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added new study on K1-like toxin
Several strains of ''Saccharomyces eubayanus'' isolated from seeds from monkey puzzle trees in Patagonia, Argentina, were found to secrete a killer toxin that kills ''Brettanomyces'' and ''Pichia''. One strain was found to produce a lot of the toxin, which is called "SeKT". ''S. cerervisiae'' strains, including strains that are sensitive to the above toxins, are not sensitive to this toxin. Mazzucco et al. (2019) found that SeKT toxin produced by this one strain of ''S. eubaynus'' in a special growth medium designed to maximize the SeKT toxin production (WUJ medium, which is "ultrafiltered" apple and pear juice) inhibited a strain of ''B. bruxellensis'' to around 50% growth after 48 hours in a wine growth medium. It also inhibited ''Pichia guilliermondii'', ''Pichia manshurica'', and ''Pichia membranifaciens'' by 50-70%. Note that the toxin was applied directly to the ''Brettanomyces'' and ''Pichia'' species, and not in a co-fermentation setting. Since ''S. cerevisiae'' strains are not effected by the SeKT toxin, it has been proposed as a way to limit ''Brettanomyces'' and ''Pichia'' in wine fermentations <ref>[https://www.ncbi.nlm.nih.gov/pubmed/30671692?dopt=Abstract Production of a novel killer toxin from Saccharomyces eubayanus using agro-industrial waste and its application against wine spoilage yeasts. Mazzucco MB, Ganga MA, Sangorrín MP. 2019. DOI: 10.1007/s10482-019-01231-5.]</ref>.
A newly discovered toxin that is related to the K1 toxin, called "K1-like" or K1L, has been identified in ''Saccharomyces paradoxus''. The ability for this species to produce this toxin is caused by a virus that binds the the DNA of the yeast cells, and spread via horizontal gene transfer. The K1L toxin has a pH optimum mostly between 4.5 and 5, with no inhibitory activity at pH 5.5. It is denatured at a temperature of 98°C. A screening of this genetic change, called “K1-like Killer Toxin” (KKT) genes, in other yeasts showed that many other species can also produce toxins similar to the K1L toxin but slightly different in effect, including ''Kazachstania africana'', ''Naumovozyma castellii'', ''Naumovozyma dairenensis'', ''Tetrapisispora phaffii'', and ''Pichia membranifaciens''. Each of the identified species could kill at least one other type of yeast with its toxin, and was immune to its own toxin, but susceptable to other K1-like toxins from other yeast species. Differences in the production of these K1-like toxins between 5 different strains of ''P. membranifaciens'' indicated that the toxins can be strain-specific, rather than species-specific. Using the genetic relatedness between the different KKT genes, the researchers concluded that this family of K1-like toxins originated outside of the ''Saccharomyces'' genus. This research uncovered a new family of K1-like antifungal killer toxins amoung many species of yeast in the Saccharomycotina subphylum <ref>[https://journals.plos.org/plosgenetics/article?id=10.1371%2Fjournal.pgen.1009341 Fredericks LR, Lee MD, Crabtree AM, Boyer JM, Kizer EA, Taggart NT, et al. (2021) The Species-Specific Acquisition and Diversification of a K1-like Family of Killer Toxins in Budding Yeasts of the Saccharomycotina. PLoS Genet 17(2): e1009341. https://doi.org/10.1371/journal.pgen.1009341]</ref>.
Various other yeast species have the ability to produce toxins that effect a range of other yeasts (but generally not 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>. The occurrence of killer strains of yeast in the wild is also wide spread. For example, out of 210 yeasts from various genera isolated from molasses, 13 of them were killer strains. Out of 1,000 isolates of various ''Candida'' species isolated from human skin, 52 were killer strains. Out of 65 strains of various yeasts isolated from fermented foods, soil samples, and spoiled fruits/vegetables, 12 were killer strains <ref name="Bajaj_2017" />. It has been hypothesized that toxin production is ubiquitous throughout nearly all genera of yeast; the more studies that have been done on a particular genus of yeast, the more likely it is that toxin production has been found by species and strains within that genus. Yeasts that produce toxins have been found on every continent and in every natural habitat of yeast, including leaf surfaces, leaf litter, tree slime fluxes, fruits, cactus stems and cladodes, insect guts, mammal feces, leaf-cutting ant nests, lake water, ocean sediment, soil, wine, bakeries, and dairy products <ref name="Boynton_2019" />.