bacterial cellulose nanocrystals for their applications in the stabilization of olive oil pickering emulsion. Food Hydrocoll. 2017;72:127–35. https://doi.org/10.1016/j.foodhyd.2017.05.044

175 

Moradi M, Tajik H, Almasi H, Forough M, Ezati P. A novel pH-sensing indicator based on bacterial cellulose nanofibers and black carrot anthocyanins for monitoring fish freshness. Carbohydr Polym. 2019;222:115030. https://doi.org/10.1016/j.carbpol.2019.115030 PubMed: http://www.ncbi.nlm.nih.gov/pubmed/31320095

176 

Kuswandi B, Asih NPN, Pratoko DK, Kristiningrum N, Moradi M. Edible pH sensor based on immobilized red cabbage anthocyanins into bacterial cellulose membrane for intelligent food packaging. Packag Technol Sci. 2020;33(8):321–32. https://doi.org/10.1002/pts.2507

177 

Kuswandi B, Maryska C. Jayus, Abdullah A, Heng LY. Real time on-package freshness indicator for guavas packaging. J Food Meas Charact. 2013;7:29–39. https://doi.org/10.1007/s11694-013-9136-5

178 

Mohammadalinejhad S, Almasi H, Moradi M. Immobilization of Echium amoenum anthocyanins into bacterial cellulose film: A novel colorimetric pH indicator for freshness/spoilage monitoring of shrimp. Food Control. 2020;113:107169. https://doi.org/10.1016/j.foodcont.2020.107169

179 

Wen Y, Liu J, Jiang L, Zhu Z, He S, He S, et al. Development of intelligent/active food packaging film based on TEMPO-oxidized bacterial cellulose containing thymol and anthocyanin-rich purple potato extract for shelf life extension of shrimp. Food Packag Shelf Life. 2021;29:100709. https://doi.org/10.1016/j.fpsl.2021.100709


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