## 4 Discussion
AAB are believed to produce numerous compounds, including aldehydes, that influence the flavor profile of cacao products [44; 45], and Benzaldehyde is known to yield an almond-like aroma and is considered a major odor-active compound commonly used as a fine-favor index [4]. The LAB _Limosilactobacillus fermentum_ and the yeast _C. metapsilosis_ were detected by culture-based methods at 48 h of spontaneous fermentation. A similar trend was observed by NGS analyses that identified LAB, such as _Limosilactobacillus fermentum_, and yeast, such as _Candida glabrata_, among the predominant microorganisms at this stage of the spontaneous fermentation process. _Candida metapsilosis_ has been identified in only a few cacao fermentation studies [46; 47], and the inoculation of this yeast yielded a significant production of benzaldehyde and butanal-3-methyl at this point of the fermentation process. Both metabolites contribute to almond and chocolate aromas, respectively [48]. However, butanoic acid was also significantly produced by _C. metapsilosis_. This volatile has been associated with undesirable, rancid aromas [49]. Similarly, _Limosilactobacillus fermentum_also contributed to the production of butanal-3 methyl in cacao beans. The formation of this aldehyde has been attributed to the degradation of amino acids by LAB during fermentation [36]. LAB showed dominance after 72 h of spontaneous fermentation, as revealed by the culture-based approach, in which _Limosilactobacillus fermentum_ was the most frequently isolated microorganism. Similarly, a high relative abundance of LAB, including _Limosilactobacillus fermentum_ and _Lacticaseibacillus zeae_, was detected by NGS at this fermentation period. The results are in agreement with previous reports showing that heteroformative LAB species are typically detected after 24-72 h of spontaneous fermentation due to the development of suitable growth conditions [1, 50, 51, 52]. The inoculation of LAB into previously fermented (72 h spontaneous fermentation) cacao beans contributed to the formation of 1-butanol-3 methyl and ethanol, 2-nitro, which can contribute to malty and chocolate aromas. The production of both compounds could be attributed to the metabolism of hexose sugars by LAB, carried out to produce a wide variety of metabolites, including higher alcohol [53]. After 96 h of spontaneous fermentation, _B. subtilis_ and _B. amyloliguefaciens_ were detected through culture-dependent techniques. Other researchers [54, 55, 50] have also found _Bacillus_ spp. at the late stages of cacao fermentation. The increased availability of oxygen at this stage of the fermentation could promote the growth of these species [1]. In contrast, NGS revealed an increment in the relative abundance of _Limosilactobacillus fermentum_ after 96 h of fermentation. The presence of LAB contributes to maintaining several metabolic pathways throughout fermentation and influences the growth of AAB [18, 56], which were also detected at this fermentation point (_A. ghanensis_ and _A. pasteurianus_). Moreover, the detection of yeast, such as _P. kudriavzevii_ and _C. metapysilosis_, observed by the NGS and culture-base methods, respectively, after 96 h of spontaneous fermentation is in agreement with recent studies reporting the presence of yeast in late fermentation stages [57, 58]. The production of volatiles, such as ethanol, benzophenone, ethylphenyl acetate, 3-Penten-2-ol, benzaldehyde, phenylvinylacetylene, caffeine, acetic acid and benzeneacetic acid, ethyl ester, was linked to the inoculation of _Bacillus_ species at this fermentation period, and was associated with floral/herbal, alcoholic and almond aromas (Table 3). Furthermore, cacao beans inoculated with _C. metapsilosis_ yielded a significant concentration of hydrocarbons, such as hexadecane and esters, including ethylphenyl acetate and B-phenylethyll formate. Several esters have been associated with floral aromas [59]. Conversely, _A. ghanensis_ and _A.