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Pediococcus

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This "ropiness" is caused by production of exopolysaccharides (EPS) in the form of β-glucans (beta glucans) by some strains ''Pediococcus'' and some other lactic acid bacteria species. The β-glucans are made up of beta 1, 3 linkages and beta 1, 2 branches composed of single units <ref name="Wade_2018" />. A small amount (20-30 mg/L <ref name="Wade_2018" />) of β-glucan is adequate enough to affect the visible viscosity of beer or wine. The gene known as "dps" has been identified with the production of β-glucan/EPS in ''P. damnosus'', and the gene "gtf" in ''P. claussenii'' <ref name="Snauwaert"></ref>. Not all strains of ''P. damnosus'' express the gene, and only ones that do will cause a beer to go ropy. Although it is not needed to survive in beer, EPS production is probably has importance in biofilm production <ref>[http://cat.inist.fr/?aModele=afficheN&cpsidt=23890699 Ethanol tolerance of lactic acid bacteria, including relevance of the exopolysaccharide gene gtf. Pittet V, Morrow K, Ziola B. 2011.]</ref>, and ''Pediococci'' that are ropy have been found to be more acid, alcohol, and SO2 tolerant than other ''Pediococci''. The thickness of the ropiness is increased with the presence of malic acid <ref name="ESP"></ref>. While strains of ''P. damnosus'' and ''P. parvulus'' are the ''Pediococcus'' species most associated with ropiness, some strains of ''P. pentosaceus'' have also been found to produce EPS <ref name="Wade_2018" />.
One study showed that the production of β-glucan coincided with the end of the growth phase of ''Pediococcus''. While small amounts of β-glucan were produced during growth, after 2 days of growth, β-glucan production increased as growth slowed. β-glucan production stopped when growth stopped. This study showed that β-glucan production is linked to ''Pediococcous'' growth, producing more towards the end of growth. This would explain why beer containing ''Pediococcus'' often goes ropy shortly after naturally carbonating in the bottle. This study found that other variables were not factors in the production of β-glucan, such as differing levels of alcohol (although alcohol interacts with the β-glucan in a way that makes the viscosity seem thicker). The study also found that the lack of agitation increased the β-glucan production (wine makers will often agitate or aerate ropy wine to cure the wine from ropiness). A higher initial pH encourages higher growth (5.5+), which increases β-glucan production. A lower initial pH (3.5), decreases growth and β-glucan production. A higher concentration of glucose increased growth and β-glucan production. Glucose is needed for β-glucan production. While fructose alone is mostly insufficient to produce ropiness, a combination of glucose and fructose was slightly more efficient than glucose alone <ref name="ESP"></ref>. The introduction of malic acid, glucose, fructose, and/or nitrogen from things like fruit that is added to sour beer or even sugar added for natural carbonation can trigger ''Pediococcus'' growth and EPS production.
Temperature and nitrogen levels also affect how much EPS is produced. One study found that at 12°C both growth and EPS production was much slower than at 25°C. After 29 days in agar media, the EPS in the 12°C samples tended to reach or slightly exceed the levels in the 25°C samples, which developed equivilant levels of EPS (or slightly less) within 7-13 days. Nitrogen levels also play a significant role, according to this study, particularly at lower fermentation temperatures. At 12°C, nitrogen was more important for the formation of EPS than glucose (although glucose was found to be the most important factor in EPS development overall, which is in agreement with the previously cited study). At 25°C nitrogen levels played a significant role in producing EPS, however less so than glucose levels. In general though, higher availability of nitrogen complimented higher levels of glucose to produce more EPS (and faster/higher cell growth) <ref>[https://www.sciencedirect.com/science/article/pii/S0168160503000606 Exopolysaccharide production by Pediococcus damnosus 2.6 in a semidefined medium under different growth conditions. Maite Dueñas, Arantza Munduate, Aidé Perea, Ana Irastorza. 2003.]</ref>. Stress in the environment such as ethanol and SO<sub>2</sub> has also been shown to induce EPS production, and a lack of available glucose has been associated with eliminating the production of EPS (e.g. malolactic fermentation by ''Pediococcus'' in wine tends not to produce EPS, perhaps due to the lack of glucose in the environment) <ref name="Wade_2018" />.

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