Changes

Many wine yeast strains are known to be "killer" yeast strains(this term has been changed to "zymocidal" in science <ref name="Stewart_2018">[https://link.springer.com/chapter/10.1007/978-3-319-69126-8_10 "Killer (Zymocidal) Yeasts." Brewing and Distilling Yeasts. Graham G. Stewart. 2018.]</ref>). In ''Saccharomyces'', killer strains are genetically determined to produce secrete toxins (in the form of extracellular proteins or glycoproteins) called 'mycocins' that kill sensitive strains. The mycocin toxins can act on sensitive strains in a number of ways: by blocking DNA synthesis and preventing cell division, inhibiting the synthesis of the cell wall component called "beta-glucans (β-1,36-glucan") that a part of their cell wall formation, and by causing ions to leak through the cell wall. In low dosages, which is typical in the natural environment, toxin triggers active cell death ([http://www.biology-pages.info/A/Apoptosis.html apoptosis]), while large dosages cause necrotic cell killing ([https://en.wikipedia.org/wiki/Necrosis necrosis]). One study in wine found that the use of killer strains to out compete sensitive strains resulted in off-flavors from yeast autolysis <ref>[http://www.nature.com/nrmicro/journal/v4/n3/full/nrmicro1347.html Yeast viral killer toxins: lethality and self-protection. Manfred J. Schmitt & Frank Breinig. 2006.]</ref><ref name="Hatoum2012">[http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3525881/ Rima Hatoum, Steve Labrie, and Ismail Fliss. 2012.]</ref>. Another study found that a lager strain that was genetically modified to secrete killer toxins was able to eliminate all cells of a sensitive ale strain within 24 hours of beer fermentation at a pitching rate of 99% sensitive ale strain to 1% killer lager strain, indicating that even a small amount of killer cells is enough to kill a larger population of sensitive cells <ref name="Stewart_2018" />. Neutral strains do not produce toxins, nor are they killed by them <ref>[https://books.google.com/books?hl=en&lr=&id=mvORN6OXHh4C&oi=fnd&pg=PA93&dq=Bussey,+H.+1981.+Physiology+of+killer+factor+in+yeast.+Adv.+Microb.+Physiol.+22:93-121&ots=jUY4T9NpgB&sig=aw-Y1um0KsDnGe6rRe5PTWIDYdI#v=onepage&q&f=false Advances in Microbial Physiology, Volume 22. Academic Press, Sep 15, 1981. Pg 94-95.]</ref>. Almost all ale and lager strains are sensitive to the toxins produced by killer strains <ref>[https://www.facebook.com/groups/MilkTheFunk/permalink/1179271825434363/?comment_id=1179424538752425&offset=0&total_comments=5&comment_tracking=%7B%22tn%22%3A%22R%22%7D Conversation with Bryan of Sui Generis Blog on MTF on Killer Factor for Saccharomyces. 11/16/2015.]</ref><ref>[http://onlinelibrary.wiley.com/doi/10.1002/j.2050-0416.1973.tb03515.x/pdf Strains of Yeast Lethal to Brewery Yeasts. A.P. Maule and P.D. Thomas. 1972.]</ref>. In ''Saccharomyces'', four toxins have been identified: K1, K2, K28, and Klus, the first three of which can only kill other strains/species of ''Saccharomyces''. The Klus toxin has been found to kill yeast from other genra, such as ''Hanseniaspora spp.'', ''Kluyveromyces lactis'', ''Candida albicans'', ''Candida dubliniensis'', ''Candida kefir'' and ''Candida tropicalis'', and the K1, K2 and K28 killer strains of ''S. cerevisiae'' <ref name="Rodriguez">[http://aem.asm.org/content/77/5/1822.long A New Wine Saccharomyces cerevisiae Killer Toxin (Klus), Encoded by a Double-Stranded RNA Virus, with Broad Antifungal Activity Is Evolutionarily Related to a Chromosomal Host Gene. Nieves Rodríguez-Cousiño, Matilde Maqueda, Jesús Ambrona, Emiliano Zamora, Rosa Esteban and Manuel Ramírez. 2011]</ref>. However, none of the toxins secreted by killer strains of ''Saccharomyces'' have been found to kill ''Brettanomyces'' <ref>[http://www.scielo.org.za/scielo.php?pid=S2224-79042015000100010&script=sci_arttext&tlng=pt Non-Saccharomyces killer toxins: Possible biocontrol agents against Brettanomyces in wine? S. Afr. J. Enol. Vitic. vol.36 n.1 Stellenbosch. 2015.]</ref>. The K1 toxin is most active between a pH of 4.6 and 4.8, while K2 and Klus are active around a pH of 4.0 to 4.3 <ref name="Rodriguez"></ref>. The activity of the toxin is greatest during the log phase of growth, and decays during the stationary phase of fermentation <ref name="Buyuksirit"></ref>.

In ''Saccharomyces'', four toxins have been identified: K1, K2, K28, and Klus, the first three of which can only kill other strains/species of ''Saccharomyces''. The Klus toxin has been found to kill all strains of ''S. cerevisiae'' (including those that produce the previous three toxins), as well as yeast from other genera, such as ''Hanseniaspora spp.'', ''Kluyveromyces lactis'', ''Candida albicans'', ''Candida dubliniensis'', ''Candida kefir'' and ''Candida tropicalis'' <ref name="Rodriguez">[http://aem.asm.org/content/77/5/1822.long A New Wine Saccharomyces cerevisiae Killer Toxin (Klus), Encoded by a Double-Stranded RNA Virus, with Broad Antifungal Activity Is Evolutionarily Related to a Chromosomal Host Gene. Nieves Rodríguez-Cousiño, Matilde Maqueda, Jesús Ambrona, Emiliano Zamora, Rosa Esteban and Manuel Ramírez. 2011]</ref>. However, none of the toxins secreted by killer strains of ''Saccharomyces'' have been found to kill ''Brettanomyces'' <ref>[http://www.scielo.org.za/scielo.php?pid=S2224-79042015000100010&script=sci_arttext&tlng=pt Non-Saccharomyces killer toxins: Possible biocontrol agents against Brettanomyces in wine? S. Afr. J. Enol. Vitic. vol.36 n.1 Stellenbosch. 2015.]</ref>. The K1 toxin is most active between a pH of 4.6 and 4.8, while K2 and Klus are active around a pH of 4.0 to 4.3 <ref name="Rodriguez"></ref>. The activity of the toxin is greatest during the log phase of growth, and decays during the stationary phase of fermentation <ref name="Buyuksirit"></ref>. Various other yeast species have the ability to produce toxins that effect a range of other yeasts and bacteria , including species from the genera ''Candida'', ''Cryptococcus'', ''Debaryomyces'', ''Hanseniaspora'', ''Hansenula'', ''Kluyveromyces'', ''Metschnikowia'', ''Pichia'', ''Ustilago'', ''Torulopsis'', ''Williopsis'', ''Zygosaccharomyces'', ''Aureobasidium'', ''Zygowilliopsis'', and ''Mrakia'' <ref name="Buyuksirit">[http://waset.org/publications/9999528/antimicrobial-agents-produced-by-yeasts Antimicrobial Agents Produced by Yeasts. T. Buyuksirit, H. Kuleasan. 2014.]</ref><ref name="Stewart_2018" />. For example, strains of the yeast species ''Candida pyralidae'' <ref name="Buyuksirit"></ref>, ''Wickerhamomyces anomalus'', ''Kluyveromyces wickeramii'', ''Torulaspora delbrueckii'' and ''Pichia membranifaciens'' have been found to produce toxin that inhibits ''Brettanomyces'' <ref name="Ciani_2016">[https://www.researchgate.net/publication/301581233_Yeast_Interactions_in_Inoculated_Wine_Fermentation Yeast Interactions in Inoculated Wine Fermentation. Maurizio Ciani, Angela Capece, Francesca Comitini, Laura Canonico, Gabriella Siesto and Patrizia Romano. 2016.]</ref>. In addition, the toxin produced by ''Wickerhamomyces anomalus'' and ''Williopsis markii'' have been found to inhibit a wide range of spoilage and pathogenic fungi <ref name="Hatoum2012"></ref>. Killer strains of ''S. cerevisiae'' and other yeast can occur naturally in the wild on fruit and can have a negative impact on other flora that are found in the same environment <ref name="Buyuksirit"></ref>. Strains of ''Torulaspora delbrueckii'' have been shown to kill killer strains of ''S. cerevisae'' (wine strains), as well as to kill ''Pichia'' species <ref name="Ciani_2016"></ref>.