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Article

Exploring Volatile Profiles in Cactus-Based Fermented Beverages: Effects of Fermentation Method

by
César Ojeda-Linares
1,*,
Alejandro Casas
2,*,
Patricia Severiano-Pérez
3,
Marcela Sandoval-Velasco
1,
Yolanda M. García-Rodríguez
2 and
Francisco J. Espinosa-García
2
1
Centro de Ciencias Genómicas, Universidad Nacional Autónoma de México, Cuernavaca 62210, Mexico
2
Instituto de Investigaciones en Ecosistemas y Sostenibilidad, Universidad Nacional Autónoma de México, Morelia 58190, Mexico
3
Facultad de Química, Departamento de Alimentos y Biotecnología, Universidad Nacional Autónoma de México, Ciudad de México 04510, Mexico
*
Authors to whom correspondence should be addressed.
Fermentation 2025, 11(5), 275; https://doi.org/10.3390/fermentation11050275
Submission received: 29 March 2025 / Revised: 3 May 2025 / Accepted: 8 May 2025 / Published: 12 May 2025
(This article belongs to the Special Issue Nutrition and Health of Fermented Foods—4th Edition)
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Abstract

:
Mexico is home to a rich variety of fermented beverages made from both wild and domesticated plant species. Fermentation practices vary, with producers using either wild or inoculated techniques to obtain culturally valued final products. It is generally assumed that wild fermentations yield a greater diversity of volatile compounds compared to inoculated fermentations, as the latter tend to reduce microbial diversity throughout the process. However, this pattern remains largely unexplored in relation to the volatile profiles of traditionally fermented cactus-based beverages. Despite this assumption, comparative studies examining these profiles across different fermentation methods are scarce, especially given that these beverages are not produced under standardized conditions. To investigate this, we used GC-MS to characterize the aroma profile of colonche, a traditional fermented beverage made primarily from Opuntia streptacantha fruits. Colonche is produced by both wild and inoculated fermentation methods. In addition, a rapid sensory evaluation using the modified Flash Profile (mFP) technique was performed to evaluate flavor differences between the fermentation methods. A total of 55 volatile compounds were identified, with wild fermentations showing greater diversity (55) than inoculated fermentations (50). Most compounds overlapped, but five were unique to spontaneous fermentations, contributing to distinct sensory profiles. The mFP results also indicate that sensory attributes vary by fermentation type, with wild fermentations being more strongly associated with positive descriptors such as taste and smell, while inoculated samples have a distinctly pungent aftertaste. These findings highlight colonche not only as a reservoir of microbial diversity in arid regions but also as a culturally significant beverage with complex sensory attributes. Recognizing and preserving these attributes is essential for safeguarding traditional foodscapes.

1. Introduction

Traditional fermented products harbor complex microbial communities consisting primarily of lactic acid bacteria (LAB), acetic acid bacteria (AAB), and both Saccharomyces and non-Saccharomyces yeasts [1]. These microbial communities convert sugars to ethanol, lactic acid, acetic acid, carbon dioxide, and volatile compounds, among other by-products. Combined with substances derived from the raw materials, these transformations contribute to the unique sensory profiles of fermented products [2,3,4,5]. In addition, the complex interactions that occur during fermentation play a critical role in shaping the characteristics of the final product and ultimately determining its cultural acceptability [6,7].
Traditional fermented products have been prepared and consumed for generations, and production knowledge has evolved continuously. This ongoing refinement has diversified fermentation practices and product types across regions of the world [8,9]. Often rooted in home kitchens, fermentation processes vary in technique but commonly rely on wild fermentation, driven by endemic microbial communities naturally associated with substrates and their environments. Alternatively, controlled fermentation involves deliberately adding a pre-established microbial community (inoculum) to initiate the process [10]. Both approaches allow for the co-occurrence of bacteria and yeast, each contributing to the distinct flavors, textures, and aromas of the final product. Wild fermentation is often considered unpredictable due to variability in microbial activity and environmental conditions, as well as limited processing technology and microbial knowledge [8,11,12]. In contrast, controlled fermentation (preferred in industrial contexts) is valued for its consistency and efficiency. Nevertheless, traditional producers also use starter cultures, storing and reusing microbial communities to maintain desirable sensory characteristics and ensure reliable fermentations over time [13,14].
Flavor and aroma are key sensory attributes of many fermented products worldwide, resulting from the complex interactions between taste, smell, and texture [15,16]. Factors influencing these attributes can be broadly categorized as those influencing taste and those influencing aroma, the latter often referred to as aroma compounds. Odor and flavor perception are mediated by volatile organic compounds (VOCs) that interact with chemoreceptors in the olfactory epithelium of the nasal cavity [15,16]. In fermented products, VOCs are generated by multiple biochemical pathways involving microbial activity during fermentation. These compounds belong to several chemical classes, including alkenes, alcohols, ketones, benzenoids, pyrazines, sulfides, and terpenes [17,18,19,20]. These VOCs give rise to the characteristic flavors and aromas of each fermented product, creating a complex volatile space that has long been valued and pursued by mankind [21].
Differences in the production of aroma compounds during fermentation are largely due to yeast and bacterial enzyme activity, which also plays an essential role in their survival [22,23]. Some of these enzymes are secreted extracellularly to break down complex compounds and polymers in the surrounding environment to provide energy and nutrients. While some enzymes are directly responsible for the conversion of sugars to ethanol, others catalyze the formation of primary and secondary aroma compounds [22,23]. In addition, the substrates themselves (whether plant or animal) contain volatile compounds or precursors that contribute to the final sensory profile [24,25].
Due to the diversity of environmental microorganisms, wild fermentations typically produce a broader spectrum of extracellular enzymes compared to fermentations using monocultures of Saccharomyces cerevisiae or other pre-selected starter cultures [26,27]. Consequently, wild fermentations are expected to yield a greater diversity of volatile compounds, reflecting the metabolic activity of a wider range of microbial species and their associated enzymatic processes. Based on this premise, wild fermentations should exhibit greater volatile diversity because they involve a more complex microbial composition, ultimately resulting in a richer array of flavors. However, because wild fermentations are less controlled, the potential for spoilage-related off-odors also increases. In contrast, inoculated fermentations are thought to produce a narrower range of odor compounds due to reduced environmental heterogeneity and microbial diversity.
Recent studies have investigated these assumptions under semi-controlled and controlled conditions in several industrial products, including wine [28], sake, coffee, and cocoa [28,29,30,31]. In these cases, wild fermentations are often favored because of the increased complexity they impart to sensory attributes, whether positive or negative. However, this hypothesis has rarely been tested in non-controlled traditional fermented beverages.
Traditional Mexican Fermented Beverages (TMFBs) are produced at the household level for personal consumption or by small producers who sell them in local markets. These beverages are produced using different fermentation practices that result in different microbial compositions, making them ideal models for studying differences in volatile compound profiles between fermentation methods [32]. A total of 16 traditional beverages have been identified in Mexico [32], but many others likely exist or have existed, with few studies addressing their volatile profiles. One example is colonche, a cactus-based fermented beverage that has been traditionally made in central Mexico using prickly pear fruits from Opuntia streptacantha, also known as tuna cardona [33]. A recent study showed that the microbial community in colonche varies depending on the fermentation method, with wild fermentations having greater microbial diversity compared to inoculated fermentations [34]. This makes colonche a suitable model to test our hypothesis that wild-fermented colonche will have a greater diversity of volatile compounds compared to inoculated samples. Therefore, this study aims to characterize the volatile profiles of colonche samples produced with different fermentation techniques and to compare the volatile compound diversity between them.

2. Materials and Methods

2.1. Study System and Sampling Method

As mentioned above, a traditional beverage made by fermenting Opuntia streptacantha fruits, known as colonche, was chosen to test our hypothesis. Colonche is wildly fermented in some localities; the fruits are peeled on site and then fermented in clay pots for 12 h. Colonche can also be made by cooking the fruit to obtain a juice that is later fermented with a starter culture obtained from a previous fermentation. In both types of fermentation, the final product has a markedly high diversity of lactic acid bacteria (LAB) and a low diversity of yeasts [34]. However, the vessels used for wild fermentation harbor a high diversity of microorganisms such as Lactobacillus, Leuconostoc, Pediococcus, and the yeast Saccharomyces, which may contribute to the final sensory characteristics of the beverage [34].
Five samples of wild colonche (WC) were collected from local vendors at the Laguna de Guadalupe, Guanajuato, Mexico. Five samples of inoculated colonche (InC) were also collected from local vendors in Mexquitic de Carmona, San Luis Potosí (Figure 1). All the fermented samples were collected in duplicate in 50 mL sterile Falcon tubes and stored at −70 °C prior to analysis.

2.2. Extraction of Volatile Compounds and Gas Chromatography–Mass Spectrometry Analyses

Volatile compounds were extracted using a liquid–liquid extraction (LLE) technique. Each sample was prepared in a clean separating funnel, to which 20 mL of CH2Cl2 (Sigma-Aldrich-Fluka Co., St. Louis, MO, USA) was added, followed by 20 mL of colonche sample. The mixture was shaken manually for 10 min and then treated with anhydrous Na2SO4 (Sigma-Aldrich-Fluka Co., St. Louis, MO, USA) to remove residual water. Phase separation was achieved by centrifugation at 500 rpm for 20 min in a refrigerated centrifuge at 4 °C, after which the organic phases were collected.
The samples were then stored at −20 °C for 12 h, filtered, and centrifuged again under the same conditions to remove inorganic residues. The organic phases were concentrated under a nitrogen gas stream to a final volume of 1 mL and stored in 5 mL clear glass autosampler vials at −20 °C until gas chromatography–mass spectrometry (GC-MS) analysis. For each sample, an aliquot (500 μL) was vortex mixed with 500 μL of a tetradecane solution (0.25 mg mL−1) and further concentrated to 250 μL under nitrogen gas.
Volatile compound analysis was performed using a gas chromatograph (Agilent 6890A, Santa Clara, CA, USA) coupled to a mass spectrometer (Agilent 5973N) with an ECWAX10 capillary column (30 m × 0.25 mm i.d. × 0.25 µm film thickness, Agilent, Santa Clara, CA, USA) at the Laboratory of Chemical Ecology and Agroecology, Institute of Research on Ecosystems and Sustainability of the National University of Mexico (IIES, UNAM, Ciudad de México, Mexico). Helium was used as the carrier gas at 7.67 psi, with a constant flow rate of 1.0 mL min−1. Instrument conditions were programmed as follows: the front inlet temperature was maintained at 200 °C, the interface temperature at 250 °C, the scan range at 35–500 m/z, the electrical potential at 70 eV ionization voltage, and the injector pressure at 5 psi. The initial oven temperature was set at 44 °C for 2 min and increased by 15 °C every minute until 80 °C was reached for 1 min.

2.3. Volatile Profile Characterization

Volatile compounds were identified by comparison of retention indices (RIs) with reference values. The structural assignment of the compounds was based on their mass spectral matching with the National Institute of Standards and Technology (NIST2011.L) [35] mass spectral library. Only compounds with a spectral identity/quality match of 90% or greater were retained for identification. The concentration of volatile and semi-volatile compounds was determined using an internal standard consisting of a tetradecane solution (0.25 mg mL−1). Only peaks with a signal-to-noise (S/N) ratio ≥ 2 were included, based on the limit of detection (LOD) criteria. Compounds with a relative abundance below 0.09% were considered traces and excluded from further quantitative analysis. Concentration values were then transformed using the Box–Cox method to normalize the data.
Kovats retention indices (RIs) were calculated based on the retention times of a series of n-alkanes (C8–C26) and compared with values reported in the literature [36]. The odor and aroma profiles of the identified compounds were characterized using data from the PubChem database [37].

2.4. Sensory Profiling

A rapid sensory evaluation technique, modified Flash Profile [38,39], was used to evaluate the colonche samples. The evaluation was performed by 25 panelists, with an average age of 22.5 years, all of whom were trained in descriptive methodology at the Sensory Evaluation Laboratory of the Faculty of Chemistry of the National University, UNAM. Although the panelists had no previous experience in evaluating colonche, they had been trained in evaluating other fermented beverages, including pulque (a fermented agave sap), yogurt, and probiotic-fermented beverages. Exclusion criteria for panelists included pregnancy, lactation, regular smoking, and having had an olfactory or gustatory impairment due to SARS-CoV-2. Attributes were generated by the panelists and selected by consensus to ensure a standardized list of attributes. Panelists then rated the attributes that were most relevant or strongest in the samples.
FIZZ software (version 2.3, Biosystems, France) was used to design the questionnaire and score the samples. A structured scale ranging from 1 to 9 was used to quantify the intensity of the attributes, with 1 representing the lowest intensity and 9 representing the highest intensity. Throughout the evaluation process, the samples were kept in a refrigerator at 4 °C.
Colonche has a short optimal consumption period, and its sensory profile requires fresh samples. Therefore, due to the inability to transport all the samples fresh, only four samples were analyzed, two wild and two inoculated. Samples (200 mL) were presented in clear plastic cups labeled with random three-digit codes and brought to room temperature 5 min before evaluation. The order of presentation was randomized, and all samples were presented simultaneously to the panelists. Water was provided for mouth rinsing between samples to minimize carryover effects. While this approach ensured the integrity of the sensory characteristics, the limited sample size may affect the generalizability of the results and should be considered a limitation of the study.
The modified Flash Profile (mFP) evaluation was performed at the Sensory Evaluation Laboratory of the Faculty of Chemistry, UNAM, in individual sensory booths according to ISO standards (ISO, 2007) [40]. The sensory evaluation sessions were designed using FIZZ software (Biosystems, version 2.3, Acquisition and Judge Module, Courtenon, France). All panelists provided written informed consent before participation. In the first modified FP session, panelists independently generated a list of non-hedonic attributes (appearance, mouthfeel, taste, and odor) to differentiate the samples. All panelists discussed the generated attributes together to ensure a common understanding; attributes with only one mention were eliminated.

2.5. Statistical Analysis

To evaluate the volatile profiles of the colonche samples under different fermentation practices, statistical analyses were performed using R software (v4.5.0, R Core Team, Vienna, Austria). The data set was normalized with the mean and standard deviation using the scale() function of the base package v3.6.2 [41]. Pearson correlation coefficients were calculated for the quantitative variables using the corrplot v0.84 package [42]. Variables with correlations <−0.85 or >0.85 were excluded to avoid potential overestimation of the results.
To assess the differences between the two fermentation practices, an ANOVA was performed to test for differences in volatile concentrations by fermentation type. A PERMANOVA was also performed to statistically test whether the total volatile profiles were significantly different between the two fermentation practices using the vegan v2.6-4 package [43]. Before the PERMANOVA, we tested for the homogeneity of the multivariate dispersion using the betadisper() function in the vegan package. The analysis showed no significant differences in dispersion between the groups, confirming that the assumption of equal dispersion was met. A Wilcoxon test was performed to contrast whether the mean concentration differed between the two fermentation methods.
Finally, a General Procrustes Analysis (GPA) was used to assess the panel consensus and to visualize the relative sensory positioning of the attributes [38]. The analysis of the modified Flash Profile (mFP) data was performed with XLSTAT 2012, Addinsoft, version 10.0, and a PCA was performed with the consensus ratings of the panelists to represent the differences between the fermentation treatments.

3. Results

3.1. Characterization of Organic Compounds

A total of 55 volatile organic compounds (VOCs) were identified in the analyzed colonche samples (Table 1). Of these, 50 compounds were common to both fermentation types, while 5 additional VOCs were detected exclusively in the wild fermentation samples. Compounds that showed the most significant differences between wild and inoculated colonche samples include alcohols such as 3-methyl-1-butanol, 1-propanol-2-methyl, and 3-heptanol-4-methyl; the alkane dodecane; the alkene 2-undecene-5-methyl; the aromatic compound ethylbenzene; and the carboxylic acid octanoic acid (Table 1). These compounds are major contributors to the sensory characteristics of the final product; however, other compounds that were present in the samples contribute to the sensory complexity.
In the colonche samples, 10 different alcohols were identified in samples from both fermentation techniques, with 1-butanol and 3-methyl being the most abundant in both. Phenol, 2,4-bis(1,1-dimethylethyl), was the only phenolic compound identified in the samples for both fermentation techniques, being more abundant in the wild fermentation. Eleven alkanes were detected in the colonche samples. However, Tridecane, 3-methyl, and Decane-2-methyl were absent in the inoculated samples. Four alkenes, two aldehydes, six aroma compounds, four carboxylic acids, three ketones, and one lactone were also identified in the colonche samples. Notably, 11 esters were identified in the colonche samples, whereas ethyl (E)-octadec-9-enoate and oleic acid were not detected in the inoculated samples.

3.2. Differences in the Composition of Organic Compounds Between the Fermentation Practices

The results of the ANOVA show significant differences in the volatile concentrations by type of fermentation in the colonche samples (Table 2). Similarly, the PERMANOVA test shows a difference in the composition of the volatile compounds in the colonche samples between the fermentation practices (Table 3).
Furthermore, the Wilcoxon test confirms significant differences in the mean concentration between the two fermentation methods, as shown in Table 1. In particular, the differences are significant for compounds such as 2-methyl, 1-propanol, 2-undecene, 5-methyl, ethylbenzene, dodecane, 1-butanol, 3-methyl, and octanoic acid. These results confirm our hypothesis of a higher number of volatiles in wild fermentations and a reduction in inoculated fermentations.

3.3. Sensory Differences Between Fermentation Practices in the Colonche Samples

A total of 95 and 56 attributes were generated for the two inoculated samples of colonche, with 84 and 62 for the wild fermented colonches. From these, synonyms and those that perceived by only one judge were eliminated. The 48 consensus sensory attributes selected for the modified Flash Profile are listed in Table 4. Inoculated colonche samples had a greater number of sensory attributes (43) compared to wild fermentations (38). Inoculated samples had more attributes related to odor and taste than wild colonche samples.
The PCA (Figure 2) shows that the first component explains 38.50% of the variance, while the second component explains 34.96%. The wild colonche was positively correlated with component 1 and had a different sensory profile than the inoculated colonche. Since these are traditionally produced beverages, similarities can be observed between O. streptacantha cactus prickly pear fruits based on their fermentation processes. However, their sensory profiles are not identical.
Wild colonche samples were characterized by their appearance, such as particulate matter, a prune aroma, and bitter and spicy notes, accompanied by a retronasal sensation and prune and honey flavors. Other wild colonche samples had a precipitated appearance and a more intense color, along with a full-bodied and pseudothermal mouthfeel. These samples also exhibited an intense alcoholic, dairy, tepache (a fermented pineapple beverage), and vinegar aroma, and flavors with alcoholic, fermented, fresh, fruity, and sweet notes.
In contrast, the inoculated samples had a thick and effervescent mouthfeel, with a fruity, punchy, acidic, and fresh aroma. Their flavor was characterized by intense red fruit notes with a slightly pungent aftertaste. Some inoculated samples were also characterized by a bright appearance, an aroma of dried fruit (such as walnut) and red fruit, and a complex flavor profile including sweet, bitter, pungent, acidic, astringent, citrus, and vinegar notes. In addition, the alcoholic sensation and the astringent aftertaste were more pronounced in the inoculated samples.

4. Discussion

4.1. Fermentation Practices and Traditional Ecological Knowledge: Different Outcomes That Can Be Tasted and Smelled

As mentioned before, the aim of this study was to characterize the volatile profiles of colonche, a traditional fermented beverage made from prickly pear (Opuntia streptacantha), and to compare the diversity and abundance of volatile organic compounds (VOCs) between wild and inoculated fermentation techniques. Our results show clear differences in the number and concentrations of volatiles produced under each fermentation condition. These results confirm the reduction in diversity and concentration of VOCs in inoculated samples.
As previously reported, colonche fermentation practices differ not only in the composition of microbial assemblages but also in the relative abundance of bacterial and fungal taxa within the fermenting communities, with wild fermentations showing the most pronounced differences [34]. In this sense, the greater microbial diversity observed in wild fermentations likely drives the production of a wider range of volatile compounds, increasing sensory complexity and potentially contributing to the traditional flavor identity of colonche. Similar patterns have been documented in other fermented products, where microbial diversity correlates with the production of volatile organic compounds (VOCs) [21,44]. In wild colonche fermentations, non-Saccharomyces yeasts were dominant during the early stages of fermentation but declined as S. cerevisiae became dominant. The presence of oxidative and weakly fermentative non-Saccharomyces species in these early stages may contribute important volatile precursors that persist in the final product and shape sensory differences between fermentation types. This succession pattern and its impact on volatile profiles has also been reported in other fermented products such as traditional beverages, where early microbial colonizers play a critical role in defining the organoleptic profile of the final beverage [45,46].
From an ethnographic perspective, fermentation is a culturally meaningful process guided by empirical observation and inherited knowledge. Although Claude Lévi-Strauss [47] classified fermented foods in his culinary triangle as “rotten” (associated with uncontrolled decay), this perspective fails to capture the intentionality and expertise behind traditional fermentation techniques. Colonche, like other fermented beverages worldwide, is the result of deliberate microbial management aimed at achieving culturally valued sensory outcomes [32,33,34]. For example, using starters reflects unconscious microbial selection, shaping the fermentative environment and sensory consistency across batches. Indeed, VOCs in the inoculated samples showed less variation between batches and also less variation in the mFP. Thus, the use of starters by producers may be a practical mechanism to achieve a consistent sensory profile for colonche. These differences could reflect the activity of different microbial consortia (yeast and bacteria) and correspond to the traditional sensory expectations of each producer, the production techniques, and the ecological knowledge embedded in local practices.

4.2. Alcohols and Organic Compounds, Metabolic By-Products, and Sensory Markers

Alcohol is part of the starter set of VOCs in several fermented beverages, highlighting the microbial metabolism that occurs during fermentation [21,48]. Among the alcohols identified, 3-methyl-1-butanol had the highest concentration in both colonche fermentation samples. It is associated with alcoholic, pungent, and molasses flavors [49]. It is also associated with fruity, banana-like aromas [49] and is a known product of Saccharomyces cerevisiae and other wine-related fermentative wild yeasts [50,51]. This compound was significantly more abundant in wild fermentation samples (p < 0.05), consistent with findings from spontaneous pulque fermentations where more diverse yeast communities resulted in richer aromatic profiles but also lower alcohol content [52,53]. Nevertheless, as mentioned in the previous section, alcohols are among the VOCs that statistically differentiate the final product across fermentation techniques. Alcohols were found at lower levels in the inoculated samples. This is consistent with findings from co-fermentation studies, where lactic acid bacteria, due to their higher stress tolerance, can dominate and inhibit yeast activity, delaying ethanol production [54]. The microbial interactions may help explain the relatively low ethanol content in some Mexican fermented beverages, such as tepache [55], compared to others, such as pulque [56] and taberna [57].
These sensory attributes were reflected in the mFP, where colonche panelists associated fruity, banana-like, and molasses aromas with high-quality beverages. Indeed, in a previous study, the producers recognized that fruity attributes in wild fermentations were critical to colonche quality [33].
Organic compounds such as 1,3-butanediol, isoprenol, (E)-2-hexenol, 1-hexanol-2-ethyl, 3-heptanol-4-methyl, and 2-phenylethyl alcohol were detected not only in red wines but also in wine and sherry vinegar. Phenylethyl alcohol, known for its rosy and honey-like aromas [49,58], was significantly more abundant in wild colonche samples but also appeared in the inoculated samples. These compounds result from yeast metabolism in wine and the activity of yeasts [59,60], LAB, and AAB in vinegars and other beverages [61]. LAB groups such as Leuconostoc, Lactococcus, and Lactobacillus genera can produce organic acids that result in intense fermented and fruity aromas that can be perceived in red wines with low alcohol content [61,62]. Higher alcohol levels typically result in a negative, solvent-like aroma, while low concentrations of higher alcohols have ideal complexity [60,61].
This characteristic has been previously recognized by producers, who do not primarily associate high-quality colonche with an alcoholic odor, but rather with a complex interplay of sensory attributes such as texture, color, and flavor [33]. In the modified Flash Profile (mFP), panelists identified characteristics such as the intense color and grainy texture of inoculated samples as contributing to the perceived differences between wild and inoculated colonche. However, the most distinctive characteristics that allowed for the differentiation between the two fermentation types were olfactory: red fruit, caramel notes, and fruit punch-like aromas were particularly associated with the inoculated samples and similarly recorded in the production of a prickly pear wine [63,64].

4.3. Phenolics and Spice-Linked Volatiles

A single phenolic compound was identified in the colonche samples analyzed, phenol, 2,4-bis(1,1-dimethylethyl), which is associated with smoky, clove, and roasted aromas and flavors. Although it was found to be more abundant in wild fermentations, its appearance in the inoculated samples may be related to the practices used by the producers to prepare the inoculated colonche, as they add clove or cinnamon during the boiling of the prickly pear cactus fruit. Cinnamon and clove have been reported to have antifungal and antimicrobial properties, mostly due to the presence of inhibitors; they have also been characterized for their flavor-enhancing qualities and added in several fermented products [65,66]. Indeed, the addition of these plants has been characterized to have impacts on the microbial community of colonche, reducing the bacterial groups [34]. The presence of this phenolic compound has been reported in the fermented juice of prickly pear but not in the fruit [67]. Nonetheless, for Mexican fermented beverages produced with agave species, such as pulque, raicilla, bacanora, tequila, and mezcal, phenolic compounds have been described as key elements involved in their authenticity [53].
The mFP results highlight the presence of cactus and prickly flavors associated with the raw material (red tuna) used to prepare colonche. Particularly spicy, woody, and nutty odor notes were more pronounced in the inoculated samples, potentially due to the inclusion of fruit seeds throughout fermentation. This finding parallels reports from tepache production, where added spices and fruit skin layers contribute to complex ester and phenolic profiles [68].

4.4. Acids and Flavor Outcomes

Acids play an important role in traditional fermented products, as they can be produced by bacterial groups and yeasts during fermentation [69]. In wine, S. cerevisiae can synthesize mainly hexanoic and octanoic acids in high amounts, but also pentanoic, decanoic, and 3-methylbutanoic acids. In the colonche samples, these groups have been previously described [34], and the presence of this compound could be related to the acid and vinegar odors and the intense and astringent taste that was also recorded in the mFP. Hexanoic acid, known for its rancid, cheesy aroma, was significantly higher in inoculated colonche (p < 0.05). This compound can be produced mainly by S. cerevisiae, but also by bacterial groups such as LAB and AAB [70,71], all previously identified in colonche fermentations [34]. Indeed, the inoculum has been previously studied and is composed of a prominent dominance of S. cerevisiae species and LAB groups such as Leuconostoc, Lactobacillus, and Pediococcus [34].
The abundance of this compound in inoculated samples may reflect differences in microbial succession induced by controlled inoculation techniques. Similar acid profiles have been reported in pulque and other low-alcohol beverages, where LAB and AAB dominate in the late fermentation stage [55,56,57].

4.5. Aldehydes and Ketones: Compounds from Raw Materials and Fermentation Processes

Aldehydes and ketones generally contribute to the aroma of the beverage, providing smooth and harmonious flavors [72]. Most aldehydes are produced by the deamination and decarboxylation of amino acids during fermentation [73]. Among them, (E)-2-hexenal was identified in both types of fermentation, but was more abundant in the sample of wild fermentations. This compound was previously detected in red prickly pear fruits from Opuntia robusta, a species closely related to O. streptacantha [67]. Thus, its presence may be related to the extraction process of prickly pear fruit juice for wild fermentation. The ketone 3-hydroxy-2-butanone, also known as acetoin, is a well-known compound in food flavoring. It is primarily produced during fermentation by Kluyveromyces spp. and is associated with buttery, milky, and caramel-like aromas [71]. In particular, Kluyveromyces spp. and Kluyveromyces marxianus have been previously isolated from colonche samples [34]. This ketone also contributes to flavor development in fermented dairy products and beverages and has been reported in traditional pulque [53].

4.6. Esters, Fruity, and Floral Complexity in Colonche Samples

Esters were abundant in both wild and inoculated colonche samples, although several esters were uniquely present in wild fermentations. Compounds such as ethyl 9-hexadecenoate, butanedioic acid esters, and (S)-2-hydroxy-propanoic acid ethyl ester contribute fruity, floral, and buttery aromas [74,75]. Although not all differences were statistically significant (p > 0.05), a general trend toward greater ester diversity was observed in wild samples. This diversity likely reflects the broader microbial consortia active in traditional wild fermentations. Esters are primarily formed through microbial esterification processes, which are influenced by both substrate availability and the diversity of fermentative microbes. Traditional fermentation vessels, such as unglazed clay, can maintain stable communities of yeast and bacteria [34], such as Lactobacillus, Leuconostoc, Pediococcus, and Saccharomyces, that enhance ester synthesis. These microbial reservoirs, shaped by longstanding fermentation practices, contribute to the complex and distinctive sensory profiles associated with wild colonche fermentations.

5. Conclusions

The production of traditional fermented beverages is often perceived as heterogeneous and unpredictable. However, producers employ a variety of strategies to achieve consistent and desirable final products. Fermentation practices serve as key methods to shape the sensory profile of these beverages, ensuring an optimal product despite inherent variability. The current results confirm that wild fermentations produce a greater diversity of volatile compounds, while inoculated fermentations show a reduction in the number of volatiles. Nevertheless, the core volatile organic compounds (VOCs) remain consistent across both fermentation methods, highlighting a common fundamental composition in colonche.
Both fermentation techniques contribute to a complex spectrum of sensory attributes, including flavor, aroma, and aftertaste, which ultimately influence the organoleptic properties of the final product. Importantly, these differences in fermentation methods extend to consumer perception, as demonstrated by the modified Flash Profile (mFP), where colonche samples could be classified based on fermentation type.
This study contributes to the growing body of research on traditional fermented beverages by integrating chemical and sensory analyses to characterize wild and inoculated colonche fermentations. It highlights the role of traditional practices in maintaining sensory diversity, demonstrating that artisanal knowledge and fermentation environments can shape the biochemical and perceptual qualities of beverages. These findings have broader implications for food heritage conservation, microbial ecology, and the valorization of traditional fermentation techniques in contemporary food systems.
Future research should explore producers’ sensory preferences and the microbial consortia associated with different fermentation environments. Studies employing metagenomic or metabolomic approaches could provide deeper insights into the microbial dynamics shaping volatile profiles. Additionally, examining how changes in fermentation vessels and Opuntia varieties affect aroma development could offer valuable knowledge for both scientific understanding and the protection of traditional fermented beverages.

Author Contributions

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

Funding

This research was funded by Becas Posdoctorales por México (SECIHTI, formerly CONAHCYT) via academic and financial support provided to the first author. This study was also supported by CONACHYT (now SECIHTI) research projects A1-S-14306 and CBF-2025-I-1572 and PAPIIT, DGAPA, UNAM research projects IN209024, IN206520, and IN224023.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

The data that support the findings of this study are available from the corresponding author, C.O.-L., upon reasonable request.

Acknowledgments

The authors thank Becas Posdoctorales por México (SECIHTI, formerly CONAHCYT) for the academic and financial support provided to the first author. Special thanks go to Cristian Cervantes and Patricia Lappe for their valuable feedback on the initial draft. Finally, we extend our gratitude to all of the local producers who preserved and shared the creativity of fermentation. Special thanks go to Gallina (Martín) and Anita Prado the coloncheros; without them, this study would not be possible. Grammarly was used to check the text. The version is 14.1235.0.

Conflicts of Interest

The authors declare no conflicts of interest.

Abbreviations

The following abbreviations are used in this manuscript:
FPFlash Profile
TMFBTraditional Mexican Fermented Beverage
VOCsvolatile organic compounds
LABlactic acid bacteria
AABacetic acid bacteria
WCwild fermented Colonche
InCinoculated Colonche
LLEliquid–liquid extraction

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Fermentation 11 00275 g001
Figure 1. The sampled study area in Mexico. The red diamond marks Laguna de Guadalupe, Guanajuato, where the wild colonche samples were collected. The green dot marks Mexquitic de Carmona, San Luis Potosí, where the inoculated colonche samples were collected from local vendors. The map was created using the open-source software QGIS v3.4 (https://qgis.org/es/site/, accessed on 1 February 2025). Source: INEGI, conjunto de datos vectoriales.
Figure 1. The sampled study area in Mexico. The red diamond marks Laguna de Guadalupe, Guanajuato, where the wild colonche samples were collected. The green dot marks Mexquitic de Carmona, San Luis Potosí, where the inoculated colonche samples were collected from local vendors. The map was created using the open-source software QGIS v3.4 (https://qgis.org/es/site/, accessed on 1 February 2025). Source: INEGI, conjunto de datos vectoriales.
Fermentation 11 00275 g001
Fermentation 11 00275 g002
Figure 2. The PCA of sensory attributes for two types of colonche fermentations. The blue lines represent appearance-related attributes, while the green lines correspond to odor-related attributes. The red lines represent mouthfeel attributes, the pink lines represent taste-related attributes, and the yellow lines represent aftertaste sensations. * highlights the tepache odor recorded in the mFP.
Figure 2. The PCA of sensory attributes for two types of colonche fermentations. The blue lines represent appearance-related attributes, while the green lines correspond to odor-related attributes. The red lines represent mouthfeel attributes, the pink lines represent taste-related attributes, and the yellow lines represent aftertaste sensations. * highlights the tepache odor recorded in the mFP.
Fermentation 11 00275 g002
Table 1. Mean concentration values and mean squares of ANOVA of volatile compounds evaluated in colonche by two types of fermentation.
Table 1. Mean concentration values and mean squares of ANOVA of volatile compounds evaluated in colonche by two types of fermentation.
Volatile CompoundtR bColonche Samples
Wild aInoculated a
1,3-butanediol8.950.53 ± 0.540.28 ± 0.31
Alcohols(E)-2-hexenol10.390.22 ± 0.570.27 ± 0.76
1-butanol5.240.29 ± 0.280.19 ± 0.26
3-methyl,1-butanol ***6.304.29 ± 4.641.39 ± 2.80
2,3-butanediol8.550.65 ± 1.000.31 ± 0.67
1-hexanol,2-ethyl13.610.08 ± 0.110.00 ± 0.01
1-Propanol,2-methyl ***6.281.02 ± 1.420.23 ± 0.45
Isoprenol8.750.02 ± 0.030.00 ± 0.01
3-Heptanol,4-methyl **10.390.16 ± 0.310.00 ± 0.01
Phenylethyl alcohol14.910.22± 0.250.12 ± 0.13
PhenolsPhenol, 2,4-bis(1,1-dimethylethyl)32.830.72 ± 0.890.42 ± 0.74
Decane-2-methyl8.550.10 ± 0.13Nd
Dodecane ***12.500.10 ± 0.100.02 ± 0.06
Dodecane, 2,6,10-trimethyl6.100.10 ± 0.220.00 ± 0.01
Alkanes Dodecane, 2,4-dimethyl12.040.14 ± 0.130.06 ± 0.10
Heptadecane19.080.10 ± 0.250.02 ± 0.02
Hexadecane19.480.04 ± 0.060.03 ± 0.05
Pentadecane13.830.10 ± 0.080.07 ± 0.08
Hexadecane, 2,6,10,14,-tetramethyl12.350.09 ± 0.150.0 ± 0.00
Hexadecane, 4-methyl19.000.40 ± 1.080.03 ± 0.05
Tridecane, 3-methyl24.450.01 ± 0.02Nd
Tridecane, 5-methyl18.120.04 ± 0.050.01 ± 0.02
Alkens1-Dodecene12.000.04 ± 0.120.00 ± 0.01
2-Undecene, 5-methyl ***4.620.46 ± 0.480.23 ± 0.45
1-Eicosense28.980.04 ± 0.090.01 ± 0.02
10-Eicosene, (E)30.450.03 ± 0.050.01 ± 0.01
Aldehydes Hexanal24.190.07 ± 0.070.01 ± 0.02
(E)-2-hexenal18.303.09 ± 7.670.64 ± 1.14
1,3-Dimethylbenzene4.980.24 ± 0.390.27 ± 0.39
Benzaldehyde, 4-methyl48.800.06 ± 0.070.05 ± 0.13
Aroma compoundsBenzyl-alcohol27.010.15 ± 0.230.20 ± 0.35
Phenylethyl alcohol32.844.08 ± 3.431.90 ± 3.36
prop-2-enyl 2-phenylacetate4.500.38 ± 0.790.06 ± 0.11
Ethylbenzene ***4.870.36 ± 0.370.18 ± 0.30
Acetic acid12.470.04 ± 0.712.64 ± 5.37
Carboxylic acidsHexanoic acid6.700.07 ± 0.081.20 ± 2.10
Decanoic acid18.410.04 ± 0.070.07 ± 0.19
Octanoic acid **33.510.77 ± 1.760.06 ± 0.13
EstersIsoamyl lactate16.380.07 ± 0.100.00 ± 0.02
ethyl (E)-octadec-9-enoate40.370.15 ± 0.34Nd
Ethyl linolelaidate47.400.20 ± 0.200.13 ± 0.19
Butanedioic acid7.971.68 ± 2.150.77 ± 1.11
Decanoic acid, ethyl ester18.610.50 ± 0.650.73 ± 2.06
Dodecanoic acid29.490.02 ± 0.040.00 ± 0.02
Ethyl 9-hexadecenoate10.261.23 ± 2.96 0.07 ± 0.15
Ethyl dl-2-hydroxycaproate40.290.30 ± 0.600.06 ± 0.09
Oleic acid48.620.56 ± 1.61Nd
Hexadecanoic acid43.761.03 ± 1.810.10 ± 0.15
Linoleic acid47.600.52 ± 0.870.07 ± 0.22
Vinyl hexanoate26.190.20 ± 0.26 0.02 ± 0.05
Propanoic acid, 2-hydroxy-, ethyl ester, (S)9.375.69 ± 9.152.34 ± 3.83
3-Hydroxy-2-butanone7.120.40 ± 0.43 0.19 ± 0.43
Ketones4-Hydroxy-4-methyl-2-pentanone24.340.06 ± 0.06 0.05 ± 0.06
7-Tridecanone11.250.05 ± 0.10 Nd
LactonesButyrolactone18.230.07 ± 0.16 0.01 ± 0.02
a Concentration of compounds in mg/L; b RT: retention time in min; Nd: not detected. Not significant * at p > 0.05; significant ** at p > 0.05; significant *** at p > 0.01.
Table 2. ANOVA test results from colonche samples.
Table 2. ANOVA test results from colonche samples.
DfSum of SqsFPrSig
Fermentation type140.215.349.56 × 10−5***
Volatile54675.14.76<2 × 10−16***
Residual10442736.7
Not significant * at p > 0.05; significant ** at p > 0.05; significant *** at p > 0.01.
Table 3. PERMANOVA test results from colonche samples.
Table 3. PERMANOVA test results from colonche samples.
DfSum of SqsR2FSig
Model10.5280.102.040.025 *
Residual184.620.89
Total195.151.00
Not significant * at p > 0.05; significant ** at p > 0.05; significant *** at p > 0.01.
Table 4. Sensory attributes identified in the colonche samples for each type of fermentation.
Table 4. Sensory attributes identified in the colonche samples for each type of fermentation.
Attributes Colonche
WildInoculated
AppearanceColor++
Brightness++
Grainy++
Precipitated++
Bubbling+−
OdorAcidic++
Alcoholic++
Sweet++
Prune++
Fruity++
Tepache++
Fresh++
Bitter++
Vinegar++
Intense++
Red fruits−+
Caramel−+
Spiced+−
Tejocote−+
Fruit punch−+
Woody−+
Nutty−+
MouthfeelFull-bodied++
Warming++
Viscous−+
Retronasal+−
Sparkling+−
Effervescent−+
FlavorAlcohol++
Sweet++
Honey++
Caramel++
Prune++
Fruity++
Acidic++
Fermented++
Fresh+−
Prickly pear fruit+−
Cacti/Biznaga+−
Punch fruit++
Astringent++
Intense++
Red fruits−+
Vinegar−+
Bitter−+
Pungent++
Citric−+
AftertasteAstringent++
Metallic++
Pungency++
Total503843
The “+” symbol represents the presence of the attributes characterized by the panelist in the mFP and the “−“ symbol represents the absence of these attributes in the samples.
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Ojeda-Linares, C.; Casas, A.; Severiano-Pérez, P.; Sandoval-Velasco, M.; García-Rodríguez, Y.M.; Espinosa-García, F.J. Exploring Volatile Profiles in Cactus-Based Fermented Beverages: Effects of Fermentation Method. Fermentation 2025, 11, 275. https://doi.org/10.3390/fermentation11050275

AMA Style

Ojeda-Linares C, Casas A, Severiano-Pérez P, Sandoval-Velasco M, García-Rodríguez YM, Espinosa-García FJ. Exploring Volatile Profiles in Cactus-Based Fermented Beverages: Effects of Fermentation Method. Fermentation. 2025; 11(5):275. https://doi.org/10.3390/fermentation11050275

Chicago/Turabian Style

Ojeda-Linares, César, Alejandro Casas, Patricia Severiano-Pérez, Marcela Sandoval-Velasco, Yolanda M. García-Rodríguez, and Francisco J. Espinosa-García. 2025. "Exploring Volatile Profiles in Cactus-Based Fermented Beverages: Effects of Fermentation Method" Fermentation 11, no. 5: 275. https://doi.org/10.3390/fermentation11050275

APA Style

Ojeda-Linares, C., Casas, A., Severiano-Pérez, P., Sandoval-Velasco, M., García-Rodríguez, Y. M., & Espinosa-García, F. J. (2025). Exploring Volatile Profiles in Cactus-Based Fermented Beverages: Effects of Fermentation Method. Fermentation, 11(5), 275. https://doi.org/10.3390/fermentation11050275

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