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Review

Exploring Exopolysaccharides Produced in Indigenous Mexican Fermented Beverages and Their Biotechnological Applications

by
Julián Fernando Oviedo-León
1,
Abril Ramírez Higuera
2,
Jorge Yáñez-Fernández
3,
Humberto Hernández-Sánchez
1 and
Diana C. Castro-Rodríguez
4,*
1
Departamento de Ingeniería Bioquímica, Escuela Nacional de Ciencias Biológicas, Instituto Politécnico Nacional, Unidad Profesional Adolfo López Mateos, Mexico City 07738, Mexico
2
Facultad de Nutrición, Universidad Veracruzana Región Veracruz, Iturbide S/N Ignacio Zaragoza, Veracruz 91700, Mexico
3
Centro de Investigación en Ciencia Aplicada y Tecnología Avanzada, CICATA, Instituto Politécnico Nacional, Unidad Legaria 694, Mexico City 11500, Mexico
4
Departamento Inmuno-Bioquímica, Instituto Nacional de Perinatología, Montes Urales 800, Mexico City 11000, Mexico
*
Author to whom correspondence should be addressed.
Fermentation 2025, 11(8), 463; https://doi.org/10.3390/fermentation11080463
Submission received: 10 July 2025 / Revised: 2 August 2025 / Accepted: 9 August 2025 / Published: 12 August 2025
(This article belongs to the Special Issue The Health-Boosting Power of Fermented Foods and Their By-Products)
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Abstract

Indigenous Mexican fermented beverages, such as pulque, colonche, tepache, and water kefir, are pillars of the country’s cultural and gastronomic heritage. Their sensory attributes and health-promoting properties arise from complex microbial consortia, in which lactic acid bacteria (LAB), mainly Lactobacillus and Leuconostoc, acetic acid bacteria (AAB), primarily Acetobacter, and yeasts such as Saccharomyces and Candida interact and secrete exopolysaccharides (EPSs). Dextran, levan, and heteropolysaccharides rich in glucose, galactose, and rhamnose have been consistently isolated from these beverages. EPSs produced by LAB enhance the viscosity and mouthfeel, extend the shelf life, and exhibit prebiotic, antioxidant, and immunomodulatory activities that support gut and immune health. Beyond food, certain EPSs promote plant growth, function as biocontrol agents against phytopathogens, and facilitate biofilm-based bioremediation, underscoring their biotechnological potential. This review integrates recent advances in the composition, biosynthetic pathways, and functional properties of microbial EPSs from Mexican fermented beverages. We compare reported titers, outline key enzymes, including dextransucrase, levansucrase, and glycosyltransferases, and examine how fermentation variables (the substrate, pH, and temperature) influence the polymer yield and structure. Finally, we highlight emerging applications that position these naturally occurring biopolymers as sustainable ingredients for food and agricultural innovation.

1. Introduction

Mexico is a country rich in traditions, including the consumption of handcrafted beverages recognized for their medicinal properties [1]. Since pre-Hispanic times, diverse natural resources have been exploited to prepare foods and drinks, an activity that became crucial as the population grew [2]. Today, traditional Mexican beverages serve not only as meal accompaniments, but also play a role in ceremonial contexts.
More than sixty indigenous peoples inhabit Mexico, each with its own cultural richness [1]. The preservation of their traditions has added value to handcrafted beverages, which are currently considered rich in nutrients as well as in microbial communities that can offer health benefits to the consumer. Mexico’s geographic location has favored the development of ecosystems rich in raw materials, such as fruits, cereals, roots, and prickly pear fruits, used to produce fermented beverages (Figure 1) [3]. More than 200 fermented products have been described [4], involving different genera and species of microbial communities that yield alcoholic, lactic, acetic, or alkaline drinks [5]. Examples include sotol, bacanora, charanda, comiteco, pulque, colonche, tascalate, pozol, chilate, bupu, agua de barranca, and tejate (Figure 1) [6].
Fermented beverages are widely appreciated for their flavors, derived from volatile compounds generated during fermentation [7]. The profile of these volatile compounds varies with the temperature, substrate concentration, raw material maturity, and the metabolism of the microbial community involved [8,9]. Non-alcoholic fermented beverages have also gained attention as sources of probiotics and prebiotics that benefit the intestinal microbiota, favoring eubiosis and thereby helping to prevent chronic diseases [10]. Among the more than 200 fermented products, some display a characteristic viscosity attributable to the production of exopolysaccharides (EPSs) by several bacterial genera [6]. Pulque, for instance, owes its viscosity to dextran produced by the genus Leuconostoc, whereas tepache and colonche are often fermented using tibicos (water kefir grains), polysaccharide matrices that embed diverse microbial communities [6,11,12,13]. The objective of this review is, therefore, to emphasize the importance of various EPS-producing genera isolated from Mexican fermented beverages, and to describe the biotechnological applications of these exopolysaccharides.

2. Tibicos (Water Kefir Grains) and Their Microbial Communities

Tibicos are gelatinous masses composed of bacteria and yeasts that develop in sugary liquids left to ferment undisturbed [14]. The origin of tibicos remains uncertain, and their bacterial and yeast composition can vary depending on the source [15,16]. Water kefir, also known as sugary kefir or tibicos tepache, contains 80–89% sucrose, 10% reducing sugars, and approximately 0.4% protein [15]. Tibicos are formed by consortia of microbial communities embedded in a polysaccharide matrix, primarily lactic acid bacteria, whose functional characteristics and safety position them as potential probiotics of significant research interest [11].

2.1. Microbial Communities Involved in the Production of Water Kefir

The microbial community is essential for water kefir production, being primarily composed of lactic acid bacteria (LAB), accompanied by acetic acid bacteria (AAB), and yeasts. LAB constitute approximately 80% of the total microbiota [17]. These bacteria synthesize bioactive compounds and markedly decrease the pH through the production of D- and L-lactic acid, thereby increasing the titratable acidity and conferring the beverage’s characteristic acidic flavor [18]. LAB also enhance intestinal health and immune function while contributing to the flavor, texture, and preservation.
ABB account for about 10% of the microbial community and produce acetic acid, which further acidifies the beverage and adds vinegar-like notes that enrich the flavor profile [19]. ABB also stabilize the microbial ecosystem and participate in the formation of polysaccharides.
Yeasts represent roughly 5% of the microbiota and ferment sugars to ethanol and carbon dioxide, generating natural carbonation [20]. They enrich aroma, flavor, and mouthfeel and provide vitamins and growth factors that support LAB and ABB growth, sustaining the symbiotic balance within the grains.
The specific LAB, AAB, and yeasts species present in each starter culture vary with the raw materials and geographic origin. Although Table 1 lists the species most frequently reported in the literature, additional, as-yet-undocumented, strains are likely to exist [17].

2.2. Techniques for Identifying the Microbial Contents in Water Kefir

State-of-the-art approaches combine culture-dependent protocols with culture-independent molecular methods, notably polymerase chain reaction (PCR) fingerprinting and next-generation sequencing (NGS) [17]. The culture-dependent methods typically start with the isolation and cultivation of microbes on selective agar media, such as de Man, Rogosa, and Sharpe (MRS) for LAB and Yeast Glucose Chloramphenicol (YGC) for yeast. After isolation, colonies are selected and subcultured, followed by DNA extraction and the use of PCR fingerprinting methods. For bacteria, techniques like (GTG)5-PCR are utilized, while for yeast, various specific methods are applied to generate strain-specific profiles. Identification may also involve the clustering of PCR fingerprints and sequencing of representative isolates [21,22]. There are also culture-independent techniques. These include extracting DNA directly from water and kefir grains without prior cultivation. Subsequently, denaturing gradient gel electrophoresis (DGGE) is employed to target specific microbial groups, such as LAB and yeast, to profile the community’s diversity. The amplicon sequencing of 16S rRNA gene regions (e.g., V1–V4 and V4–V5) using NGS platforms allows for a comprehensive characterization of the microbial diversity (Table 2). Additionally, metagenomic approaches provide insights into whole microbiomes, including non-culturable species [23].

3. Exopolysaccharides

LAB produce a variety of metabolites, including lactic acid, carbon dioxide, and diacetyl and acetoin, which enhance the flavor and texture and help preserve fermented foods. They also synthesize polysaccharides during the cell wall remodeling of peptidoglycan [24,25]. These polymers are termed capsular polysaccharides (CPS) when they remain attached to the cell surface and as exopolysaccharides (EPSs) when secreted into the surrounding medium [26]. This section focuses on EPSs, which show diversity in the sugar composition, glycosidic linkages, branching, and substitution patterns [27]. A comparative overview of representative EPSs produced by Mexican strains isolated from pulque, pozol, aguamiel, and other traditional matrices is provided in Table 3. Their production acts as a protective mechanism, maintaining microbial cell integrity under adverse conditions, such as desiccation, osmotic stress, pH fluctuations, and exposure to antimicrobial factors, including phagocytosis, reactive agents, antibiotics, and ethanol [26,28].

3.1. Classification, Chemical Composition, and Synthesis

EPSs are secondary metabolites synthesized by diverse microbial communities, including bacteria, filamentous fungi, yeasts, and microalgae, mainly during the late exponential or stationary phase as a stress response [36,37]. These high-molecular-weight sugar polymers play multiple roles: they promote biofilm formation, enable surface adhesion, protect cells against desiccation and antimicrobials, and in some pathogens, contribute to virulence [38].
According to the monomer composition, EPSs are classified as homopolysaccharides (HoPs), built from a single type of sugar, or heteropolysaccharides (HePs), composed of two or more different sugars [38]. HoPs may be branched or unbranched chains of glucose or fructose [39] and are further subdivided by the linkage type: α-glucans (e.g, dextran α-D-Glc(1→6), mutan: α-D-Glc(1→3), alternan alternating α-D-Glc(1→6)/(1→3), β-glucans (β-D-Glc(1→3) with (1→2) side chains), β-fructans (levan β-D-Fru(2→6), and inulin β-D-Fru(2→1)) (Figure 2) [39,40].
The biosynthesis of EPSs, specifically α-glucans and β-fructans, primarily relies on sucrose as the essential donor substrate and involves specialized extracellular glycoside hydrolase enzymes. For α-glucans, the enzyme glucansucrase, which belongs to the glycoside hydrolase GH-70 family, is essential. Glucansucrases catalyze the hydrolysis of sucrose and sequentially transfer the released glucose monomers to form a growing polymeric α-glucan chain. Similarly, the synthesis of β-fructans is mediated by fructansucrases, classified within the GH-68 glycoside hydrolase family. Fructansucrases cleave sucrose to release fructose units, then transfer these residues to elongate the β-fructan polymer [41,42]. These steps are summarized in the biosynthetic pathway diagram (Figure 3).
Unlike HoPs, HePs are complex EPSs composed of repeating units that include several monosaccharides, mainly glucose, galactose, and rhamnose, as well as less common residues such as N-acetylglucosamine, N-acetylgalactosamine, and organic or inorganic substituents like glucuronic acid, acetyl, glycerol, or phosphate groups (Figure 4) [43]. Backbone monomers are linked by glycosidic bonds and may carry side branches, generating HePs with distinctive physicochemical properties and solubility that underpin a wide range of technological and biological applications [43,44]. Due to this structural diversity, the biosynthetic machinery is correspondingly complex. In LAB, the genes required for HePs production are organized in eps clusters, which show an operon-like arrangement [45,46]. These clusters typically encode transcriptional regulators, multiple glycosyl-transferases for assembling the repeating unit, and proteins that determine the chain length, polymerization, and extracellular export [47].
The biosynthesis of HePs begins in the cytoplasm, where intracellular glycosyltransferases generate the activated sugar nucleotide donors, uridine diphosphate glucose (UDP-Glc) and uridine diphosphate galactose (UDP-Gal), that fuel polymer assembly [36]. These enzymes transfer monosaccharide residues to the lipid carrier undecaprenyl phosphate, initiating the repeat unit that forms the polymer backbone [36]. As synthesis progresses, the growing oligosaccharide is flipped across the cytoplasmic membrane and is further elongated or exported through dedicated ATP-binding cassette (ABC) transporters or Wzy-dependent transmembrane complexes [39].
The expression of the EPS gene cluster is finely regulated by environmental cues such as a nutrient limitation, osmotic stress, and temperature shifts. Two-component signal transduction systems, quorum-sensing circuits, and LuxR/AraC-type transcription factors modulate the transcription of genes encoding glycosyltransferases, glycoside hydrolases, and chain-length-determining proteins [39]. Additional layers of control, including post-translational modifications, protein–protein interactions, and the feedback inhibition of sugar nucleotide pools, further modulate enzyme activity, ensuring the precise control of the HeP molecular weight and branching patterns [40].
As outlined above, the two EPS classes also differ in size: HoPs typically exceed 1 × 106 Da, whereas HePs span a broader range of approximately 1 × 104 to 6 × 106 Da [36]. Consequently, the biological activities of EPSs depend closely on their structural attributes, including the monosaccharide composition, molecular weight, type and spatial arrangement of glycosidic linkages, and overall chain conformation [48,49].

3.2. Extraction, Purification, and Characterization of Exopolysaccharides

The production of EPSs by LAB is fundamental to optimizing the biotechnological process and achieving the highest possible yields. Selecting a suitable strain, or a synergistic co-culture, is, therefore, essential [38]. The efficiency of EPS biosynthesis depends largely on correctly configuring culture parameters, including a medium composition, micronutrient profile, and fermentation conditions [48], all tailored to the specific physiological requirements of the chosen strain [50,51].
EPS synthesis is strongly influenced by physicochemical variables, particularly the pH and temperature, which are strain-dependent [52]. Optimal production is generally reported within a pH range of 5.0–7.0; active pH control can double yields compared with uncontrolled fermentation [48,53,54,55], e.g., Streptococcus thermophilus produced 1029 mg L−1 of EPSs at a constant pH 5.5 versus 491 mg L−1 without a control [54]. Temperature effects are similar: yields increase when cultures are grown a few degrees below their optimum growth temperature, whereas temperatures above that range reduce production [33,43,53,54]. In Lactococcus lactis, Lacticaseibacillus rhamnosus, and Lactiplantibacillus plantarum, the optimal window is 18–25 °C [56,57,58], while for S. thermophiles, it is 32–42 °C [54,59,60]. Preventing enzymatic degradation during fermentation, triggered by endogenous or exogenous hydrolases and accentuated by an extreme pH or temperature, is critical to maximize net EPS accumulation [48,61].
After biosynthesis, EPSs are typically recovered from the culture in three steps. First, the microbial cells are removed by centrifugation. Next, the EPSs in the supernatant are precipitated by adding organic solvents, such as ethanol or acetone. Finally, the precipitated material is dried using either spray-drying or freeze-drying [36,48,49,62].
The structural characterization of EPSs relies on nuclear magnetic resonance (NMR), Fourier transform infrared spectroscopy (FTIR), scanning electron microscopy (SEM), and transmission electron microscopy (TEM) [26,33,36,62]. The physicochemical characterization test includes rheological profiling and the determination of the monosaccharide composition, solubility, water absorption capacity, and moisture content [33,63]. Such comprehensive characterization is indispensable for understanding EPS functional properties and for evaluating their application in diverse industrial and biotechnological sectors.

4. Functional Roles and Emerging Applications of Exopolysaccharides Derived from Microbial Community in Indigenous Mexican Fermented Beverages

Exopolysaccharides, produced by LAB, AAB, and yeasts isolated from pulque, pozol, sotol, aguamiel, and other indigenous Mexican fermented beverages, offer biocompatibility, biodegradability, and remarkable structural versatility. These traits support their growing use as natural texturizers, prebiotic fibers, and bioactive coatings in local food products, while also creating opportunities in pharmaceutical, biomedical, agricultural, and environmental technologies. In this section, we contextualize each functional role, texture engineering, antioxidant and immunomodulatory activity, plant growth promotion, and bioremediation, through case studies involving Mexican strains or beverages, highlighting how their EPSs can replace or outperform synthetic polymers.

4.1. Applications of Exopolysaccharides in the Food Industry

Exopolysaccharides synthesized by LAB, AAB, and yeasts from aguamiel, pulque, stool, and pozol provide an illustrative benchmark of how traditional Mexican fermentations can inspire cutting-edge food design. Their ability to enhance viscosity, stabilize emulsions and confer bioactivity parallels the long-recognized performance of industrial dextran, xanthan, and gellan, but with the added value of cultural provenance and sustainability. Recent advances in the culture-medium design and the optimization of carbon and nitrogen sources have boosted EPS yields and functionality, facilitating their transition from laboratory research to large-scale industrial application [64].
Recent studies on aguamiel and pulque derived from Agave species have highlighted the nutritional and functional value of their microbial exopolysaccharides. EPSs isolated from these sources demonstrated a notable radical-scavenging capacity, antioxidant activity, and selective stimulation of beneficial probiotic microorganisms [65]. Aguamiel itself promotes the growth of probiotic strains like Bacillus mojavensis, which presents notable probiotic potential and antimicrobial activity against pathogens such as Escherichia coli, Staphylococcus aureus, and Candida albicans [66].
Both HoPs and HePs of a microbial origin are extensively utilized in the food industry as thickening and stabilizing agents, owing to their exceptional ability to modify the rheological properties of food systems. These polysaccharides enhance the viscosity, improve texture, and prevent phase separation in products such as sauces, dairy items, and beverages. For instance, dextran from lactic acid bacteria isolated from aguamiel forms hydrated networks that immobilize water and oil droplets, promoting stability in emulsions and beverages like pulque [65]. Their functional effects are primarily attributed to the formation of hydrated networks that trap water molecules and immobilize dispersed particles or oil droplets, thereby promoting product stability and extending the shelf life [67].
As illustrated in Figure 5, dextran forms a hydrated three-dimensional network through the entanglement and branching of its glucose chains. This network effectively retains water and encapsulates oil droplets, thereby enhancing the viscosity and stability of emulsions. Pulque fermentation by microorganisms like Leuconostoc mesenteroides notably involves dextran biosynthesis, contributing significantly to pulque’s characteristic viscosity and probiotic functionality [68]. This concept is further supported by studies demonstrating the stabilization of β-carotene emulsions using ovalbumin–dextran conjugates, which significantly enhance the emulsion stability and interfacial activity. [69]. Although this mechanism is characteristic of many HoPs with branched or high-molecular-weight structures (such as levan and pullulan), it can vary depending on the chemical composition and structural architecture of the specific polysaccharide [67].
HePs such as xanthan, gellan, and bacterial alginates can function through either similar or distinct mechanisms. Xanthan gum increases viscosity by forming entangled molecular networks, whereas gellan and alginate require ions such as Ca2+ to gel and create elastic structures [70]. These differences offer tailored textural properties and specific applications in food formulations. For example, HePs produced during the fermentation of pozol by lactic acid bacteria, including Weissella confusa and Streptococcus infantarius, not only enhance the texture, but also increase the lactic acid yield and fermentation efficiency through microbial synergism [71].
Advancement in HoPs production lie in optimizing fermentation conditions and exploring diverse carbon sources. For example, EPS biosynthesis in sotol fermentation (Dasylirion sp.) is influenced by the microbial composition, where the genera Pichia and Saccharomyces contribute significantly to the biosynthesis of metabolites affecting flavor and polysaccharide profiles, thus expanding their potential use in tailored food applications [72]. Recent multi-omic analyses have also revealed genetic and metabolic regulation sensitive to substrate variation, opening avenues for metabolic engineering to produce tailored polysaccharides [73].
HoPs such as dextran and scleroglucan are widely employed in the food industry as texture enhancers, stabilizers, and water retention agents, especially in bakery goods, dairy products, and plant-based formulations. Scleroglucan’s resistance to heat and salt makes it suitable for thermally processed foods like soups and sauces. Moreover, advances in strain engineering and the utilization of low-cost substrates, such as agro-industrial byproducts, lower production costs and support large-scale commercialization [67,73].
Table 4 presents a comprehensive overview of key microbial exopolysaccharides, their respective producing microorganisms, and primary industrial applications. These biopolymers function as thickeners, stabilizers, gelling agents, water retention enhancers, and edible film materials, roles determined by distinct structural characteristics. Recent advances in statistical optimization techniques, such as a Plackett–Burman design and response surface methodology (RSM), have significantly improved the yield of HePs, enabling the fine-tuning of functional properties such as the emulsification capacity and thermal stability [74]. Lactiplantibacillus paraplantarum and Weissella paramesenteroides, isolated from pulque and aguamiel, have been investigated for their HePs, exhibiting antioxidant activity, rheological functionality, and cholesterol-lowering effects [75].
HePs have demonstrated significant health-promoting properties, including prebiotic effects and lipid-modulating activity, underscoring their promise in functional food applications. For example, EPS produced by Lactiplantibacillus paraplantarum NCCP 962 reduced cholesterol levels by up to 46% in vitro by trapping cholesterol in micelles and modulating gene expression—downregulating HMGCR and upregulating LDLR in human HepG2 cells. Likewise, EPS from Weissella paramesenteroides MYPS51 exhibited antitumor activity in human HT-29 colorectal cancer cells and showed antioxidant and hepatoprotective effects in vivo in a CCl4-induced liver injury mouse model [74,75].
Thus, while HoPs are primarily utilized as texture enhancers and rheological agents, thanks to their cost-effectiveness, HePs are increasingly valued in functional food formulations. Their appeal lies in the ability to combine textural and stability improvements with added health benefits, reflecting a growing trend toward more sophisticated and personalized food products [74,76].

4.2. Recent Advances in the Health Benefits of Microbial Exopolysaccharides

Although EPSs were first prized for improving the texture and stability of fermented foods, those produced by LAB, and other microbial communities, are now recognized for their diverse bioactive properties. Evidence shows that EPSs isolated from dextran-rich pulque, levan-containing aguamiel, and HePs recovered from pozol act as prebiotics, selectively stimulating Bifidobacterium and Lactobacillus populations, while favoring short-chain fatty acid production highlights their role as prebiotics, which can modulate the gut microbiota and support intestinal health [66,71]. In addition, EPSs have demonstrated antioxidant, immunomodulatory, and antibacterial activities, positioning them as promising candidates for the development of next-generation functional foods and nutraceuticals. This section reviews recent advances in understanding the health-promoting effects of microbial EPSs, with a focus on new mechanisms of action and innovative applications that expand their potential benefits for human health [85,86].

4.2.1. Exopolysaccharides as Next-Generation Prebiotics

Among traditional Mexican fermented beverages, pulque stands out as the most promising source of bioactive EPSs with prebiotic potential. Its microbial community, including Leuconostoc mesenteroides, Lactobacillus spp., and Zymomonas mobilis, produces dextran-rich EPSs that contribute to its viscosity and have been shown to selectively stimulate Bifidobacterium and Lactobacillus, while enhancing short-chain fatty acid (SCFA) production [87]. Other traditional beverages, such as pozol, fermented with Lactobacillus plantarum and Weissella spp., and colonche, which involves EPS-producing Lactobacillus and Leuconostoc strains, also exhibit significant EPSs biosynthesis with potential prebiotic effects [88]. Similarly, aguamiel contains native levan-producing bacteria such as Fructobacillus and Acetobacter, which initiate EPS production even before fermentation. Collectively, these beverages represent a valuable source of microbial EPSs capable of modulating the gut microbiota and promoting intestinal health [87,89].
Notably, research on Mexican fermented beverages has already described the structural diversity and unique features of their exopolysaccharides. For example, the dextran synthesized by Leuconostoc mesenteroides strains isolated from pulque and agave [33] shares the α-1,6-glucan backbone with commercial dextran, but often exhibits distinctive branching patterns and molecular weights, which may confer specific prebiotic functions. Likewise, exopolysaccharides isolated from pozol (L. plantarum, Weissella spp.) and colonche (Lactobacillus, Leuconostoc), are emerging as promising candidates for modulating the gut microbiota, although in vivo functional studies remain limited. In the case of aguamiel, the presence of high-molecular-weight levan produced by Fructobacillus and Acetobacter spp. prior to fermentation suggests untapped potential for indigenous prebiotics with unique immunomodulatory profiles [89].
Among the expanding array of dietary fibers, four microbial exopolysaccharides—dextran, β-(2→6) levan, kefiran, and pullulan—have emerged as the most intensively studied next-generation prebiotics. Of these, dextran and levan are naturally present in pulque and aguamiel, respectively, as direct products of the Mexican microbial consortia described above. Kefiran, though more widely associated with kefir, is also produced in Mexican tibicos (water kefir), another traditional fermented beverage [90].
Long-chain dextran from Weissella cibaria increased the diversity of beneficial gut taxa such as Bifidobacterium and Faecalibacterium in an ex vivo human feces model, confirming its bifidogenic potential [91]. Levan-type fructans not only promote commensal growth, but also exert molecular-weight-dependent effects on innate immune receptors, attenuating pro-inflammatory TLR5/8 while moderately activating TLR2/4. These effects were demonstrated in vitro using human THP-1 macrophages and HEK-Blue™ reporter cell lines expressing individual TLRs, where high-molecular-weight levans induced a balanced immunomodulatory profile marked by the downregulation of TLR5/8 responses and the selective, moderate activation of TLR2/4 signaling pathways [90]. In vivo, rice-derived kefiran shifted the murine microbiota towards Bacteroides and Alistipes, increased fecal acetate, and alleviated diet-induced obesity and steatosis [92]. Finally, a cellulose–pullulan matrix effectively delivered Lacticaseibacillus rhamnosus GG as a stable biofilm to the large intestine, where the released pullulan concurrently served as a selective carbon source. This was demonstrated in vivo using BALB/c mice, with oral administration enabling targeted intestinal delivery and prolonged colonization [93].
Owing to their structural complexity, many EPSs are selectively metabolized by commensal genera such as Lactobacillus and Bifidobacterium, which in turn produce SCFAs like acetate, propionate, and butyrate. These SCFAs acidify the intestinal lumen, repress virulence genes in pathogens such as Salmonella and E. coli, and strengthen intestinal tight junctions. A recent study revealed that Streptococcus salivarius establishes an EPS-SCFA axis that modulates host energy metabolism and displaces opportunistic pathogens in murine models. In SPF mice colonized with S. salivarius M18, its exopolysaccharide promoted butyrate production by resident microbiota, resulting in improved intestinal energy homeostasis, the upregulated expression of tight-junction proteins, and reduced Clostridioides difficile colonization [94].
Their selectivity for probiotic, rather than pathogenic, guilds stems from the ecology of enzymes and the architecture of polymers. Dextran’s predominantly α-1,6-linked backbone is degradable by GH49 and GH66 dextranases, which are widely encoded in Bifidobacterium and lactobacilli, producing isomalto-oligosaccharides that fuel the bifid shunt; these enzymes are rarely found in enterobacteria [95,96]. Levan relies on GH32 β-fructofuranosidases, abundant in butyrate-producing Parabacteroides, but scarce in most Proteobacteria; its high-molecular-weight fractions also interact with TLRs to modulate host immunity [90,95]. Kefiran and pullulan, characterized by mixed α/β linkages, require multi-domain pullulanases and α-glucosidases, which are enriched in lactobacilli genomes, establishing a metabolic niche that excludes common pathogens. Thus, polymer chemistry regulates access to specialized carbohydrate-active enzymes (CAZymes) and links microbial cross-feeding (acetate, lactate, and butyrate) with receptor-level immunomodulation, an emerging paradigm in precision nutrition fibers [97,98]. As illustrated in Figure 6, the molecular architecture of each polysaccharide, dextran (α-1,6-glucan) and kefiran (mixed α/β glucogalactan), dictates the specificity of the degrading CAZymes (GH49/GH66 dextranases versus multi-domain pullulanases/α-glucosidases). The oligosaccharides released are then selectively fermented by probiotic guilds, generating acetate, propionate, and butyrate that strengthen epithelial barrier integrity and restrict the overgrowth of pathobionts [95].
Although most mechanistic studies have used international strains or commercial exopolysaccharides, those produced by Mexican traditional beverages represent a largely untapped resource with comparable or even greater structural and functional diversity. The isolation and functional characterization of exopolysaccharides from pulque, pozol, colonche, aguamiel, and tibicos will not only help validate these mechanisms in a local context, but could also reveal novel molecules with region-specific prebiotic, immunomodulatory, or bioactive profiles, thereby expanding the global repertoire of next-generation prebiotics [89].

4.2.2. Antioxidant and Anti-Inflammatory Properties

The oxidative stress-inflammatory response axis is bidirectional; the exacerbation of one intensifies the other, perpetuating tissue damage. EPSs appear to disrupt this cycle through both chemical mechanisms, such as electron donation or the chelation of pro-oxidant metals, and biological pathways that reset intracellular signaling and reshape the gut microbiota [99].
In vitro, dextrans, levans, and kefiran neutralize DPPH and ABTS radicals and sequester Fe2+ or Cu2+ ions, thanks to exposed hydroxyl and uronic groups on their branches [100]. However, their high molecular weight limits systemic absorption; the most consistent in vivo effects stem from the activation of cytoprotective pathways. A study using EPS from Lactiplantibacillus plantarum and L. delbrueckii demonstrated Nrf2 disinhibition, the increased expression of HO-1, NQO1, and glutathione, and a 40% reduction in intracellular ROS in PC12 cells exposed to hydrogen peroxide. Thus, EPSs act as conditional antioxidants and they exhibit modest local chemical activity, but primarily enhance endogenous defenses [101].
EPS production from Leuconostoc pseudomesenteroides cultivated on agro-industrial by-products, such as beet residues, has shown promising antioxidant properties. EPS synthesized by these strains exhibited strong radical-scavenging and metal-chelating capacities (IC50 = 0.78 mg mL−1 for DPPH). Although these findings originate from non-Mexican strains, the presence of native Leuconostoc isolates in traditional Mexican beverages like pulque and colonche opens a sustainable avenue for exploiting local agro-industrial residues. Beet waste, abundant in regions such as Guanajuato and Zacatecas, could be used as a low-cost substrate for EPS synthesis, aligning with circular economy principles to generate valuable antioxidant biopolymers while adding value to regional agricultural by-products [102].
Simultaneously, the colonic fermentation of EPS yields postbiotic metabolites that reinforce the epithelial barrier and improve the mitochondrial efficiency. The high-molecular-weight dextran generated by Apilactobacillus waqarii serves as an example in a mouse model of metabolic syndrome, resulting in decreased weight gain, cholesterol levels, and blood glucose, along with reduced indicators of lipid peroxidation [103].
EPSs interact with pattern recognition receptors (TLR2/4, Dectin-1, SR-A), attenuating the MyD88→NF-κB/MAPK cascade. The administration of EPS from L. plantarum NMGL2 to DSS-induced colitic mice decreased TNF-α and IL-1β, increased IL-10, strengthened tight junction proteins (ZO-1, occludin), and suppressed NF-κB p65 phosphorylation [104]. Complementarily, pre-supplementation with kefiran or kefir water reduced neutrophilic infiltration and levels of TNF-α, KC, and MCP-1 in the poly(I:C)-induced respiratory challenge, while elevating regulatory cytokines IL-10 and IL-27, resulting in reduced lung damage [105]. Knowledge gaps remain, particularly regarding human pharmacokinetics and the effective doses of purified EPSs.

4.2.3. Antibacterial Activity of Exopolysaccharides

EPSs produced by LAB can inhibit biofilm formation, suppress virulence gene expression, and, in some cases, compromise the structural integrity of pathogenic cell envelopes [106].
Among the most compelling findings is EPS 715 from Lactiplantibacillus plantarum PC715, which disperses up to 78% of Hafnia alvei biofilms as well as those formed by other Gram-positive and Gram-negative foodborne bacteria [106]. Similarly, EPS-cn2 from Lactobacillus casei NA-2 reduces Escherichia coli O157:H7 adhesion to plastic surfaces and Caco-2 cells by 50%, an effect linked to an altered membrane hydrophobicity and surface charge, along with the silencing of LEE pathogenicity island genes [107].
In the gastrointestinal tract, several LAB-derived EPSs have shown efficacy against classical enteropathogens. EPSs from Lacticaseibacillus rhamnosus GG attenuated Salmonella Typhimurium-induced enteritis in murine models, restoring the ileal barrier function and suppressing the TLR4/NF-κB/MAPK cascade more effectively than penicillin [108]. A heteropolysaccharide (EPS 7–4) from Lactobacillus crispatus limited Salmonella proliferation and epithelial invasion by destabilizing the bacterial cell wall and limiting ASC-inflammasome-mediated pyroptosis, thereby reducing translocation and inflammation. In vitro, EPS 7–4 suppressed Salmonella invasion in Caco-2 and HT-29 cells and downregulated virulence genes invA and hilA. In vivo, oral administration in BALB/c mice lowered the pathogen load in the spleen and liver and attenuated systemic inflammation by reducing IL-6 and TNF-α levels [109]. Additional evidence supports antibacterial activity against Helicobacter pylori and Clostridioides difficile, indicating a broad-spectrum effect that spans both clinically and food-relevant enteropathogens [110,111].
The antibacterial actions of exopolysaccharides encompass a broad spectrum of mechanisms, including the inhibition of bacterial adhesion, direct disruption of the cell envelope, downregulation of virulence genes, and sequestration of quorum-sensing molecules. Table 5 provides a comparative overview of these processes, detailing the primary mechanisms, specific modes of action, representative producing strains, and the corresponding effects on gastrointestinal pathogens.
EPSs help construct a favorable microenvironment for beneficial microbiota, restricts ecological niches available to pathogens, and, when necessary, directly inhibits or eliminates them. Their biocompatibility and low selective pressure make them promising candidates for antibiotic-free strategies in food preservation and gastrointestinal health, paving the way for precision formulations tailored to specific pathogens and microbial ecosystems [114].

4.2.4. Emerging Health Avenues and Research Gaps

Microbial EPSs produced by bacteria found in traditional Mexican fermented beverages hold promising yet underexplored therapeutic applications beyond their well-documented antioxidant, anti-inflammatory, and antibacterial properties. For instance, studies have indicated that certain EPSs could influence the gut–brain axis, potentially modulating neurological health through microbiota-mediated pathways. A notable example is EPSs from heat-killed Lactobacillus fermentum PS150, which exhibited hypnotic properties by reducing sleep latency in animal models. These effects correlated with significant shifts in the gut microbiota composition, implying potential psychobiotic roles [115]. However, similar investigations with EPSs from microbial strains native to traditional Mexican fermented beverages remain unexplored, representing a significant knowledge gap.
Furthermore, EPSs show considerable promise in reprogramming metabolic homeostasis, particularly regarding obesity and insulin resistance management. EPSs from Lactiplantibacillus plantarum KX041 demonstrated beneficial effects against diet-induced obesity in animal studies, including reductions in insulin resistance and hepatic inflammation markers [116]. Additionally, EPSs from Weissella cibaria PFY06-EPS improved glucose tolerance and modulated leptin and TNF-α levels in high-fat-diet-fed mice [117].
Beyond their direct biological activities, microbial EPSs have potential applications as biocompatible matrices for drug delivery and probiotic protection. Xanthan gum-based hydrogels have been successfully engineered for the controlled release of antibiotics such as ciprofloxacin, showing sustained antibacterial effects and cytocompatibility [118].
In Mexico, a study using traditional fermented beverages, such as pulque, demonstrated a significant reduction in academic stress and improvement in the gut microbiota composition among medical students. This effect was attributed to lactic acid bacteria present in the beverage rather than isolated EPS directly [89]. Thus, while these beverages have shown beneficial impacts on health, the specific contributions of isolated EPSs have not yet been clearly delineated, highlighting another critical research gap.

4.3. Agro-Biotechnological Potential of Microbial Exopolysaccharides

EPS-producing LAB and AAB, together with kefir-associated yeasts, constitute versatile biotools that can be integrated into soil management, crop protection, and low-input agronomy. EPS-overproducing plant-growth-promoting rhizobacteria (PGPR) play a crucial role in reinforcing soil physical architecture by improving aggregation, nutrient retention, and moisture availability. For instance, recent studies conducted in the Cuatro Ciénegas Basin, a unique arid ecosystem in northern Mexico, revealed that bacterial communities associated with the rhizosphere of Agave lechuguilla significantly enhance soil aggregation and stability compared to surrounding bulk soil. These beneficial effects were strongly associated with EPS-producing bacteria naturally selected by the plant roots, underscoring the potential of locally adapted microbial strains as bioinoculants to improve plant nutrition, drought tolerance, and overall soil health in arid agricultural systems [119].
In lettuce microcosm trials, two native Bacillus isolates (Z23 and Z39) secreted high-molecular-weight polymers that cemented micro-aggregates and increased water-stable macro-aggregates by 22%. They reduced Cd2+ and Pb2+ accumulation in edible tissues by >40% [120]. Beyond these mechanical effects, specific LAB polymers act as chelators. Paraburkholderia phytofirmans PsJN mutants defective in EPS synthesis lost their ability to mobilize Fe and Zn in the rhizosphere. They failed to rescue pea seedlings from drought stress, underscoring a direct link between EPSs, micronutrient bioavailability, and root development [121].
EPSs also function as biocontrol effectors. A neutral, dextran-like polymer (EPS_B3) purified from Leuconostoc mesenteroides inhibited eight bacterial pathogens. It dispersed ≥75% of Listeria monocytogenes biofilms at 2.5 mg mL−1, demonstrating direct antibiofilm activity relevant to phyllospheric and post-harvest diseases [122]. Likewise, a cell-free EPS fraction from Lactobacillus fermentum O1.1, isolated from citrus waste, suppressed the growth of Alternaria alternata and Fusarium oxysporum in tomato fruits, reducing visible lesions by ~60% during storage [123]. Beyond direct pathogen inhibition, several EPS preparations enhance plant innate immunity, as evidenced by the accumulation of phenylpropanoid phytoalexins and pathogenesis-related proteins in treated leaves.
Thanks to their polyanionic, highly hydrated matrices, bacterial EPSs display an exceptional metal-binding capacity. In pot trials with contaminated farmland soils, EPS-rich inoculants not only immobilized Cd2+ and Pb2+, but also shifted the resident microbiome toward taxa that enhance aggregate stability and heavy metal precipitation [120,124]. Translational research is now turning EPSs into bespoke agricultural inputs. For instance, EPSs from Mexican strains of Leuconostoc mesenteroides isolated from aguamiel exhibited effective water retention properties and showed promise as soil amendments under drought conditions, facilitating the emergence and early growth of maize seedlings. This highlights the potential of native Mexican microbial resources in addressing local agricultural challenges, especially in semiarid regions [125]. Pullulan/PHB seed coatings doped with antagonistic microbes accelerated barley germination by 15% and protected seedlings against Rhizoctonia without the use of synthetic fungicides [126]. The domain-swap engineering of a Leuconostoc dextransucrase (variant CZ8) produced narrowly dispersed 40 kDa dextran in a single step, ideal for controlled-release agro-hydrogels [127]. The genome mining of Liquorilactobacillus mali T6–52 revealed a compact EPS gene cluster whose promoter can be up-regulated to tune the chain length and branching, offering a chassis for a climate-specific polymer design [128].

5. Conclusions and Future Perspectives

Mexico’s traditional fermented beverages represent a valuable source of bioactive compounds and have significant commercial potential. However, their use on an industrial scale faces major challenges related to process standardization and quality and safety assurance. In this context, scientific research and the development of appropriate technologies are essential not only to improve their production, but also to promote the functional benefits associated with their consumption.
Reviews such as the present one highlight the relevance of these beverages as natural reservoirs of microbial strains with the capacity to synthesize biopolymers, particularly exopolysaccharides, which have attracted increasing interest worldwide. This attention is due to their safety, their multiple biological activities—including immunomodulatory, antioxidant, and antimicrobial properties—as well as their advantages over other natural compounds in industrial and therapeutic applications.
However, the study of EPSs still presents important challenges. Their structure is highly complex and variable, which makes the detailed analysis of their structure–activity relationship difficult, and so far, their mechanism of action has not been fully elucidated. In addition, factors such as the incubation time, temperature, and pH of the medium can significantly influence their composition, structure, and functionality. Therefore, the controlled manipulation of these parameters represents a promising strategy to optimize and diversify the applications of EPSs in different sectors.

Author Contributions

Conceptualization, J.F.O.-L. and D.C.C.-R.; methodology, J.F.O.-L. and D.C.C.-R.; software, J.F.O.-L. and D.C.C.-R.; validation, D.C.C.-R.; formal analysis, J.F.O.-L. and D.C.C.-R.; investigation, J.F.O.-L., A.R.H., J.Y.-F., H.H.-S. and D.C.C.-R.; resources, D.C.C.-R.; data curation, J.F.O.-L., A.R.H., J.Y.-F., H.H.-S. and D.C.C.-R.; writing—original draft preparation, J.F.O.-L., A.R.H., J.Y.-F., H.H.-S. and D.C.C.-R.; writing—review and editing, D.C.C.-R.; visualization, J.F.O.-L. and D.C.C.-R.; supervision, D.C.C.-R.; project administration, D.C.C.-R.; funding acquisition, D.C.C.-R. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Acknowledgments

The authors have reviewed and edited the output and take full responsibility for the content of this publication.

Conflicts of Interest

The authors declare no conflicts of interest.

Abbreviations

The following abbreviations are used in this review:
ABTS2,2-azinobis-(3-ethylbenzthiazoline-6-sulfonic acid)
AABAcetic Acid Bacteria
CAZymesCarbohydrate-active enzymes
CPSCapsular Polysaccharides
DNADeoxyribonucleic Acid
DPPH2,2-diphenyl-1-picrylhydrazyl
EPSExopolysaccharides
FTIRFourier Transform Infrared
HePsHeteropolysaccharides
HoPsHomopolysaccharides
LABLactic Acid Bacteria
MRSMan, Rogosa, and Sharpe
NGSNext-Generation Sequencing
NMRNuclear Magnetic Resonance
PCRPolymerase Chain Reaction
ROSReactive Oxygen Species
RSMResponse Surface Methodology
SCFAsShort-Chain Fatty Acids
SEMScanning Electron Microscopy
TEMTransmission Electron Microscopy
YGCYeast Glucose Chloramphenicol

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Figure 1. Geographic distribution of native beverages in Mexico. Each color indicates the principal traditional fermented or distilled beverage associated with a Mexican state, together with their main ingredients. The map illustrates the regional diversity of beverages such as pulque, pozol, chilate, mezcal, and others, emphasizing their cultural relevance across the country. State boundaries based on Instituto Nacional de Estadística y Geografía (INEGI) shapefile 2020; map generated in R.
Figure 1. Geographic distribution of native beverages in Mexico. Each color indicates the principal traditional fermented or distilled beverage associated with a Mexican state, together with their main ingredients. The map illustrates the regional diversity of beverages such as pulque, pozol, chilate, mezcal, and others, emphasizing their cultural relevance across the country. State boundaries based on Instituto Nacional de Estadística y Geografía (INEGI) shapefile 2020; map generated in R.
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Figure 2. Representative EPS-producing microorganisms, principal sugar donor substrate, and main homopolysaccharides (HoPs) synthesized. Structures adapted from ChEBI (Chemical Entities of Biological Interest) and KEGG (Kyoto Encyclopedia of Genes and Genomes) databases.
Figure 2. Representative EPS-producing microorganisms, principal sugar donor substrate, and main homopolysaccharides (HoPs) synthesized. Structures adapted from ChEBI (Chemical Entities of Biological Interest) and KEGG (Kyoto Encyclopedia of Genes and Genomes) databases.
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Figure 3. Extracellular biosynthetic pathway of dextran in lactic acid bacteria. Sucrose is cleaved by dextransucrase (GH70) anchored at the cell surface, releasing fructose and forming a covalent Glc–GH70 intermediate. Sequential transfer of glucose units builds an α-(1→6) dextran backbone, with occasional α-(1→3) branches. The liberated fructose can be transported into the cytoplasm and used as a carbon source for cellular metabolism. For levan-type fructans, the same substrate (sucrose) is polymerized by levansucrase (GH68), yielding a β-(2→6) fructan backbone with β-(1→2) branches.
Figure 3. Extracellular biosynthetic pathway of dextran in lactic acid bacteria. Sucrose is cleaved by dextransucrase (GH70) anchored at the cell surface, releasing fructose and forming a covalent Glc–GH70 intermediate. Sequential transfer of glucose units builds an α-(1→6) dextran backbone, with occasional α-(1→3) branches. The liberated fructose can be transported into the cytoplasm and used as a carbon source for cellular metabolism. For levan-type fructans, the same substrate (sucrose) is polymerized by levansucrase (GH68), yielding a β-(2→6) fructan backbone with β-(1→2) branches.
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Figure 4. Representative EPS-producing microorganisms, principal sugar donor substrate, and main heteropolysaccharides (HePs) synthesized. Structures adapted from ChEBI and KEGG databases.
Figure 4. Representative EPS-producing microorganisms, principal sugar donor substrate, and main heteropolysaccharides (HePs) synthesized. Structures adapted from ChEBI and KEGG databases.
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Figure 5. Thickening and stabilization mechanism of dextran in food systems. Dextran forms a hydrated three-dimensional network through α-(1→6) and α-(1→3) glycosidic linkages, effectively trapping water molecules and surrounding oil droplets. This network increases viscosity and prevents phase separation, thereby enhancing the stability of emulsions and other food products.
Figure 5. Thickening and stabilization mechanism of dextran in food systems. Dextran forms a hydrated three-dimensional network through α-(1→6) and α-(1→3) glycosidic linkages, effectively trapping water molecules and surrounding oil droplets. This network increases viscosity and prevents phase separation, thereby enhancing the stability of emulsions and other food products.
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Figure 6. CAZyme-driven utilization of EPS by probiotics and their impact on SCFA generation and host physiology. Select probiotic strains degrade complex exopolysaccharides through specific CAZymes (e.g., GH13, GH49/66, and GH32), releasing fermentable mono- and disaccharides. These are selectively utilized, leading to short-chain fatty acid (SCFA) production and host benefits.
Figure 6. CAZyme-driven utilization of EPS by probiotics and their impact on SCFA generation and host physiology. Select probiotic strains degrade complex exopolysaccharides through specific CAZymes (e.g., GH13, GH49/66, and GH32), releasing fermentable mono- and disaccharides. These are selectively utilized, leading to short-chain fatty acid (SCFA) production and host benefits.
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Table 1. Representative microbial species identified in water kefir grains.
Table 1. Representative microbial species identified in water kefir grains.
GroupSpecies Identified a
LABL. kefiranofaciens, Len. hilgardii, Ll. nagelii, Lp. plantarum, Lim. fermentum, Lac. casei, Lac. paracasei, Lim. reuteri, Len. diolivorans, Fr. farraginis, Ll. satsumensis, Lig. harbinensis, Lim. fermentum, Lp. plantarum, Api. kunkeei, Leuc. mesenteroides, Fr. hordei, Lev. brevis, Leuc. citreum
AABA. aceti, A. lovaniensis, A. tropicalis, A. indonesiensis, G. oxydans, A. pasteurianus, A. fabarum, A. orientalis, G. liquefaciens
YeastsS. cerevisiae, C. kefyr, C. guilliermondii, C. colliculosa, R. mucilaginosa, P. membranifaciens, P. occidentalis, H. valbyensis, Zt. florentina, D. bruxellensis, Z. lentus, T. delbrueckii, Lc. fermentati
a Abbreviations refer to the following genera: A. = Acetobacter; Api. = Apilactobacillus; C. = Candida; D. = Dekkera; Fr. = Fructilactobacillus; G. = Gluconobacter; H. = Hanseniaspora; Lac. = Lactcaseibacillus; Lc. = Lachancea; Len. = Lentilactobacillus; Leuc. = Leuconostoc; Lev. = Levilactobacillus; Lig. = Ligilactobacillus; Lim. = Limosilactobacillus; Ll. = Liquorilactobacillus; Lp. = Lactiplantibacillus; P. = Pichia; R. = Rhodotorula; S. = Saccharomyces; T. = Torulaspora; Z. = Zygosaccharomyces; Zt. = Zygotorulaspora.
Table 2. Techniques used to identify microbial communities in water kefir.
Table 2. Techniques used to identify microbial communities in water kefir.
Technique aPurpose/UseAdvantagesDisadvantages
Culture-dependent isolationGrowth on selective media (e.g., MRS agar for LAB); colony morphology, Gram stain, catalase test- Inexpensive and simple
- Can isolate live strains for downstream use		- Misses non-culturable or slow-growing microbes
- Biased toward dominant organisms
SDS-PAGE of whole-cell proteinsPhenotypic typing of LAB and related bacteria- Provides strain-level fingerprints
- Good for grouping isolates		- Labor-intensive
- Low resolution for phylogeny
(GTG)5-PCR fingerprinting (Rep-PCR)Genotyping of LAB strains- High discriminatory power
- Fast and reproducible for strain typing		- Requires reference patterns
- Not ideal for unknown taxa
pheS gene sequencingPrecise identification of LAB to species level using phenylalanyl-tRNA synthase gene- More specific than 16S rRNA
- Good for closely related species		- Limited databases
- Requires sequencing expertise
16S rRNA gene sequencing (amplicon)Culture-independent community profiling (mainly for bacteria)- Detects both culturable and non-culturable species
- Taxonomic assignment to genus/species level		- Low resolution for some genera
- Bias from primers and PCR amplification
Shotgun metagenomicsComprehensive analysis of total DNA from the microbial community- Strain-level resolution
- Can detect functional genes
- Detects bacteria, archaea, yeasts, and viruses		- High cost and computational resources
- DNA extraction must be efficient for all organisms
DGGE (Denaturing Gradient Gel Electrophoresis)Community fingerprinting using 16S rDNA V3 region- Detects dominant species
- Allows visual comparison across samples		- Cannot resolve all species
- Limited to most abundant populations
HPLC/GC (e.g., volatile/aromatic compounds)Analyzes fermentation metabolites: organic acids, ethanol, and esters- Indicates metabolic activity
- Helps correlate microbes with functional traits		- Indirect method
- Requires correlation with microbiota
Microscopy (light, SEM)Visualizes kefir grain structure and microbial distribution- Observes biofilms, morphology
- Supports structural characterization		- Not taxonomically informative
- Requires complementary methods
Co-culturing (e.g., Transwell system)Assess microbial interaction (LAB/yeast mutualism)- Reveals ecological relationships
- Simulates natural consortia		- Not taxonomically specific
- Complex to set up
ᵃ Complex to set DGGE, denaturing gradient gel electrophoresis; GC, gas chromatography; HPLC, high-performance liquid chromatography; LAB, lactic acid bacteria; PCR, polymerase chain reaction; SEM, scanning electron microscopy; SDS-PAGE, sodium dodecyl sulfate–polyacrylamide gel electrophoresis.
Table 3. Mexican microbial isolates reported to synthesize exopolysaccharides in traditional fermented beverages and cereal-based foods.
Table 3. Mexican microbial isolates reported to synthesize exopolysaccharides in traditional fermented beverages and cereal-based foods.
Microbial Strain (Isolate)Source BeverageMonosaccharide Composition/EPS typePotential
Applications		Reference
Leuconostoc mesenteroides N06Pozol (maize)Dextran–glucose
(α-1→6 backbone, α-1→3 branches)		Food thickener; prebiotic carrier[29]
Leuconostoc kimchii EPSAPulqueDextran
(α-1→6 main, α-1→2 and α-1→3 branches)		Bio-film matrix; functional food additive[30]
Leuconostoc kimchii EPSBPulqueCell-bound dextran + levan (β-2→6 fructan)Encapsulation; viscosity modifier[30]
Leuconostoc citreum/
L. kimchii		PulqueFructans
(levan/inulin-type)		Prebiotics; viscosity enhancer[31]
Ustilago maydisCorn tejuino/native corn brewβ-(1→3/1→6) glucans (enzymatically confirmed)Texture modifier; pharma precursor[32]
Leuconostoc mesenteroides (SF-type)Aguamiel (Agave salmiana)Dextran
(α-1→6 main, α-1→2 and α-1→3 branches)		Prebiotic fiber; food texture modifier[33]
Zymomonas mobilis B-14023Aguamiel/PulqueLevan–fructose
(β-2→6)		Thickener; prebiotic; edible films[34]
Weissella
confusa WCP-3a		PozolDextran
(α-1→6 main, α-1→3 branches); high glucosyltransferase activity		Prebiotic soluble fiber; viscosity modifier[35]
Leuconostoc citreum (pozol isolate)PozolInulin-type fructan (β-2→1)Low-calorie sweetener; prebiotic[29]
Table 4. Industrial applications of microbial exopolysaccharides in food systems.
Table 4. Industrial applications of microbial exopolysaccharides in food systems.
Type aMain monomersEPSMicroorganismApplicationProductRef.
HoPsGlucose (α-1,6; α-1,3 branches)DextranLeuconostoc mesenteroides, LactobacillusThickener, texture enhancer, stabilizerBakery, beverages, additives, confectionery[33,76]
HoPsFructose (β-2,6; β-2,1 branches)LevanBacillus subtilis, Zymomonas mobilisSweetener, bulking agent, prebioticPastries, beverages, dietary products[76]
HoPsGlucose (α-1,4; α-1,6 linkages)PullulanAureobasidium pullulansEdible films, coatings, encapsulationFruits, candies, biodegradable films[77,78]
HoPsGlucose (β-1,3 linkages)CurdlanAgrobacterium spp.Thermostable gels, stabilizerSurimi, sausages, instant soups[79,80]
HoPsGlucose (β-1,4 linkages)Bacterial celluloseKomagataeibacter xylinusTexture reinforcement, encapsulation matricesDesserts, kombucha, low-calorie products[81]
HoPsGlucose (β-1,3 backbone, β-1,6 branches)ScleroglucanSclerotium rolfsiiViscosity agent, stabilizerSauces, dressings, soups[73]
HePsGlucose and galactose (1:1)KefiranLactobacillus kefiranofaciensViscosity, texture in fermented beveragesKefir, yogurt[82]
HePsGlucose, rhamnose, glucuronic acidGellanSphingomonas elodeaGelling agent, stable matricesDesserts, confectionery, vegan gels[83]
HePsGlucose, mannose, glucuronic acidXanthanXanthomonas campestrisThickener, stabilizer, viscositySauces, dressings, gluten-free baking[70]
HePsβ-D-mannuronic acid, α-L-guluronic acidBacterial alginatePseudomonas, AzotobacterEncapsulation, gel formationMinimally processed fruits, molecular caviar[84]
a Abbreviations: HoPs, homopolysaccharides; HePs, heteropolysaccharides.
Table 5. Antibacterial mechanisms of microbial EPSs against gastrointestinal pathogens a,b.
Table 5. Antibacterial mechanisms of microbial EPSs against gastrointestinal pathogens a,b.
Main MechanismMode of ActionEPS (Strain)Outcome Against GI PathogensReference
Adhesion interferenceCompete with bacterial lectins or modify surface charge/hydrophobicity, preventing initial attachment to epithelial cells or plasticsEPS-cn2
(Lactobacillus casei NA-2)		↓ adhesion and ↓ early biofilm formation by E. coli O157:H7[107]
Direct structural disruptionChelate Ca2+/Mg2+ and disorganize LPS or peptidoglycan via highly charged, branched conformationsEPS 7–4
(Lactobacillus crispatus); EPS from L. plantarum PC715		Loss of membrane integrity and lysis of Salmonella Typhimurium[109]
Gene-expression modulation/aggregationDown-regulation of biofilm, motility and secretion-system genes; promotion of auto-aggregation that limits epithelial invasionEPS-cn2
(L. casei NA-2); EPS from Lacticaseibacillus rhamnosus GG		↓ LEE and other virulence genes in E. coli; aggregate formation and reduced invasion by S. Typhimurium[107,108]
Quorum-sensing signal sequestrationCaptures or blocks autoinducers (e.g., AI-2, AHL), reducing toxin production and biofilm maturationKefiran; EPS/extract from L. plantarum Z057↓ reporter bioluminescence, ↓ toxins and thinner biofilm in Vibrio parahaemolyticus and other pathogens[112,113]
a Symbols: ↓ = decrease/inhibition. b Abbreviations: EPSs, exopolysaccharides; LPS, lipopolysaccharide; LEE, locus of enterocyte effacement; AI-2, auto-inducer-2; AHL, N-acyl-homoserine lactone.
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Oviedo-León, J.F.; Higuera, A.R.; Yáñez-Fernández, J.; Hernández-Sánchez, H.; Castro-Rodríguez, D.C. Exploring Exopolysaccharides Produced in Indigenous Mexican Fermented Beverages and Their Biotechnological Applications. Fermentation 2025, 11, 463. https://doi.org/10.3390/fermentation11080463

AMA Style

Oviedo-León JF, Higuera AR, Yáñez-Fernández J, Hernández-Sánchez H, Castro-Rodríguez DC. Exploring Exopolysaccharides Produced in Indigenous Mexican Fermented Beverages and Their Biotechnological Applications. Fermentation. 2025; 11(8):463. https://doi.org/10.3390/fermentation11080463

Chicago/Turabian Style

Oviedo-León, Julián Fernando, Abril Ramírez Higuera, Jorge Yáñez-Fernández, Humberto Hernández-Sánchez, and Diana C. Castro-Rodríguez. 2025. "Exploring Exopolysaccharides Produced in Indigenous Mexican Fermented Beverages and Their Biotechnological Applications" Fermentation 11, no. 8: 463. https://doi.org/10.3390/fermentation11080463

APA Style

Oviedo-León, J. F., Higuera, A. R., Yáñez-Fernández, J., Hernández-Sánchez, H., & Castro-Rodríguez, D. C. (2025). Exploring Exopolysaccharides Produced in Indigenous Mexican Fermented Beverages and Their Biotechnological Applications. Fermentation, 11(8), 463. https://doi.org/10.3390/fermentation11080463

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