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(In progress) For a general description and information on '''lactic acid''' see [https://en.wikipedia.org/wiki/Lactic_acid the Wikipedia entry]. ==Amounts in Sour Beer==* [https://www.facebook.com/groups/MilkTheFunk/permalink/2139922666035936/ Summary of measured amounts of lactic acid in various type of sour beers.]

It has recently been discovered that both L-lactic acid and D-lactic acid produced by lactic acid bacteria can manipulate a genetic trait in yeast that dictates how yeast ferments different types of sugars, including some strains (but not all) of ''Saccharomyces cerevisiae'' and ''Brettanomyces bruxellensis'' <ref name="Garcia_2016">[https://elifesciences.org/content/5/e17978 A common bacterial metabolite elicits prion-based bypass of glucose repression. David M Garcia, David Dietrich, Jon Clardy, Daniel F Jarosz. 2016.]</ref>. Normally ''Saccharomyces'' and many other types of yeast will preferentially metabolize glucose when glucose is present and they ignore other sugars such as maltose and maltotriose until the glucose is completely consumed(or until 40-50% of glucose is consumed in the case of wort and in anaerobic conditions <ref>Stewart, G. G. (2006). Studies on the uptake and metabolism of wort sugars during brewing fermentations. Tech. Q. Master Brew. Assoc. Am. 43:265-269. </ref>). This is called "glucose repression". Recently it has been identified that lactic acid produced by lactic acid bacteria in the presence of some yeast strains turns off this "glucose repression" in yeast, allowing them to simultaneously ferment all types of sugars. This has a side effect of limiting attenuation in wine, and It has been one of the identified causes of proposed by some researchers to explain stuck fermentations in wine fermentations (it has been observed as far back as Louis Pasteur that stuck wine fermentations often contain lactic acid bacteria) , but this claim has been disputed by Ramakrishnan et al. (2016) who showed that yeast that have been effected this way can still ferment wine adequately and is not directly the cause of stuck wine fermentations, partly because in natural conditions (juice fermentations versus lab growth media) only 50% of the total yeast population changes in this way, leaving plenty of yeast that still demonstrate normal glucose repression. They (and others) were also able to show that the presence of acetic acid might also play a role in triggering this state in yeast <ref name="cross-kingdom">[http://weitzlab.seas.harvard.edu/files/weitzlab/files/2014_cell_jarosz.pdf Cross-Kingdom Chemical Communication Drives a Heritable, Mutually Beneficial Prion-Based Transformation of Metabolism. 2014. Daniel F. Jarosz, Jessica C.S. Brown, Gordon A. Walker, Manoshi S. Datta, W. Lloyd Ung, Alex K. Lancaster, Assaf Rotem, Amelia Chang, Gregory A. Newby,David A. Weitz, Linda F. Bisson, and Susan Lindquist. Cell. 2014 Aug 28;158(5):1083-93.]</ref><ref name="Ramakrishnan_2016">[https://www.frontiersin.org/articles/10.3389/fevo.2016.00137/full Inter-Kingdom Modification of Metabolic Behavior: GAR+ Prion Induction in Saccharomyces cerevisiae Mediated by Wine Ecosystem Bacteria. Vidhya Ramakrishnan, Gordon A. Walker, Qingwen Fan, Minami Ogawa, Yan Luo, Peter Luong, C. M. Lucy Joseph and Linda F. Bisson. 2016. DOI: https://doi.org/10.3389/fevo.2016.00137.]</ref>.  The ability for yeast to bypass glucose repression and ferment multiple types of sugars simultaneously is controlled a by a protein-based genetic [https://en.wikipedia.org/wiki/Prion prion] called '''<nowiki>[</nowiki>GAR⁺<nowiki>]</nowiki>'''. These genetic "prions" are not the same as DNA in genes but are rather misfolded proteins contained in the cytoplasm of the cell. These proteins are dominant over <nowiki>[</nowiki>gar^-<nowiki>]</nowiki>, and are passed to the offspring of the cell during cell division. This type of passing of genetic material from mother cell to daughter cell is much more frequent than genetic mutations and probably exists to help yeast populations quickly adapt to rapidly changing conditions in their environment. Normally in brewers yeast only a small number of cells are <nowiki>[</nowiki>GAR⁺<nowiki>]</nowiki> if any at all. In the brewing environment where there is no competition from other yeasts, brewers yeast benefits from consuming glucose first. In the wild, however, many more strains have been found to be <nowiki>[</nowiki>GAR⁺<nowiki>]</nowiki>. This is thought to be an adaptive advantage for wild yeast depending on the environments in which they live; such yeasts can "hedge their bets" towards consuming other types of sugars, with the side effect of allowing bacteria to produce compounds such as lactic acid that may inhibit competing yeasts <ref name="Jarosz_2014">[http://www.cell.com/cell/abstract/S0092-8674(14)00974-X An Evolutionarily Conserved Prion-like Element Converts Wild Fungi from Metabolic Specialists to Generalists. Daniel F. Jarosz, Alex K. Lancaster, Jessica C.S. Brown, Susan Lindquist. Cell. Volume 158, Issue 5, p1072–1082, 28 August 2014]</ref><ref name="cross-kingdom">[http://weitzlab.seas.harvard.edu/files/weitzlab/files/2014_cell_jarosz.pdf Cross-Kingdom Chemical Communication Drives a Heritable, Mutually Beneficial Prion-Based Transformation of Metabolism. 2014. Daniel F. Jarosz, Jessica C.S. Brown, Gordon A. Walker, Manoshi S. Datta, W. Lloyd Ung, Alex K. Lancaster, Assaf Rotem, Amelia Chang, Gregory A. Newby, David A. Weitz, Linda F. Bisson, and Susan Lindquist. Cell. 2014 Aug 28;158(5):1083-93.]</ref>.

The Not only do <nowiki>[</nowiki>GAR⁺<nowiki>]</nowiki> yeast have the ability for yeast to bypass glucose repression and ferment multiple types of other sugars simultaneously is controlled a at the same time as glucose, but it has been proposed by a protein-based genetic some researchers that they produce less alcohol (for example in the referenced study, the <nowiki>[https:</nowiki>gar<sup>-</en.wikipedia.orgsup><nowiki>]</wiki/Prion prion] called nowiki> ''Saccharomyces''cells fermented grape must into a 12% ABV wine, and the <nowiki>[</nowiki>GAR⁺<nowiki>]</nowiki>''Saccharomyces''cells fermented the same wine must into an 8% ABV wine <ref name="cross-kingdom"/>). These genetic "prions" are not the same as DNA Viability over time is also increased in genes, but yeast cells that are rather misfolded proteins contained in the cytoplasm <nowiki>[</nowiki>GAR⁺<nowiki>]</nowiki> versus those that aren't. In wild fermentation of grapes, the cell. These proteins are dominant over wild <nowiki>[</nowiki>garGAR^-+<nowiki>]</nowiki>, and are passed to ''S. cerevisiae'' strains thrived over the offspring other types of fungi that were found on the cell during cell divisionwild grapes. This type While the induction of passing <nowiki>[</nowiki>GAR⁺<nowiki>]</nowiki> can occur in yeast without the presence of genetic material from mother cell to daughter cell is much more frequent than genetic mutationslactic acid, and probably exists to help yeast populations quickly adapt to rapidly changing conditions in their environment. Normally in brewers yeast only a small number the presence of lactic acid greatly increases the occurrence of cells are <nowiki>[</nowiki>GAR⁺<nowiki>]</nowiki> if any at all. In the brewing environment where there is no competition from other yeasts, brewers in yeast benefits from consuming glucose first. In the wildcells, however, many with higher concentrations of lactic acid producing more strains have been found to be occurrences of <nowiki>[</nowiki>GAR⁺<nowiki>]</nowiki>cells. This is thought The lactic acid merely has to be an adaptive advantage present for wild yeast depending on the environments in which they live; such yeasts can "hedge their bets" towards consuming other types of sugarsthis to happen, with and the side effect of allowing bacteria yeast's ability to produce compounds such as metabolize lactic acid or not does not have an effect. It is thought that may inhibit competing yeasts this benefits both the <ref name="Jarosz_2014"nowiki>[http:</nowiki>GAR<sup>+</www.cell.comsup><nowiki>]</cell/abstract/S0092-8674(14)00974-X An Evolutionarily Conserved Prion-like Element Converts Wild Fungi from Metabolic Specialists to Generalistsnowiki> ''S. Daniel F. Jaroszcerevisiae'' and the lactic acid bacteria, Alex Kwhich are often found together in the wild during fermentation of fruit. Lancaster The bacteria isn't killed by higher alcohol levels, Jessica C.S. Brown, Susan Lindquistand the yeast has a broader food source. Cell. Volume 158, Issue 5, p1072–1082, 28 August 2014]</ref>This effect that lactic acid bacteria have on yeast is known as ''cross kingdom chemical communication'' <ref name="cross-kingdom"/>[http:<ref name="Garcia_2016" //weitzlab>.seas Ramakrishnan et al.harvard.edu(2016) disputed this claim by showing that <nowiki>[</filesnowiki>GAR<sup>+</weitzlabsup><nowiki>]</files/2014_cell_jarosz.pdf Cross-Kingdom Chemical Communication Drives a Heritablenowiki> yeast can still ferment sugars adequately, Mutually Beneficial Prion-Based Transformation of Metabolism. 2014. Daniel F. Jarosz, Jessica C.S. Brown, Gordon A. Walker, Manoshi S. Datta, W. Lloyd Ung, Alex K. Lancaster, Assaf Rotem, Amelia Chang, Gregory A. Newby,David A. Weitz, Linda F. Bissonand showed that these yeast were slower at depleting nitrogen and oxygen, thus leaving these resources for the bacteria and Susan Lindquist. Cell. 2014 Aug 28;158(5):1083-93.]creating an environment that stresses the yeast which results in the reported stuck wine fermentations <ref name="Ramakrishnan_2016" /ref>.

Not In a previous study by Jarosz et al. (2014), it was observed that only do certain bacteria species had the effect of inducing <nowiki>[</nowiki>GAR⁺<nowiki>]</nowiki> in some strains of yeast have the ability to ferment other sugars at the same time as glucose. These bacteria included ''P. damnosus'', ''Lactobacillus kunkeei'', and species from genres ''Staphylococcus'', ''Micrococcus'', ''Bacillus'', ''Listeria'', ''Paenibacillus'', ''Gluconobacter'', ''Sinorhizobium'', ''Escherichia'', but they produces less alcohol (for example in the referenced ''Serriatia''. In this study, the <nowiki>[</nowiki>gar^-<nowiki>]</nowiki> ''SaccharomycesL. brevis'', ''L. hilgardii'' cells fermented grape must into a 12% ABV wine, and the ''L. plantarum'' did not appear to induce <nowiki>[</nowiki>GAR⁺<nowiki>]</nowiki> ''Saccharomyces'' cells fermented the same wine must into an 8% ABV wine in yeast <ref name="cross-kingdom"/>). Viability over At the time is also increased in yeast cells of this study, it was not understood that are lactic acid was an inducer of <nowiki>[</nowiki>GAR⁺<nowiki>]</nowiki> versus those that aren't. In wild fermentation of grapesAdditionally, the wild <nowiki>[</nowiki>GAR⁺<nowiki>]</nowiki> ''Smethod they used to discover this was simply to streak bacteria next to yeast on a plate, and see if it grew on a medium that would show whether or not they bypassed glucose repression. cerevisiae Therefore, it is possible that not enough lactic acid was produced, or that they didn'' strains thrived over t give the other types of fungi that were found bacteria enough time to have an effect on the wild grapesyeast. While the induction of <nowiki>[</nowiki>GAR⁺<nowiki>]</nowiki> can occur in yeast without the presence of More work would need to be done to show that indeed all lactic acid, the presence of bacteria that produce lactic acid greatly increases the occurrence of have this effect on yeast, and also it is not clear whether other bacterial metabolites may influence this phenomenon <nowikiref name="preiss">[<https://www.facebook.com/groups/MilkTheFunk/nowiki>GAR<sup>+<permalink/sup><nowiki>]<1500240613337481/nowiki> in yeast cells, Conversation with higher concentrations of lactic acid producing more occurrences of <nowiki>[<Richard Preiss on MTF. 12/nowiki>GAR<sup>+<7/sup><nowiki>2016.]</nowikiref> cells. The lactic acid merely has to be present for this to happen, and the yeast's ability to metabolize lactic acid or not does not have an effectRamakrishnan et al. It is thought (2016) more recently confirmed that this benefits both the induction of <nowiki>[</nowiki>GAR⁺<nowiki>]</nowiki> ''S. cerevisiae'' and the lactic acid bacteriais not only species dependent, which are often found together in the wild during fermentation of fruit. The bacteria isn't killed by higher alcohol levels, and the yeast has a broader food source. This effect that lactic acid bacteria have on yeast is known as ''cross kingdom chemical communication'' but strain dependent within species <ref name="cross-kingdom" /><ref name="Garcia_2016Ramakrishnan_2016" />.

In a previous study by Jarosz et albeer, this might explain other observations as well. (2014) For example, it was observed that only certain bacteria species had the effect of inducing <nowiki>[</nowiki>GAR⁺<nowiki>]</nowiki> in Yakobson reported higher attenuation with some strains of yeast. These bacteria included ''P. damnosusBrettanomyces bruxellensis''(WLP650, ''Lactobacillus kunkeei''BSI Drie, CMY001, and species from genres WY5526) and one strain of ''Staphylococcus'', ''Micrococcus'', ''Bacillus'', ''Listeria'', ''Paenibacillus'', ''Gluconobacter'', ''Sinorhizobium'', ''Escherichia'', ''SerriatiaB. anomalus''(WY5151) demonstrated a trend of increased attenuation with increasing concentrations of lactic acid <ref>[http://brettanomycesproject.com/dissertation/pure-culture-fermentation/impact-of-initial-concentration-of-lactic-acid/ "The Brettanomyces Project". In this study, ''LChad Yakobson. brevis'', ''L 2011. hilgardii'', ''L. plantarum'' did not appear to induce <nowiki>[< Retrieved 12/nowiki>GAR<sup>+<7/sup><nowiki>2016.]</nowiki> in yeast <ref name="cross-kingdom" />. At the time In mixed fermentations of beers such as lambic and American sour ales, attenuation is often slower, but typically eventually reaches a high degree of attenuation. Some strains of ''S. cerevisiae'' are more tolerant of acidic conditions than others. Although this studymight answer some questions, it was not understood that lactic acid was an inducer mixed fermentation is a complex thing with many other variables and more work needs to be done to identify whether all or just some strains of yeast/bacteria have the effect of inducing <nowiki>[</nowiki>GAR⁺<nowiki>]</nowiki>. Additionally, the method they used to discover this was simply to streak bacteria next to yeast on a plate, and see if it grew on a medium that would show whether or not they bypassed glucose repression. Therefore, it is possible how that not enough lactic acid was produced, or that they didn't give might affect the bacteria enough time to have an effect on the yeast. More work would need to be done to show that indeed all lactic acid bacteria that produce lactic acid have this effect on yeast, and also it is not clear whether other bacterial metabolites may influence this phenomenon fermentation profile of various types of beers <ref name="preiss">[https://www.facebook.com/groups/MilkTheFunk/?comment_tracking=%7B%22tn%22%3A%22R4%22%7D Conversation with Richard Preiss on MTF. 12/7/2016.]</ref>.

In beer, this might explain other observations as well. For example, Yakobson reported higher attenuation with some strains of ''Brettanomyces bruxellensis'' (WLP650, BSI Drie, CMY001, and WY5526) and one strain of ''B. anomalus'' (WY5151) demonstrated a trend of increased attenuation with increasing concentrations of lactic acid <ref>See also [httphttps://brettanomycesprojectwww.facebook.com/dissertationgroups/pure-culture-fermentationMilkTheFunk/impact-of-initial-concentration-of-lactic-acidpermalink/ "The Brettanomyces Project". Chad Yakobson. 2011. Retrieved 121500240613337481/7/2016.]</ref>. In mixed fermentations this MTF thread by Richard Preiss of beers such as lambic and American sour ales, attenuation is often slower, but typically eventually reaches a high degree Escarpment Labs on speculating impacts of attenuation. Some strains of ''S. cerevisiae'' are more tolerant of acidic conditions than others. Although this might answer some questions, mixed fermentation is a complex thing with many other variables and more work needs to be done to identify whether all or just some strains of yeastin sour/bacteria have the effect of inducing <nowiki>[</nowiki>GAR⁺<nowiki>wild beer]</nowiki>, and how that might effect the fermentation profile of various types of beers <ref name="preiss"/>.