Brettanomyces

Brettanomyces - Milk The Funk Wiki 11/05/2023 21:07
Brettanomyces
Brettanomyces, also referred to by brewers as "Brett" or "Bretta", is Greek for "British Fungus" and is a yeast that was originally thought
of as an important yeast for producing the character of some 17th century and prior English ales. Since the wide adoption of pure cultures
of Saccharomyces cerevisiae and S. pastorianus in the brewing and wine industries starting in the late 1800's, Brettanomyces has been
mostly viewed as a spoilage yeast, except in Belgian lambic, Flanders red/brown beers, and a handful of styles of wine. More recently
Brettanomyces has gained popularity in the United States (and subsequently the brewing industries of other countries) as a yeast that can
contribute desirable and novel characteristics to beer and other alcoholic beverages. The genus name Dekkera is used interchangeably
with Brettanomyces, as it describes the teleomorph or spore-forming form of the yeast, although this form is extremely rare or perhaps
even non-existent [1][2]. Known for its barnyard, fecal, horsey, metallic or Band-Aid flavors, Brettanomyces continues to be unwelcome
in many breweries and most wineries [3]. However, Brettanomyces also produces high levels of fruity esters that are desirable in some Brettanomyces
styles like saison, lambic, and American sour beers. Brettanomyces can also form a pellicle during fermentation. See Lactobacillus,
Pediococcus, Saccharomyces, Mixed Cultures, Kveik, and Nonconventional Yeasts and Bacteria charts for other commercially available
cultures.
Contents
1 Introduction of History, Characteristics, and Taxonomy
1.1 Taxonomy
1.2 Morphology
1.3 Culturing
1.4 Environment and Survival
1.4.1 Sulfur Tolerance
1.4.2 Biofilm Brett Aroma Wheel by Dr. Linda
1.4.3 UV Light Bisson and Lucy Joseph at UC Davis
2 Brettanomyces Metabolism
2.1 Carbohydrate Metabolism and Fermentation Temperature
2.1.1 Glycosides and Beta-Glucosidase Activity
2.1.2 Hop Biotransformation
2.2 Nitrogen Metabolism
2.3 Secondary Metabolites
2.3.1 Ester Production
2.3.2 Phenol Production
2.3.3 Acid Production
2.3.4 Biogenic Amines
2.3.5 Glycerol
2.3.6 Other Compounds
3 Commercial Cultures
3.1 Bootleg Biology/Spot Yeast
3.2 Brewing Science Institute
3.3 Community Cultures Yeast Lab
3.4 Craft Cultures
3.5 East Coast Yeast
3.6 Escarpment Laboratories
3.7 Fermentis
3.8 Fermmento Labs (Brazil - CLOSED)
3.9 Imperial Organic Yeast
3.10 Inland Island Yeast Laboratories
3.11 GigaYeast
3.12 Jasper Yeast
3.13 Mainiacal Yeast (CLOSED)
3.14 Omega Yeast Labs
3.15 Propagate Lab
3.16 RVA Yeast Labs
3.17 The Yeast Bay
3.18 White Labs
3.19 Wyeast
3.20 Smaller Labs
4 Brett Blends (Brett only)
5 Using Brettanomyces
5.1 Primary versus Secondary Fermentation
5.2 Starter Information
5.2.1 Two Approaches to Starters
5.2.2 Pitching Rate Calculators
5.2.3 MYPG Growth Substrate and Other Laboratory Substrates
5.2.4 Cell Counting
5.2.5 Example of a Home Lab Orbital Shaker
5.3 Storing Brett
5.4 Fermentation Methods
5.4.1 Fermenting Under Head Pressure
5.5 Tips From Brewers
5.5.1 The Yeast Bay
5.5.2 Escarpment Laboratories
5.6 Pasteurization
5.7 Catching/Bioprospecting Wild Brettanomyces
6 See Also
https://www.milkthefunk.com/wiki/Brettanomyces Page 1 sur 35

6.1 Additional Articles on MTF Wiki

6.2 External Resources
7 References
Introduction of History, Characteristics, and Taxonomy

Closely related to Saccharomyces, Brettanomyces diverged from its cousin yeast more than 200 million years ago, around the same time that the first mammals emerged [4].
Both genera evolved independently to ferment sugar and produce ethanol [5][6]. Although first isolated in 1889 and again in 1899 by scientists at Guinness, the discovery of
Brettanomyces was first publicly published by the Director of laboratory of the New Carlsberg Brewery, Hjelte Claussen, in 1904 after he cultured it in 1903 from English
beers that exhibited a sluggish secondary fermentation [7][8]. At the time of discovery, Claussen was aiming to recreate the flavor profile of traditional English ales by
fermenting them with pure cultures of Saccharomyces, and either pitching pure cultures of his newly discovered Brettanomyces yeast along with Saccharomyces, or as he
preferred, after the primary fermentation of Saccharomyces [9]. Brettanomyces, along with dry hop creep, was identified as the source of secondary fermentation during long
aged ales, contributing to their lasting high carbonation [10][11][12](8 minutes in). Beer historian, Ron Pattinson (https://barclayperkins.blogspot.com/search?
q=brettanomyces), has stated that Brettanomyces was typically present in 1800's English aged beers such as stock ales, pale ales, porters, and barrel aged IPA's that were
shipped to India, and it was considered an important component of both the flavor profile of these beers and in protecting beer from contaminants via Brettanomyces
fermenting the majority of residual sugars [13] (~56 and 59 mins in )[14] (45 minutes in).
Following the discovery of this yeast by Claussen, isolates of Brettanomyces were discovered in Belgian lambic beers in the 1920's. At this time, Brettanomyces was
proposed as the genus name. The species name 'bruxellensis', meaning 'Brussels' in Latin, became the proposed species name for B. bruxellensis.. This yeast species was then
isolated from other industrial fermentations such as wine, cider, kombucha, kefir, olives, and bioethanol production. Custers was the first to attempt to describe the rest of the
genus using phenotypic characteristics in 1940. In 1960, J. van der Walt observed some species of Brettanomyces formed ascospores, and this form of Brettanomyces was
named Dekkera. However, after the initial discovery of sporulating strains of Brettanomyces, this behavior has not been reported since, therefore some scientists prefer to use
the term "Brettanomyces" to refer to this genus. Originally, a total of 9 species were attributed to the genus Brettanomyces, but after gene technology was invented, some of
these species were changed (see Taxonomy below) [15].
Although Claussen and others saw the character from Brettanomyces as a desirable character in English ales and identified its character as a hallmark quality of traditional
English ales, as pure cultures of Saccharomyces were introduced in English brewing in the early 20th century, Brettanomyces became identified as a contaminate in both
wineries and breweries due to some of the phenols, acids, and haze that it sometimes produces. These phenols and acids have generally been described as "barnyard", "burnt
plastic", "wet animal", "fecal", and "horse sweat", although some tasters describe these flavors with different terminology because they perceive certain flavor compounds
differently while some other tasters simply cannot detect certain flavor compounds at all [16][5][17]. The general viewpoint of brewers (other than Lambic brewers, Flanders
red/brown brewers, and certain Trappist brewers in Belgium, as well as Berliner Weisse brewers in Berlin, Germany) and vintners became that Brettanomyces is primarily a
spoilage organism, and this still holds in most cases today. More recently, however, the positive flavor components that have been identified in Brettanomyces beer such as
"pineapple", "stone fruits", and to some degree acetic acid, have regained popularity with brewers outside of Belgium. Some winemakers and wine tasters have also described
wines with certain flavor compounds derived from Brettanomyces as positive characteristics of some wines [18]. It is important to keep in mind that individual tasters on
tasting panels describe some flavor compounds as "negative" while others describe them as "positive" (and sometimes a mixed response is given by a taster in regards to a
certain flavor compound). This discrepancy in acceptability of flavor characteristics derived from Brettanomyces appears to be based on personal preference and experience.
For example, in some cases and for some drinkers low levels of vinyl phenols produced by Brettanomyces contribute positively to wine, while higher amounts contribute
negatively. Thus, a lower intensity of some flavor compounds can be seen as more desirable by some producers or consumers. Overall, the enjoyment or displeasure of the
various flavor compounds produced by Brettanomyces and at certain levels is largely subjective [17][19].
See also:
A collection of early 1900's papers by Claussen and Alfred Chapman; MTF post by Cory Widmayer. (https://www.facebook.com/groups/MilkTheFunk/permalink/3364
694553558735/)
Yvan de Baets believes that Brettanomyces probably played an important role in historical saison. (https://beerandbrewing-com.cdn.ampproject.org/c/s/beerandbrewing.
com/amp/brasserie-de-la-sennes-yvan-de-baets-explains-saisons-greatest-myth-the)
Presentation by Ron Pattinson on the role of Brettanomyces in historical English beer; hosted by Odd Breed Wild Ales on 02/15/2022. (https://www.facebook.com/odd
breedwildales/videos/483806376609910)
Taxonomy
It is common in scientific literature to see the names Dekkera and Brettanomyces used as the genus name, with Dekkera being the teleomorph (https://en.wikipedia.org/wiki/T
eleomorph,_anamorph_and_holomorph) version and Brettanomyces being the anamorph (https://en.wikipedia.org/wiki/Teleomorph,_anamorph_and_holomorph). There are
five species within the genus of Brettanomyces: B. anomalus, B. bruxellensis, B. custersianus, B. nanus, and B. naardenensis (one study on the genetics of B. nanus from
1990 classified B. nanus as belonging to another genus of yeast called Eeniella, however this has not been agreed upon in more recent studies [20][21][22]). The species
previously known as B. intermedius and B. lambicus have recently been genetically analyzed and reclassified as strains of B. bruxellensis [23]. Of these five species, only B.
anomalus and B. bruxellensis have been identified to have a teleomorph version. In their teleomorph version they are referred to as Dekkera anomala and Dekkera
bruxellensis [16][5][24][25]. All of the other names such as the ones often used by yeast labs (e.g. "claussenii") are derived from old nomenclature that is no longer used
scientifically (click here (http://www.sciencedirect.com/science/article/pii/S0168160515001865#t0005) for a table that lists old and new taxonomical nomenclature). Most
Brettanomyces cultures from brewer's yeast labs are classified genetically as B. bruxellensis or B. anomalus.
Recently a new species of Brettanomyces has been proposed, although classification has not been fully established. The proposed name is Brettanomyces acidodurans sp.
nov. Two strains of B. acidodurans were isolated from olive oil from Spain and Israel; however, its presence in olive oil has been described as "rare" because only two strains
were found after searching dozens of olive oils. Its closest relation is to B. naardenesis by 73% of its genetic makeup. No teleomorph form was observed. This species is a
strong acetic acid producer, and it is very tolerant of acetic acid in its environment. It can consume lactose and cellobiose but does not consume maltose. it is unknown but a
possibility that this species contributes to the vinegary taste of spoiled olive oils, although this has generally been attributed to acetic acid bacteria [26].
A genetic survey of 145 different strains of B. bruxellensis from 29 countries, 5 continents, and 9 different fermentation niches was conducted in 2018 by Avramova et al.
They found that these strains formed roughly 6 genetic groups with mostly separate ancestral lineages, and 1 group with a mixed ancestral lineage: 3 wine groups, 1 beer
group, 1 kombucha group (most distantly related to the beer group), as well as 1 tequila/ethanol group that has multiple ancestral lineages [2]. These groups are partially
determined by the identification of at least two hybridization events that happened during the evolution of B. bruxellensis, similar to the hybridization events that created the
Saaz and Frohberg subgroups of S. pastorianus (the parents of these hybridization events in B. bruxellensis, whether from different species or not, has yet to be determined

and will require whole genome sequencing of species closely related to B. bruxellensis) [6]. This was expressed mostly in the ploidy level of each group (the number of sets of
chromosomes), with 2 of the wine groups, the tequila group, and the beer group containing more sets of chromosome pairs than the other groups (diploid vs triploid; this is
thought to encourage adaption and hybridization). Additionally, the triploid wine group was generally more tolerant of SO2 than the diploid wine groups [2].
Colomer et al (2020) (https://www.frontiersin.org/articles/10.3389/fmicb.2020.00637) surveyed the whole genomes of 64 strains of B. bruxellensis, 14 strains of B. anomalus,
3 strains of B. custersianus, and 3 strains of B. naardenensis. They broke the B. bruxellensis beer group into two clades: a "farmhouse" clade which comprised of strains of B.
bruxellensis isolated from commercial craft beer breweries, and a "lambic" clade which comprised of strains isolated from spontaneously fermented Belgian lambic beers.
There was also a subdivision of the lambic clade which comprised of strains of B. bruxellensis identified with various natural origins; ethanol plants, barrel-aged beers, and
matured wines. This subclade was called "wild/wood" by the researchers. See Figure 2 (family tree) (https://www.frontiersin.org/files/Articles/495404/fmicb-11-00637-HTM
L-r1/image_m/fmicb-11-00637-g002.jpg) from the study [27].
Science has also begun to explore targeted gene manipulation of B. bruxellensis via CRISPR, which will eventually lead to a better understanding of the Brettanomyces
bruxellensis genome [28].
Morphology
The morphology of Brettanomyces can vary immensely from strain to strain (and species to species). Some strains can look similar in size and shape to S. cerevisiae under a
microscopic image, while others are elongated or much smaller. This makes it difficult to identify Brettanomyces without DNA analysis (see PCR). Morphologies of
Brettanomyces grown on agar plates can also be different from strain to strain. For example, Devin Henry found that a sample of WLP648 that contained two closely related
strains of B. bruxellensis grew completely differently on the same growth media. At first, larger, slightly off-white colonies grew on the plates (this was the first strain), and
then a few days later the second strain grew as many smaller white-colored colonies. Other strains may appear as glossy or matted with jagged edges, etc. Morphology on
agar plates can change depending on the type of growth media [29][30][31]. While genetic (PCR) identification is required for any kind of confident identification of
Brettanomyces, specialized selective media can also help identify Brettanomyces; see Selective Media.
See also:
Brettanomyces morphology examples from Remi Bonnart. (http://brettanomycesproject.com/2010/06/brettanomyces-yeast-cell-images/)

"Brett Trois – A riddle, wrapped in a mystery, inside an enigma," Sui Generis Blog; an example of S. cerevisiae appearing like Brettanomyces cells under a microscope.
(http://suigenerisbrewing.com/index.php/2014/12/15/brett-trois-a-riddle-wrapped-in-a-mystery-inside-an-enigma/)
Morphology examples on Bootleg Biology's website. (https://bootlegbiology.com/diy/microbe-portrait-gallery)
Culturing
See Laboratory Techniques.
Environment and Survival

Brettanomyces has been thought to occur naturally on the skins of fruit such as apples and grapes. However, there are only a handful of reports of Brettanomyces being
identified on the skins of fruit, and in some cases where Brettanomyces has been found, its abundance is extremely minimal [32][33][34]. In contrast, there are also studies that
indicate Brettanomyces only being found during or after food processing, which indicates that the processing equipment may be the primary source for the Brettanomyces. In
addition, Brettanomyces has been isolated in abundance from the surfaces of equipment/processed materials in wineries and breweries [16][5][35][24][36] (Table 1). For
example, an ongoing survey of wild yeasts in most of the US which isolated nearly 2,000 isolates with 262 unique species has not yet found a single occurrence of
Brettanomyces in the wild (so far they have only surveyed non-human inhabited wild areas of the US and Alaska; substrates sampled included leaves, soil, bark, moss,
mushrooms, needles, pine cones, twigs/wood, and other plant matter) [37]. It is therefore unclear that Brettanomyces found on grape skins originated there or from the
industrial processing where it is more abundant. It is also thought to disperse via fruit-flies (called "vectors" in the scientific literature), similar to how Saccharomyces travels,
although direct evidence for this has only been reported rarely and only on fruit-flies in wineries that are likely to come into contact with equipment/processed material that is
already contaminated with Brettanomyces [38][34][24][36][35]. Brettanomyces is known to be difficult to grow in a lab due to slow growth, specific nutrient requirements, or
perhaps because of a "VBNC" state (see Wild Brettanomyces for more information), which may account for the lack of evidence for fruit being the primary natural habitat for
Brettanomyces. More recently, techniques have been invented to more easily isolate and grow Brettanomyces [34][33]. There is also significant evidence that the natural habitat
of Brettanomyces might actually be the root systems of certain plants, known as the "rhizosphere" (https://www.nature.com/scitable/knowledge/library/the-rhizosphere-roots-
soil-and-67500617/). The rhizosphere refers to the complex symbiotic community of microbe populations that live on and around the root system of plants. Wild strains of
Brettanomyces have been found in the root systems of dill, common beans, sunflowers, maize, corn, jute, cassava, and grey mangroves found in the estuaries of Indonesia
[39][40][41][42][43][44][45]. See Dr. Bryan Heit's video "Where (Do) The Wild Brettanomyces Roam?" (https://www.youtube.com/watch?v=G2nhUM5PIrg) and his comments
in Milk The Funk (https://www.facebook.com/groups/MilkTheFunk/posts/5940213029340195), as well as "Philip Poole. Plant Control of the Rhizosphere Microbiome" (http
s://www.youtube.com/watch?v=BrR7G_YyfmA). For documented isolation attempts from plant rhizospheres, see Wild Yeast Isolation.
The occurrence of Brettanomyces has been more commonly identified in industrial food processing areas (wine, beer, kombucha, soft drinks, dairy products, tea, sourdough,
etc.) [46]. For example, B bruxelensis, B. anomala, and B. custersianus have mostly been isolated from wine or beer production, while B. naardenensis has mostly been
isolated from soda production [47]. Brettanomyces is not considered to be airborne; however, one study has demonstrated a very small amount of cells in the air at wineries
where wine with Brettanomyces in it was being handled (most of the yeasts found in the air were Aureobasidium and Cryptococcus, which aren't considered spoilage
organisms in beer and wine). This set of studies also determined that very specific methodology was needed in order capture Brettanomyces from the air, and indicated that
the yeast was "stressed". While it is possible for Brettanomyces to be briefly carried by gusts of air, it only happens in the vicinity where the Brettanomyces beer or wine is
being bottled (more so) or is actively fermenting (less so) [48]. Good cleaning and sanitation and cold temperatures should be employed to keep Brettanomyces from
contaminating other equipment; however, flying insects are also a potential cause for contamination of Brettanomyces (although evidence for this is lacking).
Brettanomyces is commonly isolated from the surface of wood structures within breweries, wineries, and sometimes cideries (although the median occurrence of
Brettanomyces in barrels may be very low to none within a given winery or brewery depending on their hygiene and other factors [49][50]). These include structures such as
wooden fermentation vessels, walls of the building, as well as the inside surface of wood barrels and actually buried within the wood of barrels. Brettanomyces has been
easily cultured from within the wood of oak barrels up to 4 mm into the wood, and occasionally as deep as 5 to 8 mm, depending on the age and variety (slightly higher
populations tend to survive in French oak over American oak, and one study found that the Brettanomyces was able to penetrate the French oak barrels up to 8 mm, while
only penetrate American oak barrels up to 4 mm) of the barrel [23][51], with the highest concentration of surviving cells being at the top staves where oxygen is more
accessible (although Cartwright et al. found the opposite was true, perhaps due to methodology of sampling or a difference in SO2 concentrations). Some strains are able to
utilize the cellulose of the wood as a carbon source, and occasionally form pseudohyphae within the wood which expands the surface area of the cells allowing them more
access to nutrients and allowing them to survive in nutrient deficient environments [51]. Ozone gas has been shown to be an effective way to kill Brettanomyces that is buried
in the wood of oak barrels, but the ozone must be applied for an adequate time to allow for the ozone to diffuse into the oak. Ozone has also been shown to be an effective

way of greatly reducing but not completely eliminating the number of Brettanomyces on wine grapes. Liquid ozone has been shown to be less effective at eliminating
Brettanomyces. Heating the inside of the oak barrels to 60°C for 20 minutes with hot water or steam has also been found to be an effective way of killing Brettanomyces
within the wood of barrels (see Barrel Sanitation for information on pasteurizing barrels) [52][53]. Although the role of Brettanomyces appears to be limited in distillation, it
has been isolated during the fermentation process of tequila making. It has also been isolated from drains, pumps, transfer hoses, and other equipment that is difficult to
sanitize. The survivability of Brettanomyces has also partly been attributed to its ability to form a biofilm (in particular B. bruxellensis). Microorganisms that can form a
biofilm are more resistant to chemical cleaning agents and sanitizers than those that don't. Brettanomyces has therefore been identified as a significant contaminate for
breweries and wineries. Oak barrels from wineries with unsanitary practices, in particular, have been identified as common contamination sites for B. bruxellensis.
Brettanomyces is also commonly found in sherry, and is found (although only rarely) in olive production, lemonade, kombucha, yogurt, pickles, and soft drinks. B. anomalus
and B. bruxellensis are generally found much more commonly than the other three species of Brettanomyces [16].
Unlike most genera of yeast, Brettanomyces has the characteristics of being very tolerant to harsh conditions, including high amounts of alcohol (up to 14.5-15% ABV
[54][23]), a pH as low as 2 [55], and environments with low nitrogen [5] and low sugar sources [56]. It has been reported that B. bruxellensis is more tolerant of high levels of
bicarbonate than compared to S. cerevisiae (levels above 100 mg/l slow the fermentation of B. bruxellensis, but do not completely inhibit it, with up to 400 mg/l being tested
in one study) [57]. It has been reported that some strains require a very low concentration of fermentable sugars (less than 300 mg/L) and nitrogen (less than 6 mg/L), which is
less than most wines contain [58]. Some strains are able to utilize ethanol, glycerol, acetic acid, and malic acid when no other sugar sources are available [56]. This capability
allows Brettanomyces to survive in alcoholic beverages such as beer, wine, and cider. In alcoholic beverages, B. bruxellensis tends to lag after the primary fermentation with
Saccharomyces. It is believed that during this lag phase, B. bruxellensis adapts to the harsh conditions of the beverage (low pH, high concentrations of ethanol, and limited
sugar/nitrogen sources). After this lag phase, B. bruxellensis can grow and survive when no other yeasts can. Brettanomyces is also more resistant to pH and temperature
changes, and tolerant of environments limited in oxygen (although Brettanomyces prefers the availability of at least a little bit of oxygen). Scientifically, which specific
nitrogen and carbon sources B. bruxellensis uses in these stressful environments has not received much research [16]. One study from Dr. Charles Edwards (https://www.wine
sandvines.com/news/article/200000/New-Tools-to-Limit-Wine-Spoilage) found that a combination of keeping wine under 54°F (12.2°C) and alcohol at or above 14%
resulted in a decline of B. bruxellensis populations for up to 100 days for two strains that were tested. The study found that neither of the strains grew well at 14% and
stopped growth completely at 16% ABV in wine, but one strain grew better than the other at 15%, demonstrating the genetic diversity of Brettanomyces. The researchers
concluded that a combination of high ethanol and cold temperatures as well as sulfur dioxide, chitosan, and filtration could be used to control Brettanomyces in winemaking.
Brettanomyces has been found to be able to grow at temperatures as low as 50°F (10°C) and as high as 95°F (35°C); see fermentation temperature for more information [59].
The genetic diversity of Brettanomyces is particularly wide. For example, one study that analyzed the whole genomes of 53 strains of B. bruxellensis found that the overall
genetic diversity between different strains of B. bruxellensis was higher than strains of S. cerevisiae (however, the entire gene set, known as the pangenome, of all the genes
among all of the strains of B. bruxellensis is much smaller than the entire gene set of S. cerevisiae) [6]. Some studies have indicated that strains of B. bruxellensis have
adapted to specific environments. For example, one study found that strains of B. bruxellensis isolated from wine had 20 genes involved in the metabolism of carbon and
nitrogen, whereas strains isolated from beer did not. This indicated that B. bruxellensis strains living in wine have adapted to the harsher environment of wine [16]. Another
study found that one out of the two strains tested that were isolated from soda could not ferment maltose, and only strains isolated from wine were able to grow in wine and
the beer/soda strains did not. The wine strains were also more resistant to sulfites, which are commonly used in the wine industry to prevent microbial contamination [46]. The
whole genome sequencing of one strain of B. naardenensis and lambic strains of B. bruxellensis found that they are missing the genes associated with nitrate utilization,
indicating that the assimilation of nitrates is not required to survive in beer, perhaps because of the abundance of nitrogen from other sources found in beer [47][27].
The addition of vitamins can have a positive impact on Brettanomyces growth. For example, while Brettanomyces does not need riboflavin (vitamin B2) or thiamine (vitamin
B1) in order to grow, the presence of either or both of these two vitamins encourages Brettanomyces growth [60]. Other vitamins such as p-aminobenzoic acid (PABA), folic
acid (vitamin B9), nicotinic acid (vitamin B3), pantothenic acid (vitamin B5) are also not required for most strains of Brettanomyces to grow. The presence of alcohol can
increase the dependence on vitamins for some strains of Brettanomyces to grow. For example, Myo-inositol (vitamin B8) and thiamine (vitamin B1) were required by two
strains of B. bruxellensis when grown in 10% ethanol but not in 0% ethanol. Biotin (vitamin B7) is the one exception, and it was found that the lack of biotin inhibited the
growth of some strains of B. bruxellensis. Other studies contradict these previously mentioned findings, showing that thiamine was not required by the strains of B.
bruxellensis tested, that pyridoxine was required, and biotin was not required. These discrepancies between scientific studies are probably due to the genetic differences
between the strains selected, the growth media chosen by the scientists, and/or the growth conditions [61].
Sulfur Tolerance
Sulfite and SO2 inhibits the growth of Brettanomyces, and is often used in the wine industry to prevent the growth of Brettanomyces (although Brettanomyces is usually seen
is a contaminant in wine, some wineries have identified small amounts of flavors from Brettanomyces as being beneficial to certain wine styles, and is said to increase the
complexity and impart an aged character in young wines [16]) [62]. However, it has been shown that wine strains of B. bruxellensis could survive dosages of up to 1 mg/L of
molecular SO2, and the very high dosage of 2.1 mg/L was needed to kill Brettanomyces in wine [23]. This dosage of molecular SO2 requires a total amount of SO2 that is
beyond legal limits (350 mg/l [63]; see this Cornell University (https://grapesandwine.cals.cornell.edu/newsletters/appellation-cornell/2012-newsletters/issue-12/article-contai
ns-sulfites/) blog post that explains the difference between free and molecular SO2) and has negative effects on wine. One study found that out of 145 strains of B.
bruxellensis, 107 of which were wine strains with the rest being from beer, tequila, kombucha, etc., 36% of them were either tolerant (lagged growth, but achieved full
growth eventually) or resistant (no lagged growth, and achieved full growth) to 0.6 mg/L of molecular SO2. 46 of the 52 resistant/tolerant strains were wine strains, thus
demonstrating that wine strains of B. bruxellensis are generally more tolerant of SO2 than strains of B. bruxellensis that are found in other types of beverages. It has been
demonstrated that the wine strains have adapted to the conditions of winemakers adding SO2 to wine [64][65]. There is also variability between wine strains in their ability to
tolerate SO2 [66]. For example, Brettanomyces strains isolated from sweet wine tend to be more tolerant of sulfur dioxide than strains isolated from dry wine [67]. In addition,
it has been proposed that SO2 and other currently unidentified environmental stress factors can induce a so-called "viable but nonculturable" (VBNC) state in Brettanomyces,
which means that Brettanomyces cells in this state cannot grow or be cultured on traditional media but can remain viable and create a low amount of phenol character (see
VBNC in Yeast) [68]. Some strains of Candida pyralidae, Wickerhamomyces anomalus, Kluyveromyces wickeramii, Torulaspora delbrueckii and Pichia membranifaciens
have been found to produce toxin that inhibits Brettanomyces, and these toxins have been proposed as an alternative to SO2 as a way to kill Brettanomyces (killer wine strains
of Saccharomyces cerevisiae do not kill Brettanomyces; see Killer Wine Yeast for more information). Kaolin silver complex (KAgC) (http://www.laboratoriosenosan.com/en/
effectiveness-of-kaolin-silver-complex/) has been found to inhibit Brettanomyces and acetic acid bacteria in wine when used in legal dosages, and has been proposed as a
replacement for SO2 or to minimize the use of SO2 [69]. Other proposed replacements for SO2 as a way to inhibit Brettanomyces in wine include high pressure processing (htt
ps://en.wikipedia.org/wiki/Pascalization) and pulsed electric fields (https://www.sciencedirect.com/science/article/abs/pii/S0255270106001929) [70][71].
A study by Cibrario et al. (2019) looked at the genome of B. bruxellensis strains in bottles of wine going back as far as the year 1909 revealed that the SO2 tolerant triploid
strains only started appearing after the year 1990, which corresponds to when the wine industry started using SO2 in most wine production (although it can also mean that the
triploid strains are not as good at surviving in bottles of wine long-term compared to the diploid strains that have been isolated from much older bottles of wine; this will be
determined to be the case or not in the future as bottles of wine from the 1990s continue to age). They also identified dozens of examples of wineries throughout France and
Italy where the same strain of B. bruxellensis was found in multiple vintages of bottles of wine going back many decades, indicating that individual B. bruxellensis strains
become long-term residents in wineries. Some identical strains have been found in different regions and even different continents, indicating that some strains have traveled

not just due to traditional vectors such as insects or birds, but also probably due to human transportation such as wine bottle imports/exports or exchanges between industrial
processes. This also indicates that while B. bruxellensis becomes rather sedentary and a constant resident in wineries, it can also adapt to the different winemaking conditions
in different regions once it's been transported (different grapes, different climates, different fermentation temperatures, etc.), including adapting to improved modern hygienic
practices such as higher SO2 treatment. Overall, the results from this study suggest that Brettanomyces is able to adapt to living alongside certain human industries and has
done so for at least a couple of centuries [72]. The genetic differences between the fermentation substrates (beer, wine, etc.) were lower but still significant, and this was
explained by the frequent cross-over of equipment such as wine barrels being used for beer fermentation. When comparing the geographic differences, they found geography
contributed only 5% of genetic differences, while geography explained more than 50% of genetic differences in non-wine strains, suggesting that beer, kombucha, and tequila
strains are more localized genetically than wine strains and that humans probably helped the wine strains travel across the globe. They also found that although one study
reported spore-forming versions of B. bruxellensis (referred to as Dekkera bruxellensis), the genetic makeup of the analyzed strains determined their ability to sporulate to be
non-existent or rare (only one study that we know of by Walt and Kerken in 1960 (https://link.springer.com/article/10.1007%2FBF02539015) has reported sporulation in
Brettanomyces only on specific agar types with vitamins added, indicating that sporulation in Brettanomyces is extremely rare) [2]. See also Richard Preiss's discussion of this
study on MTF (https://www.facebook.com/groups/MilkTheFunk/permalink/2022801681081369/). A study by Bartel et al. (2021) (https://academic.oup.com/femsyr/advance-
article-abstract/doi/10.1093/femsyr/foab036/6293842) used laboratory adaptive evolution methods to demonstrate that some strains of B. bruxellensis strains did in fact adapt
to SO2 usage in wineries [64].
See also:
Calculations for SO2 additions (http://www.foodsci.purdue.edu/research/labs/enology/Winemaking%20Calculations%20Spring%20Workshop%204_29_2011.pdf?fbcli

d=IwAR3WnDfo6k9hqfo4EvnWrp4GwFiB7qHoexifJBfLnIEIhLwRtC2Hm7qkUug) and associated MTF discussion (https://www.facebook.com/groups/59256031743
8853/?comment_tracking=%7B%22tn%22%3A%22R2%22%7D).
Viable But Nonculturable Yeast.
Biofilm
Brettanomyces has the ability to form a biofilm. Biofilm formation is a survival mechanism induced by stress whereby the
cells adhere to non-living surfaces such as plastic and stainless steel [75]. After adhesion to the surface, the cells produce a
protective layer of proteins and polysaccharides that help protect the organism from cleaning and sanitizing agents.
Joseph et al. (2007) tested 36 wine strains of B. bruxellensis for biofilm formation over a 10 day period. They found that
just under half of the strains formed a biofilm, and about half of those formed considerable and consistent biofilms
throughout the tests. Almost all strains tested (95%) adhered to a surface with 0.1% glucose within 6 hours of contact (the
same conditions that get Saccharomyces cerevisiae to adhere to a surface; longer contact with surfaces and higher
residual sugar could encourage Brettanomyces to adhere more readily to surfaces). A juice-based growth media in the
range range of 2 - 4.5 pH was tested for biofilm formation and 3-4 for cell adhesion to a surface, and for most of the
strains they formed stronger biofilms and adhered better in the higher pH growth media (4.5 pH being the highest tested).
Under a pH of 3.5 significantly dropped biofilm formation and adherence, indicating that something about pH affects the First evidence of possible (unconfirmed [73])
cells ability to attach themselves. The researchers concluded that winemakers should keep wine in the lower end of the chlamydospore cell structures of B. bruxellensis, found
pH range (3.5). Six different types of cleaners were tested to see how well they removed the biofilms: keytones + in a biofilm. Photo by Lebleux et al. (2019) (https://ww
surfactant detergent, quaternary ammonia + surfactant detergent, sodium hydroxide (caustic soda), sodium carbonate w.sciencedirect.com/science/article/abs/pii/S016816051
(soda ash), sodium hydroxide + surfactant (alkaline detergent), and chlorine (sanitizer, not a detergent). They found that 9303952) [74].
only caustic soda was consistently efficient at removing the biofilm. The chlorine, while it did not remove the biofilm,
still killed all of the Brettanomyces cells, and it was presumed that the other cleaners might have killed the
Brettanomyces, but that was not tested for. They also tested to see if cells that were adhered to a surface could be cleaned. Again, the caustic soda performed consistently the
best, but the ammonia + surfactant cleaner and the quaternary ammonia + surfactant detergent also effectively removed adhered cells. The other cleaners varied in how well
they removed adhered cells from a surface [76].
Dimopoulou et al. (2019) studied Brettanomyces bruxellensis biofilms from each of the genetic branches of B. bruxellensis. They found that for the wine strains biofilms
formed more readily when grown in wine must rather than YPD media; however, the beer strains grew biofilms equally well in wine must and YPD media. The biofilms
contained a large portion of saturated fatty acids and a smaller portion of monounsatured fatty acids. The amount of exopolysaccharide produced varied widely across the
strains tested per cell population, with some wine strains producing little EPS (40 mg/L/OD), beer strains producing moderate amounts, and the other wine group producing a
high amount (100 mg/L/OD). Additionally, the different strains displayed a varying degree of negative cell wall charges, with the beer and tequila strains being more
negatively charged than wine strains, which could help them adhere to surfaces and form biofilm [77].
Lebleux et al. (2019) measured biofilm density for 12 strains across 5 of the genetic groups of B. bruxellensis. All of the strains produced a biofilm when in contact with a
surface (polystyrene and stainless steel, in the case of this study), and the thickness of the biofilm was proportional to the cell size of each strain. The biofilms contained
filamentous cells that started from the base of the biofilm and extended upward, indicating multiple layers. The biofilms also contained exopolysaccharides (EPS), but the
makeup of the EPS was not analyzed and this was identified as a goal for further study. The average thickness was only 9.45 µm which is much thinner than other biofilm-
forming yeast species (Candida and a biofilm-producing strain of S. cerevisiae [78]). They found that one or two strains were less dense (contained fewer cells) than the
average. A couple of the strains grew a biofilm a little slower than average. Two of the strains in biofilm form were added to wine; each of the biofilms released cells into the
wine, although one strain released more cells than the other. Introduction to the wine first led to cell death for some cells due to the harsh environment of the wine, but after
several days the B. bruxellensis strains began to re-grow in the wine. It was observed that for one of the strains, the cells appeared larger than normal, round, and had thicker
cell walls, possibly forming what is known as chlamydospore cell structures (https://en.wikipedia.org/wiki/Chlamydospore). It was not confirmed in the study whether these
cells were actually chlamydospores, and their structure could be due to relatively insignificant reasons [73]. Chlamydospore cell structures are known to help certain species of
non-yeast fungi survive harsh environments; however, it has not yet been established that yeast with chlamydospore cell structures helps them survive harsh conditions, and
this was also identified in the study as an area for further research [74].
See also:
Quality Assurance
Pellicle
UV Light
There is some evidence that Brettanomyces can be sensitive to high levels of light. Catrileo et al. (2021) (https://www.frontiersin.org/articles/10.3389/fmicb.2021.747868/full
) showed that under laboratory conditions, Brettanomyces bruxellensis was not able to grow when exposed to a 2500 lux and 4000 lux light source. For reference, the lux of
indirect daylight is around 10,000 - 25,000 and the lux of office lighting is usually between 350 and 500 [79]. However, when p-coumaric acid, a phenolic precursor that is

present in plants and fruits (including malted barley and wheat), is present, certain genes are expressed during the growth of B. bruxellensis that allow it to adapt to the high
light exposure conditions. While this study does not show at what level light begins to affect B. bruxellensis (the lowest light intensity that they tested was 2500 lux),
Woodward et al. (1978) (https://journals.asm.org/doi/abs/10.1128/jb.133.2.692-698.1978) demonstrated that Saccharomyces cerevisiae growth is unaffected by light until
about 1,250 lux, at which point it begins to inhibit growth and the transfer of nutrients across the cell membrane [80][81].
As a follow up question within Milk The Funk group on Facebook regarding if lower levels of light could impact Brettanomyces growth, Richard Preiss of Escarpment Labs
performed an in-house experiment to grow Brettanomyces in the presence of standard fluorescent lights and reported finding no impact of the lights on Brettanomyces growth
[82].
Brettanomyces Metabolism
Like Saccharomyces, Brettanomyces is Crabtree (https://en.wikipedia.org/wiki/Crabtree_effect) positive (produces
alcohol in the presence of oxygen and high sugar concentration), and is petite (https://en.wikipedia.org/wiki/Petite_mutati
on) positive (unable to grow without carbon sources, and forms small colonies when able to grow on growth media) [16].
Perhaps the most differentiating characteristic of Brettanomyces is its preference to ferment glucose in the presence of
oxygen to produce ethanol and acetic acid, which is the opposite preference in Saccharomyces where the presence of
oxygen inhibits fermentation (dubbed the "Pasteur effect (https://en.wikipedia.org/wiki/Pasteur_effect)"). Also opposite
of most yeasts including Saccharomyces, in a completely anaerobic environment Brettanomyces ceases alcoholic
fermentation for about 7-8 hours before adapting to the anaerobic conditions [23]. This was initially dubbed the "negative
Pasteur effect" by Custers, and later the "Custers effect" by W. A. Scheffers [83][84]. A notable exception to this is the
species B. naardenensis, which only produces ethanol when oxygen is limited [47]. Dr. Custers memorial. Photo by Jan Beekaa Lemmens (h
ttps://www.facebook.com/groups/MilkTheFunk/permali
Despite the Custers effect in Brettanomyces, this genus is not classified as an "oxidative yeast" but rather as a nk/3061430667218460/).
"fermentative yeast" since oxidative yeasts produce little to no ethanol in the presence of glucose, and only grow as scum
on the surface of a liquid rather than within the liquid [85][86][87][88][89].
The Custers effect is abolished under anaerobic conditions when nitrate (https://en.wikipedia.org/wiki/Nitrate) is available. Under conditions where there is no oxygen, as
long as nitrates are available, it has been shown that Brettanomyces can produce ethanol just as capably as S. cerevisiae [90][91]. Brewers who prefer the character of
Brettanomyces in their beer can take advantage of this ability by limiting oxygen and providing a food source for Brettanomyces (aka beer or wort). See Nitrogen Metabolism
below.
Carbohydrate Metabolism and Fermentation Temperature

Brettanomyces is able to ferment a wide range of sugars. All strains can ferment glucose, and many strains can ferment sucrose, fructose, and maltose, although at a slower
rate than glucose. The ability of Brettanomyces to produce invertase enzyme which breaks sucrose down into glucose and fructose has been attributed to horizontal gene
transfer from an unknown bacteria at some point in the evolution of Brettanomyces [92]. Some strains can also ferment galactose, mannose, ethanol, acetic acid, malic acid,
and glycerol, although historically there are some contradicting studies in science regarding the specifics (more recent studies tend to use better methods), probably due to the
genetic diversity of Brettanomyces species, and many previously published studies do not specify whether testing conditions were aerobic or anaerobic even though the
availability of oxygen effects whether or not certain sugars can be fermented by a given strain of Brettanomyces [24][16][56]. For example, the species B. naardenensis can
ferment a wide range of carbon sources, including galactose, maltose, xylose, trehalose, cellobiose, rhamnose, and arabinose [47]. Acetic acid, glycerol, succinic acid, and
ethanol are only consumed if oxygen is present [16]. The addition of H+ acceptors such as acetaldehyde, acetone, pyruvic acid, and other carbonyl compounds, stimulates
anaerobic fermentation. Small amounts of oxygen also stimulate fermentation [83]. The presence of small amounts of oxygen can allow some strains of Brettanomyces to
utilize certain carbon sources. For example, several strains of B. bruxellensis can consume ethanol, glycerol, and acetic acid as food sources only when at least a low amount
of oxygen is present (semi-aerobic conditions) and no other sugar is available. Acetic acid and glycerol are used as food sources by some strains only under fully aerobic
conditions, but not under semi-aerobic or anaerobic conditions. It has been hypothesized that acetic acid and glycerol are only consumed by Brettanomyces when ethanol and
other food sources are no longer available [56].
Brettanomyces strains may possess both alpha and beta-glucosidases. Beta-glucosidase is intracellular (works on sugars that are passed into the cell through the cell wall),
while alpha-glucosidase is both intracellular and extracellular (released into the environment by the cell). [93][94] These enzymes allow Brettanomyces strains to break down a
broad range of sugars, including long-chain carbohydrate molecules (polysaccharides, dextrins, and cellulose/cellobiose), and to liberate glycosidically bound sugars which
are unfermentable to Saccharomyces yeasts. [24][95].
Extracellular and intracellular alpha-glucosidase activity has been shown to break down sugars up to 9-12 chain carbons in one strain of B. lambicus (now classified as B.
bruxellensis), which is partly responsible for the slow, over-attenuation of wort that some strains of Brettanomyces an achieve in beers such as lambic and American sour
beers [83][16]. Alpha-glucosidases are the enzymes that allow them to break down maltose, turanose, melezitose, and trehalose, as well as dextrins such as maltotetraose and
maltopentaose. These enzymes work by cleaving off glucose that can be directly consumed by the cell, leaving a shorter chain sugar behind which is then further broken
down. In the case of extracellular alpha-glucosidase activity, this breakdown of complex sugars occurs outside of the cell and may benefit other microorganisms if present
such as lactic acid bacteria. These dextrins are left over after a normal Saccharomyces fermentation [24]. Some other polysaccharides can be fermented by Brettanomyces,
including starch, laminarin, and pectin [54]. The more complex the starch or sugar, the slower it is hydrolyzed by the alpha-glucosidase enzymes. The optimal pH for the
alpha-glucosidase enzyme produced by one strain of B. bruxellensis was 6 and at a temperature of 39-40°C (102-104°F), and its activity was greatly reduced below a pH of
4.5 and above 8 (although citric acid was used as a buffer, and its effects on the enzyme was not compared to other acids), which might contribute to slower Brettanomyces
fermentation in acidic beers [94]. A survey of 84 strains from several species of Brettanomyces showed that there is a wide variability in the ability of different strains to
ferment maltose, with some strains not being able to ferment it at all and others fermenting it very slowly, suggesting that alpha-glucosidase is not functional or poor in some
strains. Additionally, when maltose is present instead of just glucose, the researchers saw an increased lag during the growth phase [27].
B. bruxellensis and B. nanus can also produce oligo-1,6-glucosidase, which hydrolyze the alpha-1,6 linkages in starch and glycogen to produce oligosaccharides, and then
further break down these oligosaccharides to produce sugars with alpha-1,4 linkages (for example, maltose, in the case of starches from malted barley [96]). These alpha-1,4
linkages (maltose) are then further broken down by the maltose enzyme by strains of Brettanomyces or other present microbes that produce this enzyme [25]. Unlike for some
domesticated diastatic S. cerevisiae strains, this ability for Brettanomyces to break down starches has occurred in the wild without domestication [92][97][98].
Beta-glucosidases can break down the beta-glycosidic bond in disaccharides (cellulose, cellobiose, and gentiobiose) [99][16], as well as glycosides. Glycosides are sugar
molecules connected to other organic compounds such as acids, alcohols, and aldehydes which are flavor and aroma inactive due to the sugar molecule attached. By cleaving
off the sugar molecule through beta-glucosidase activity, Brettanomyces species can liberate these compounds (called aglycones) into their aroma-active and flavor-active

states, or states that may become flavor and aroma active through further modification [100]. Therefore some Brettanomyces strains are believed to be able to produce novel
flavors and aromas from hops, fruits, and fruit pits that Saccharomyces yeasts cannot produce. In addition, the liberated aroma and flavor active compounds may be further
processed by Brettanomyces through ester production or destruction pathways. See Beta-Glucosidase Activity for more information.
There is a highly genetic diversity between strains of Brettanomyces species, both in a genotypic and phenotypic (http://www.diffen.com/difference/Genotype_vs_Phenotype)
sense [54]. Not all species are capable of consuming the same types of sugars. For example, B. anomalus (aka claussenii) are generally able to ferment lactose, but B.
bruxellensis is generally not. Different strains within the same species may not be able to ferment the same types of sugars [101][102]. For example, some strains are not able to
ferment maltose (often B. anomalus strains), which is almost half the sugar content of wort [103]. Such strains would not be a good choice for 100% Brettanomyces
fermentation.
The ability of a given Brettanomyces strain to ferment different types of sugars might be at least partially linked to its source of isolation. For example, in one study a strain of
B. bruxellensis isolated from a soft drink could not ferment the disaccharides maltose, turanose, or the trisaccharide melezitose, whereas all of the other B. bruxellensis strains
isolated from beer and wine could ferment these disaccharides/trisaccharide. The beer strains, however, were unable to ferment cellobiose or gentiobiose, as well as arbutin
and methyl-glucoside. The wine strains were able to ferment these disaccharides, perhaps because they were adapted to the environment in which they were isolated (wine
barrels). Further studies are needed to see if this is a trend throughout the species [54]. Daenen et al. (2007) found that none of the B. bruxellensis strains isolated from lambic
that they tested could utilize cellobiose (see glycosides below). This data point challenges the belief that Brettanomyces lives in wooden barrels because it is able to consume
the cellobiose of the wood. A study by Tyrawa et al. from Escarpment Laboratories agreed that wine isolated strains were generally better at fermenting cellobiose than
strains isolated from beer at 15°C (59°F), however at 22.5°C (72.5°F) some of the beer strains started to utilize cellobiose, indicating that temperature plays a role in whether
Brettanomyces can ferment certain sugars [104]. Colomer et al. (2020) surveyed the whole genome of 64 strains of B. bruxellensis and found that beer strains are more likely
to be able to ferment maltose (alpha-glucosidase) but not cellobiose (beta-glucosidase) and wine/wild strains tend to have the opposite tendency, indicating that B.
bruxellensis strains have adapted to the environments in which they live over time [27].
Currently, research into how well Brettanomyces strains ferment the trisaccharide maltotriose has not been explored much by science. However, one study found that B.
custersianus can ferment maltotriose. Another study found that all 7 strains of B. bruxellensis tested could ferment maltotriose, but not the trisaccharide raffinose. More
investigation into this possibility is needed [105][54].
Just like in other yeast species, the temperature has a direct effect on the rate of fermentation for Brettanomyces. The optimal fermentation rate temperature range for
Brettanomyces is between 22-32°C (77-90°F). However, one study by Tyrawa et al. found that several strains of B. bruxellensis fermented at 30°C "smelled terrible" of
aromas typical of sulfur and autolysis [104][106]. At 20°C (68°F) fermentation rate is about half as slow. Brettanomyces will still grow at temperatures as low as 15°C (59°F)
with about a third of strains being able to grow as low as 10°C (50°F) [107][108] but growth will be much slower. However, one study showed a slightly higher viability during
the full-time period of fermentation at 15°C as opposed to the optimal growth and fermentation temperature range of 20-32°C. The growth rate at 15°C, while still slowly
active, varies from strain to strain with some strains growing very poorly. Carbohydrates are consumed much slower, with cellobiose metabolizing ceasing for some strains
(although phenol production stayed the same between 15°C and 22.5°C) [104]. At a temperature of 35°C (95°F), fermentation is greatly inhibited due to cell death for most
strains of B. bruxellensis, with about a third of strains able to grow as high as 37°C (98.6°F) [107], and complete elimination in wines at 50°C for 5 minutes (see also Barrel
Sanitizing and Pasteurization) [109][110]. B. naardenensis is less tolerant to extreme temperatures, and it has been demonstrated that this species cannot grow at 30°C or higher
[47]. The primary byproducts of Brettanomyces fermentation, which are ethanol, acetic acid, and CO2 are produced both during growth but also during fermentation after
growth has stopped. At the more optimal fermentation temperatures of 25-32°C, ethanol and acetic acid are produced faster from fermentation, but the amounts of ethanol
and acetic acid produced from fermentation are not affected by temperature (i.e. higher temperatures do not produce more ethanol and acetic acid from the same amount of
sugar, they are just produced faster at warmer temperatures because fermentation is faster) [111]. The warmer temperature ranges that are ideal for Brettanomyces fermentation
rates and growth rates may still produce unfavorable flavors such as higher alcohols; however, this has not been analyzed as far as we know [112]. For more information on
how fermentation temperature affects the flavor compounds of 100% Brettanomyces fermentation, see Impact of Fermentation Temperature.
The below table is an example of the variety of sugar types that different strains/species of Brettanomyces banked at the National Collection of Yeast Cultures (https://catalog
ue.ncyc.co.uk) can ferment under semi-aerobic fermentation and aerobic growth (the semi-aerobic fermentation value is probably more useful for brewers since oxygen
availability is limited during fermentation in normal brewing practices):
Cellobiose Soluble Starch

Species NCYC Glucose (Semi- Sucrose (Semi- Maltose (Semi- Lactose (Semi- Lactic
[102] (Semi- (Semi- Glycerol* Ethanol*
Num Aerobic/Aerobic) Aerobic/Aerobic) Aerobic/Aerobic) Aerobic/Aerobic) Acid*
Aerobic/Aerobic) Aerobic/Aerobic)
NCYC 2
(https://c
atalogue.
Weak or
D. anomala ncyc.co.u +/+ +/+ +/+ Weak or Latent/+ +/+ -/- - -
Latent
k/dekker
a-anomal
a-2)
NCYC
749 (http
s://catalo
gue.ncyc. Weak or
B. anomalus +/+ +/+ -/- +/+ +/+ -/- Latent -
co.uk/bre Latent
ttanomyc
es-anoma
lus-749)
NCYC
615 (http
s://catalo
gue.ncyc. Unknown/Weak or
B. anomalus +/+ Weak or Latent/+ Weak or Latent/+ +/+ +/+ + Unknown Unknown
co.uk/bre Latent
ttanomyc
es-anoma
lus-615)
NCYC
449 (http
s://catalo
gue.ncyc.
D. anomala co.uk/de +/+ -/+ -/- Unknown/+ Weak or Latent/+ Unknown/Unknown Unknown Unknown Unknown
kkera-an
omala-44

9)
NCYC
2818 (htt
ps://catal
ogue.ncy
B.
c.co.uk/b +/+ +/+ +/+ -/- -/- -/- - + -
bruxellensis
rettanom
yces-bru
xellensis-
2818)
NCYC
2263 (htt
ps://catal
Weak or
D. ogue.ncy Weak or Weak or
+/Weak or Latent +/Weak or Latent +/Weak or Latent Latent/Weak or -/- -/Weak or Latent -
bruxellensis c.co.uk/d Latent Latent
Latent
ekkera-br
uxellensi
s-2263)
NCYC
1559 (htt
ps://catal
D. ogue.ncy
+/+ Weak or Latent/+ Weak or Latent/- Weak or Latent/+ -/- -/- + + -
bruxellensis c.co.uk/d
ekkera-br
uxellensi
s-1559)
NCYC
823 (http
s://catalo
D. gue.ncyc.
+/+ +/+ +/+ Unknown/- -/- -/- - - -
bruxellensis co.uk/de
kkera-bru
xellensis-
823)
NCYC
395 (http
s://catalo
D. gue.ncyc.
+/+ +/+ +/+ -/Unknown -/- Unknown/Unknown Unknown Unknown Unknown
bruxellensis co.uk/de
kkera-bru
xellensis-
395)
NCYC
370 (http
s://catalo
gue.ncyc.
B.
co.uk/bre +/+ +/+ Weak or Latent/+ Unknown/- Unknown/- Unknown/Unknown Unknown Unknown Unknown
bruxellensis
ttanomyc
es-bruxel
lensis-37
0)
NCYC
362 (http
s://catalo
gue.ncyc.
B.
co.uk/bre +/+ Weak or Latent/+ Weak or Latent/+ Unknown/- -/- Unknown/Unknown Unknown Unknown Unknown
bruxellensis
ttanomyc
es-bruxel
lensis-36
2)
NCYC
3450 (htt
ps://catal
ogue.ncy
B. Weak or Weak or
c.co.uk/b Weak or Latent/+ -/ Unknown -/- -/+ -/- -/- -
naardenensis Latent Latent
rettanom
yces-naar
denensis-
3450)
NCYC
3015 (htt
ps://catal
ogue.ncy
B.
c.co.uk/b +/+ -/- -/Weak or Latent -/+ -/- -/+ - + +
naardenensis
rettanom
yces-naar
denensis-
3015)
NCYC
924 (http
s://catalo
gue.ncyc.
B. Weak or
co.uk/bre +/+ -/- -/+ Unknown/+ -/- -/Weak or Latent - +
naardenensis Latent
ttanomyc
es-naarde
nensis-92
4)
NCYC
899 (http

s://catalo
B. gue.ncyc. Unknown/Weak or Weak or Weak or
co.uk/bre Weak or Latent/+ -/- -/Weak or Latent -/- -/- -
naardenensis Latent Latent Latent
ttanomyc
es-naarde
nensis-89
9)
NCYC
813 (http
s://catalo
gue.ncyc.
B. Weak or
co.uk/bre Weak or Latent/+ -/- -/+ Unknown/+ -/+ -/- -/Unknown +
naardenensis Latent
ttanomyc
es-naarde
nensis-81
3)
* Measured only under aerobic utilization and growth because these compounds can only be metabolized under aerobic conditions [16].
Glycosides and Beta-Glucosidase Activity
Glycosides are flavorless compounds often found in plants/fruits that are composed of a molecule (often a flavor active compound) bound to a sugar molecule. The glycosidic
bond can be broken, releasing the sugar molecule and the potential flavor active compound. These bonds can be broken with exposure to acid, as well as specific enzymes
(beta-glucosidase) which can be added synthetically or produced naturally by some microorganisms, including some strains of Brettanomyces that have beta-glucosidase
enzyme activity (mostly B. anomalus strains) [113]. The release of flavor molecules from glycosides is thought to contribute to the flavor development of aging wines, as well
as kriek (cherry) lambic [114]. It is speculated that flavor compounds from hops can also be released from glycosides [93]; however, at least one study has shown no significant
difference in a blind taste test between hopped beer exposed to the beta-glucosidase enzymes and hopped beer that was not exposed to the enzyme [115].
Beta-glucosidase also allows the breakdown of cellobiose and cellotriose [116][117]. This has been believed to be a mechanism in which Brettanomyces can survive in barrels;
however, most strains of Brettanomyces found in lambic do not seem to have the ability to produce beta-glucosidase nor utilize cellobiose. Daenen et al. (2007) found that
none of the B. bruxellensis strains isolated from lambic could utilize cellobiose, but strains of B. anomalus and B. custersianus isolated from lambic could utilize cellobiose
[93][117]. Additionally, a study by Tyrawa et al. from Escarpment Laboratories agreed that wine isolated strains were generally better at fermenting cellobiose than strains
isolated from beer at 15°C (59°F). However, at 22.5°C (72.5°F) most of the beer strains started to utilize cellobiose after a few days of incubation (they preferred other food
sources such as glucose and maltose), indicating that temperature plays a role in whether Brettanomyces can ferment certain sugars [104], and the table from the NCYC
Brettanomyces strains suggests that fermenting cellobiose is generally rare for B. bruxellensis. This also suggests that not only are B. bruxellensis strains that are isolated
from beer generally unable to break down glycosides, but they are probably also unable to utilize the cellobiose in wooden barrels as a food source (although higher
temperature might allow some beer strains to start fermenting cellobiose).
See the Glycosides page for more details.
Hop Biotransformation
Colomer et al. (2020) was the first to examine the effects of Brettanomyces on hops during fermentation. The researchers selected 4 strains of B. bruxellensis and 1 strain of
B. anomalus that had the genetic markers for producing beta-glucosidase enzyme, and fermented them as a primary fermenter in a non-dry hopped beer, and also performed a
second experiment where they inoculated the Brettanomyces strains in a dry hopped beer that was first fermented with S. cerevisiae. Monoterpene alcohols were measured
before and after inoculation with the 5 strains of Brettanomyces. In both beers, they found a decrease in geraniol and a rise in beta-citronellol after the inoculation with
Brettanomyces. Beta-citranellol reached a level of 31.5 μg/L with one of the strains, which is a much higher level of beta-citronellol than anything that has been reported with
S. cerevisiae, suggesting that some strains of Brettanomyces might be better at converting monoterpenes from hops than Saccharomyces. See Figure 5 (https://onlinelibrary.wi
ley.com/doi/full/10.1002/jib.610) in the open access study. Interestingly, the strains with the highest beta-glucosidase activity produced the lowest amount of beta-citranellol,
indicating that there is no link between beta-glucosidase activity in Brettanomyces and hop biotransformation. This might be due to the fact that most of the beta-glucosidase
enzyme is produced within the cell and is not released outside of it. The researchers hypothesized that this conversion could be due to two proteins referred to as BbHye2 and
BbHye3 that can be present in Brettanomyces metabolism [118].
For more information on glycosides, see the Glycosides page. For more information on hop biotransformation in general, see the Hops page.
Nitrogen Metabolism
Other than sugars, nitrogen in the form of amino acids is an essential nutrient for yeast. One significant source for nitrogen in wort is boiling hops and dry hopping [119][120].
Brettanomyces can survive in environments that are very low in nitrogen, with one report being as low as 6 mg N/L of yeast assimilable nitrogen (YAN) which is less than
most finished wines contain [121]. While nitrogen usage for S. cerevisiae is well understood, the general utilization of nitrogen by Brettanomyces and its preferred sources for
nitrogen under the stressful conditions of fully fermented beer and wine are not yet well known. However, it is known that Brettanomyces can use a wide range of sources for
nitrogen, and its requirements for nitrogen as a nutrient are extremely low when oxygen is available. When oxygen is not present, nitrogen is required for the survival and
growth of Brettanomyces. Preferred sources of nitrogen include the amino acids glutamine (aerobically and anaerobically), glutamate and aspartate (anaerobically) [122], as
well as possible secondary sources such as lysine, histidine, arginine, asparagine, aspartic cid, glutamic acid, and alanine [16]. Some strains of Brettanomyces can metabolize
other nitrogen sources, such as the amino acids proline and arginine [54]. Ammonium nitrates may also be utilized by some strains of B. bruxellensis. Although studies have
been contradictory and some have not documented whether conditions were aerobic or anaerobic (these contradictions might also be due to strain differences between the B.
bruxellensis strains that were used in different studies), it appears as though some strains of B. bruxellensis might be able to take advantage of trace amount of amino acids
that S. cerevisiae does not use during fermentation, and nitrates and nitrites that S. cerevisiae is not able to consume, as well as amino acids from yeast autolysis (proline,
leucine, tryptophan, and gamma aminobutyric acid) [16]. Other compounds from Saccharomyces autolysis may also be used by Brettanomyces, such as glucose, fatty acids,
nucleotides, polysaccharides, polypeptides, and other proteins [123][124]. The role that oxygen plays in the ability of B. bruxellensis to uptake nitrogen from various sources
might be an important one, and something that should be examined in science going forward [16].
Secondary Metabolites

Secondary metabolites are compounds that are not essential to the life of an organism [125]. Brettanomyces will use a range of secondary metabolites to produce many of the
fruity and funky esters, phenols, and acids that this genus of yeast has become known for. Brettanomyces has also been observed anecdotally to produce thin beer when
fermented on its own, and this has at least partially been attributed to the lack of glycerol production by Brettanomyces. The lack of glycerol production has been attributed to
a genetic predisposal to prefer pyruvate production over glycerol production during fermentation, and it has been speculated that this gives Brettanomyces an adaptive
advantage [126][83]. The major secondary metabolites of B. bruxellensis fermentation have been identified in one study as the ethyl phenols (4EP and 4EG), the alcohols
isoamyl alcohol, 2-methyl-butanol, 2-ethylhexanol, phenethyl alcohol, and an ester ethyl 2-methyl butyrate. Many other compounds are considered minor secondary
metabolites and are produced in varying degrees or not at all based on the strain of Brettanomyces, but may still be produced in high enough concentrations to contribute to
the flavor and/or aroma in beers fermented with Brettanomyces. The types and amounts of flavor compounds produced by Brettanomyces cover a wide spectrum, and many
factors such as species/strain, amino acid precursors, the presence of oxygen, and other nutrients, play a large role in the production of these compounds. In one study on
Brettanomyces in wine, some strains rated as being perceived positively if the strains metabolized certain compounds slower and produced other compounds slower,
indicating that the age of the fermented beverage also plays a large role in how beverages fermented with Brettanomyces are perceived [17]. Some strains of B. bruxellensis
also produce the amines cadaverine, hexylamine, phenylethylamine, putrescine and spermidine, under wine-model conditions [23].
Ester Production
Brettanomyces is capable of synthesizing several ethyl esters from ethanol and fatty acids, as well as other types of esters from various alcohol types (methanol, for example).
Among the most prolific of these are ethyl acetate (synthesized from ethanol and acetic acid), ethyl lactate (synthesized from ethanol and lactic acid), phenethyl acetate, ethyl
caproate, ethyl caprylate, ethyl deconoate [104], along with the hydrolysis (breakdown) of isoamyl acetate. Esters have been found to attract fruit flies and other flying insects,
which help many species of yeast transfer from one food source to another (namely 2-phenyl-ethanol, 3-methyl-1-butanol, ethyl acetate, 2-methyl-1-butanol, and 3-methyl-3-
butenol). Some of these esters are also released by blooming flowers and it is thought that the attraction to flowers by insects is also driven by these same esters [127]. During
non-mixed fermentations where lactic acid is minimal to none, insignificant amounts of ethyl lactate esters are produced, whereas ethyl caprylate and ethyl caproate have a
general increase. With the addition of lactic acid, ethyl lactate levels are greatly increased although may still not reach the flavor threshold level of 250 mg/L (strain
dependent), and ethyl acetate is generally slightly increased. The amounts of esters produced vary widely based on species and strain [128]. A similar but slower evolution of
esters has been seen in a long-term study on examining how Belgian lambic from Cantillon ages in bottles. The study found that lactic acid (produced by lactic acid bacteria)
and ethyl lactate increased as bottles aged, while ethyl decanoate and isoamyl acetate decreased, all presumably from Brettanomyces metabolism over time [129].
Ester production peaks towards the end of growth and is influenced by temperature, aeration/agitation, and pH. Spaepen and Verachtert found in one study that the optimal
temperature for growth and thus ester production was 28°C (77°F), although they did not test higher temperatures. This study also found that continuously shaken samples
produced relatively fewer esters, as well as samples that were not exposed to oxygen at all. The highest ester production was found under conditions of limited oxygen supply
(semi-aerobic versus aerobic or anaerobic), no agitation, held at a temperature of 28°C (77°F), and young cells produced more esters than older cells. It also found that
esterase activity (esterase is the enzyme that facilitates ester production and destruction) increases as pH rises until a pH of 7.6 is reached, after which it begins to decline
again. It was shown that the ester formation/degradation was indeed caused by enzymatic activity of any Brettanomyces species/strain, and not caused by chemical reactions
or from Saccharomyces or Kloeckera activity [130]. Another study by Tyrawa et al. found that all strains of B. bruxellensis tested produced above threshold levels of ethyl
caproate, ethyl caprylate, and ethyl deconoate esters at 15°C versus 22.5°C, but for some strains the higher fermentation temperature of 22.5°C produced significantly more
of these esters than the lower 15°C temperature (other strains produced similar levels of esters at both temperatures, although they fermented slower at 15°C) [104].
Pitching rate of Brettanomyces may have a slight effect on ester production levels, but the differences caused by pitching rate probably do not have a significant impact on the
sensory character of the beer [131]. Brettanomyces produces higher levels of esters when fermented without competition from S. cerevisiae, and this correlates with higher
Brettanomyces cell growth when not in competition with S. cerevisiae (see 100% Brettanomyces Fermentation) [132]. The aromatic amino acids phenylalanine, tryptophan,
and tyrosine have been associated with higher ester formation [17].
High levels of bicarbonate can affect the ester production of B. bruxellensis, as well as the production of acids and phenols. One study reported that levels of 100 mg/l
produced significantly higher ethyl acetate, but there was less of an effect on other esters. High amounts of bicarbonate over 100 mg/l in the 100% B. bruxellensis
fermentations produced significantly lower amounts organic acids (hexanoic, octanoic, and decanoic acid) and lower amounts of vinyl phenols [57]. See also The Brü Lab
Podcast with Dr. Thompson-Witrick (http://thebrulab.libsyn.com/episode-087-impact-bicarbonates-have-on-brettanomyces-fermentations-w-dr-katherine-thompson-witrick).
Esters are also broken down via a process called hydrolysis. Hydrolysis breaks the esters down using the same esterase enzyme within the Brettanomyces cells that are used
to create esters. In general, all acetate based esters, except for phenethyl acetate and methyl acetate, are broken down faster than non-acetate esters by Brettanomyces. In
lambic brewing, sometime after the primary fermentation finishes, Pediococcus begins to produce lactic acid. The formation of lactic acid by Pediococcus coincides with the
appearance and growth of Brettanomyces, which produces more acetic acid. After another 2-3 months, the ester content of the lambic beer changes and reaches an
equilibrium. Ethyl acetate and ethyl lactate are greatly increased, while isoamyl acetate is greatly decreased, reaching an equilibrium of these esters. Given a static amount of
acetic acid, Brettanomyces reaches an equilibrium of ethyl acetate within 24 hours, while ethyl lactate equilibrium takes longer and is much more complex. In lambic, the
majority of ester production and breakdown occurs within 1-3 months after lactic acid production by Pediococcus begins, and at a pH of around 3.5 and a temperature of
around 15°C or less [130]. Pitching rate of Brettanomyces has an effect on the breakdown of isoamyl acetate with higher pitching rates breaking down this ester at a faster rate
[131]. As far as we are aware, ethyl acetate is not metabolized further by Brettanomyces, and the level of ethyl acetate will not be hydrolized over time (although levels can
continue to increase over time with more oxygen oxposure, since oxygen exposure encourages acetic acid synthesis by Brettanomyces and acetic acid bacteria, and acetic acid
and ethanol are then metabolized into ethyl acetate by Brettanomyces).
See also:
Brettanomyces secondary fermentation experiment.

Understanding Esterification, Sour Beer Blog. (http://sourbeerblog.com/understanding-esterification/)

Flavor/Odor Molecular
Ester Precursors Notes
Threshold Formula
Amyl octanoate (spicy, Amyl alcohol and C13H26O2

Also known as pentyl octanoate, it is a flavoring agent [134].
clove, chemical, plastic) [17] caprylic acid [133]
Ethyl acetate (fruity, 33ppm (odor), C4H8O2

Acetic Acid and High flavor threshold; pineapple or pear-like in low amounts and nail polish in high amounts. Increases production
pineapple, pear, solventy, 100ppm
nail polish remover)
ethanol
(flavor)
[135] with higher temperatures and oxygen. Also produced by Saccharomyces species [132].
Ethyl butyrate (pineapple,

Butyric Acid and 0.4ppm (flavor) C6H12O2 Low levels of production by some species of Brettanomyces; production decreases with higher acidity [140]. Also
mango, tropical fruit [136], [138] [139]
ethanol known as ethyl butanoate [139].
juicy fruit gum [137])
Ethyl caproate (sweet, fruity,

pineapple, banana, apple or Caproic acid and 0.2ppm (flavor) C8H16O2 Also known as Ethyl hexanoate, Ethyl butyl acetate, and butylacetate [144]. Can also be produced by Saccharomyces
aniseed) ethanol [141] [142] [143] species [132].
Ethyl caprylate (Sweet, Caprylic acid

waxy, fruity and pineapple (contained in
15ppb (flavor) C10H20O2
with creamy, fatty, buckwheat; produced [147] [148] Also known as ethyl octanoate [148][149].
mushroom and cognac notes by yeast autolysis)
[145]) and ethanol [146]
Ethyl Decanoate/Ethyl Decanoic acid (Capric

Acid) and ethanol
0.5ppm (flavor C12H24O2 Also known as Ethyl caprate, Ethyl caprinate, and Capric acid ethyl ester [152]. Can also be produced by
Caprate (brandy, fruity, oily,
grape) [150] in water) [151] [150] Saccharomyces species [132]>
Ethyl hexanoate [17] (Apple Hexanoic acid and 0.2ppm (flavor

C8H16O2 Also produced by both ale and lager yeast; it is a key flavor note in many beers [153]. High amounts Produced by two
or aniseed [153]) ethanol [154]. in beer) [153] strains out of 9 B. bruxellensis in one study [17].
Ethyl isobutyrate [17] Also known as Ethyl 2-methylpropanoate, it is found in many alcoholic beverages as well as fruits such as apple,
(Pungent, etherial and fruity Isobutyric acid and 0.1 ppb (odor
C6H12O2 banana, orange, wine grapes, strawberry, and nectarine, and is used as a flavoring agent [158]. Produced in significant
with a rum-and egg nog-like ethanol [156] in water) [157]
amounts by 3 out of 9 strains of B. bruxellensis tested in one study [17].
nuance [155])
C7H14O2
Ethyl isovalerate (fruity, Also found in pineapple, orange juice and peel oil, bilberry, blueberry, strawberry, Swiss cheese, other cheeses,
(same as
sweet, berry-like with a ripe, Isovaleric Acid and 30ppm (flavor) cognac, rum, whiskey, sherry, grape wines, cocoa, passion fruit, mango, and mussels [159]. Also known as Ethyl 3-
[159] ethyl
pulpy fruit nuance, artificial ethanol
valerate) methylbutanoate [160]. Not identified as a major product of B. bruxellensis, but is produced in large quantities by
grape [159]) [160][161][17] [159] some strains [17].
0.2 ppm-1.66
Ethyl lactate (fruity, creamy, Lactic Acid and C5H10O3
ppm (odor) Increases production with higher amounts of Lactic Acid [166]
rum [162][163]) ethanol [164]
[165]
Ethyl valerate (Sweet, fruity,

Valeric acid quantities found in beer are minimal (0-1 ppm) and below odor threshold [169][170], and is probably also
acidic, pineapple, apple, Valeric Acid
green, berry, tropical, 1500-5000 ppm C7H14O2 the case for Ethyl valerate. Ethyl valerate is also known as ethyl pentanoate [167]. Also found in apples, bananas,
(pentanoic acid) and
bubblegum [17][167]) ethanol (odor) [168] [167] guava, stawberry, cheeses, rum, whiskey, cider, sherry, grape wines, cocoa, coffee, honey, and passion fruit [168]. Not
[160][161] identified as a major product of B. bruxellensis, but is produced in large quantities by some strains [17].
Ethyl-2-methyl butyrate
(minty, menthol, citrus, Ethanol, methanol, C7H14O2 Also known as ethyl 2-methylbutanoate [171]. Found in bilberry, and in many other fruits, e.g.raw and cooked apple,
green apple) [17]
and butyric acid apricot, orange, grapefruit. Used as a fruit flavor additive [172]. Identified as a major product of B. bruxellensis [17].
Acetic Acid and 1.1ppm (flavor) C7H14O2 Produced by certain Saccharomyces strains but concentrations are generally reduced by Brettanomyces.
Isoamyl acetate (banana) [173]
Isoamyl alcohol [174] Brettanomyces produces only very small amounts itself [130]
Octyl butyrate (spicy,

Octanol and butyric C12H24O2
eucalyptus, floral, plastic) [175]
[17] acid
Pentyl formate (artificial Pentanol and formic

C6H12O2 Also known as amyl formate [178].
fruit, candy, chemical) [17] acid [176][177]
Acetyl-CoA and 2- 3-5ppm (odor),
Phenethyl acetate (sweet,
honey, rose flower like)
phenylethanol 5-10ppm C10H12O2 Produced in very small amounts in Lambic [130][182]. Can also be produced by Saccharomyces species [132]
[179][180] (flavor) [181]
Phenethyl formate (artificial
floral, perfume, wild flower, 2-phenylethanol and C9H10O2
formic acid
solvent) [17]
Phenol Production
Phenols (https://www.britannica.com/science/phenol) such as 4-vinylphenol (4VP; barnyard, medicinal) and 4-vinylguaiacol (4-VG; clove) can be produced in beer through
the decarboxylation of hydroxycinnamic acids (HCAs) by yeast, and also in small amounts by long boils with a portion of the wort coming from wheat (3+ hours resulted in
0.3 ppm of 4VG). HCAs, such as ferulic acid and p-coumaric acid, are present in the non-starch polysaccharide arabinoxylan in malted barley and wheat. They are released
into wort during mashing at levels that are far below their flavor thresholds (greater than 500ppm flavor threshold) [183][184]. Some strains of Oenococcus oeni and
Lactobacillus, as well as some strains of yeast such as Pichia spp, have been found to produce HCA's via cinnamoyl esterase activity in wine, although when these strains
have been used in wine to increase the HCA levels, the final phenol levels produced by Brettanomyces were the same as wine that did not have an increase in HCA levels (the
precursors in wine that lead to HCA's are different than they are in beer) [185]. The esters in grape must that contain HCA's (ethyl ferulate and ethyl coumarate) can also be
formed by acidic hydrolysis which occurs at the low pH of wine, and HCA's can then be released from these esters. This formation of esters is a slow process in wine, with
one study reporting ~0.03 ppm of ethyl ferulate and ~0.4 ppm of ethyl coumarate at the end of primary fermentation and ~0.09 ppm of ethyl ferulate and ~1.4 ppm of ethyl
coumarate after 10 months of barrel aging [186]. We are not aware of any studies that have reported an increase in HCA's from acidic hydrolysis over time in beer; however,
this is a standard laboratory technique for forcing the release of HCA's from barley (although this lab technique uses a lower pH then that of sour beer). In addition, it has
been demonstrated that spent yeast (S. cerevisiae collected after beer fermentation) contains a small fraction of phenols and polyphenols absorbed from wort during
fermentation [187]. It is therefore conceivable that HCA levels could increase in sour beer over time.

While both Saccharomyces (only by "phenolic off flavor positive/POF+" strains) and Brettanomyces strains have varying capabilities based on strain of converting
hydroxycinnamic acids to their vinyl derivatives [188][189], Brettanomyces is also able to reduce these vinyl phenol derivatives to ethyl phenol derivatives. Phenolic acid
decarboxylase (PAD) is the enzyme that converts the HCAs into vinyl phenols. Vinyl reductase (VA) is the enzyme that reduces vinyl phenols to ethyl phenols [190]. Phenol
production has been observed to occur shortly after inoculation of Brettanomyces and has been hypothesized to play a large role in replenishing NAD+ to alleviate the initial
lag growth phase in Brettanomyces [191]. While almost all strains of Brettanomyces produce ethyl phenols, one strain of Brettanomyces anomalus has been found that has lost
the genetic capability to produce phenols [27].
These vinyl derivatives have similar tastes to the ethyl derivatives but have lower flavor thresholds. Levels of all compounds produced vary depending on species and strain
of Brettanomyces. Although the production of ethyl phenols has been identified to occur higher in substrates with more available sugars, and this has also correlated with
higher growth [192], some data supports that pitching rate does not have an effect on how much phenol content is produced by Brettanomyces[131]. Additionally, Curtin et al.
(2013) showed that while both cell growth and attenuation was inhibited in anaerobic conditions in wine, phenol production was not (in fact, the phenol production was
inhibited by aerobic conditions). They also showed that each of the three strains of B. bruxellensis tested all produced the same amount of phenols, while other flavor
compounds such as esters were produced at different levels by each of the strains [193]. Riley Humbert's Bachelors thesis (https://ir.library.oregonstate.edu/downloads/gh93h6
31p) also reported no correlation between fermentation rate and phenol production in several strains of B. bruxellensis [194]. Perhaps growth itself is not as much of a factor in
producing phenols, but having sugars available for metabolism is. This contradicts the somewhat popular belief that under-pitching Brettanomyces produces more "funky"
flavors. Additionally, perhaps some strains are perceived as "funkier" than others due to less ester production and more fatty acid production (isobutyric acid, for example),
rather than more phenol production.
Another study by Tyrawa et al. found that fermentation temperature also did not have a significant effect on phenol production in 9 strains of B. bruxellensis. Given the same
wort composition, strains of B. bruxellensis produced similar levels of phenols at both 15°C and 22.5°C. The ester production was affected by this temperature difference in
some strains but not others (see Esters above). Assuming that phenols contribute the "funky" flavor characteristics, this suggests that perhaps a lower balance of esters to
phenols produces a more "funky" tasting beer more so than a beer with more phenol content. If so, a lower fermentation temperature may be one way to emphasize phenols
over fruity esters [104]. Both Tyrawa and Humbert reported that there was no correlation between flavor profiles from phenol production of different strains of Brettanomyces
bruxellensis and their origin [194].
It has been hypothesized that the production and destruction of various phenols by Brettanomyces is connected with the redox balance (https://en.wikipedia.org/wiki/Redox);
however, this has not been demonstrated. Ethyl phenols have been shown to be highly attractive to fruit flies, and it has also been proposed that these aromatics allow
Brettanomyces to travel from food source to food source and by doing so increasing its chances of survival in the wild [195][16]. Phenols have been shown to have positive
effects on decreasing protein glycation, a complication associated with type 2 diabetes [196].
It has been demonstrated in wine that some phenols can be masked by other flavor compounds, especially lower levels of phenols. Schmuaker et al. (2018) showed that wines
that were spiked with 4EG, 4EP, and isobutyric acid were preferred more when additionally spiked with whiskey lactone (oaky flavor). The oaky flavor at least partially
masked the perception of phenols on the palate [197]. It has been hypothesized by some members of Milk The Funk that higher esters could also mask the perception of
phenols in beer (see 100% Brettanomyces fermentation).
See also:
Effects of Saccharomyces Strain Selection and Staggered vs Co-Pitch

Brettanomyces secondary fermentation experiment.
See the Phenol Explorer website for more information on sources of precursors. (http://phenol-explorer.eu/foods)
Food sources of Hydroxycinnamic acids (p-Coumaric acid, ferulic acid, caffeic acid, etc.). (http://phenol-explorer.eu/classifications/compounds/15/15)
Brü Lab Podcast Episode 030 | Phenolic Compounds In Beer w/ Dr. Mike Lentz. (https://thebrulab.libsyn.com/episode-030-phenolic-compounds-in-beer-w-dr-mike-len
tz)

Phenol Flavor/Odor Molecular

Phenol Precursors Notes
Type Threshold Structure
4-
Vinylphenol
[198] [199] Production level is different across species/strains of Brettanomyces [202]. Coumaric acid levels vary greatly between barley
0.2 ppm
Vinyl p-Coumaric varieties; for example, between 320 µg/kg to 950 µg/kg in different varities of barley husks and 73 µg/kg to 657 µg/kg in
(Musty, (flavor; in beer) C8H8O [201]
phenol Acid [200] different varities of barley malt [187]. Coumaric levels are generally higher in barley malt than they are in wheat malt.
Medicinal,
Band-aid, Coumaric acid is stable through the wort boiling process [183].
Plastic)
Produced by some strains of S. cerevisiae (see Saccharomyces) [205]. Some Brettanomyces species/strains may also be able to
produce this compound at varying levels [160][206][202]. Organic malts have been linked to higher levels of 4VG, vanillan, and
their malt precursor ferulic acid [207]. Ferulic acid is released during mashing, with an optimal mash temperature of 40-45°C
(104-113°F) and a mash pH of 5.7-5.8 (enyzmatic release of ferulic acid is optimal at a pH of 7.5, but this high of a pH is
C9H10O2. difficult to achieve during mashing and would cause other enzymatic problems during the later steps of the mash) [205][208].
Some studies have found that ferulic acid is generally more efficiently extracted from a combination of 70% barley malt and
4- Also known
0.3 ppm 30% wheat malt (not raw wheat), despite studies showing that barley malt often contains more ferulic acid than wheat malt
Vinylguaiacol Vinyl as 2-
[198][199] Ferulic Acid (flavor; in beer) (see this MTF thread (https://www.facebook.com/groups/MilkTheFunk/permalink/2053354874692716/) that explains why this
phenol [203] methoxy-4-
(Clove) vinyl phenol is) [209][205][184][210][211]. A more recent studies disagreed and found a linear increase soluble ferulic acid correlated with
[204]. higher percentages of wheat malt [183]. Malting parameters also affect the levels of ferulic acid in malt; for example, wheat
malt with higher germination temperatures (24-26°C versus 12-18°C) were shown to form more ferulic acid in one study that
looked at the impact of germination temperature and aeration during germination of barley and wheat malt [183]. Ferulic acid is
also There is also a correlation between how dark a malt is (or how highly kilned it is and how much melanoidin content it has)
and how much ferulic acid the malt has; the darker the malt, the more ferulic acid (however, roasted malts were not tested in
the referenced study) [212]. Ferulic acid is stable through the wort boiling process [183].
4-
Vinylcatechol
[198][199] Vinyl C8H8O2
phenol
Caffeic Acid [213] Production level is difference across species/strains of Brettanomyces [202].
(Plastic,
Bitter,
Smokey)
4-
Ethylphenol
[198][199]
(Barnyard, Also known as 1-Ethyl-4-hydroxybenzene and P-Ethylphenol [216]. Identified as a major product of B. bruxellensis [17].
Horsey, Ethyl 0.3 ppm (odor; C8H10O Richard Preiss of Escarpment Laboratories describes pure 4EP as the following, "barnyardy with a slight solvent edge at low
4-vinylphenol
Spicy, phenol in beer) [215] [216] concentrations, and full on hospital antiseptic/bandaid/barnyard at high concentrations. Really quite complex, but maybe not in
Smoky, a good way." [217]
Medicinal,
Band-Aid
[214])
4-
Ethylguaiacol
0.13 ppm Also known as 4-Ethyl-2-methoxyphenol [219]. Identified as a major product of B. bruxellensis [17]. Richard Preiss of
[198][199] Ethyl 4- C9H12O2 Escarpment Laboratories describes pure 4EP as the following, "4EG is a lot more pleasant (than 4EP), very light woodsmoke
(odor; in beer) [219]
(Smokey, phenol vinylguaiacol [215] at low concentrations and heavier smoke and vanilla-like (actually "tonka bean" is closer but not as universally understood) at
Spicy, Clove high concentrations." [217]
[218])
4-
Ethylcatechol
[198][199] Ethyl 4- C8H10O2
[220] Also known as 4-ethylbenzene-1,2-diol [220].
(Band-aide, phenol Vinylcatechol
Medicinal,
Barnyard)
Another way to read the table above is to list the order in which the precursors are converted into phenol metabolites:
p-Coumaric Acid (found in malt and other ingredients) converts to 4-Vinylphenol by POF+ strains of Saccharomyces and Brettanomyces, which converts to 4-
Ethylphenol by Brettanomyces.
Ferulic Acid (found in malt and other ingredients) converts to 4-Vinylguaiacol by POF+ strains of Saccharomyces and Brettanomyces, which converts to 4-
Ethylguaiacol by Brettanomyces.
Caffeic Acid (found in malt and other ingredients) converts to 4-Vinylcatechol by POF+ strains of Saccharomyces and Brettanomyces, which converts to 4-
Ethylcatechol by Brettanomyces.
Acid Production
In the presence of oxygen, Brettanomyces species produce acetic acid as a byproduct of respiratory metabolism. The more oxygen that is present, the more acetic acid is
produced and the less ethanol is produced by Brettanomyces [221][222][4][223][224]. In an environment with oxygen present, Brettanomyes switches to respiratory metabolism.
Sugar is reduced to pyruvate within the cell and is then broken down into acetaldehyde which is then enzymatically oxidized into acetic acid or ethanol (dubbed the Custers
effect). The acetate that is produced by Brettanomyces under aerobic conditions is an important requirement for the cells to fully metabolize certain types of sugars like
galactose [225]. This is thought to be a defensive tactic against competing microorganisms (e.g. Brettanomyces has been shown to produce more acetic acid when co-
fermented with S. cerevisiae, and S. cerevisiae has been shown to have less viability over time in the presence of acetic acid and ethanol) [226][132]. Depending on the brewer's
palate and the degree of acetic production, this can be a desirable or undesirable trait. The degree of acetic acid production varies among different Brettanomyces species and
strains, and it is limited by limiting oxygen exposure (see aging mixed fermentation beer for practical tips on limiting oxygen exposure). For example, B. naardenensis and B.
custersianus produce less acetic acid than other species of Brettanomyces [27][47]. Acetic acid produced by Brettanomyces is also used in the synthesis of acetate esters such
as ethyl acetate, perhaps as a mechanism to protect itself after hindering other microbes via the acetic acid precursor.
Brettanomyces has been shown to produce enough fatty acids in anaerobic fermentation to drop the pH to 4.0, which can also be esterified (see the ester table above) [140].
Many of these acids can have an unpleasant rancid odor and/or taste, which may be noticeable in young Brettanomyces beers before these acids are esterified. Some strains
can also produce succinic acid as a byproduct of fermentation under semi-aerobic conditions, but not anaerobic conditions [56].

Michael Lentz and Chad Harris tested whether or not the hydroxycinnamic acids (HCAs) inhibit the growth of Brettanomyces. They found that high levels of
hydroxycinnamic acids (HCAs), which includes ferulic acid, p-coumaric acid, and caffeic acid, do inhibit the growth of Brettanomyces. Ferulic acid is the strongest inhibitor
of these three HCAs with most strains tested not being able to grow in wort that contained 12 mM (millimolar) of ferulic acid. Caffeic acid was generally shown to be the
weakest inhibitor of the three HCAs tested. Levels of 25 mM p-coumaric acid inhibited the growth of all strains tested, and levels of 30 mM of caffeic acid inhibited all
strains tested. The ability of HCAs to inhibit growth is different from strain to strain of Brettanomyces. Inhibition does not appear to be species dependent. Some strains
display a lag time and grow more slowly in the presence of high amounts of HCA's, but still eventually achieve maximum growth compared to if they were grown without
exposure to HCAs, while others lag and then stop growing before reaching maximum growth [189].
The amount of HCAs varies widely from plant to plant, and the amount that is found in must or wort also varies on how the raw ingredients are treated. These measurements
are generally not a consideration for maltsters or grape growers [189]. The one exception to this is the ferulic acid rest that German brewers have used to create more clove-
like flavors in certain beer styles.
Acids Produced Precursors Notes
Acetic Acid (Vinegar, hard

Oxygen Increased production with higher levels of oxygen exposure [140].
boiled egg)
Associated with supplements of
Butyric Acid (Vomit, bile) Fatty acid.
phenylalanine or tyrosine [17]
Capric acid (Barnyard animal
Fatty acid. Also found in milk, coconut oil, and seed oils [227]. Also referred to as Decanoic acid [228].
odor/taste) [140][17]
Caproic acid (Fatty, cheesy,

Fatty acid. Also known as hexanoic acid [229].
waxy, barnyardy) [140][17]
Caprylic acid (Rancid-like
Fatty acid. Also found in milk. Gives a waxy/oily mouthfeel. Flavor is more intense at low pH levels. Also called octanoic acid.[230]
smell and taste [140][17]
Heptanoic acid (sweaty, Associated with phenylalanine or

Fatty acid. Also known as enanthic acid [231].
solvent, rotten, barnyard) tyrosine [140][17]
Isovaleric Acid (Feety,

Leucine Commonly described as a "spoilage" acid produced by Brettanomyces in wine, but also appears in beer.
parmesan) [232][233]
Lauric acid (faint odor of bay
Fatty acid. Also known as dodecanoic acid and dodecylic acid [234].
oil or soap) [140]
Nonanoic acid (Rancid odor)

[140] Fatty acid. Also known as pelargonic acid [235].
Produced by some strains of b. bruxellensis under semi-aerobic conditions (small amounts of available oxygen), but not anaerobic (no
Succinic acid
available oxygen), and only during the stationary phase (after growth has finished) [56].
Undecanoic acid (coconut,
Associated with tyrosine [17]. Fatty acid. Also known as undecylic acid [236].
balsam, oily, vanilla) [140]
Biogenic Amines
Biogenic amines are produced by all living things and are present in many fermented beverages. High dosages can lead to health issues such as vomiting, headache, asthma,
hypotension, and cardiac palpitation. Thus, biogenic amines have been studied intensely. Levels of histamine below 50 mg/kg in the US and 400 mg/kg in the UK are
considered safe for human consumption (these levels are usually regulated for meets and fish; see reference) [237][238]. For more information, see "Fact or Fiction – Biogenic
Amines in Beer" by Dr. Bryan Heit (http://suigenerisbrewing.com/index.php/2019/01/22/biogenic-amines/).
The production of biogenic amines by Brettanomyces is strain dependent. Relatively low levels of biogenic amines can be produced by various strains. Vigentini et al. (2008)
found that 5 strains of B. bruxellensis were able to produce detectable levels of putrescine, cadaverine and spermidine. Most strains produced 0.4 mg/L or below of these
biogenic amines, while one strain produced 1.2-1.3 mg/L of spermidine at 60 days, which reduced 0.70-0.85 mg/L at 95 days. In general, the maximum biogenic amines were
produced between 40-60 days [239].
Agnolucci et al. (2009) found similar results with 7 strains of B. bruxellensis. None of the strains produced histamine, synephrine, tyramine or spermine. All of the strains
produced under 1 mg/L of most biogenic amines, with hexylamine being produced more so in the range of 1.02-3.92 mg/L [240]:
Source: Agnolucci M, Vigentini I, Capurso G, Merico A, Tirelli A, Compagno C,

Foschino R, Nuti M (2009) Genetic diversity and physiological traits of
Brettanomyces bruxellensis strains isolated from Tuscan Sangiovese wines. Int J
Food Microbiol 130:238–244 (https://www.sciencedirect.com/science/article/pii/S0
168160509000488)

Filipe-Ribeiro et al. (2019) used a statistical model to look for a correlation between biogenic amine levels and the presence of Brettanomyces in different vintages of
Portuguese wines. They did this by looking at the levels of ethyl phenols in the wines, which is a strong indicator of the presence of Brettanomyces. They found no statistical
evidence that the presence of Brettanomyces in wine had any effect on the levels of biogenic amines, positive or negative [241].
For more information on biogenic amines, see the following:
Biogenic amines in spontaneous fermentation.

Biogenic amine production in Pediococcus.
"Spontaneous fermentation and biogenic amines" by Dr. Dave Janssen; review of several studies that looked at the levels of biogenic amines in different beers and their
production, and their potential flavor contribution. (http://www.horscategoriebrewing.com/2019/01/spontaneous-fermentation-and-biogenic.html)
"Fact or Fiction – Biogenic Amines in Beer" by Dr. Bryan Heit; an analysis of biogenic amines in spontaneously fermented beer and associated health concerns. (http://
suigenerisbrewing.com/index.php/2019/01/22/biogenic-amines/)
Glycerol
Brettanomyces is known for not producing much glycerol in beer. Glycerol (https://en.wikipedia.org/wiki/Glycerol) is a colorless, sweet-tasting, viscous liquid that is thought
to be an important contributor to the mouthfeel of beer. Glycerol is produced as a stress response by a wide range of microbes, including S. cerevisiae, and various species
and strains of Debaryomyces, Candida, Lachancea, and Zygosaccharomyces. Despite not producing amounts of glycerol that are perceivable in beer, some strains of
Brettanomyces bruxellensis actually produce glycerol which is stored inside of their cells as a response to osmotic stress. They can also uptake glycerol into their cells. Doing
so allows the cells to survive osmotic pressure [242][243]. It is currently not known how many strains are capable of producing glycerol internally, or if this amount of glycerol
has any impact on perceived mouthfeel of a beer if a substantial amount of Brettanomyces cells eventually autolyze (see this MTF thread (https://www.facebook.com/groups/
MilkTheFunk/permalink/2003626776332193/)). The role of glycerol in creating mouthfeel is debatable in the wine world [244].
Other Compounds

Compound Chemical
Precursors Threshold Notes
Produced Type
2-Methoxyphenethyl
Associated
alcohol (white glue,
Alcohol with ferulic
plastic, mimeograph
acid [17]
sheet)
Associated
with caffeic
2-Ethyl-1-hexanol
acid, ferulic
(fake floral, Alcohol
acid, tyrosine, Produced by many strains of B. bruxellensis. Identified as a major product of B. bruxellensis [17].
chemical, fusel oil)
and
phenylalanine
Associated
with ferulic
2-Methyl-1-butanol acid,
(oxidized/canned Alcohol phenylalanine, Identified as a major product of B. bruxellensis [17].
fruit, plastic, sulfur) and tyrosine
[17].
Bisabolene (spicy, Associated

tropical, toasty, Terpene with caffeic
wood resin, minty) acid [17].
Decanol [17] Alcohol Also known as decyl alcohol [245].
3 One of several isomers of Amyl alcohol; also known as 3-methyl-1-butanol. It is a major higher chain alcohol produced in
Isoamyl alcohol [17] Alcohol
Methylbutanal fermentation [246]. Commonly produced by many strains of B. bruxellensis. Identified as a major product of B. bruxellensis [17].
Nerilidol [17] Terpene Often found in bitter gourd and is a component of many essential oils. It is often used as a flavoring agent [247].
Nonanal (lemon Associated

furniture polish, air Aldehyde with tyrosine
fresener, oily [17]
Associated
Ocimene (sweet with
floral, hyacinth, Terpene phenylalanine
jasmine) [17]
A colorless, slightly viscous liquid used as a defoaming or wetting agent, and is found naturally as a part of esters in some
Octanol [17] Alcohol
essential oils [248].
Phenethyl alcohol
[17] (floral, dried Alcohol Also known as phenethanol. Identified as a major product of B. bruxellensis [17].
rose [249])
Phenylacetaldehyde
[17] (honey, floral
rose, swaat,
Aldehyde
powdery, chocolate
with a slight earthy
nuance [250])
Tetrahydropyridine
L-Lysine,
(Cheerios®, mousy,
Ketone [251] ethanol, and Varies See the Tetrahydropyridine page for more details.
urine, cracker
oxygen
biscuit, corn chips)
Chalcogen
Hydrogen sulfide
(rotten egg)
hydride gas 4 µg/l in beer [253] Produced in high amounts by B. custersianus and B. naardenensis [27].
[252]
0.1–0.2 ppm in lager

Produced by all strains and all species of Brettanomyces except B. naardenensis; the amount that it is produced varies widely
and 0.1–0.4 ppm in
Vicinal ales, although flavor and not much is known about what determines diacetyl levels produced from Brettanomyces [27]. Brettanomyces generally
Diacetyl/butanedione diketone thresholds as low as produce very low amounts of diacetyl (0.019 - 0.048 mg/L). It is hypothesized that Brettanomyces can reduce diacetyl during its
(butter) [254] 17 ppb and 14–61 maturation phase in a similar way to Saccharomyces species, but this has not been investigated that we are aware of. It has been
ppb have been reported that diacetyl reduction is faster at a lower pH of around 3.5, which is a typical pH range for sour beer and might be one
reported [255]. of the contributing factors to a lack of anecdotal reports of diacetyl in sour beer [256][255].
Commercial Cultures
Pure cultures. In cooperation with Funk Factory (http://www.funkfactorygeuzeria.com/2013/06/brett-strain-guide.html).
Bootleg Biology/Spot Yeast

Common Species Synonym

Lab/Package Flavor/Aroma Source Note
Name Name Name
West Flanders,
Belgium
brewery
This rare, and commercially unavailable yeast isolate, produces pungent horse blanket and fresh leather aromas. Perfect
Bruxellensis Dekkera Brettanomyces Funk Weapon specializing in
[257] for breaking out the funk in farmhouse-style beers. This is the first release in the Dusty Bottoms Collection’s ongoing
bruxellensis bruxellensis #1 BB0034A funky, sour,
Funk Weapon Series of unique, rare Brettanomyces and Brett-like wild yeast cultures. Single isolate [258].
mixed-
fermentation
beers.
This rare, and commercially unavailable yeast isolate is perfect for 100% Brettanomyces fermentations, especially Brett Family-owned
IPAs. Amplifies citrus and tropical fruit-forward hop flavors and aromas into a punchy ripeness. Great for maintaining the brewery
Bruxellensis Dekkera Brettanomyces Funk Weapon nuance of hops in beer with greater keeping qualities than a Brewer’s Yeast fermentation. You can’t make every style of producing
[257] bruxellensis bruxellensis #2 BB0035C beer with one or two strains of brewer’s yeast, so why would you only use only one or two strains for your funky beers? Gueuze in West
This is the second release in the Dusty Bottoms Collection’s ongoing Funk Weapon Series of unique, rare Brettanomyces Flanders,
and Brett-like wild yeast cultures. Single isolate [258]. Belgium.
Dry-hopped,
Unblended
Funk Weapon #3 is a versatile culture that creates wildly different flavor and aroma profiles depending on the age of Lambic
fermentation. Young fermentations produce mild musty funk and ripe tropical fruit, while older and bottle conditioned Produced by a
Bruxellensis Dekkera Brettanomyces Funk Weapon
[257] ferments show off unique flavors and aromas of strawberry, cherry and tropical candy. This commercially unavailable Traditional
bruxellensis bruxellensis #3 BB0022 yeast isolate is ideal for 100% Brettanomyces fermentations or as a secondary strain along with a phenolic Brewer’s Yeast Lambic
culture. Single isolate [258]. Brewery in
Brussels,
Belgium.
Brewing Science Institute

Name Name Name
Thought to be the same as White Labs WLP648; however,

Dekkera Brettanomyces Brettanomyces Medium intensity Brett character. Classic strain used in secondary side by side sensory comparisons seem to indicate that they
Bruxellensis
bruxellensis bruxellensis bruxellensis fermentation for Belgian style beers and lambics.
might not be the same [259]. Commercial pitches only.
Dekkera Brettanomyces Brettanomyces Thought to be the same as White Labs. Commercial pitches
Claussenii Low intensity Brett character.
anomala anomalus clausenii only.
Dekkera Brettanomyces Chad Yakobson's mutation of BSI Drie. Commercial pitches

CMY1 bsi
bruxellensis bruxellensis only.
Brettanomyces Isolate from Drie Fonteinen; Pro-Brewers only.

Dekkera Brettanomyces Highly aromatic of tropical fruit come with aging. Works well with citrus Recommended fermentation levels to get the most character
Drie bruxellensis var.
bruxellensis bruxellensis and fruity hops.
Drei out of Drie include 75-77°F [260]. Commercial pitches only.
High intensity Brett character. Know to produce the “horsey” aroma
Dekkera Brettanomyces Brettanomyces
Lambicus characteristic of Brettanomyces yeast. Classic strain used in secondary Commercial pitches only.
bruxellensis bruxellensis lambicus
fermentation for Belgian style beers and lambics. Same as White Labs.
Community Cultures Yeast Lab
Common Species Synonym Source

Lab/Package Flavor/Aroma
Name Name Name Note
Dekkera Brettanomyces Brettanomyces A little less "Bretty" and a little more fruity. Flocculation: Low. Alcohol Tolerance: Medium-High (8-12%).
Claussenii
anomala anomalus claussenii Fermentation Temperature: 85F.
Dekkera Brettanomyces Brettanomyces For secondary fermentation and Belgians, with classic "Bretty" characters. Flocculation: Low. Alcohol Tolerance:
Bruxellensis
bruxellensis bruxellensis bruxellensis Medium-High (8-12%). Fermentation Temperature: 85F.
Craft Cultures

Name Name Name
Isolated from a wild rasberry in Michigan's Upper Peninsula along

CCYL410 Razz Great for use in secondary fermentation for Belgian style beers and
Unknown Unknown Unknown the shore of the Portage Lake near Houghton Michigan [261].
Brettanomyces™ lambic style beers.
Commercial pitches only.
CCYL411
Dekkera Brettanomyces
Bruxellensis Brettanomyces Exhibits medium intensity Brett character [261]. Commercial pitches only.
bruxellensis bruxellensis
bruxellensis
CCYL412 Low intensity Brett character. The Brett flavors produced are more 75-85% attenuation, low flocculation, 8-12% alcohol tolerance,
Claussenii Brettanomyces subtle than CCYL410. More aroma than flavor contribution. Fruity, 67-74°F fermentation temperature [261]. Commercial pitches
anomala anomalus
claussenii pineapple like aroma [261]. only.
CCYL413 High intensity Brett character. Defines the "Brett character": Smoky
Dekkera Brettanomyces and spicy flavors. This strain is often found in Lambic, Flanders, and
Lambicus Brettanomyces Commercial pitches only.
lambicus sour brown style beers [261].
East Coast Yeast


Name Name Name Note
beer -
Dekkera Brettanomyces This species displays strong esters of juicy-fruit gum, moderate funk and gradual acidity over time. A super-attenuator, up
Anomala ECY-04 Adelaide,
anomala anomalus to 100% [262]
Australia
isolated
Dekkera Brettanomyces from
Bruxellensis
ECY-05 funky with barnyard notes accompanied by some fruit. May no longer be available [262]. Belgian
stout.
bottled beer
Brettanomyces Brettanomyces This species of Brettanomyces reveals a phenolic, spicy, saison-like profile with none to low esters. Some earthiness and
Nanus ECY-16 - Kalmar,
nanus nanus tartness may develop over time. Slow to attenuate fully alone, may be best in fermenting with other yeast.
Sweden
Bantu beer
Brettanomyces Brettanomyces
Custersianus ECY-19 Light fruit and hay. May no longer be available [262]. brewery,
custersianus custersianus
South Africa
Strawberry, honey, ripe fruit with a tart, citrusy acidity after 6mo of aging. Creates an abundance of acidity quickly and Isolated
Brettanomyces Brettanomyces may initially display a mousy-like tainted flavor but dissipates over time leading to ester production with hints of
Naardenensis ECY-30 from Dr.
naardenensis naardenensis
strawberry. Best used with Saccharomyces; aging up to 6 months is recommended [262]. Pepper
Isolated
Dekkera Brettanomyces from bottle
Bruxellensis
ECY35 Funky with barnyard notes accompanied by ripe tropical fruit [262]. of Belgian
gueuze.
Anomala ECY36 Notes of pineapple esters with acidity over time, no funk or barnyard present. Versatile [262].
anomala anomalus
Lambicus
ECY37 Cherry-stone and strong barnyard [262].
Escarpment Laboratories
"How to Choose a Brett Strain," a guide to Escarpment Laboratories Brettanomyces products on their blog. (https://escarpmentlabs.com/blogs/resources/how-to-choose
-a-brett-strain-for-beer)

Name Name Name Note
Isolated
Dekkera Brettanomyces A classic Brettanomyces bruxellensis strain, typically used in secondary fermentations. Attenuation: 70-85% // Optimum from a
Bruxellensis Brett B
bruxellensis bruxellensis Temp: 26ºC+ (80F+) // Alcohol tolerance: Medium-high // Flocculation: Low [263]. bottle of
Orval [264].
A strain of Brettanomyces clausenii, now understood to be among the Brettanomyces anomalus species. Minimal funk, tends
Dekkera Brettanomyces to exhibit fruity pineapple or mango notes. Pairs well with fruit and/or hops. Recommended for secondary or co-fermentation
Claussenii Brett C as attenuation is variable. Attenuation: Highly Variable // Optimum Temp: 18-25ºC (64.4-77F) // Flocculation: Medium-low.
anomala anomalus
Potentially no longer available (no longer listed on website) [263].
This strain of Brettanomyces bruxellensis is a notoriously vigorous fermentor, suitable for primary fermentation of 100%
Brett beers or secondary fermentation where some extra funk is desired. Attenuation: 80+% // Optimum Temp: 20-25ºC (68-
Bruxellensis
Brett D 77ºF) // Alcohol tolerance: 12%+ // Flocculation: Medium-low [263]. See this MTF thread (https://www.facebook.com/groups
/MilkTheFunk/permalink/2331941676834033/?comment_id=2331958926832308&comment_tracking=%7B%22tn%22%3A
%22R1%22%7D) on experiences using it for 100% fermentation.
Unknown Unknown Unknown Brett M This strain offers balanced funk fast. Suited for use in sour beers, saisons, and other barrel-aged treats [263]. Michigan
Originally
isolated
from a
This strain typically completes primary fermentation within one month, and is also suitable for secondary aging of a wide barrel-aged
Bruxellensis Brett Q range of beer styles where subtle Brett character is desired. Tasting notes include ripe strawberry, pear, apple, with underlying sour beer
funk. Attenuation: 80%+ // Optimum Temp: 20-25ºC // Alcohol tolerance: High // Flocculation: Medium-low from
Quebec
[263].
This Brettanomyces bruxellensis strain displays a balanced selection of fruity characteristics, with testers noting plum, red
Dekkera Brettanomyces berry, citrus, and red apple, alongside subtle acidity. It is suitable for primary or secondary fermentation, but does shine in
Bruxellensis Brussels Brett secondary with extended aging, where it displays prominent funky flavours. Attenuation: 80+% // Optimum Temp: 22-25ºC //
Alcohol tolerance: 12%+ // Flocculation: Medium-low [263].
Dekkera Brettanomyces Isolated from a bottle of Schultheiss Berliner Weisse, this B. bruxellensis strain produces notes of orchard fruit and pineapple.
Bruxellensis Berliner Brett II
bruxellensis bruxellensis Complements Berliner Brett I nicely to produce a complete classic Berliner profile [263].
Fermentis
Common
Species Name Synonym Name Lab/Package Flavor/Aroma Source Note
Name
Dekkera Brettanomyces SafBrew™ BR- First known dried Brettanomyces product. See the product page (https://fermentis.com/en/product/s
Bruxellensis
bruxellensis bruxellensis 8 afbrew-br-8/).
Fermmento Labs (Brazil - CLOSED)
Common Name Species Name Synonym Name Lab/Package Flavor/Aroma Source Note
Bruxellensis Dekkera bruxellensis Brettanomyces bruxellensis FB2 Heavy Funkies
Imperial Organic Yeast


Name Name Name Note
Suburban Brett is Brettanomyces yeast that works great as a secondary aging strain. It really shines when used in wood barrels
W15 Brettanomyces
Dekkera and will produce complex and balanced aromas of sour cherry and dried fruit. It can also be used for as a primary strain for Brett
Suburban bruxellensis W15
bruxellensis [265] only beers. Temp: 64-74F, 18-23C // Flocculation: Low // Attenuation: 75-80% [266]. MTF thread on experiences with this
Brett
culture. (https://www.facebook.com/groups/MilkTheFunk/permalink/1735063539855186/)
Inland Island Yeast Laboratories

Name Name Name Note
Isolated from a brewery in Brussels this particular Brettanomyces strain is known for producing aromatics reminiscent of
INIS-901
Dekkera Brettanomyces horse, barnyard, sweat, and goat. It is highly attenuative and will take up to 6 months to fully finish fermentation. It is Brewery in
Bruxellensis Brettanomyces
bruxellensis bruxellensis suggested that this strain be used with another primary fermentation strain. 90% + Attenuation. Low Flocculation. 60-75 F Brussels
bruxellensis I
Temperature Range. High Alcohol Tolerance.
INISBC-902 This strain is capable of fermenting a beer without the use of a Saccharomyces cerevisiae. This yeast produces a beer with a
Bruxellensis Brettanomyces complex flavor profile, containing both fruity and sour notes. 85%+ Attenuation. Low Flocculation. 70-85 F Temperature
bruxellensis II Range. Medium-High Alcohol Tolerance.
Small
INISBC-903 Isolated from a small brewery just outside of Brussels. Produces an aromatic profile that is more mild than INISBC-901 with
Dekkera Brettanomyces Brewery just
Bruxellensis Brettanomyces increased tropical fruitiness. Able to ferment a beer without added Saccharomyces c. Mixed with lactobacillus this strain will
bruxellensis bruxellensis outside of
bruxellensis III create a wonderful sour beer. 90% + Attenuation. Low Flocculation. 60-75 F Temperature Range. High Alcohol Tolerance.
Brussels
Isolated from
INISBC-913 Strain is able to ferment without added help from another Saccharomyces c. strain. Produces mild acidity and tropical fruit a famous
Unknown Unknown Unknown Brett Barrel notes. Leaves the beer with very little mouthfeel. See recipe recommendations for fermentation additions to boost mouthfeel. American
Yeast III 90% + Attenuation. Low Flocculation. 60-75 F Temperature Range. High Alcohol Tolerance. wild ale
brewery.
INISBC-920 Originally isolated from spontaneously fermenting beers, this strain will give your beer textbook Brett flavors like horse
Lambicus Brettanomyces blanket and spice, but also fruity notes of pineapple. Fermentation will take well over a month to fully attenuate if used as a
lambicus primary strain. 85%+ Attenuation. Low Flocculation. 70-85 F Temperature Range. Medium-High Alcohol Tolerance.
INISBC-930 Possibly the least “Bretty” of the Brettanomyces strains, Claussenii adds great fruity pineapple aroma without the traditional
Claussenii Brettanomyces flavors that turn some off from Brett fermented beers. An excellent choice for adding a little funk to your barrel aged
anomala anomalus
claussenii favorite. 85%+ Attenuation. Low Flocculation. 85 F+ Temperature Range. Medium-High Alcohol Tolerance.
Isolated from
INIS-950 a craft
NA NA NA Brettanomyces This unique yeast will add notes of barnyard and ripe fruit to beer. brewery in
"Copenhagen" Denmark
[267].
GigaYeast

Name Name Name Note
Brussels
Dekkera Brettanomyces Produces classic Brett “Barnyard” characteristics plus some subtle fruity aroma and moderate acidity. Adds a tart
Bruxellensis Bruxellensis
bruxellensis bruxellensis complexity to any beer.
GB001
Dekkera Brettanomyces Tart Cherry Brett Produces some Brett Barnyard funk plus stone fruit and cherry-like esters. This Strain also produces a moderate
Bruxellensis
bruxellensis bruxellensis GB002 amount of acid that adds a tart complexity to the brew.
Dekkera Brettanomyces Sweet Flemish
Bruxellensis Produces a sweet, slightly fruity profile with just a hint of barnyard and spicy phenolics
bruxellensis bruxellensis Brett GB144
Jasper Yeast

Name Name Name Note
JY033 - Brett Amsterdam,

Dekkera Brettanomyces Netherlands
Bruxellensis brux Slow growing, good for conditioning in the style of Orval
bruxellensis bruxellensis [268]
Amsterdam
Dekkera Brettanomyces JY044 - Brett California,

Bruxellensis Comparable to JY033, but slightly more funky, tart and aggressive.
bruxellensis bruxellensis brux Rosa USA [268]
JY87 is a Brettanomyces yeast isolated by Jasper Yeast LLC. Fast-growing, it has almost Saccharomyces-like doubling times.
Shows great fermentation as primary strain in a variety of beers. Ferments fast to 60% attenuation, after which fermentation West-
Dekkera Brettanomyces JY061 - Brett slows down and more flavor and aroma is produced. Strong pineapple and stonefruit aroma after prolonged fermentations (3-9 Flanders,
Bruxellensis Belgium.
bruxellensis bruxellensis brux Chateaux months). Great companion to beers that could use some funk, and complements hoppy beers perfectly. Flocculation is low,
strain will form a pellicle when oxygen is present. Sequencing of ITS regions indicated Brettanomyces bruxellensis. [268]
Micrograph of JY87 cells coming soon.
Dekkera Brettanomyces JY062 - Brett California,

Bruxellensis Mild aromatics and phenolics, but quite tart and high tendency to produce acetic acid in the presence of oxygen.
bruxellensis bruxellensis brux Abbey USA [268]
Dekkera Brettanomyces JY157 - Brett Good for Brettanomyces beers where an intense character is preferred. Prolonged aging can produce almost a cherry like Belgium
Lambicus [268]
bruxellensis bruxellensis brux lambicus sourness. Optimum temperature: 80-90°F (27-32°C)
Dekkera Brettanomyces Optimum temperature: 80-90°F (27-32°C). Pineapple aroma, but can also develop a leather character. This yeast is not Belgium
Claussenii JY196 - Brett c [268]
anomala anomalus suitable to be used on its own in malt based worts because of its limited fermentation capacity.
Mainiacal Yeast (CLOSED)


Name Name Name
Isolated from a 5 year old bottle of Orval.

Depending on how this strain will dictate its outcome. In primary fermentation, it will lend 65-80°F fermentation temperature with 65-
Dekkera Brettanomyces stone fruit notes and a very thin mouthfeel. Although in secondary it can produce cherry
Bruxellensis MYOrval 100% attenuation [269]. Commercial
bruxellensis bruxellensis notes all the way to barnyard like "funk". We find to obtain this funk quality you will need a
pitches only; occasionally available to
decent amount of aged hops in the boil alongside a secondary fermentation with this strain.
homebrewers.
isolated from a red wine barrel that was used
It produces a stone fruit forward Brett quality when used in primary fermentation and like by a winemaker. 65-80°F fermentation
Bruxellensis MYLBB1 many Bretts it does not produce high amounts of glycerol so mouthfeel can be expected to be
thin. Used in secondary it can lend notes of cherries and a light barnyard like aroma. temperature with 65-100% attenuation [269].
Commercial pitches only.
Depending on how this strain is used will determine the outcome. In primary fermentation it American Spontaneous Producer, isolated
Dekkera Brettanomyces produces light stone fruits with a hint of barnyard like hay. You can also expect a thin from a bottle. 60-80°F fermentation
Bruxellensis MYLBB2
bruxellensis bruxellensis mouthfeel due to the low glycerol production. In secondary it will produce a more heavy temperature with 75-95% attenuation [269].
barnyard like aroma with hints of cherries and raspberries. Commercial pitches only.
This strain of Brett b is better suited for either co-fermentation with a Sacc strain or used in Isolated from a Belgian spontaneous
Dekkera Brettanomyces secondary. If used for primary fermentation it can produce significant sulfur like aromas. This producer. 55-78°F fermentation temperature
Bruxellensis MYLBB3
bruxellensis bruxellensis strain can produce a significant amount of funk if using aged hops at a similar rate as with 75-100% attenuation [269].
traditional Lambics. It also produces notes of pineapples. Commercial pitches only.
This Brett strain is described as 'unique'. It's not great for primary fermentation, especially in
Isolated from an apple orchard in Maine. 50-
Dekkera Brettanomyces kettle sours as it throws very odd smells off. However, it works better as a secondary Brett
Bruxellensis MYLBB4 80°F fermentation temperature with 70-90%
bruxellensis bruxellensis strain producing peach and light fresh apple notes, sometimes also producing a spice
character. attenuation [269]. Commercial pitches only.
Omega Yeast Labs

Name Name Name Note
Contributes more Brett aroma than flavor. Fruity, pineapple-like aroma. Flocculation: low, Attenuation: 70-85%, Temp: >85°F,
Brettanomyces
Dekkera Brettanomyces Alcohol Tolerance: medium-high, compares to WLP645. Can ferment lactose [270]. Pro brewers only. Recommended to use in
Claussenii claussenii
anomala anomalus conjunction with brewers yeast; not recommended for 100% Brettanomyces fermentation as it doesn't attenuate wort on its own
OYL-201
very well [271].
Brettanomyces Medium intensity Brett character. Classic strain used in secondary fermentation for Belgian style beers and lambics.
Bruxellensis bruxellensis Flocculation: low, Attenuation: 70-85%, Temp: >85°F, Alcohol Tolerance: medium-high, compares to WLP650. Pro brewers
OYL-202 only.
Brettanomyces This strain has been described as producing horsey, smoky and spicy flavors. As the name suggests, this strain is found most
Lambicus lambicus OYL- often in Lambic style beers. Flocculation: low, Attenuation: 70-85%, Temp: >85°F, Alcohol Tolerance: medium-high, compares
203 to WLP653. Pro brewers only.
White wine character and light funk, and develops its character rather quickly. Brett character will be apparent within a few
Brettanomyces weeks of reaching terminal gravity and will continue to develop if given additional conditioning time. Flocculation: low, A US
Bruxellensis bruxellensis Attenuation: 70-85%, Temp: 68-80°F, Alcohol Tolerance: medium-high. Potentially the same B. bruxellensis strain as their C2C Northwest
OYL-216 brewery.
blend [272]. Pro brewers only.
Propagate Lab


Name Name Name
Used for secondary fermentation in Belgian-style beers such as lambics, this strain creates a medium-intensity, (Editor's note: this is likely
Dekkera Brettanomyces MIP-701 Brett earth-forward character in finished beer. A historic brewery in Belgium uses this yeast in secondary
Bruxellensis to be a strain isolated from
bruxellensis bruxellensis Brux I
fermentation and bottling to produce the signature flavor of its beer [273]. Orval).
Dekkera Brettanomyces MIP-702 Brett a strain used for secondary fermentation in Belgian-style beers such as lambics. It creates a medium-intensity,
Bruxellensis
bruxellensis bruxellensis Brux II earth-forward character in finished beer. Balanced between barnyard and fruit [273].
Dekkera Brettanomyces MIP-703 Brett A strain used for secondary fermentation in Belgian-style beers such as lambics. Characterized as "fruity with
Bruxellensis
bruxellensis bruxellensis Brux III tropical fruit dominating the aroma" [273].
In collaboration with Wild Pitch Yeast. High attenuation, produces a lemony/tart/subtle “Brett” aroma and
flavors characterized as lemon, limoncello, and mint with a slight undertone of the more typical Brett barnyard Isolated from a
Bruxellensis YH169 funk. Can be used at a wide variety of temperatures (68-99 F) with a potential flavor sweet spot of 80-85 F. spontaneous fermentation
in Indianapolis, IN.
Would work well in a saison or Brett IPA [273].
This particular strain was

MIP-710 Brett isolated from a Colorado
Unknown Unknown Unknown strain used for secondary fermentation in Belgian-style beers such as lambics; characterized as "fruity". Brewery and produces
Stave I
intense fruit notes [273].
This strain was isolated

MIP-714 Brett from a bottle of beer
Unknown Unknown Unknown Produces barnyard like aromas and is a very aggressive strain. originally produced in the
Stave IV
Netherlands [273].
A strain used for secondary fermentation in Belgian-style ales and English Old Ales. The yeast can be fairly
Dekkera Brettanomyces MIP-720 Brett neutral in aroma and works well by itself. When it is paired with a phenol producing yeast it will create
Clausenii
anomala anomalus clausenii
barnyard like aromas [273].
Isolated from a bottle of
MIP-750 Brett European beer fermented
Unknown Unknown Unknown An aggressive strain that produces strong barnyard aroma. Works well as a blend.
tool
with fruit [273].
MIP-760 Brett Isolated from a famous

Unknown Unknown Unknown A highly fruity strain that will ferment well by itself or pitched into a blend.
phantom saison producer [273].
Isolated from a natural

Characterized as sweet tarts, ripe peach skin, clementine, white grapes, and a hint of candied strawberry with a wine from Mendocino
Unknown Unknown Unknown BTN-70 Feints
backbone of soft Brett funk.
County, California [273].
BTN-81 Yellow Isolated from a yellow

Unknown Unknown Unknown Characterized as Brett-forward with notes of sweet tarts, ripe peach skin, dried lime peel, tangerine pith.
Jacket jacket insect [273].
RVA Yeast Labs

Name Name Name Note
A medium-intensity Brettanomyces yeast strain. Will add a bit of funk when added during the secondary. Typically used in
Bruxellensis RVA 502 Belgian-style beers, especially lambic. A famous Trappist brewery produces its unique beer with this yeast during secondary
fermentation.
Dekkera Brettanomyces A low-intensity strain. Contributions from this strain are mostly aromas of pineapple and fruit. This strain prefers higher
Claussenii RVA 501
anomala anomalus temperatures (85º F), but will produce nice aroma and subtle flavor at normal ale fermentation termperatures (68-72º F).
Dekkera Brettanomyces High-intensity “Brett” strain. Very spicey with “smoky” and “horseblanket” flavors and aromas. This strain is used mostly in
Lambicus RVA 503
bruxellensis bruxellensis Lambics and Flanders sour beers.
Produces some amazing aromas of pears, and other fruit esters. We highly recommend this strain for Belgian Dubbels. This This strain
strain also makes a very nice cider. A highly flocculating, medium-high attenuating strain adds nice complexity to stouts, and originates
Unknown Unknown Unknown RVA 804
Belgian Ales and Specialty Belgian Ales. Flocculation: Medium, Attenuation: 78-85%, Suggested Temp Range: 65-72°F, from local
Alcohol Tolerance: 14%. fruit trees.
The Yeast Bay


Name Name Name
This strain is the first single strain Brettanomyces isolate we are releasing all on its own because it deserves that distinct
Isolated from a
honor. Isolate TYB184 is literally the 184th isolate of yeast/bacteria that has made it through primary fermentation trials,
rustic farmhouse
was assigned an isolate number and carried into larger scale fermentation evaluation since our conception in 2013. This
style beer
Dekkera Brettanomyces isolate is attenuative, produces a moderate acidic-like character and an ester profile of lemon/pineapple. Another notable
Bruxellensis TYB184 produced in the
bruxellensis bruxellensis characteristic of this isolate is the mild barnyard character it produces that doesn't take over the profile; rather, it balances
Northeastern
the ester profile. The unique character balance in this strain is what makes it well suited for use on its own, in both
United States
primary and secondary fermentation. Approximately 15 billion cells/vial. Temperature: 72 - 85 ºF. Attenuation: [275].
82%-88%. Flocculation: Medium-Low. Recommended by Nick for darker sours [274].
Isolated from a
This isolate exhibits good attenuation, and produces a moderate acidic-like character and an ester profile the combination Belgian-inspired
Dekkera Brettanomyces of which produces a character reminiscent of sweet tarts. It's a fruity, funky tartness that's refreshing and crisp. This strain brewery in the
Bruxellensis TYB207 Northeastern
bruxellensis bruxellensis is well-suited for primary and secondary fermentation. Approximately 15 billion cells/vial. Temperature: 70 - 82ºF ,
Attenuation: 80%-82%, Flocculation: Medium-Low. United States
[276].
This Brettanomyces isolate exhibits a mild tartness and soft funk with a solid backbone of tropical fruit esters (papaya,
guava, pineapple, guinep). It's great primary fermenter, but really shines in secondary fermentation following up a
primary fermentation that produces a lot of flavor compounds. This strain is a true flavor modulator, and the more raw
Dekkera Brettanomyces material it has to work with the greater the complexity that will be achieved in the finished beer. Sequencing results
Bruxellensis TYB261
bruxellensis bruxellensis revealed it's Brettanomyces bruxellensis. This strain will produce a massive, thick krausen, so be sure to use a blowoff
tube or reserve plenty of fermentor headspace! Isolated from a unique mixed fermentation beer produced in the
Midwestern United States [277].
Isolated from a
California
brewery that
utilizes an
This strain exhibits a lemony-tartness with hints of hay and mild barnyard funkiness, and has a crisp and dry finish. extensive and
Dekkera Brettanomyces diverse array of
Bruxellensis TYB307 TYB307 is suitable for primary fermentation and extended aging. Fermentation temperature: 70-80ºF. Attenuation: 80-
bruxellensis bruxellensis organisms in the
84%. Flucculation: low.
production of
their
wild/sour/funky
beers [278].
Isolated from a
brewer of all
This strain exhibits a strong profile of complex tropical fruit that is dominated by pineapple with a noticeable earthiness things sour and
Bruxellensis TYB415 that adds a unique complexity and depth of character to the beer. TYB415 is suitable for primary fermentation and wild in the
extended aging. Fermentation temperature: 70-80ºF. Attenuation: 82-86%. Flucculation: low. Mountain West
[279].
White Labs
See the public Yeastman database (https://www.yeastman.com/Login/Public/Report/PublicLabQCResult.aspx) for exact cell counts for specific lot numbers.
See attenation rates based on pitch rate from this White Labs data sheet (http://www.whitelabs.com/sites/default/files/R%26D%20Wild%20Yeast%20and%20Bacteria
%20Experiments_2.pdf).

Name Name Name
Barnyard. Prominent Leathery

Dekkera Brettanomyces aroma and flavor. Low levels of Not the same as WY's Brux. Approx. 500 million cells per mL; homebrew vials are approx. 17.5 billion
Bruxellensis WLP650
bruxellensis bruxellensis supporting tropical fruit in the cells at 35 mL [281].
aroma and flavor. [280]
The vrai (true, in French) Brettanomyces bruxellensis Trois. The infamous strain used for all-
Pear. Overly ripe stone fruit and Brettanomyces fermentations, has a robust, complex sour character with aromas of pear. Best used as a
pineapple, with hints of citrus in primary fermentation strain. Might be the same as BSI Drie? Profile is very similar to BSI Drie [282]
the aroma. Pineapple, overly ripe [283]. Ale of the Riverwards sensory analysis (http://www.alesoftheriverwards.com/2015/12/brettanomyc
Bruxellensis Dekkera Brettanomyces tropical stone fruit, lemon and es-drei-vs-brettanomyces-vrai.html) suggests they may be different strains. Approx. 500 million cells per
WLP648
Trois Vrai bruxellensis bruxellensis lime rind flavor. Little to no mL; homebrew vials are approx. 17.5 billion cells at 35 mL [281].
"funk". Less complex overall
compared to other Brettanomyces Bryan of Sui Generis blog and Devin Henry found that this culture contains two closely related B.
cultures from WL [280]. bruxellensis strains with very different flavor profiles [30][284].
Typical barnyard funk with some

Dekkera Brettanomyces fruitiness; claimed that it can be
Anomalus WLP640
anomala anomalus used for primary fermentation but
a starter may be necessary.
Fruity, pineapple. Wine grape- Approx. 500 million cells per mL; homebrew vials are approx. 17.5 billion cells at 35 mL [281]. See also
like aroma, with light wood-like, this MTF thread (https://www.facebook.com/groups/MilkTheFunk/permalink/1385144124847131/?matc
floral, and citrus aromas. More h=YXR0ZW51YXRpb24sYXR0ZW51YXRlZCxjbGF1c3Nlbmlp) and this MTF thread (https://www.fa
Claussenii WLP645 cebook.com/groups/MilkTheFunk/permalink/1309024112459133/?comment_id=1310955328932678&c
anomala anomalus fruit forward in the flavor, clean
aftertaste with little to no "funk" omment_tracking=%7B%22tn%22%3A%22R1%22%7D) which discuss the purity of this culture, and
[280]. references Yakobson's data (http://brettanomycesproject.com/dissertation/pure-culture-fermentation/imp
act-of-pitching-rate/) that indicates that it does not attenuate wort efficiently when purely isolated.
Horsey, Smoky, Spicy. High
amount of ripe pineapple and
Dekkera Brettanomyces overly ripe stone fruit in the Different from WY's "lambicus". Approx. 500 million cells per mL; homebrew vials are approx. 17.5
Lambicus WLP653
bruxellensis bruxellensis aroma and flavor, with mild billion cells at 35 mL [281].
levels of blue cheese, leather, and
spicy phenol in the flavor [280].
Wyeast


Name Name Name
bottled stout - Burton on

Trent, England.
Dekkera Brettanomyces Discontinued in 2007 due
Anomalus Wyeast 5110 to being misclassified [285].
anomala anomalus
Cell count: 7.5 x 108
cells/mL [286].
Not the same as WL's Brux.
Bruxellensis Wyeast 5112 "sweaty horse blanket" Cell count: 7.5 x 108
cells/mL [286].
Notes of tropical fruit, pineapple and, to a lesser extent, peach and blueberry round out a classic Brett profile. Private collection for
Produces “horse blanket,” leathery, and smoky character, but at lower level than other Brett strains. Can be Spring 2015/Summer 2016.
Claussenii Wyeast 5151 used as the primary strain for fermenting, but is often used after a primary fermentation with an S. cerevisiae
anomala anomalus Cell count: 7.5 x 108
strain, and in blends to produce sour beers. It is highly attenuative, given proper time to fully ferment out, and
is known to create a pellicle during fermentation. cells/mL [286].
Different from WL's

Dekkera Brettanomyces "lambicus". Cell count: 7.5
Lambicus Wyeast 5526 Pie-cherry
x 108 cells/mL [286].
Smaller Labs
Mfg Package Taxonomy Notes
BKYeast Brett X1 Suspected Brettanomyces Anomalus
BKYeast Brett C1 Isolate from Cantillon Iris
BWY-001 Berliner Brett I (http://black This Brettanomyces yeast was isolated from a Willner-Brauerei-Berlin Berliner Weisse. A Brettanomyces yeast strain that develops
Blackwell
wellbrewery.ch/product/bwy-001-berlin a rather subtle flavour profile but leads to a super dry finish. The perfect yeast to create a classical interpretation of a Brettanomyces
Brewery
er-brett/) bottle conditioned Berliner Weisse.
DCYeast DCY01
Brettanomyces yeast from a Walloon Trappist brewery that gives an earthy aroma to the beer. Recommended for secondary
Saccharolicious Brett I
fermentation after primary fermentation with Trappist O.
Fruity Brettanomyces yeast strain with an aroma that reminds of French cider. originates from Brasserie à Vapeur in Pipaix,
Saccharolicious Brett II
Belgium, and was isolated from a bottle of Cochonne.
Brett Blends (Brett only)

Manufacturer Package Notes
Twelve (12) different isolates of Brettanomyces exhibiting high production of barnyard "funk" and esters. Dryness, ripe fruit, and acidity will be encountered over a period
ECY34 Dirty of months and over time (>1 yr), may display gueuze-like qualities in complexity. Contains various isolates from lambic-producers, B. bruxellensis, B. anomalus, B.
East Coast Yeast Dozen Brett lambicus, and B. naardenensis. For those who want the most from Brett yeast, whether a 100% Brett fermentation is desired or adding to secondary aging projects.
Blend Suggested fermentation temperature: 60-74 F. Attenuation high. See this MTF thread for fermentation tips for 100% and mixed fermentations (https://www.facebook.com/
groups/MilkTheFunk/permalink/1258857067475838/).
Escarpment Berliner Brett A blend of Berliner Brett 1 (Brettanomyces anomalus) and Berliner Brett 2 (Brettanomyces bruxellensis) for balanced, subtle Brett character in traditional Berliner Weisse
Laboratories Blend and beyond. Flavour profile includes citrus, white wine, and peach. Secondary pitch rates only.
Escarpment MOTHERSHIP This blend typically contains 10 individual strains. The character is highly dependent on fermentation conditions, but tends toward balanced, medium to high intensity
Laboratories Brett Blend Brett character.
Fermmento Labs FB1 Tropical

Contains B. custersianus and 'B. anomalus.
(Brazil) funky Bugs
Brux Blend
GigaYeast A blend of Brettanomyces yeast that produces stone fruit esters and a hint of barnyard. Creates a moderate amount of acid that adds a tart complexity to the brew.
(GB156)
This will be an evolving blend comprised of nearly every Brettanomyces strain in our collection (inaugural release will contain 12 strains). When used in secondary,
Omega Yeast All The Bretts expect high attenuation and a fruity and funky complexity developing over time. Attenuation: 85+%; Flocculation: low; Temperature: 68F-85F. Homebrew pitch contains
Labs OYL-218
~70 billion cells [287].
Hurricane Brett
Mainiacal Yeast Mix of all 70 Brettanomyces strains at Mainiacal Yeast. Commercial brewery pitches always available, and occasionally available to homebrewers [269].
Blend
MIP-765
Blend of Brettanomyces isolated from a number of natural sources. Isolates are from plums, natural wines, wild plants, and spontaneous fermentation. This blend will add
Propagate Lab Native Brett
Blend complexity to any beer [273].
Not overly funky but there is a sweaty note hanging behind lemon and citrus fruits, nice blend of subtle funk and citrus/fruit. All strains were identified as B. bruxellensis
The Yeast Bay Beersel [288]. Making a starter is fine despite the instructions advising against it on the vial and will not greatly effect the character of the final beer [289].
Similar to Beersel but with more funk in aroma and less fruit, complex barnyard character. All strains were identified as B. bruxellensis [288]. Making a starter is fine
The Yeast Bay Brussels
despite the instructions advising against it on the vial and will not greatly effect the character of the final beer [289].
Smells of Iris C2, probably the same, subtle blend with some delicate fruit, strawberry. All strains were identified as B. bruxellensis [288]. Making a starter is fine despite
The Yeast Bay Lochristi
the instructions advising against it on the vial and will not greatly effect the character of the final beer [289].
Amalgamation
6 Brett blend to create a dry beer with a bright and complex fruit-forward flavor and aroma, accompanied by some funk. All strains were identified as B. bruxellensis [288].
The Yeast Bay Brett Super
Blend Making a starter is fine despite the instructions advising against it on the vial and will not greatly effect the character of the final beer [289].
Amalgamation II is a blend of 5 Brettanomyces isolates: Brettanomyces bruxellensis - Strain TYB184, Brettanomyces bruxellensis - Strain TYB207, Brettanomyces
bruxellensis - Strain TYB261, and both Beersel Brettanomyces Blend isolates. The balanced funk of the Beersel isolates and TYB184, the sweet tart character of TYB207,
Amalgamation and the tropical bouquet of the combined ester profile of lemon/pineapple/guava/mango/papaya contributed by all the isolates. Expect beers fermented with this blend to
The Yeast Bay II Brett Super finish crisp, dry, tart and fruity with just a touch of funk on the finish. This blend produces noticeable character only 3-4 weeks into fermentation and is well suited for
Blend faster turnaround beers. Amalgamation II shines as a primary or secondary fermenter. Making a starter is fine despite the instructions advising against it on the vial and
will not greatly effect the character of the final beer [289]. Reportedly a fast fermenter [290]
Using Brettanomyces
Primary versus Secondary Fermentation
Brettanomyces can be pitched into a beer at many points in the beer's fermentation life cycle. If used as the primary fermenter, the beer that is produced is often perceived as
fruit forward and not very "funky". A large cell count will be needed (somewhere between an ale and lager pitching rate). See the 100% Brettanomyces Fermentation page for
more information on pitching rates for 100% Brettanomyces fermentation, as well as the fermentation characteristics of 100% Brettanomyces fermentation. If pitched into a
beer that has already been fermented by Saccharomyces or if co-pitched with Saccharomyces, a wider range of flavors including the funkier flavors can be produced by
changing some of the flavor compounds produced during the Saccharomyces fermentation into other flavor compounds (see the Brettanomyces Metabolism section above),
although Brettanomyces can also produce funky phenols on its own (de novo) without phenolic precursors produced from Saccharomyces. A small cell count of
Brettanomyces is plenty for creating these flavors if pitched after and co-pitched with Saccharomyces, and normally a starter is not necessary unless pitching Brettanomyces
as the primary fermenter. See the Mixed Fermentation, Brettanomyces and Saccharomyces Co-fermentation, and Brettanomyces secondary fermentation experiment pages for
more information on using Brettanomyces in secondary.
Starter Information
When pitching just Brettanomyces from a commercial pure or blended culture and no other microbes, it is recommended to make a starter for the culture. If the
Brettanomyces is being pitched into secondary, no starter is necessary unless the brewer suspects that the Brettanomyces has lost a lot of viability due to age, heat exposure,
etc., or prefers higher cell count pitches (current information suggests that there is no significant flavor difference between high and low pitching rates in secondary pitches of
Brettanomyces; see Brettanomyces secondary fermentation experiment).
Starter wort made with dried malt extract at around 1.040 starting gravity is adequate for most strains of B. bruxellensis. For Brettanomyces strains such as many B. anomalus
strains that don't ferment maltose, a mixture of 50% DME and 50% table sugar or dextrose at 1.040 starting gravity should be adequate. Yeast nutrient at the manufacturer's
recommended rate can be added if sufficient growth is not observed [291].
Just like in other yeast species, temperature has a direct effect on the rate of growth for Brettanomyces. The optimal growth rate temperature range for Brettanomyces is
between 25-32°C (77-90°F). Growth is about half as slow at 20°C (68°F). Brettanomyces will still grow at temperatures as low as (and maybe lower than) 15°C (59°F) and
will be much slower, however one study showed a slightly higher viability during the full-time period of fermentation at 15°C as opposed to the optimal growth temperature
range of 20-32°C. At a temperature of 35°C (95°F), both growth and viability over time are greatly inhibited [111].
For information on mixed culture starters, see Mixed Culture Starters.
Two Approaches to Starters

There are generally two approaches to handling Brettanomyces starters. The first is to use a stir plate set to a medium-high RPM with tin foil on top of the flask for 7-8 days,
cold crash for a few days, and then decant the beer before pitching the sedimented yeast. The second approach is to use an orbital shaker set to 80 RPM to create a semi-
aerobic environment (this means that the oxygen levels are low, but also not non-existent) for 7-8 days as described in The Brettanomyces project [292], cold crashing can be
skipped, and the entire starter is pitched into the wort. An alternative to the second approach is to use a stir plate on a very low setting so that only a very small "dimple" of a
vortex is formed [293]. If a stir plate is not available, give the starter an initial dosage of pure O2, and then cover it with foil so that oxygen can slowly diffuse into the starter,
and gently agitate as often as possible [294].
Oxygen levels are an important factor to consider when deciding which of the above two methods to use for a Brettanomyces starter. Brettanomyces creates acetic acid in the
presence of oxygen, potentially leading to higher levels of ethyl acetate, which is considered an off flavor in higher amounts. As the amount of oxygen increases, cell growth
increases, but so does acetic acid production. The amount of acetic acid produced is species/strain dependent, so some strains may benefit from more aeration without having
the negative effect of creating too much acetic acid. Other strains may need a less aerobic starter (semi-aerobic) in order to produce the highest cell count with minimal acetic
acid [295][296][297]. In addition to acetic acid production, it has been observed that some Brettanomyces strains grown under aerobic conditions continue to produce THP when
transferred to anaerobic conditions. See THP for details.
This presents a sort of "catch 22" when growing Brettanomyces in a starter. The brewer must weigh the pros and cons of how much aeration to provide. If the Brettanomyces
is going to be used in a 100% Brettanomyces Fermentation, for example, then a stir plate with foil covering the flask is the best choice. If the Brettanomyces is instead being
pitched in secondary with the intention of long aging, then having a high cell count isn't as necessary and the risk of adding more acetic acid/ethyl acetate to an aging beer is
greater. If a lot of acetic acid is produced during the starter, then they can opt to cold crash and decant the starter. Brettanomyces can have a difficult time flocculating and
settling out, even when cold crashed. The brewer may need to allow a few days for the cells to fully sediment [298]. Additionally, Brettanomyces that is cold crashed may be
slower to begin fermentation. If the brewer believes that the amount of acetic acid produced was insignificant, then cold crashing can be skipped and the entire starter can be
pitched. Even if the starter has a lot of acetic acid, the amount of acetic acid in the volume of a starter is fairly insignificant once diluted into a full batch of wort or beer. If the
starter is not going to be used within a month, then an aerobic starter is not the best option since the presence of a lot of acetic acid will slowly kill the Brettanomyces over
time. In this case, the starter should be lightly shaken (or occasionally manually stirred), and an airlock put in place on the flask in order to keep out most of the oxygen.
Although more experiments are probably needed, agitation is believed to be an important factor for any species of microbe (yeast and bacteria). Gentle stirring on a stir plate
or orbital shaker, or frequent gentle manual agitation leads to faster growth and a higher number of organisms. Agitation keeps the microbes in solution. It also maximizes the
microbes' access to nutrients and disperses waste evenly. In a non-agitated starter, the microbes are limited to the diffusion rate of nutrients, leading to a slower and more
stressful growth [299].
Maintaining a temperature of 77°-86°F/25°-30°C results in faster growth than lower temperatures and is recommended [300]. Brettanomyces cell growth typically takes about
7-8 days to reach it's maximum growth [301], however some strains may grow at faster rates and finish in 3-4 days [302][56]. When the starter turns a rich creamy color, it
should be done within a few hours after this visual indication occurs [300]. Each step of a starter for Brett should be 7-8 days (or 3-4 days for faster growing strains).
For more information regarding aeration and agitation effects on Brettanomyces growth, see Mark Trent's Brettanomyces Propagation Experiment.
Pitching Rate Calculators
Current yeast pitching calculators for brewers are not adequate for determining Brettanomyces pitching rates based on starter volume size because the maximum cell density
of Brettanomyces per mL of wort is 3 to 6 times the cell density of Saccharomyces. For example, a given Saccharomyces strain may reach a cell density of 130 million cells
per mL in a 1.040 wort (different Saccharomyces strains can have different cell densities as well, although they are a lot lower than Brettanomyces overall). Different
Brettanomyces strain cell densities have been reported to be 600 to 885 million cells per mL in 1.040 wort depending on the species/strain [301][303]. Since yeast calculators
are based on S. cerevisiae or S. pastorianus cell density, using one of these tools for Brettanomyces starters will create an unexpectedly high cell count in reality. There is not
currently enough data to accurately determine starter volumes for Brettanomyces, particularly because each strain and species have a different maximum cell density per mL
of wort. However, pitching around 500-600 mL of a Brettanomyces starter for 5 gallons of 1.060 SG wort will achieve a pitching rate that is similar to lager yeast pitching
rates, which has been recommended for 100% Brettanomyces Fermentation. Omega Yeast Labs is currently working on a project to create a more accurate Brettanomyces
pitching rate calculator (it will also contain pitching rate calculations for specific strains of Saccharomyces, which is something that current yeast pitching calculators do not
include) [303].
Given this information, many brewers historically have been using the lager pitching rate settings in online yeast pitching calculators for Brettanomyces starters (around 2000
mL for 5 gallons, for example). Effectively, this means they have been pitching around 4 to 5 times the amount of Brettanomyces cells that they thought they were pitching.
However, if this very high pitching rate is giving good results for brewers, it should continue to be used. Exploration of Brettanomyces pitching rates for 100% Brett
fermentations is something to be desired once we know what our pitching rates actually are, and many brewers have been pitching 4-5 times the pitching rate for lagers if
they use an online yeast pitching rate calculator instead of counting the cells under a microscope.
See also 100% Brettanomyces fermentation.
MYPG Growth Substrate and Other Laboratory Substrates
For yeast laboratories, "Malt Yeast Peptone Glucose" growth substrate has been shown to be a better substrate than wort for initially growing Brettanomyces from a plate or
slant. When grown in wort, Brettanomyces will often go through a 24 hour lag phase, a growth phase, another lag phase, and a second growth phase (all within 7-8 days).
When grown in MYPG substrate, there is only a single growth phase and no lag phase, which has been reported by Yakobson to produce a larger cell count in the same
amount of time [304]. Cells grown in MYPG also are better adapted to grow in wort [305]. Practical instructions for making this substrate can be found on Jason Rodriguez's
blog, "Brew Science - Homebrew Blog (http://sciencebrewer.com/2011/04/29/wild-yeast-project-mypg-culture-media/)". Unfortunately, growing Brettanomyces pitches in
MYPG for breweries isn't very practical due to needing almost 4 times the amount of MYPG versus wort to get the same pitching rate. In a brewery or homebrewery, using
wort for Brettanomyces starters is more practical [306].
For other suggested substrates for growing Brettanomyces and potentially other yeasts, see Laboratory Techniques.
Cell Counting
The use of methylene blue, although popular in breweries, has been shown to be inaccurate when counting cells of Brettanomyces. Trypan blue staining has been shown to
give more accurate cell counting results than methylene blue [132][307].

One of the problems with cell counting for Brettanomyces is that they tend to form pseudohyphae (elongated cells that are attached to one another) during growth. This makes
counting the cells difficult. Martyniak et al. (2017) has proposed a new way of counting cells for Brettanomyces by using the fluorescence capability of a Nexcelom X2 image
cytometer (https://www.nexcelom.com/nexcelom-products/cellometer-fluorescent-viability-cell-counters/cellometer-x2-fluorescent-automated-viability-cell-counter/) with
stains created from acridine orange (AO) and propidium iodide (PI). The AO-PI stains only the nuclei of the individual cells, allowing them to be counted easily by the
cytometer regardless of pseudohyphae formation. During their research, they found that a "clausenii" strain of B. anomalus formed more pseudohyphae than two strains of B.
bruxellensis (one of which was refered to as a "lambicus" strain), and this corresponded with higher viability over time. It
was therefore hyopthesized that pseudohyphae might play a role in the longer survival of some strains of Brettanomyces
over others [308][309][310]. It has been observed that pseudohyphae might be less common in Brettanomyces cultures that
have been grown in wort versus lab media, making them easier to count [311], but this method makes it easy to count cells
which are attached via pseudohyphae. This method has been criticized for its relatively high cost (both for the fluorescent
microscope and the dyes) [312].
See also:
MBAA Podcast interview with Leo Chan on automating cell counting with of Brettanomyces psuedohyphae. (http:/
/masterbrewerspodcast.com/092-automated-brettanomyces-cell-counts)
Example of a Home Lab Orbital Shaker
Mark Trent's shaker platform (obtained from a used equipment outlet in Gilroy, CA called "Outback Equipment" ) used to
create a semi-aerobic environment for Brettanomyces. Mark built an insulated box for it, and added temperature control.
He can propagate up to 7 liters. This is running at 80 RPM as described in The Brettanomyces project [292][313].
Bright-field and fluorescent images of B. clausenii, B.
bruxellensis, and B. lambicus. The fluorescent images
IMG 0599 show the counting of the yeasts with and without
declustering the bud and pseudohyphae. Source:
https://link.springer.com/article/10.1007/s10295-016-
1861-4. Used with permission from Brian Martyniak.
Storing Brett
Major yeast labs will often store yeast in a -80°C laboratory freezer in a media/glycerol solution, although this option is generally not practical for brewers [314]. The next
best option for long-term storage of Brettanomyces is freezing with 10% glycerol in a home freezer. However, the effects of storing yeast at such a high and often variable
temperature have not been evaluated scientifically. Traditionally Saccharomyces yeast has been stored on slants held in a refrigerator and can provide storage for a few
months up to 2+ years, depending on the type of slant used (using mineral oil in slants has been shown to extend the life of stored Saccharomyces). Homebrewers, however,
have reported poor survival of Brettanomyces on slants. Data from a MTF member showed promising results by buffering the slant media. In this data, Brettanomyces has
stored well for up to 100 days on the buffered media. It is not known for how long viability will remain high on buffered slants. For instructions on how to make slants at
home capable of storing any microbe for potentially 2+ years, see Bryan's video on Sui Generis Brewing (requires a pressure cooker) (http://suigenerisbrewing.blogspot.com/
2015/11/easy-home-yeast-banking-and-video.html). Agar plates are the least effective solution and have been observed anecdotally to reduce the viability of Brettanomyces
over a few months [315][316].
Perhaps the best method for storing Brettanomyces long term is in sterilized (autoclaved or pressure cooked) wort or MYPG. Although not as ideal as freezing with glycerol
at -80°C, this is the most practical way to store Brettanomyces for brewers without a lab freezer. Regarding temperature, it has been shown that cold storage for as long as a
month is better than room temperature. However, after one month Brettanomyces appears to be more viable when stored at room temperature. More data is required before
assuming this is the case with all strains of Brettanomyces. Chad Yakobson noted that after storing Brettanomyces in a refrigerated environment (we don't know how Chad
was storing the Brettanomyces cultures when he observed this, for example on agar plates or slants or something else.), after 6 months the Brettanomyces would die. If
Brettanomyces is stored cold, it will be very sluggish and slow to start fermentation. Making a starter is highly recommended if the Brettanomyces culture has been stored
cold [317].
In order to explore Yakobson's anecdotal observations in a more controlled manner, Mark Trent performed an experiment on storing one strain of Brettanomyces in wort,
MYPG, buffered wort (buffered to prevent a drop in pH), and buffered MYPG, and compared storage of the Brettanomyces in each of the storage solutions at room
temperature versus cold temperatures for 100 days. This single Brettanomyces strain survived best in unbuffered MYPG at room temperature, and second best in unbuffered
wort at room temperature, and survived less in cold storage conditions for all media. See the Brettanomyces Storage Survival Experiment for more details. Therefore, when
storing Brettanomyces for one month or less in wort (or perhaps beer), it should be stored refrigerated. However, if the Brettanomyces will be stored for more than one month
in wort (or perhaps beer), it should be stored at room temperature (until more data improves our understanding). Note that at best these storage techniques will decrease
viability greatly (80%+) within 3 months, and a starter should be used to try and revive the culture before use [318].
Occasional feeding has been shown to keep Brettanomyces alive in beer for brewers who do not have a lab; however, many variables may come into play as far as how
effective this will be for individual strains and in different environments. Although no research has been done to indicate what the best practices are for feeding
Brettanomyces to keep it alive in beer, we recommend trying this method: every 3-6 months swirl the vessel so as to suspend all of the yeast and then decant 70-90% of the
beer and suspended yeast slurry, and replace it with a 1.040 starter wort with yeast nutrients. This method will discard a lot of the old yeast cells, while retaining enough
living cells for replication [319]. Some strains may survive extended periods of aging in beer; however, their viability and vitality will be greatly reduced over time.
Interestingly, Brettanomyces remains more viable over time if it was co-fermented with S. cerevisiae than if it was fermented without the presence of S. cerevisiae; i.e. 100%
Brettanomyces beers or Brettanomyces and Lactobacillus [132].

Another method for storing Brettanomyces has reportedly worked for MTF member Justin Amaral. This method involves storing the culture in isotonic sodium chloride.
Brettanomyces cultures have been reported by Amaral to survive at least for 6-7 months. This includes other microbes as well (RVA Orchard Brett, ECY Dirty Dozen, Bright
Yeast Labs Brett Chateaux, T. delbrueckii, L. plantarum isolated from goodbelly, Omega Lacto blend, Pediococcus damnosus, Bootleg Biology Sour Weapon, and Funk
Weapon 2 and 3, and a Brettanomyces isolate from Yeast Bay). For more information on this method, see this Eureka Brewing blog article (https://eurekabrewing.wordpress.c
om/2012/09/05/yeast-banking-3-isotonic-sodium-chloride/) [320].
Fermentation Methods
Mixed Fermentation
Brettanomyces and Saccharomyces Co-fermentation
100% Brettanomyces Fermentation
Spontaneous Fermentation
Fermenting Under Head Pressure
Tips From Brewers
The Yeast Bay
Both Ed Coffey (http://www.alesoftheriverwards.com/2015/03/revisiting-yeast-bay-brett-blends.html) and Amos Browne (http://www.browneandbitter.com/2015/03/ta

sting-notes-lochristi-peach-sour.html?utm_source=feedburner&utm_medium=email&utm_campaign=Feed%3A+BrowneAndBitter+%28Browne+and+Bitter%29)
have noted that the Lorchristi blend ages particularly well.
To make a starter for the Lochristi blend, run it semi-aerobic for 4-6 days in the 70's and then let it settle at room temp and decant what you can if the starter is large
[321].
MTF thread on The Yeast Bay Brett/Sacch flavor profiles. (https://www.facebook.com/groups/MilkTheFunk/permalink/1585625541465654/)
Escarpment Laboratories
"How to Choose a Brett Strain For Beer." (https://escarpmentlabs.com/blogs/resources/how-to-choose-a-brett-strain-for-beer)
Pasteurization
Brettanomyces has complete thermal death at 122°F (50°C) for 5 minutes [110][109] . See also Barrel Sanitizing and Pasteurization.
Catching/Bioprospecting Wild Brettanomyces

See Isolating Wild Brettanomyces.
See Also
Additional Articles on MTF Wiki
Brettanomyces Propagation Experiment
Brettanomyces secondary fermentation experiment
Brettanomyces Storage Survival Experiment
100% Brettanomyces Fermentation
Crooked Stave Artisan Beer Project
Scientific Publications
Mixed Cultures
Mixed Fermentation
Brettanomyces and Saccharomyces Co-fermentation
Debaryomyces
Saccharomyces
External Resources
Family tree of Brettanomyces, by Eureka Brewing Blog. (https://eurekabrewing.wordpress.com/2012/05/26/family-tree-of-brettanomyces/)

Insight into the Brettanomyces Mitochondrial Genome, by Eureka Brewing Blog. (https://eurekabrewing.wordpress.com/2013/01/19/brettanomyces-genome-blasting/)
National Collection of Yeast Cultures in the UK - Database on what compounds different species/strains can ferment. (https://catalogue.ncyc.co.uk/catalogsearch/result/
?q=brettanomyces)
The Brettanomyces Project - Chad Yakobon's Brett research. (http://www.brettanomycesproject.com/)
The Mad Fermentationist - Commercial Brettanomyces, Lactobacillus, and Pediococcus Descriptions (http://www.themadfermentationist.com/p/commercial-cultures.ht
ml)
The Mad Fermentationist - Comparison between English Ale yeast and Belgian Ale yeast primary fermentations, and Brett in secondary (http://www.themadfermentati
onist.com/2014/11/phenols-and-brett-initial-results.html?m=1)
Blog Article on Brett and Glycosides by Cy Wood. (http://www.centralstatebrewing.com/blog/2015/7/1/whats-that-smell-eraroma-compound)
Brettanomyces in Brewing the horse the goat and the barnyard, presentation by Chad Yakobson. (http://www.mbaa.com/districts/michigan/events/Documents/2011_01
_14BrettanomycesBrewing.pdf)
Insight into the Dekkera anomala YV396 genome by Samuel Aeschlimann; self published on Eureka Brewing Blog. (http://www.scribd.com/doc/277758178/Insight-int
o-the-Dekkera-anomala-YV396-genome)
Esters - Table of esters and their smells. (https://jameskennedymonash.files.wordpress.com/2013/12/table-of-esters-and-their-smells.jpg)
See the Phenol Explorer website for more information on sources of precursors. (http://phenol-explorer.eu/foods)
Scientific research on Brettanomyces by Mike Lentz et al., funded by the Oregon Wine Board. (http://industry.oregonwine.org/research/)
Old and new taxonomical classification of Brettanomyces and Dekkera. (http://www.sciencedirect.com/science/article/pii/S0168160515001865#t0005)
"Brettanomyces", Steve Piatz in Brew Your Own Magazine, October 2005. (http://byo.com/stories/issue/item/262-brettanomyces)

"100 year old Czech beer," by Dave Janssen (musings on the presence of Brettanomyces and high amounts of acetic and lactic acid in a 100 year old lager). (http://ww
w.horscategoriebrewing.com/2017/06/100-year-old-czech-beer.html)
"Brewing with Brettanomyces Yeast and Mixed Cultures" presentation by Chad Yakobson at BC Craft Brewers Guild, 2019. (https://www.youtube.com/watch?v=qaX7
tiFQ2iA)
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Brettanomyces - Milk The Funk Wiki 11/05/2023 21:07

https://www.milkthefunk.com/wiki/Brettanomyces Page 1 sur 35

6.1 Additional Articles on MTF Wiki

Introduction of History, Characteristics, and Taxonomy

https://www.milkthefunk.com/wiki/Brettanomyces Page 2 sur 35

Brettanomyces morphology examples from Remi Bonnart. (http://brettanomycesproject.com/2010/06/brettanomyces-yeast-cell-images/)

Environment and Survival

https://www.milkthefunk.com/wiki/Brettanomyces Page 3 sur 35

https://www.milkthefunk.com/wiki/Brettanomyces Page 4 sur 35

Calculations for SO2 additions (http://www.foodsci.purdue.edu/research/labs/enology/Winemaking%20Calculations%20Spring%20Workshop%204_29_2011.pdf?fbcli

https://www.milkthefunk.com/wiki/Brettanomyces Page 5 sur 35

Carbohydrate Metabolism and Fermentation Temperature

https://www.milkthefunk.com/wiki/Brettanomyces Page 6 sur 35

Cellobiose Soluble Starch

https://www.milkthefunk.com/wiki/Brettanomyces Page 7 sur 35

https://www.milkthefunk.com/wiki/Brettanomyces Page 8 sur 35

Glycosides and Beta-Glucosidase Activity

See the Glycosides page for more details.

https://www.milkthefunk.com/wiki/Brettanomyces Page 9 sur 35

Brettanomyces secondary fermentation experiment.

https://www.milkthefunk.com/wiki/Brettanomyces Page 10 sur 35

Amyl octanoate (spicy, Amyl alcohol and C13H26O2

Ethyl acetate (fruity, 33ppm (odor), C4H8O2

Ethyl butyrate (pineapple,

Ethyl caproate (sweet, fruity,

Ethyl caprylate (Sweet, Caprylic acid

Ethyl Decanoate/Ethyl Decanoic acid (Capric

Ethyl hexanoate [17] (Apple Hexanoic acid and 0.2ppm (flavor

Ethyl valerate (Sweet, fruity,

Octyl butyrate (spicy,

Pentyl formate (artificial Pentanol and formic

https://www.milkthefunk.com/wiki/Brettanomyces Page 11 sur 35

Effects of Saccharomyces Strain Selection and Staggered vs Co-Pitch

https://www.milkthefunk.com/wiki/Brettanomyces Page 12 sur 35

Phenol Flavor/Odor Molecular

https://www.milkthefunk.com/wiki/Brettanomyces Page 13 sur 35

Acids Produced Precursors Notes

Acetic Acid (Vinegar, hard

Caproic acid (Fatty, cheesy,

Heptanoic acid (sweaty, Associated with phenylalanine or

Isovaleric Acid (Feety,

Nonanoic acid (Rancid odor)

Source: Agnolucci M, Vigentini I, Capurso G, Merico A, Tirelli A, Compagno C,

https://www.milkthefunk.com/wiki/Brettanomyces Page 14 sur 35

For more information on biogenic amines, see the following:

Biogenic amines in spontaneous fermentation.

https://www.milkthefunk.com/wiki/Brettanomyces Page 15 sur 35

Bisabolene (spicy, Associated

Decanol [17] Alcohol Also known as decyl alcohol [245].

Nonanal (lemon Associated

0.1–0.2 ppm in lager

Bootleg Biology/Spot Yeast

https://www.milkthefunk.com/wiki/Brettanomyces Page 16 sur 35

Common Species Synonym

Brewing Science Institute

Common Species Synonym

Thought to be the same as White Labs WLP648; however,

Dekkera Brettanomyces Chad Yakobson's mutation of BSI Drie. Commercial pitches

Brettanomyces Isolate from Drie Fonteinen; Pro-Brewers only.

Community Cultures Yeast Lab

Common Species Synonym Source

Common Species Synonym

Isolated from a wild rasberry in Michigan's Upper Peninsula along

East Coast Yeast