# A catalog of ethanol-producing microbes in humans

Babacar Mbaye, Reham Magdy Wasfy, Maryam Tidjani Alou, Patrick Borentain, Rene Gerolami, Jean-Charles Dufour, Matthieu Million

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Babacar Mbaye, Reham Magdy Wasfy, Maryam Tidjani Alou, Patrick Borentain, Rene Gerolami, et al.. A catalog of ethanol-producing microbes in humans. 2025. hal-04608799

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Distributed under a Creative Commons Attribution 4.0 International License
# A Repertoire of Ethanol-Producing Human Microbes

**Babacar MBAYE**

Aix-Marseille Université, IRD, APHM, IHU Méditerranée Infection

**Reham MAGDY WASFY**

Aix-Marseille Université, IRD, APHM, IHU Méditerranée Infection

**Maryam TIDJANI ALOU**

Aix-Marseille Université, IRD, APHM, IHU Méditerranée Infection

**Patrick BORENTAIN**

Assistance Publique-Hôpitaux de Marseille AP-HM, Hôpital de la Timone, Unité d'hépatologie

**Rene GEROLAMI**

Aix-Marseille Université, IRD, APHM, IHU Méditerranée Infection

**Jeans Charles DUFOUR**

INSERM, IRD, ISSPAM, Aix Marseille University

**Matthieu MILLION**

matthieumillion@gmail.com

Aix-Marseille Université, IRD, APHM, IHU Méditerranée Infection

## Research Article

## Keywords:

**Posted Date:** October 3rd, 2023

**DOI:** https://doi.org/10.21203/rs.3.rs-3378268/v1

**License:** © This work is licensed under a Creative Commons Attribution 4.0 International License. Read Full License

**Additional Declarations:** No competing interests reported.

**Version of Record:** A version of this preprint was published at Future Microbiology on May 3rd, 2024. See the published version at https://doi.org/10.2217/fmb-2023-0250.
Abstract

Background

Studies of the relationship between the human microbiota and the host, particularly in the maintenance of health and the onset of disease, is still a topical issue. The microbiota is home to a number of microorganisms, including bacteria, yeasts, viruses and archaea, which play a crucial role in the body's proper functioning. In the presence of risk factors, mainly antibiotics, unbalanced diet and so on, a microbiota disequilibrium known as dysbiosis can set in, eventually leading to the onset of several diseases. Several mechanisms have been described for this dysbiosis, but the one linked to endogenous ethanol production is under-investigated.

Results

The aim of this review is to provide an inventory of all ethanol-producing microorganisms in humans. Over 36,000,000 articles were searched on pubmed with our query, of which 7275 were retained. Of these, 5397 contributed to the review, enabling us to identify 85 ethanol-producing microorganisms in humans, including 61 bacterial species and 24 yeasts.

Conclusion

Our results establish an important role for endogenous ethanol in the onset of certain diseases. They will also enable us to better understand the origins of diseases linked to this mechanism, and to improve diagnosis and therapeutic management in the future.

Introduction

Under physiological conditions, small quantities of endogenous ethanol are formed in the body, particularly in the colon, during sugar metabolism [1]. This endogenously produced ethanol is rapidly eliminated by intestinal and hepatic dehydrogenases [Adh], catalases and the microsomal ethanol oxidation system [1]. When its concentration exceeds the internal metabolic capacity, ethanol accumulates and can lead to complications and even the onset of certain diseases [2]. This abnormal situation is mainly due to an imbalance in the intestinal microbiota, favoring the proliferation of microorganisms that produce ethanol from sugars. The first case of this endogenous ethanol production was described in 1948 by Ladkin and Davies in a 5-year-old African child suffering from a hypogastric tumor [3]. People suffering from this disorder present symptoms similar to those associated with acute ethanol intoxication. Various diseases have been associated with this abnormal production of endogenous ethanol, namely autobrewery syndrome [4], diabetes [5, 6], obesity [6] and NASH [7–10]. In addition to the associated pathologies, this endogenous ethanol can lead to medico-legal problems linked to drunk driving despite non-consumption of alcohol. The aim of our review is therefore repertorize the comprehehsing human ethanol producing microbes for the first time all ethanol-producing microorganisms in the human gut microbiota, in order to improve our knowledge of the pathophysiology, diagnosis and management of this problem.

Bibliographic strategy

The aim of this review was to repertoires all human ethanol-producing microorganisms. Only in vitro detection of ethanol was considered to define an ethanol-producing microorganism [EtOH-producing microbe], regardless of the atmosphere or medium used for culture. Articles reporting the presence of enzymes such as alcohol dehydrogenase in the genome which did not measure ethanol production in vitro were excluded. For this purpose, a computer program was used to search PubMed for all articles related to ethanol production for each bacterial species. For the script, we used : ""Bacteria name""[tw] OR ""Synonyms""[tw] AND [ethanol[mesh]] AND [produc*[tw] OR endogenous[tw] OR synthes*[tw] OR secret*[tw]] NOT ""forensic toxicology""[mesh] NOT mutant[tw] NOT ""DNA, recombinant""[MESH] NOT biotransformation[tw] NOT biofuels[tw] NOT
bioreactor[tw] NOT disinfect*[tw]" consisting the name of the species, or a synonym, followed by the keywords used to detect all the articles related to ethanol production. The search was carried out without date or language restrictions. For this purpose, we used all the species present in the NCBI taxonomy browser database [https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi], and for each species a query was used to extract the articles linked to its ethanol production. These queries yielded a total of 7,275 articles. These articles were screening initially according to their titles and abstract. A second screening was conducted by reading the full-text of the selected articles to confirm the ethanol production of each microorganism. The MS Access® database (Access 2007, Microsoft, France) was used to validate the ethanol production of all microorganisms. The output file obtained after validation is in Excel 2007 format (Microsoft), containing the taxonomic rank for each microorganism, its ethanol production (yes or no), the number of items (Query_PMID_Count) for each species and the PMID of each item. We then separated the microorganisms into three groups (bacteria, yeasts and archaea). Filtering according to ethanol production detection, 5,397 articles were included. These articles were all screened to create our repertoire of human bacteria and fungi which were proven to produce ethanol *invitro*.

## Overview of ethanol-producing microorganisms

Among the microorganisms investigated, 37.16% (346/931) were identified as ethanol producers, including 51.73% (179/346) fungi, 46.82% (162/346) bacteria and 1.45% (5/346) archaea. Microorganisms known to be present in humans accounted for 14%, 13.02% and 0% of fungi, bacteria and archaea while non-human microorganisms represented 86.03%, 86.98% and 100% respectively. Ultimately, we found 85 ethanol-producing human microorganisms (61 bacteria and 24 fungi) (see Table 1), none of which were human archaea. Among ethanol-producing bacteria, *Pseudomonadota* (9 species and 846 hits) and *Bacillota* (46 species and 434 hits) were the most represented at the phylum level whereas at the family level the most represented were *Enterobacteriaceae* (5 species and 837 hits), *Lactobacillaceae* (18 species and 126 hits), *Streptococcaceae* (4 species and 73 hits), *Clostridiaceae* (9 species and 67 hits), *Bifidobacteriaceae* (5 species and 18 hits), *Oscillospiraceae* (4 species and 15 hits) and *Lachnospiraceae* (5 species and 5 hits). At the genus level, the most represented were *Klebsiella* (4 species 69 hits), *Clostridium* (7 species and 61 hits), *Lactococcus* (2 species 45 hits), *Limosilactobacillus* (3 species and 32 hits), *Streptococcus* (2 species and 28 hits), *Leuconostoc* (3 species 20 hits), *Pediococcus* (3 species and 16 hits), *Bifidobacterium* (5 species and 14 hits), *Ruminococcus* (3 species and 14 hits) and *Lentilactobacillus* (3 species 13 hits) and. For yeasts, *Ascomycota* (23 species and 2593 hits) was the most represented phylum, with *Saccharomycetaceae* (4 species and 2304 hits), *Debaryomycetaceae* (7 species and 158 hits), *Phaffomycetaceae* (4 species and 40 hits) and *Pichiaceae* (4 species and 66 hits) being the most represented of the 9 identified families. Moreover, 15 genera were reported as including ethanol-producing species, the most represented being *Saccharomyces* (2 species and 2143 hits), *Candida* (6 species and 149 hits), *Pichia* (3 species and 66 hits), *Cyberlindnera* (2 species and 9 hits) and *Meyerozyma* (2 species and 9 hits).

## Human associated ethanol-producing yeast

Ethanol production though fermentation was initially described in yeast [11], and specifically the species *Saccharomyces cerevisiae* has been used for ethanol production. The most described ethanol producing microorganism is *S. cerevisiae* which has been described in cases of autobrewery syndrome [12–14] but is also the most widely used probiotic [15, 16]. However, ethanol can be produced by several other species some of which are opportunistic human pathogens. For instance, our query retrieved 27 yeast species, distributed into two phyla: *Ascomycota* 96% and *Basidiomycota* 6%, which are known to be able to produce ethanol. Thirteen genera were listed, the most represented being: *Candida* (5 species), *Pichia* (3 species), *Saccharomyces* (2 species), *Meyerozyma* (2 species), and *Cyberlindnera* (2 species) (Table 1). The *Candida* genus was the most represented, with five distinct species: *Candida sanyaensis*, *Candida sojae*, *Candida albicans*, *Candida parapsilosis* and *Candida tropicalis* (Supplementary table 2). Some species of this genus have been associated with a number of diseases, including the auto-brewery syndrome [17–20], diabetes [5, 21–23] and NASH [8] (Table 2). Their involvement in these diseases has been strongly linked to their ability to produce endogenous ethanol. The *Pichia* genus was the second most represented ethanol producer, with 3 different species: *Pichia kudriavzevii*, *Pichia membranifaciens*, and *Pichia fermentans*. Some *Pichia* species have been associated with NASH [15] and NAFLD [24]; and *Pichia kudriavzevii* has been shown to produce high
quantities of ethanol in NASH patients [15]. Yeasts generally use the same pathway for ethanol synthesis, which is the "pyruvate fermentation to ethanol II" pathway according to metacyc database (https://metacyc.org/META/substring-search?type=Nil&object=ethanol+). This pathway consists in metabolizing carbohydrates until a simple sugar is obtained. The latter enters the glycolysis cycle or the Entner-Doudoroff pathway which end-product to produce puruvate. Puruvate decarboxylase further decarboxylates puruvate into acetaldehyde and CO2. Alcohol dehydrogenase then catalyzes the reduction of acetaldehyde to ethanol (Supplementary Fig. 1).

## Human-associated ethanol producing bacteria.

We found 61 bacterial species capable of producing ethanol, divided into 4 phyla, 15 families and 30 genera. The most represented were *Bacillota* (46 species), *Pseudomonadota* (9 species) at the phylum level, *Lactobacillaceae* (18 species), *Clostridiaceae* (9 species), *Enterobacteriaceae* (5 species), *Bifidobacteriaceae* (5 species) at the family level. At the genus level, *Clostridium* (7 species), *Bifidobacterium* (5 species), *Klebsiella* (4 species) and *Lactococcus* (4 species) were the most represented.

## Bacillota

### Lactobacillales

This group comprises 44 species divided into three families *Lactobacillaceae* 63.63%, *Streptococcaceae* 25.58% and *Enterococcaceae* 10.79% and 15 genera. The most represented genera are *Streptococcus* 60%, *Enterococcus* 33.3%, *Pediococcus* and *Lactobacillus* 26.6% and *Limosilactobacillus*, *Lentilactobacillus*, and *Leuconostoc* at 20% (supplementary Table 1). These species use the heterolactic pathway for ethanol formation. Sugars are degraded via phosphoketolase (the pentose phosphate pathway) to D-xylose-5-phosphate, which is then converted to D-glyceraldehyde-3-phosphate (G3P) and acetyl phosphate. The resulting G3P is converted into lactic acid and the acetyl phosphate into ethanol or acetate (supplementary Fig. 2). Beyond heterolactic fermentation, some species of the streptococcus genus can use the hexitol pathway to produce lactate, acetate formate and ethanol. This pathway leads to the use of sugar substitutes such as sugar alcohols, for example sorbitol and mannitol (supplementary Fig. 1). These streptococci are bacteria generally associated with the oral microbiota. Their endogenous ethanol production has been correlated with their abundance in NAFLD patients [10]. In this review, nine species of the genus were shown to produce ethanol (supplementary Table 1), two of which *S. mutans* and *S. mitis*, are major causes of dental caries [25].

These two species have previously been reported in a case of oral auto-brewery [26], with higher ethanol production in co-culture with *Klebsiella pneumoniae* [18]. Streptococci ferment carbohydrates, producing mainly lactic acid, with minor amounts of acetic and formic acids, CO2 and ethanol. The Embden-Meyerhof pathway of glycolysis operates under both aerobic and anaerobic conditions, and the pyruvate resulting from these reactions is converted into different products depending on the oxygen level. In anaerobic conditions, streptococci produce formate, acetate and ethanol via the pyruvate formate lyase pathway (supplementary Fig. 2). In aerobic conditions, this puruvate lyase is inactive and replaced by pyruvate oxidase in some streptococci and pyruvate dehydrogenase in others. In case of sugar overload, they also form lactate catalyzed by lactate dehydrogenase. The abundance of *S. mutans* in NASH patients has already been described by Naka et al [27], and a specific strain of the same species (TW871) has been shown to induce NASH after administration in a mouse model [28]. In the *Enterococcus* genus, five species have been shown to produce ethanol (Supplementary table 1). Ethanol production by the *Enterococcus* genus has been described in the autobrewery syndrome [4] as well as in post-mortem samples [29]. Cytolysin-secreting. *E. feacalis* has been identified as the causative agent of liver damage in alcoholic hepatitis [30], and *E. gallinarum* has been identified as one of the bacterial species specific to primary sclerosing cholangitis [23]. The other genera represented were *Pediococcus* (5 ethanol-producing species), *Lactobacillus* (4 ethanol-producing species) and the ethanol-producing group, comprising just 3 species in each of the following genera: *Limosilactobacillus*, *Lentilactobacillus*, and *Leuconostoc* (supplementary Table 1). Among the latter, the genus *Limosilactobacillus*, recently reclassified by Zheng et al., seems the most relevant, as several species of the genus have been associated with several pathologies. In this review,
only three species of this genus are capable of producing ethanol: *Limosilactobacillus fermentum*, *Limosilactobacillus reuteri* and *Limosilactobacillus panis*. *L. fermentum* has previously been associated with NASH [10, 31] and liver enlargement in animals [32] whereas *L. reuteri* has been associated with metabolic syndrome and obesity [33, 34]. The other genera not represented are those whose ethanol production involved less than two species [Table 1 and supplementary Table 1).

## Lachnospiraceae

Our search revealed five species of the ethanol-producing family in three genera: *Blautia* (*B.coccoides* and *B.hydrogenotrophica*, *Enterocloster* (*E. bolteae* and *E.aspargiformis*) and *Mediterraneibacter* (*M.torques* and *M. gnavus*). This family is one of the most represented bacterial groups in the human intestinal microbiota, and various studies have demonstrated its beneficial and harmful effects on human health. For species of the *Blautia* and *Enterocloster* genus, the main pathway used is the Embden-Meyerhof pathway, which metabolizes carbohydrates into pyruvate, which is then oxidized to acetyl-CoA by pyruvate dehydrogenase [35]. This acetyl-CoA undergoes two successive reactions, first to acetaldehyde and then to ethanol, thanks to the bifunctional enzyme alcohol-acetaldehyde dehydrogenase (AdhE). In addition, the *Enterocloster* species described in this review can also convert acetyl-coA to acetate via the phosphotransacetylase enzyme; the latter is then converted to acetaldehyde by aldehyde dehydrogenase and then to ethanol by alcohol dehydrogenase [36]. It has been shown that *M. torques* and *M. gnavus* can produce ethanol from a variety of substrates [37], using the same pathway as above, from pyruvate to acetyl CoA and then to ethanal via the bifunctional enzyme alcohol-acetaldehyde dehydrogenase (supplementary Fig. 1).

## Clostridiaceae

This family comprises 10 ethanol-producing species in 3 genera: *Clostridium* [8], *Paraclostridium* [1] and *Sarcina* [1]. The Clostridium group includes *C.amylolyticum*, *C. butyricum*, *C.cochlearium*, *C. sporogenes*, *C. tyrobutyricum*, *C. beijerinckii*, *C. malenominatum* and *C. asparagiforme*; these bacteria can metabolize complex sugars (inulin, starch, etc.) into simple sugars (glucose, fructose) for use as a carbon source. In their ethanol production process, these bacteria use the acetonobutyl pathway. This pathway consists of the metabolism of sugars by glycolysis (EMP pathway); the formation of organic acids [acetic acid and butyric acid] following a series of pyruvate metabolism reactions; and the fermentation of these organic acids, leading to the synthesis of ethanol, acetone and butanol. They can also produce ethanol from pyruvate by transforming it into acetyl coa via the enzyme purvate synthase. This acetyl coa is then transformed into acetaldehyde-by-acetaldehyde dehydrogenase, which is finally converted into ethanol by alcohol dehydrogenase [supplementary Fig. 1]. In this pathway, organic acids are not synthesized. The species *Paraclostridium bifermentans* and *Sarcina ventriculi* use the bifunctional enzyme alcohol-acetaldehyde dehydrogenase (AdhE) to produce ethanol from acetyl CoA.

## Oscillospiraceae

Three species of the family have been shown to produce ethanol, all belonging to the genus *Ruminococcus*. These are *R. albus*, *R. flavefaciens* and *R. bromii*. The first species of the genus described for ethanol production was *R.albus* by Wolin *et al*, from starch [38] and *R. bromii* has been shown to produce ethanol, acetate and formate in pure culture with fructose [39]. Isotope labeling analysis of carbohydrate fermentation products has shown that the Embden-Meyerhof-Parnas (EMP) pathway is the main metabolic pathway for bacteria of the genus *Ruminococcus*. [40]. The pyruvate obtained from sugar metabolism is converted to acetyl CoA and then to ethanol by the bifunctional enzyme alcohol-acetaldehyde dehydrogenase. [41] ; among other things, acetate, formate, dihydrogen and carbon dioxide are produced.

## Pseudomonadota

In this phylum we found nine species capable of producing ethanol, divided into four families and six genera, with a predominance of the *Klebsiella* genus (2/8), see Supplementary Table 1. They are bacteria that can live in both aerobic and anaerobic conditions, using different types of sugars as carbon sources [42]. Whatever the carbon source, the metabolism will
be carried out until pyruvate is obtained [34] and it is from the metabolism of the latter that different metabolites are produced. These bacteria use two types of fermentation: mixed acid fermentation and butylene glycolytic fermentation. In the former, a complex mixture of several acids is produced: formate, lactate, acetate, succinate and ethanol. The ethanol produced results from the transformation of pyruvate into Acetyl CoA and formate by the enzyme puruvate formate lyase; Acetyl Coa in turn is then transformed into Acetaldehyde by acetaldehyde deshydrogenase and then into ethanol by alcohol dehydrogenase. (suppleme,tary Fig. 2). The second reaction is characterized by the formation of 2,3-butanediol via acetoine in addition to the products obtained with mixed fermentation (supplementary Fig. 1). Yuan et al have shown that the 2,3-butanediol pathway is one of the pathways involved in high ethanol production in enterobacteria in both anaerobic and aerobic conditions, particularly in *Klebsiella pneumoniae* [9].

## Actinomycetota

In this phylum only the genus *Bifidobacterium* is represented, this genus comprises 118 species and 18 sub-species, some of which are considered probiotics [43] and often used in dairy products. Species in the genus represent 10 9 to 10 10 and play an important role in the metabolism of complex sugars in the colon [44]. Five species of the *Bifidobacterium* genus were identified as a result of our inquiry as being capable of producing ethanol. These species are *Bifidobacterium adolescentis*, *Bifidobacterium bifidum*, *Bifidobacterium breve*, *Bifidobacterium longum* and *Bifidobacterium animalis*. In their metabolic process, bifidobacteria tend to depolymerize complex sugars (oligosaccharides and polysaccharides) into monomers (simple sugars), which in turn are fed into the fructose-6-phosphate shunt to give lactic and acetic acids as end products, formic acid and ethanol as minorities. The ethanol production process involves breaking down sugars into two-carbon and three-carbon molecules, in this case acetyl phosphate and glyceraldehyde 3-phosphate. Acetyl phosphate is converted to acetate by an acetate kinase, and glyceraldehyde 3-phosphate to pyruvate. The phosphoroclastic reaction of pyruvate leads to the formation of acetylphosphate and formate. The resulting acetylphosphate is reduced to ethanol at the expense of acetate production (supplementary Fig. 2).

## Number of articles according to microorganism at genus and species level

Using articles that contributed to the review, we made a graphical representation evaluating the number of articles found for microorganisms at different taxonomic ranks (genera and species level). For bacteria the figure at genus level shows that *Escherichia* and *Staphylococcus* genera are the most cited in ethanol production, while other genera are also represented, namely the lactic acid bacteria group (*Pediococcus*, *Limosolactobacillus*, *Leuconostoc*, *Enterococcus*) and certain genera from the Proteobacteria group (*Klebsiella*, *Raoutella*, *Hafnia* and *Sarcina*). In terms of species, Escherichia coli is the most represented, followed by *Lactococcus lactis*, *Klebsiella pneumoniae* and *Staphylococcus aureus*, with other species also represented (see Fig. 2). For yeasts, the most represented genera are *Saccharomyces*, *Kluyveromyces*, *Candida* and *Pichia*. In terms of species, *Saccharomyces cerevisiae*, *Candida albicans*, *Pichia Kudriavzevii* and *Kluyveromyces marxianus* are the most frequently cited in articles, but other species are also represented (see Fig. 2).

Taking into account all the microorganisms analyzed, we can see that the most frequently cited in ethanol production include both bacteria and yeasts, namely *Saccharomyces cerevisiae* (2139 hits), *Escherichia coli* (765 hits), *Kluyveromyces marxianus* (*Candida kefir* 143 hits), *Staphylococcus aureus* (110 hits), *Candida albicans* (96 hits) and *Pichia kudriavzevii* (57 hits) other species are also represented, (see Fig. 3) (see Tables 3 and 4).

## Discussion

The end goal of our study was to establish a repertory of known human microorganisms capable of producing ethanol, using a computer program capable of querying Pubmed. A total of 7275 contributions were received, enabling us to identify 85 ethanol-producing microorganisms. These microorganisms were classified according to their taxonomy, and for some of them, name changes were also taken into account following validation on LPSN or Taxonomy browser. The human gut is home to a range of microorganisms with different roles in health and human disease [45]. These microorganisms produce various molecules that contribute to digestive and host homeostasis and, in the case of dysbiosis, can lead to the onset of certain diseases. Ethanol, which is one of the metabolites produced by the microbiota even in healthy individuals, is known as
"physiological ethanol" [46], but at high concentrations in the body it can lead to the onset or complications of a number of diseases, including diabetes [21, 22], auto-brewery syndrome [12, 19, 26], NASH [7, 10, 31, 47], obesity [17], and in some cases even behavioral disorders [19]. To our knowledge, this review is the first to examine ethanol-producing microorganisms in humans; most of studies have focused on endogenous ethanol-related complications [48]. This repertory is consistent with the literature data, as most of our microorganisms were at least cited in one or more articles related to endogenous ethanol production. With regard to NASH, several species have already been described, in particular *K. pneumoniae* [8, 9, 49], *L. fermentum* [10, 31], *L.lactis* [31]. Yuan et al. even showed that administration of a strain of *K.pneumoniae* [ ethanol hyperproducer isolated from a NASH patient] to a mouse led to the onset of the disease and antibiotic intake improved the liver in the mice [47]. Various species have been described for autobrewery syndrome, including *S.cerviseae* [12], *C.albicans* [50], *C.parapsilosis* [51] and *P.kudriavzevi* [48]. On the other hand, a closely related species to *S.cerevisea* [which is known to produce ethanol] *S.boulardii* have been described in the literature as probiotics for NASH patients [52]. This repertoire will help to understand the potential involvement of ethanol-producing microorganisms in the onset of certain diseases. It can be used by clinicians in their diagnostic process following the isolation of one or more ethanol-producing microorganisms from a patient. One of the limitations of this review may be linked to the use of a single database, in this case Pubmed, but also to its temporality, as only publications submitted before January 28, 2023, were taken into account.

# Declarations

## Funding Statement

This research received no external funding.

## Author Contributions

Conceptualization, M.M and J.C.D; methodology, B.M. and. J.C.D.; software, B.M and. J.C.D.; formal analysis, B.M. and R.M.W data curation, B.M.; writing-original draft preparation, B.M., Writing, review and editing: M.M, R.G, P.B, and M.T.A; supervision, M.M. and P.B.

## Ethics approval and Consent to participate.

The study did not require ethical approval and consent for participation.

## Data Availability Statement

Data supporting reported results can be found in the supplementary files.

## Conflicts of Interest

The authors declare no conflict of interest.

## Acknowledgments

we would like to thank all the authors for taking part in this study.

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Tables
<table><caption>Table 1
Ethanol producing human microbes.</caption><thead><tr><td>Taxonomic rank</td><td>References</td></tr></thead><tbody><tr><td>FUNGI</td><td></td></tr><tr><td>Ascomycota</td><td></td></tr><tr><td>Saccharomycetaceae</td><td></td></tr><tr><td>Saccharomyces</td><td></td></tr><tr><td>Saccharomyces cerevisiae</td><td>[53]</td></tr><tr><td>Saccharomyces kudriavzevii</td><td>[54]</td></tr><tr><td>Kluyveromyces</td><td></td></tr><tr><td>Kluyveromyces marxianus</td><td>[55, 56]</td></tr><tr><td>Nakaseomyces</td><td></td></tr><tr><td>Nakaseomyces glabratus</td><td>[53, 57]</td></tr><tr><td>Debaryomycetaceae</td><td></td></tr><tr><td>Candida</td><td></td></tr><tr><td>Candida sanyaensis</td><td>[58]</td></tr><tr><td>Candida sojae</td><td>[59, 60]</td></tr><tr><td>Candida albicans</td><td>[20, 61]</td></tr><tr><td>Candida parapsilosis</td><td>[29, 61]</td></tr><tr><td>Candida tropicalis</td><td>[53, 62]</td></tr><tr><td>Meyerozyma</td><td></td></tr><tr><td>Meyerozyma guilliermondii</td><td>[63, 64]</td></tr><tr><td>Meyerozyma caribbica</td><td>[65]</td></tr><tr><td>Dipodascaceae</td><td></td></tr><tr><td>Geotrichum</td><td></td></tr><tr><td>Geotrichum candidum</td><td>[66, 67]</td></tr><tr><td>Metschnikowiaceae</td><td></td></tr><tr><td>Clavispora</td><td></td></tr><tr><td>Clavispora lusitaniae</td><td>[56, 68]</td></tr><tr><td>Phaffomycetaceae</td><td></td></tr><tr><td>Cyberlindnera</td><td></td></tr><tr><td>Cyberlindnera fabianii</td><td>[69]</td></tr><tr><td>Cyberlindnera saturnus</td><td>[70, 71]</td></tr><tr><td>Wickerhamomyces</td><td></td></tr><tr><td>Wickerhamomyces anomalus</td><td>[72, 73]</td></tr><tr><td>Barnettozyma</td><td></td></tr></tbody></table>
<table><thead><tr><th>Taxonomic rank</th><th>References</th></tr></thead><tbody><tr><td>[Candida] norvegica</td><td>[74]</td></tr><tr><td>Pichiaceae</td><td></td></tr><tr><td>Pichia</td><td></td></tr><tr><td>Pichia kudriavzevii</td><td>[8, 75]</td></tr><tr><td>Pichia fermentans</td><td>[76, 77]</td></tr><tr><td>Pichia membranifaciens</td><td>[78, 79]</td></tr><tr><td>Saturnispora</td><td></td></tr><tr><td>Saturnispora silvae</td><td>[80]</td></tr><tr><td>Saccharomycetales incertae sedis</td><td></td></tr><tr><td>Diutina</td><td></td></tr><tr><td>Diutina rugosa</td><td>[81]</td></tr><tr><td>Trichomonascaceae</td><td></td></tr><tr><td>Wickerhamiella</td><td></td></tr><tr><td>Wickerhamiella pararugosa</td><td>[82]</td></tr><tr><td>Basidiomycota</td><td></td></tr><tr><td>Sporidiobolaceae</td><td></td></tr><tr><td>Rhodotorula</td><td></td></tr><tr><td>Rhodotorula mucilaginosa</td><td>[83]</td></tr><tr><td>BACTERIA</td><td></td></tr><tr><td>Bacillota</td><td></td></tr><tr><td>Bacillaceae</td><td></td></tr><tr><td>Weizmannia</td><td></td></tr><tr><td>Weizmannia coagulans</td><td>[84]</td></tr><tr><td>Clostridiaceae</td><td></td></tr><tr><td>Clostridium</td><td></td></tr><tr><td>Clostridium amylolyticum</td><td>[85]</td></tr><tr><td>Clostridium butyricum</td><td>[86]</td></tr><tr><td>Clostridium cochlearium</td><td>[87]</td></tr><tr><td>Clostridium sporogenes</td><td>[88]</td></tr><tr><td>Clostridium tyrobutyricum</td><td>[89]</td></tr><tr><td>Clostridium beijerinckii</td><td>[90, 91]</td></tr><tr><td>Clostridium malenominatum</td><td>[87]</td></tr><tr><td>Clostridium asparagiforme</td><td>[36]</td></tr><tr><td>Paraclostridium</td><td></td></tr></tbody></table>
<table><thead><tr><td>Taxonomic rank</td><td>References</td></tr></thead><tbody><tr><td>Paraclostridium bifermentans</td><td>[92, 93]</td></tr><tr><td>Sarcina</td><td></td></tr><tr><td>Sarcina ventriculi</td><td>[94]</td></tr><tr><td>Enterococcaceae</td><td></td></tr><tr><td>Enterococcus</td><td></td></tr><tr><td>Enterococcus faecalis</td><td>[95, 96]</td></tr><tr><td>Enterococcus mundtii</td><td>[97]</td></tr><tr><td>Lachnospiraceae</td><td></td></tr><tr><td>Blautia</td><td></td></tr><tr><td>Blautia coccoides</td><td>[36]</td></tr><tr><td>Blautia</td><td>[98]</td></tr><tr><td>Enterocloster</td><td></td></tr><tr><td>Enterocloster bolteae</td><td>[41]</td></tr><tr><td>Enterocloster asparagiformis</td><td>[41]</td></tr><tr><td>Oscillospiraceae</td><td></td></tr><tr><td>Ruminococcus</td><td></td></tr><tr><td>[Ruminococcus] torques</td><td>[99]</td></tr><tr><td>Ruminococcus albus</td><td>[100, 101]</td></tr><tr><td>Ruminococcus flavefaciens</td><td>[102]</td></tr><tr><td>Ruminococcus bromii</td><td>[35, 103]</td></tr><tr><td>[Ruminococcus]gnavus</td><td>[35]</td></tr><tr><td>Lactobacillaceae</td><td></td></tr><tr><td>Fructilactobacillus</td><td></td></tr><tr><td>Fructilactobacillus florum</td><td>[104]</td></tr><tr><td>Lacticaseibacillus</td><td></td></tr><tr><td>Lacticaseibacillus casei</td><td>[105, 106]</td></tr><tr><td>Lentilactobacillus</td><td></td></tr><tr><td>Lentilactobacillus parabuchneri</td><td>[107-109]</td></tr><tr><td>Lentilactobacillus buchneri</td><td>[107, 109, 110]</td></tr><tr><td>Lentilactobacillus senioris</td><td>[107, 109]</td></tr><tr><td>Leuconostoc</td><td></td></tr><tr><td>Leuconostoc mesenteroides</td><td>[111, 112]</td></tr><tr><td>Leuconostoc lactis</td><td>[113]</td></tr><tr><td>Leuconostoc pseudomesenteroides</td><td>[59]</td></tr></tbody></table>
<table><thead><tr><th>Taxonomic rank</th><th>References</th></tr></thead><tbody><tr><td>Levilactobacillus</td><td></td></tr><tr><td>Levilactobacillus brevis</td><td>[115, 116]</td></tr><tr><td>Limosilactobacillus</td><td></td></tr><tr><td>Limosilactobacillus reuteri</td><td>[117-119]</td></tr><tr><td>Limosilactobacillus fermentum</td><td>[120]</td></tr><tr><td>Limosilactobacillus panis</td><td>[121, 122]</td></tr><tr><td>Pediococcus</td><td></td></tr><tr><td>Pediococcus inopinatus</td><td>[123]</td></tr><tr><td>Pediococcus acidilactici</td><td>[124-126]</td></tr><tr><td>Pediococcus pentosaceus</td><td>[127]</td></tr><tr><td>Schleiferilactobacillus</td><td></td></tr><tr><td>Schleiferilactobacillus perolens</td><td>[128]</td></tr><tr><td>Schleiferilactobacillus harbinensis</td><td>[72]</td></tr><tr><td>Weissella</td><td></td></tr><tr><td>Weissella confusa</td><td>[129]</td></tr><tr><td>Paenibacillaceae</td><td></td></tr><tr><td>Paenibacillus</td><td></td></tr><tr><td>Paenibacillus polymyxa</td><td>[130]</td></tr><tr><td>Paenibacillus macerans</td><td>[131, 132]</td></tr><tr><td>Staphylococcaceae</td><td></td></tr><tr><td>Staphylococcus</td><td></td></tr><tr><td>Staphylococcus aureus</td><td>[95]</td></tr><tr><td>Streptococcaceae</td><td></td></tr><tr><td>Streptococcus</td><td></td></tr><tr><td>Streptococcus mutans</td><td>[133, 134]</td></tr><tr><td>Streptococcus agalactiae</td><td>[135]</td></tr><tr><td>Lactococcus</td><td></td></tr><tr><td>Lactococcus lactis</td><td>[136-138]</td></tr><tr><td>Lactococcus cremoris</td><td>[139]</td></tr><tr><td><strong>Actinomycetota</strong></td><td></td></tr><tr><td>Bifidobacteriaceae</td><td></td></tr><tr><td>Bifidobacterium</td><td></td></tr><tr><td>Bifidobacterium adolescentis</td><td>[140, 141]</td></tr><tr><td>Bifidobacterium bifidum</td><td>[142, 143]</td></tr></tbody></table>
<table><thead><tr><th>Taxonomic rank</th><th>References</th></tr></thead><tbody><tr><td>Bifidobacterium breve</td><td>[144, 145]</td></tr><tr><td>Bifidobacterium longum</td><td>[146]</td></tr><tr><td>Bifidobacterium animalis</td><td>[147, 148]</td></tr><tr><td><strong>Proteobacteria</strong></td><td></td></tr><tr><td>Enterobacteriaceae</td><td></td></tr><tr><td>Klebsiella</td><td></td></tr><tr><td>Klebsiella variicola</td><td>[149]</td></tr><tr><td>Klebsiella aerogenes</td><td>[150]</td></tr><tr><td>Klebsiella oxytoca</td><td>[151, 152]</td></tr><tr><td>Klebsiella pneumoniae</td><td>[8, 47]</td></tr><tr><td>Escherichia</td><td></td></tr><tr><td>Escherichia coli</td><td>[102, 103]</td></tr><tr><td>Raoultella</td><td></td></tr><tr><td>Raoultella planticola</td><td>[154, 155]</td></tr><tr><td>Aeromonadaceae</td><td></td></tr><tr><td>Aeromonas</td><td></td></tr><tr><td>Aeromonas hydrophila</td><td>[156]</td></tr><tr><td>Erwiniaceae</td><td></td></tr><tr><td>Pantoea</td><td></td></tr><tr><td>Pantoea agglomerans</td><td>[157]</td></tr><tr><td>Hafniaceae</td><td></td></tr><tr><td>Hafnia</td><td></td></tr><tr><td>Hafnia alvei</td><td>[158]</td></tr></tbody></table>
<table><caption>Table 2
Diseases linked to endogenous ethanol production.</caption><thead><tr><td>Diseases</td><td>References</td></tr></thead><tbody><tr><td>Autobrewery syndrome</td><td>[12, 26, 49]</td></tr><tr><td>Urinary case</td><td>[13]</td></tr><tr><td>Gut case</td><td>[4, 6, 14]</td></tr><tr><td>Oral case</td><td>[150]</td></tr><tr><td>Diabetic</td><td></td></tr><tr><td>Auto-brewery syndrome: Ethanol pseudo-toxicity in diabetic and hepatic patients</td><td>[21]</td></tr><tr><td>High urine ethanol and negative blood and vitreous ethanol in a diabetic woman: a case report, retrospective case survey, and review of the literature</td><td>[22]</td></tr><tr><td>Endogenous ethanol production in patients with diabetes mellitus as a medicolegal problem</td><td>[5]</td></tr><tr><td>A Rare Diagnosis of Gut Fermentation/Auto-Brewery Syndrome in the Setting of Diabetes and Obesity</td><td>[6]</td></tr><tr><td>Crohn's disease</td><td></td></tr><tr><td>Auto-brewery syndrome in the setting of long-standing Crohn's disease: a case report and review of the literature</td><td>[159]</td></tr><tr><td>Obesity</td><td></td></tr><tr><td>Endogenous ethanol production and hepatic disease following jejunoileal bypass for morbid obesity</td><td>[17]</td></tr><tr><td>NASH</td><td></td></tr><tr><td>Endogenous Ethanol and Triglyceride Production by Gut Pichia kudriavzevii, Candida albicans and Candida glabrata Yeasts in Non-Alcoholic Steatohepatitis</td><td>[8]</td></tr><tr><td>Characterization of gut microbiomes in nonalcoholic steatohepatitis [NASH] patients: A connection between endogenous alcohol and NASH</td><td>[7]</td></tr><tr><td>Fatty Liver Disease Caused by High-Alcohol-Producing Klebsiella pneumoniae</td><td>[47]</td></tr><tr><td>Endogenous ethanol produced by intestinal bacteria induces mitochondrial dysfunction in non-alcoholic fatty liver disease</td><td>[160]</td></tr><tr><td>Microbiome-derived ethanol in nonalcoholic fatty liver disease</td><td>[10]</td></tr><tr><td>SIBO</td><td></td></tr><tr><td>Endogenous ethanol production in a patient with chronic intestinal pseudo-obstruction and small intestinal bacterial overgrowth</td><td>[161]</td></tr><tr><td>Others findings</td><td></td></tr><tr><td>Drunk Without Drinking: A Case of Auto-Brewery Syndrome</td><td>[19]</td></tr><tr><td>Rupture of stomach in African child</td><td>[3]</td></tr><tr><td>Auto-Brewery Syndrome in a Child with Short Gut Syndrome: Case Report and Review of the Literature</td><td>[4]</td></tr><tr><td>Endogenous ethanol fermentation in a child with short bowel syndrome</td><td>[18]</td></tr><tr><td>Endogenous alcohol production by intestinal fermentation in sudden infant death</td><td>[20]</td></tr><tr><td>Endogenous Ethanol Production Levels in Saudi Arabia Residents</td><td>[162]</td></tr></tbody></table>
<table><caption>Table 3
Most frequently cited yeasts for ethanol production in the literature</caption><thead><tr><td>Species</td><td>Number of Hits</td></tr></thead><tbody><tr><td>Saccharomyces cerevisiae</td><td>2139</td></tr><tr><td>Kluyveromyces marxianus OR Candida kefyr</td><td>143</td></tr><tr><td>Candida albicans</td><td>96</td></tr><tr><td>Pichia kudriavzevii</td><td>57</td></tr><tr><td>Candida tropicalis</td><td>39</td></tr><tr><td>Wickerhamomyces anomalus</td><td>30</td></tr><tr><td>Candida glabrata</td><td>18</td></tr><tr><td>Clavispora lusitaniae OR Candida lusitaniae</td><td>11</td></tr><tr><td>Candida parapsilosis</td><td>11</td></tr><tr><td>Rhodotorula mucilaginosa</td><td>10</td></tr></tbody></table>

<table><caption>Table 4
Most frequently cited bacteria for ethanol production in the literature</caption><thead><tr><td>Species</td><td>Number of hits</td></tr></thead><tbody><tr><td>Escherichia coli</td><td>765</td></tr><tr><td>Staphylococcus aureus</td><td>110</td></tr><tr><td>Lactococcus lactis</td><td>44</td></tr><tr><td>Clostridium beijerinckii</td><td>43</td></tr><tr><td>Klebsiella pneumoniae</td><td>39</td></tr><tr><td>Enterococcus faecalis</td><td>28</td></tr><tr><td>Streptococcus mutans</td><td>27</td></tr><tr><td>Lacticaseibacillus casei</td><td>23</td></tr><tr><td>Limosilactobacillus fermentum</td><td>23</td></tr><tr><td>Leuconostoc mesenteroides</td><td>16</td></tr><tr><td>Klebsiella oxytoca</td><td>16</td></tr><tr><td>Levilactobacillus brevis</td><td>15</td></tr><tr><td>Klebsiella aerogenes</td><td>13</td></tr><tr><td>Lentilactobacillus buchneri</td><td>11</td></tr></tbody></table>

Figures
<figure><img src="image_3.png" /><figcaption>Figure 1</figcaption></figure>

Flow chart of literature review
Bacteria

Genus level

<figure><img src="image_4.png" /></figure>

Species level

<figure><img src="image_5.png" /></figure>

Fungi

Genus level

<figure><img src="image_6.png" /></figure>

Species level

<figure><img src="image_7.png" /><figcaption>Figure 2</figcaption></figure>

Number of articles mentioning ethanol production for bacteria (left) and yeast (right) at genus and species level.
<figure><img src="image_8.png" /><figcaption>Figure 3</figcaption></figure>

The species most frequently cited in the literature for ethanol production.

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