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==Effects on Yeast==
===Stress on Yeast===
The presence of lactic acid has been identified as a yeast stressor. However, yeast can be acclimated to lactic acid and low pH conditions by growing them in media that has a slowly increasing amount of lactic acid. See [[Saccharomyces#Fermentation_Under_Low_pH_Conditions|''Saccharomyces'' Fermentation Under Low pH]]. ''[[Brettanomyces]]'' tends to be more acid tolerant than ''S. cerevisiae'', and the presence of lactic acid will change the way ''Brettanomyces'' ferments wort and the esters that it will produce. See [[Brettanomyces#Ester_Production|''Brettanomyces'' ester production]] for more information.
===Genetic Manipulation===
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 ''S. 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 only ferment 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<sup>+</sup><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<sup>-</sup><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<sup>+</sup><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<sup>+</sup><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>.
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<sup>+</sup><nowiki>]</nowiki> in Yakobson reported higher attenuation with some strains of yeast. These bacteria included ''P. damnosusBrettanomyces bruxellensis''(WLP650, BSI Drie, ''Lactobacillus kunkeei''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<sup>+</sup><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 even though this does seem to actually be the case 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>.
==See Also==
* [[Lactobacillus]]
* [[Pediococcus]]
* [[Saccharomyces#Fermentation_Under_Low_pH_Conditions|''Saccharomyces'' Fermentation Under Low pH]]
===External Resources===
==References==