Microbiology and Biochemistry: Difference between revisions

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# An enteric stage, starting around three days after the boil and ending around 30-90 days, in which enteric bacteria dominate.

# ~~Saccromyces~~ dominance, lasting from three to about thirty weeks, in which ~~Saccromyces~~ cerveciae and other ~~Saccromyces~~ species dominate the beer's yeast flora.

# [[Saccharomyces]] dominance, lasting from three to about thirty weeks, in which Saccharomyces cerveciae and other Saccharomyces species dominate the beer's yeast flora.

# Pediococcus dominance, lasting from 2 months onward, in which Pediococcus and other [[lactic acid]] bacteria dominate the bacterial flora.

# Brettanomyces dominance, lasting from 4-8 months onward.

Line 25:

MNova Users Meeting, SMASH – Atlanta, GA, September 7, 2014</ref> Significant changes to the concentration of acetic acid should not occur until the ethanol has a chance to oxidize in aging in the bottle over many years or even decades.<ref name=Vanderhaegen1> B. Vanderhaegen, H. Neven, H. Verachtert, G. Derdelinckx [http://www.google.com/url?sa=t&rct=j&q=&esrc=s&source=web&cd=1&ved=0CCAQFjAA&url=http%3A%2F%2Fwww.researchgate.net%2Fprofile%2FGuy_Derdelinckx%2Fpublication%2F222839054_The_chemistry_of_beer_aging__a_critical_review%2Flinks%2F0c960523339c4b25a6000000.pdf&ei=Tq3IVKmfFcGyogSs_YLQCA&usg=AFQjCNFaBrvqDGjqEV2I9uQ73dYh_ParXg&sig2=Z8dY4iDHozbT1eb9JeAdrw&bvm=bv.84607526,d.cGU| The chemistry of beer aging – a critical review], 2006</ref> The pellicle that forms on the top of the wort forms around ___ days, and may be the product of acetobacteria during the enteric phase,[9] though most other sources inidcate that the pellicle is the result of Brettanomyces (with Pichia and Candida).<ref name="Guinard">Jean-Xavier Guinard, [[Books#Classic Beer Styles: Lambic|Classic Beer Styles: Lambic]], 1990</ref>

Low pH (below ~4.5) and an ethanol concentration higher than ~2% by volume is a hostile environment to the enterobacteria, and ~~Saccromyces~~ species are able to dominate the flora in the wort once these conditions occur around 30-60 days into fermentation.

Low pH (below ~4.5) and an ethanol concentration higher than ~2% by volume is a hostile environment to the enterobacteria, and Saccharomyces species are able to dominate the flora in the wort once these conditions occur around 30-60 days into fermentation.

==~~Saccromyces~~ dominance==

==Saccharomyces dominance==

After the pH falls below ~4.5 and the alcohol content rises over ~2%, ~~Saccromyces~~ species take over as the dominant organisms in the wort, though ~~Saccromyces~~ is present in large numbers well prior to the dissapearance of the enterobacteria. ~~Saccromyces~~ will remain dominant until at least 6-8 months into fermentation, and will maintain a presence, though no longer active, throughout fermentation. Despite ~~Saccromyces~~' importance to the fermentation of Lambic, its concentrations remain below 10^7 cells per mL of wort, which is considerably lower than the 10^8 cells/mL found in commercial beers.<ref name=Oevelen77 >D. Van Oevelen, M. Spaepen, P. Timmermans and H. Verachtert, [http://onlinelibrary.wiley.com/doi/10.1002/j.2050-0416.1977.tb03825.x/abstract|MICROBIOLOGICAL ASPECTS OF SPONTANEOUS WORT FERMENTATION IN THE PRODUCTION OF LAMBIC AND GUEUZE], 1977</ref><ref name="Guinard">Jean-Xavier Guinard, [[Books#Classic Beer Styles: Lambic|Classic Beer Styles: Lambic]], 1990</ref>

After the pH falls below ~4.5 and the alcohol content rises over ~2%, [[Saccharomyces species]] take over as the dominant organisms in the wort, though Saccharomyces is present in large numbers well prior to the dissapearance of the enterobacteria. Saccharomyces will remain dominant until at least 6-8 months into fermentation, and will maintain a presence, though no longer active, throughout fermentation. Despite Saccharomyces' importance to the fermentation of Lambic, its concentrations remain below 10^7 cells per mL of wort, which is considerably lower than the 10^8 cells/mL found in commercial beers.<ref name=Oevelen77 >D. Van Oevelen, M. Spaepen, P. Timmermans and H. Verachtert, [http://onlinelibrary.wiley.com/doi/10.1002/j.2050-0416.1977.tb03825.x/abstract|MICROBIOLOGICAL ASPECTS OF SPONTANEOUS WORT FERMENTATION IN THE PRODUCTION OF LAMBIC AND GUEUZE], 1977</ref><ref name="Guinard">Jean-Xavier Guinard, [[Books#Classic Beer Styles: Lambic|Classic Beer Styles: Lambic]], 1990</ref>

As in controlled fermentation, ~~Saccromyces~~ is responsible for most ethanol production and attenuation in lambic. The yeasts consume all the major sugars found in lambic wort (glucose, maltose, and some maltotriose). By the end of the ~~Saccromyces~~ phase around 8 months, the ethanol content of the beer stabilizes at 5-7% by volume and will remain around that value until the end of fermentation[5,7]. Attenuation after ~~Saccromyces~~ fermentation reaches 60-65%, which is known as the "attenuation limit" for conventional beers. Despite being responsible for most of the ethanol in Lambic, yeasts of the ~~Saccromyces~~ genus are not responsible for most of the aroma and flavor compounds that give Lambic its distinct sensory characteristics.<ref name="Guinard">Jean-Xavier Guinard, [[Books#Classic Beer Styles: Lambic|Classic Beer Styles: Lambic]], 1990</ref>

As in controlled fermentation, Saccharomyces is responsible for most ethanol production and attenuation in lambic. The yeasts consume all the major sugars found in lambic wort (glucose, maltose, and some maltotriose). By the end of the Saccharomyces phase around 8 months, the ethanol content of the beer stabilizes at 5-7% by volume and will remain around that value until the end of fermentation[5,7]. Attenuation after Saccharomyces fermentation reaches 60-65%, which is known as the "attenuation limit" for conventional beers. Despite being responsible for most of the ethanol in Lambic, yeasts of the Saccharomyces genus are not responsible for most of the aroma and flavor compounds that give Lambic its distinct sensory characteristics.<ref name="Guinard">Jean-Xavier Guinard, [[Books#Classic Beer Styles: Lambic|Classic Beer Styles: Lambic]], 1990</ref>

Other bacteria and yeasts also thrive during the ~~Saccromyces~~ phase. Kloeckera and ____ both survive in considerable numbers alongside ~~Saccromyces~~. In the case of Kloeckera, the growth begins prior to ~~Saccromyces~~ in the first days of fermentation, but are overgrown by ~~Saccromyces~~ in the following weeks.

Other bacteria and yeasts also thrive during the Saccharomyces phase. Kloeckera and ____ both survive in considerable numbers alongside Saccharomyces. In the case of Kloeckera, the growth begins prior to Saccharomyces in the first days of fermentation, but are overgrown by Saccharomyces in the following weeks.

==Pediococcus dominance==

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==The Brettanomyces stage==

Brettanomyces inherits the role of most prominant yeast genera from ~~Saccromyces~~ around 8 months, and continues consuming sugar in the wort. Final attenuation can reach over 80% in lambic through the continued action of Brettanomyces, which is often referred to as "overattenuation" or "superattenuation". This is greater than is usually possible with ~~saccromyces~~ alone, as brettanomyces is able to metabolise sugars that ~~saccromyces~~ cannot, generally known as "dextrins".

Brettanomyces inherits the role of most prominant yeast genera from Saccharomyces around 8 months, and continues consuming sugar in the wort. Final attenuation can reach over 80% in lambic through the continued action of Brettanomyces, which is often referred to as "overattenuation" or "superattenuation". This is greater than is usually possible with Saccharomyces alone, as brettanomyces is able to metabolise sugars that Saccharomyces cannot, generally known as "dextrins".

Brettanomyces has been implicated in producing most of the aroma compounds in Lambic.<ref name="Guinard">Jean-Xavier Guinard, [[Books#Classic Beer Styles: Lambic|Classic Beer Styles: Lambic]], 1990</ref> Sensory-significant quantities of ethyl acetate and ethyl lactate form at this time from ethanol entering into an ester bond with acetic and lactic acid, respectively. The esterization process is greatly helped by the enzyme esterase provided by Brettanomyces. However, the enzymatic esterization is highly reversible and esters found in high concentrations in the lambic prior to the presence of the esterase will achieve a lower equilibrium. This is the case with iso-amyl acetate, which is produced by ~~Saccromyces~~ and is a characteristic odor compound in most other beers.[ref] Tetrahydropyridines (THPs) produced by Brettanomyces give Lambic its horse-like aroma, though the concentration and thus smell of THPs is variable.[ref, Heresztyn, 1986]

Brettanomyces has been implicated in producing most of the aroma compounds in Lambic.<ref name="Guinard">Jean-Xavier Guinard, [[Books#Classic Beer Styles: Lambic|Classic Beer Styles: Lambic]], 1990</ref> Sensory-significant quantities of ethyl acetate and ethyl lactate form at this time from ethanol entering into an ester bond with acetic and lactic acid, respectively. The esterization process is greatly helped by the enzyme esterase provided by Brettanomyces. However, the enzymatic esterization is highly reversible and esters found in high concentrations in the lambic prior to the presence of the esterase will achieve a lower equilibrium. This is the case with iso-amyl acetate, which is produced by Saccharomyces and is a characteristic odor compound in most other beers.[ref] Tetrahydropyridines (THPs) produced by Brettanomyces give Lambic its horse-like aroma, though the concentration and thus smell of THPs is variable.[ref, Heresztyn, 1986]

=Refermentation in the bottle=

Gueuze and other lambic bottled with either some residual sugar left unfermented at the time of bottling or added priming sugars will undergo significant fermentation in the bottle, though all unpasteurized lambic will continue to ferment to some degree there. For lambic bottled after about 8 months without additional sugar, the fermentation in the bottle progresses much as an extension of the Brettanomyces stage of fermentation and negligible carbon dioxide and ethanol production occurrs, leaving most of this lambic still. For lambic bottled younger and those with additional fermentable sugars added at bottling, considerable fermentation by ~~saccromyces~~ occurs in the bottle, causing marked increases in ethanol and the production of carbon dioxide, leading to a carbonated product [ref].

Gueuze and other lambic bottled with either some residual sugar left unfermented at the time of bottling or added priming sugars will undergo significant fermentation in the bottle, though all unpasteurized lambic will continue to ferment to some degree there. For lambic bottled after about 8 months without additional sugar, the fermentation in the bottle progresses much as an extension of the Brettanomyces stage of fermentation and negligible carbon dioxide and ethanol production occurrs, leaving most of this lambic still. For lambic bottled younger and those with additional fermentable sugars added at bottling, considerable fermentation by Saccharomyces occurs in the bottle, causing marked increases in ethanol and the production of carbon dioxide, leading to a carbonated product [ref].

The reactivation of ~~saccromyces~~ fermentation at the addition of additional sugar indicates that the dominance of brettanomyces is brought about by brettanomyces ability to ferment sugars that ~~saccromyces~~ cannot. Once ~~saccromyces~~ has consumed most of the available ______ and ______ in the wort, it goes dormant, though does not die completely, and brettanomycs is free to assume the role of primary yeast in the wort. Upon the re-introduction of _____ and _____, the faster-growing ~~saccromyces~~ once again flourishes, until the again the fermentable sugar is consumed and brettanomyces and its other associated yeasts can once again resume the slow procress of the final fermentation.

The reactivation of Saccharomyces fermentation at the addition of additional sugar indicates that the dominance of brettanomyces is brought about by brettanomyces ability to ferment sugars that Saccharomyces cannot. Once Saccharomyces has consumed most of the available ______ and ______ in the wort, it goes dormant, though does not die completely, and brettanomycs is free to assume the role of primary yeast in the wort. Upon the re-introduction of _____ and _____, the faster-growing Saccharomyces once again flourishes, until the again the fermentable sugar is consumed and brettanomyces and its other associated yeasts can once again resume the slow procress of the final fermentation.

==Continued aging==

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==Spontaneous fermentation in other traditional drinks and foods==

Other spontaneous fermentations exist as well, such as Chicha, a corn-based lightly alcoholic drink indigenous to the Andes. Much like the spontaneous fermentation of Lambic, Chicha posesses its own set of yeasts, dominated by strains of ~~Saccromyces~~ responsible for its unique characteristics.<ref name=Vallejoa >J. A. Vallejoa, P. Mirandaa, J. D. Flores-Félixb, F. Sánchez-Juanesc, J. M. Ageitosa, J. M. González-Buitragoc, E. Velázquezb, T. G. Villaa, [http://www.sciencedirect.com/science/article/pii/S0723202013001513|Atypical yeasts identified as Saccharomyces cerevisiae by MALDI-TOF MS and gene sequencing are the main responsible of fermentation of chicha, a traditional beverage from Peru], 2013</ref>

Other spontaneous fermentations exist as well, such as Chicha, a corn-based lightly alcoholic drink indigenous to the Andes. Much like the spontaneous fermentation of Lambic, Chicha posesses its own set of yeasts, dominated by strains of Saccharomyces responsible for its unique characteristics.<ref name=Vallejoa >J. A. Vallejoa, P. Mirandaa, J. D. Flores-Félixb, F. Sánchez-Juanesc, J. M. Ageitosa, J. M. González-Buitragoc, E. Velázquezb, T. G. Villaa, [http://www.sciencedirect.com/science/article/pii/S0723202013001513| Atypical yeasts identified as Saccharomyces cerevisiae by MALDI-TOF MS and gene sequencing are the main responsible of fermentation of chicha, a traditional beverage from Peru], 2013</ref>

Brettanomyces and other wild organisms, despite often being spoilage organisms in wine, are responsible for some major sensory characteristics of some wine styles.<ref name=Tristezza> M. Tristezza, C. Vetrano, G. Bleve, G. Spano, V. Capozzi, A. Logrieco, G. Mita, F. Grieco [http://www.sciencedirect.com/science/article/pii/S0740002013001408 | Biodiversity and safety aspects of yeast strains characterized from vineyards and spontaneous fermentations in the Apulia Region, Italy], 2013</ref><ref name=Medina> K. Medina, E. Boido, L. Fariña, O. Gioia, M.E. Gomez, M. Barquet, C. Gaggero, E. Dellacassa, F. Carrau [http://www.sciencedirect.com/science/article/pii/S0308814613005025 | Increased flavour diversity of Chardonnay wines by spontaneous fermentation and co-fermentation with Hanseniaspora vineae], 2013</ref> Other minor yeasts found in Lambic, such as Pichia and Hanseniaspora have also been found in unspoiled wine.<ref name=Tristezza> M. Tristezza, C. Vetrano, G. Bleve, G. Spano, V. Capozzi, A. Logrieco, G. Mita, F. Grieco [http://www.sciencedirect.com/science/article/pii/S0740002013001408 | Biodiversity and safety aspects of yeast strains characterized from vineyards and spontaneous fermentations in the Apulia Region, Italy], 2013</ref><ref name=Medina> K. Medina, E. Boido, L. Fariña, O. Gioia, M.E. Gomez, M. Barquet, C. Gaggero, E. Dellacassa, F. Carrau [http://www.sciencedirect.com/science/article/pii/S0308814613005025 | Increased flavour diversity of Chardonnay wines by spontaneous fermentation and co-fermentation with Hanseniaspora vineae], 2013</ref>

@@ Line 5: / Line 5: @@
 # An enteric stage, starting around three days after the boil and ending around 30-90 days, in which enteric bacteria dominate.
-# Saccromyces dominance, lasting from three to about thirty weeks, in which Saccromyces cerveciae and other Saccromyces species dominate the beer's yeast flora.
+# [[Saccharomyces]] dominance, lasting from three to about thirty weeks, in which Saccharomyces cerveciae and other Saccharomyces species dominate the beer's yeast flora.
 # Pediococcus dominance, lasting from 2 months onward, in which Pediococcus and other [[lactic acid]] bacteria dominate the bacterial flora.
 # Brettanomyces dominance, lasting from 4-8 months onward.
@@ Line 25: / Line 25: @@
 MNova Users Meeting, SMASH – Atlanta, GA, September 7, 2014</ref> Significant changes to the concentration of acetic acid should not occur until the ethanol has a chance to oxidize in aging in the bottle over many years or even decades.<ref name=Vanderhaegen1> B. Vanderhaegen, H. Neven, H. Verachtert, G. Derdelinckx [http://www.google.com/url?sa=t&rct=j&q=&esrc=s&source=web&cd=1&ved=0CCAQFjAA&url=http%3A%2F%2Fwww.researchgate.net%2Fprofile%2FGuy_Derdelinckx%2Fpublication%2F222839054_The_chemistry_of_beer_aging__a_critical_review%2Flinks%2F0c960523339c4b25a6000000.pdf&ei=Tq3IVKmfFcGyogSs_YLQCA&usg=AFQjCNFaBrvqDGjqEV2I9uQ73dYh_ParXg&sig2=Z8dY4iDHozbT1eb9JeAdrw&bvm=bv.84607526,d.cGU| The chemistry of beer aging – a critical review], 2006</ref> The pellicle that forms on the top of the wort forms around ___ days, and may be the product of acetobacteria during the enteric phase,[9] though most other sources inidcate that the pellicle is the result of Brettanomyces (with Pichia and Candida).<ref name="Guinard">Jean-Xavier Guinard, [[Books#Classic Beer Styles: Lambic|Classic Beer Styles: Lambic]], 1990</ref>
-Low pH (below ~4.5) and an ethanol concentration higher than ~2% by volume is a hostile environment to the enterobacteria, and Saccromyces species are able to dominate the flora in the wort once these conditions occur around 30-60 days into fermentation.
+Low pH (below ~4.5) and an ethanol concentration higher than ~2% by volume is a hostile environment to the enterobacteria, and Saccharomyces species are able to dominate the flora in the wort once these conditions occur around 30-60 days into fermentation.
-==Saccromyces dominance==
+==Saccharomyces dominance==
-After the pH falls below ~4.5 and the alcohol content rises over ~2%, Saccromyces species take over as the dominant organisms in the wort, though Saccromyces is present in large numbers well prior to the dissapearance of the enterobacteria. Saccromyces will remain dominant until at least 6-8 months into fermentation, and will maintain a presence, though no longer active, throughout fermentation. Despite Saccromyces' importance to the fermentation of Lambic, its concentrations remain below 10^7 cells per mL of wort, which is considerably lower than the 10^8 cells/mL found in commercial beers.<ref name=Oevelen77 >D. Van Oevelen, M. Spaepen, P. Timmermans and H. Verachtert, [http://onlinelibrary.wiley.com/doi/10.1002/j.2050-0416.1977.tb03825.x/abstract|MICROBIOLOGICAL ASPECTS OF SPONTANEOUS WORT FERMENTATION IN THE PRODUCTION OF LAMBIC AND GUEUZE], 1977</ref><ref name="Guinard">Jean-Xavier Guinard, [[Books#Classic Beer Styles: Lambic|Classic Beer Styles: Lambic]], 1990</ref>
+After the pH falls below ~4.5 and the alcohol content rises over ~2%, [[Saccharomyces species]] take over as the dominant organisms in the wort, though Saccharomyces is present in large numbers well prior to the dissapearance of the enterobacteria. Saccharomyces will remain dominant until at least 6-8 months into fermentation, and will maintain a presence, though no longer active, throughout fermentation. Despite Saccharomyces' importance to the fermentation of Lambic, its concentrations remain below 10^7 cells per mL of wort, which is considerably lower than the 10^8 cells/mL found in commercial beers.<ref name=Oevelen77 >D. Van Oevelen, M. Spaepen, P. Timmermans and H. Verachtert, [http://onlinelibrary.wiley.com/doi/10.1002/j.2050-0416.1977.tb03825.x/abstract|MICROBIOLOGICAL ASPECTS OF SPONTANEOUS WORT FERMENTATION IN THE PRODUCTION OF LAMBIC AND GUEUZE], 1977</ref><ref name="Guinard">Jean-Xavier Guinard, [[Books#Classic Beer Styles: Lambic|Classic Beer Styles: Lambic]], 1990</ref>
-As in controlled fermentation, Saccromyces is responsible for most ethanol production and attenuation in lambic. The yeasts consume all the major sugars found in lambic wort (glucose, maltose, and some maltotriose). By the end of the Saccromyces phase around 8 months, the ethanol content of the beer stabilizes at 5-7% by volume and will remain around that value until the end of fermentation[5,7]. Attenuation after Saccromyces fermentation reaches 60-65%, which is known as the "attenuation limit" for conventional beers. Despite being responsible for most of the ethanol in Lambic, yeasts of the Saccromyces genus are not responsible for most of the aroma and flavor compounds that give Lambic its distinct sensory characteristics.<ref name="Guinard">Jean-Xavier Guinard, [[Books#Classic Beer Styles: Lambic|Classic Beer Styles: Lambic]], 1990</ref>
+As in controlled fermentation, Saccharomyces is responsible for most ethanol production and attenuation in lambic. The yeasts consume all the major sugars found in lambic wort (glucose, maltose, and some maltotriose). By the end of the Saccharomyces phase around 8 months, the ethanol content of the beer stabilizes at 5-7% by volume and will remain around that value until the end of fermentation[5,7]. Attenuation after Saccharomyces fermentation reaches 60-65%, which is known as the "attenuation limit" for conventional beers. Despite being responsible for most of the ethanol in Lambic, yeasts of the Saccharomyces genus are not responsible for most of the aroma and flavor compounds that give Lambic its distinct sensory characteristics.<ref name="Guinard">Jean-Xavier Guinard, [[Books#Classic Beer Styles: Lambic|Classic Beer Styles: Lambic]], 1990</ref>
-Other bacteria and yeasts also thrive during the Saccromyces phase. Kloeckera and ____ both survive in considerable numbers alongside Saccromyces. In the case of Kloeckera, the growth begins prior to Saccromyces in the first days of fermentation, but are overgrown by Saccromyces in the following weeks.
+Other bacteria and yeasts also thrive during the Saccharomyces phase. Kloeckera and ____ both survive in considerable numbers alongside Saccharomyces. In the case of Kloeckera, the growth begins prior to Saccharomyces in the first days of fermentation, but are overgrown by Saccharomyces in the following weeks.
 ==Pediococcus dominance==
@@ Line 44: / Line 44: @@
 ==The Brettanomyces stage==
-Brettanomyces inherits the role of most prominant yeast genera from Saccromyces around 8 months, and continues consuming sugar in the wort. Final attenuation can reach over 80% in lambic through the continued action of Brettanomyces, which is often referred to as "overattenuation" or "superattenuation". This is greater than is usually possible with saccromyces alone, as brettanomyces is able to metabolise sugars that saccromyces cannot, generally known as "dextrins".
+Brettanomyces inherits the role of most prominant yeast genera from Saccharomyces around 8 months, and continues consuming sugar in the wort. Final attenuation can reach over 80% in lambic through the continued action of Brettanomyces, which is often referred to as "overattenuation" or "superattenuation". This is greater than is usually possible with Saccharomyces alone, as brettanomyces is able to metabolise sugars that Saccharomyces cannot, generally known as "dextrins".
-Brettanomyces has been implicated in producing most of the aroma compounds in Lambic.<ref name="Guinard">Jean-Xavier Guinard, [[Books#Classic Beer Styles: Lambic|Classic Beer Styles: Lambic]], 1990</ref> Sensory-significant quantities of ethyl acetate and ethyl lactate form at this time from ethanol entering into an ester bond with acetic and lactic acid, respectively. The esterization process is greatly helped by the enzyme esterase provided by Brettanomyces. However, the enzymatic esterization is highly reversible and esters found in high concentrations in the lambic prior to the presence of the esterase will achieve a lower equilibrium. This is the case with iso-amyl acetate, which is produced by Saccromyces and is a characteristic odor compound in most other beers.[ref] Tetrahydropyridines (THPs) produced by Brettanomyces give Lambic its horse-like aroma, though the concentration and thus smell of THPs is variable.[ref, Heresztyn, 1986]
+Brettanomyces has been implicated in producing most of the aroma compounds in Lambic.<ref name="Guinard">Jean-Xavier Guinard, [[Books#Classic Beer Styles: Lambic|Classic Beer Styles: Lambic]], 1990</ref> Sensory-significant quantities of ethyl acetate and ethyl lactate form at this time from ethanol entering into an ester bond with acetic and lactic acid, respectively. The esterization process is greatly helped by the enzyme esterase provided by Brettanomyces. However, the enzymatic esterization is highly reversible and esters found in high concentrations in the lambic prior to the presence of the esterase will achieve a lower equilibrium. This is the case with iso-amyl acetate, which is produced by Saccharomyces and is a characteristic odor compound in most other beers.[ref] Tetrahydropyridines (THPs) produced by Brettanomyces give Lambic its horse-like aroma, though the concentration and thus smell of THPs is variable.[ref, Heresztyn, 1986]
 =Refermentation in the bottle=
-Gueuze and other lambic bottled with either some residual sugar left unfermented at the time of bottling or added priming sugars will undergo significant fermentation in the bottle, though all unpasteurized lambic will continue to ferment to some degree there. For lambic bottled after about 8 months without additional sugar, the fermentation in the bottle progresses much as an extension of the Brettanomyces stage of fermentation and negligible carbon dioxide and ethanol production occurrs, leaving most of this lambic still. For lambic bottled younger and those with additional fermentable sugars added at bottling, considerable fermentation by saccromyces occurs in the bottle, causing marked increases in ethanol and the production of carbon dioxide, leading to a carbonated product [ref].
+Gueuze and other lambic bottled with either some residual sugar left unfermented at the time of bottling or added priming sugars will undergo significant fermentation in the bottle, though all unpasteurized lambic will continue to ferment to some degree there. For lambic bottled after about 8 months without additional sugar, the fermentation in the bottle progresses much as an extension of the Brettanomyces stage of fermentation and negligible carbon dioxide and ethanol production occurrs, leaving most of this lambic still. For lambic bottled younger and those with additional fermentable sugars added at bottling, considerable fermentation by Saccharomyces occurs in the bottle, causing marked increases in ethanol and the production of carbon dioxide, leading to a carbonated product [ref].
-The reactivation of saccromyces fermentation at the addition of additional sugar indicates that the dominance of brettanomyces is brought about by brettanomyces ability to ferment sugars that saccromyces cannot. Once saccromyces has consumed most of the available ______ and ______ in the wort, it goes dormant, though does not die completely, and brettanomycs is free to assume the role of primary yeast in the wort. Upon the re-introduction of _____ and _____, the faster-growing saccromyces once again flourishes, until the again the fermentable sugar is consumed and brettanomyces and its other associated yeasts can once again resume the slow procress of the final fermentation.
+The reactivation of Saccharomyces fermentation at the addition of additional sugar indicates that the dominance of brettanomyces is brought about by brettanomyces ability to ferment sugars that Saccharomyces cannot. Once Saccharomyces has consumed most of the available ______ and ______ in the wort, it goes dormant, though does not die completely, and brettanomycs is free to assume the role of primary yeast in the wort. Upon the re-introduction of _____ and _____, the faster-growing Saccharomyces once again flourishes, until the again the fermentable sugar is consumed and brettanomyces and its other associated yeasts can once again resume the slow procress of the final fermentation.
 ==Continued aging==
@@ Line 103: / Line 103: @@
 ==Spontaneous fermentation in other traditional drinks and foods==
-Other spontaneous fermentations exist as well, such as Chicha, a corn-based lightly alcoholic drink indigenous to the Andes. Much like the spontaneous fermentation of Lambic, Chicha posesses its own set of yeasts, dominated by strains of Saccromyces responsible for its unique characteristics.<ref name=Vallejoa >J. A. Vallejoa, P. Mirandaa, J. D. Flores-Félixb, F. Sánchez-Juanesc, J. M. Ageitosa, J. M. González-Buitragoc, E. Velázquezb, T. G. Villaa, [http://www.sciencedirect.com/science/article/pii/S0723202013001513|Atypical yeasts identified as Saccharomyces cerevisiae by MALDI-TOF MS and gene sequencing are the main responsible of fermentation of chicha, a traditional beverage from Peru], 2013</ref>
+Other spontaneous fermentations exist as well, such as Chicha, a corn-based lightly alcoholic drink indigenous to the Andes. Much like the spontaneous fermentation of Lambic, Chicha posesses its own set of yeasts, dominated by strains of Saccharomyces responsible for its unique characteristics.<ref name=Vallejoa >J. A. Vallejoa, P. Mirandaa, J. D. Flores-Félixb, F. Sánchez-Juanesc, J. M. Ageitosa, J. M. González-Buitragoc, E. Velázquezb, T. G. Villaa, [http://www.sciencedirect.com/science/article/pii/S0723202013001513| Atypical yeasts identified as Saccharomyces cerevisiae by MALDI-TOF MS and gene sequencing are the main responsible of fermentation of chicha, a traditional beverage from Peru], 2013</ref>
 Brettanomyces and other wild organisms, despite often being spoilage organisms in wine, are responsible for some major sensory characteristics of some wine styles.<ref name=Tristezza> M. Tristezza, C. Vetrano, G. Bleve, G. Spano, V. Capozzi, A. Logrieco, G. Mita, F. Grieco [http://www.sciencedirect.com/science/article/pii/S0740002013001408 | Biodiversity and safety aspects of yeast strains characterized from vineyards and spontaneous fermentations in the Apulia Region, Italy], 2013</ref><ref name=Medina> K. Medina, E. Boido, L. Fariña, O. Gioia, M.E. Gomez, M. Barquet, C. Gaggero, E. Dellacassa, F. Carrau [http://www.sciencedirect.com/science/article/pii/S0308814613005025 | Increased flavour diversity of Chardonnay wines by spontaneous fermentation and co-fermentation with Hanseniaspora vineae], 2013</ref> Other minor yeasts found in Lambic, such as Pichia and Hanseniaspora have also been found in unspoiled wine.<ref name=Tristezza> M. Tristezza, C. Vetrano, G. Bleve, G. Spano, V. Capozzi, A. Logrieco, G. Mita, F. Grieco [http://www.sciencedirect.com/science/article/pii/S0740002013001408 | Biodiversity and safety aspects of yeast strains characterized from vineyards and spontaneous fermentations in the Apulia Region, Italy], 2013</ref><ref name=Medina> K. Medina, E. Boido, L. Fariña, O. Gioia, M.E. Gomez, M. Barquet, C. Gaggero, E. Dellacassa, F. Carrau [http://www.sciencedirect.com/science/article/pii/S0308814613005025 | Increased flavour diversity of Chardonnay wines by spontaneous fermentation and co-fermentation with Hanseniaspora vineae], 2013</ref>