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[[File:Pedio.jpg|thumb|200px|right|[https://www.instagram.com/wildandsour/ Pediococcus - picture taken by Per Karlsson]]]
'''''Pediococcus''''' (often referred to by brewers as "Pedio") are Gram-positive lactic acid bacteria (LAB) used in the production of Belgian style beers where additional acidity is desirable. They are native to plant material and fruits <ref name="ucdavis">[http://wineserver.ucdavis.edu/industry/enology/winemicro/winebacteria/pediococcus_damnosus.html Viticulture & Enology. UC Davis website. Pedioccous damnosus. Retrieved 07/28/2015.]</ref>, and often found in [[Spontaneous_Fermentation|spontaneously fermented]] beer as the primary source of lactic acid production (with ''P. damnosus'' being the only species identified in [[Lambic]]) <ref>[http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0095384 The Microbial Diversity of Traditional Spontaneously Fermented Lambic Beer. Freek Spitaels, Anneleen D. Wieme, Maarten Janssens, Maarten Aerts, Heide-Marie Daniel, Anita Van Landschoot, Luc De Vuyst, Peter Vandamme. April 18, 2014.]</ref><ref>[[Scientific_Publications#Lambic_and_Spontaneous_Fermentation|Multiple Scientific publications linked on MTF.]]</ref>. It is also seen as a major source of beer contamination in commercial breweries due to its ability to adapt to and survive in beer. The ability to grow in beer is strain dependent rather than species dependent, however, genetic differences indicate that ''P. damnosus'' and ''P. claussenii'' are better adapted to surviving in beer than ''P. pentosaceus'' <ref name="Snauwaert">[http://www.biomedcentral.com/content/pdf/s12864-015-1438-z.pdf Comparative genome analysis of Pediococcus damnosus LMG 28219, a strain well-adapted to the beer environment. Isabel Snauwaert, Pieter Stragier, Luc De Vuyst and Peter Vandamme. 2015.]</ref>. Like many bacteria, pediococci have the ability to [https://en.wikipedia.org/wiki/Horizontal_gene_transfer transfer genes horizontally] without reproduction <ref name="Snauwaert"></ref>. They are generally considered to be facultative anaerobes, which means they grow anaerobically but can also grow in the presence of oxygen <ref>[http://textbookofbacteriology.net/lactics.html Lactic Acid Bacteria. Todar's Online Texbook of Bacteriology. Kenneth Todar, PhD. Pg 1. Retrieved 08/09/2015.]</ref>. Some species/strains (including individual strains of ''P. damnosus'') can have their growth and acid production inhibited by oxygen <ref name="NAKAGAWA">[https://www.jstage.jst.go.jp/article/jgam1955/5/3/5_3_95/_article TAXONOMIC STUDIES ON THE GENUS PEDIOCOCCUS. ATSUSHI NAKAGAWA, KAKUO KITAHARA. 1959.]</ref>, while some will have better growth and produce more acid in the presence of oxygen (microaerophilic) <ref>[http://www.ncbi.nlm.nih.gov/pmc/articles/PMC357257/ THE NUTRITION AND PHYSIOLOGY OF THE GENUS PEDIOCOCCUS. Erling M. Jensen and Harry W. Seeley. 1954.]</ref><ref>[http://www.microbialcellfactories.com/content/8/1/3#B5 Pediocins: The bacteriocins of pediococci. Sources, production, properties and applications. Maria Papagianni and Sofia Anastasiadou. 2009.]</ref>. Strains found in beer are hop tolerant <ref>[http://www.biomedcentral.com/1471-2164/16/267 Comparative genome analysis of Pediococcus damnosus LMG 28219, a strain well-adapted to the beer environment. Isabel Snauwaert, Pieter Stragier, Luc De Vuyst and Peter Vandamme. April 2015.]</ref>. Due to their continued metabolism of longer chain polysaccharides, acid production will increase with storage time. ''Pediococcus'' can form a [[pellicle]].
''Pediococcus'' may also cause “ropiness” (also called a "sick beer") due to the production of exopolysaccharides when exposed to a fresh sugar source. "Ropy" or "sick" beer is more viscous and, in extreme circumstances, can form strands. Sickness effects mostly the mouthfeel and appearance of the beer, and may have no influence on the flavor. It is considered a temporary flaw in sour beer. Some brewers, including Vinnie Cilurzo from Russian River Brewing and some Belgian lambic producers, claim that after the ropiness goes away (generally in 3-6 months <ref name="ropy_time"></ref>) it produces a deeper acidity and mouthfeel, and is viewed as a positive process in the production of sour beer <ref>[http://www.xxlbrewing.com/hb/sour_beer/img_09.html Cilurzo, Vinnie. AHA Sour Beer presentation. 2007.]</ref>. For other brewers, ropy beer is seen as a nuisance due to the beer needing to be aged for a longer period of time, especially when it occurs shortly after bottling. ''Pediococcus'' species can also produce diacetyl with extended storage time <ref name="Garcia-Garcia" />. ''[[Brettanomyces]]'' can break down exopolysaccharides and diacetyl produced by ''Pediococcus'' and the two are often used together.
! Name !! Mfg# !! Taxonomy !! Note
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| [[Bootleg Biology]]/[[Spot Yeast]] || Sour Weapon P (''Pediococcus pentosaceus'' Blend) || ''P. pentosaceus'' blend || Perfect for acidifying unhopped wort quickly for kettle or “quick” sours. At 98F, it’s capable of achieving a pH of 3.3 within 18 hours. At 84F, it can reach a pH of 3.5 within 24 hours. With more time, a terminal pH of 3.1 may be reached. ''P. pentosaceus'' can also be used for long-term sours. It is capable of growing and producing lactic acid in worts with IBUs as high as 30, though it is recommended for unhopped worts as IBUs over 10 may prevent significant quick souring. At ~30 IBU, souring occurs in 2-3 months <ref>[https://www.facebook.com/groups/MilkTheFunk/permalink/1302981419730069/?comment_id=1870460696315469&comment_tracking=%7B%22tn%22%3A%22R0%22%7D Justin Amaral and Per Karlsson on Bootleg biology Sour Weapon hop tolerance. milk The Funk Facebook group. 11/2/2017.]</ref>. This culture may produce antimicrobials called bacteriocins or pediocins. These can inhibit and kill similar species of bacteria like Lactobacillus and other Pediococcus species in mixed-culture fermentations. Read Bootleg Biology's [https://www.facebook.com/BootlegBiology/photos/a.148869931970401.1073741829.124634287727299/465185997005458/?type=1&theater Facebook post] regarding bacteriocins for more info. No signs of ropiness (exopolysaccharides) have occurred in testing <ref>[http://bootlegbiology.com/product/sour-weapon-pediococcus-pentosaceus-blend/ Bootleg Biology website. Retrieved 05/06/2016.]</ref>. It is still unknown how hops will affect souring in a long term scenario. Bootleg Biology is still researching long term effects and awaiting peoples feedback as of 5/23/2016.
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| [[East Coast Yeast]] || ECY33 || ''P. parvulus'' || Isolated from lambic which was refermented with grapes, this strain of ''Pediococcus'' produces lactic acid, diacetyl, and may cause ropiness in beer. Always add ''Brettanomyces'' where ''Pediococcus'' is used <ref name="ecy_website">[http://www.eastcoastyeast.com/wild-stuff.html "Wild Yeast / Brettanomyces / Lactic Bacteria". East Coast Yeast website. Retrieved 04/27/2018.]</ref>.
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" />.
The presence of beta-glucans from barley have been observed to extend both the growth and the viability of ''Lactobacillus'' species in probiotics, indicating that ropiness might be a stress response <ref>[http://www.mdpi.com/1422-0067/15/2/3025/htm Barley β-Glucans-Containing Food Enhances Probiotic Performances of Beneficial Bacteria. Mattia P. Arena, Graziano Caggianiello, Daniela Fiocco, Pasquale Russo, Michele Torelli, Giuseppe Spano, and Vittorio Capozzi. 2014.]</ref><ref>[http://www.mdpi.com/1422-0067/13/5/6026/htm Beta-Glucans Improve Growth, Viability and Colonization of Probiotic Microorganisms. Pasquale Russo, Paloma López, Vittorio Capozzi, Pilar Fernández de Palencia, María Teresa Dueñas, Giuseppe Spano, and Daniela Fiocco. 2012.]</ref>. One study looked at this effect in beta-glucans produced by ''Pediococcus parvulus'' and found that ''L. plantarum'' had a longer viability in a fermented medium with no additional food source when that medium was first fermented with ''P. parvulus'' and EPS was produced. The ''L. plantarum'' strain that was tested did not ferment the beta-glucans. This suggests that there is an interspecies simbiotic relationship between lactic acid bacteria that produce EPS and those that don't, and when EPS is produced (beta-glucans are present) the bacteria survive longer. The study also observed that more EPS was produced in an oat based wort and a rice based wort, while no EPS was produced in a barley based wort, suggesting that different food sources influence whether or not EPS is produced <ref>[http://www.mdpi.com/1422-0067/18/7/1588/htm In Situ β-Glucan Fortification of Cereal-Based Matrices by Pediococcus parvulus. Adrián Pérez-Ramos, María Luz Mohedano, Paloma López, Giuseppe Spano, Daniela Fiocco, Pasquale Russo, and Vittorio Capozzi. 2017.]</ref>. Pittet et al. (2011) found that the presence of the EPS genes in lactic acid bacteria did not correspond with the ability to survive beer-level ethanol levels or higher, which led to the hypothesis that perhaps EPS provides LAB a way to assist in the formation of [[Quality_Assurance#Biofilms|biofilm]], although this has yet to be demonstrated scientifically <ref>[https://www.tandfonline.com/doi/abs/10.1094/ASBCJ-2011-0124-01 Ethanol Tolerance of Lactic Acid Bacteria, Including Relevance of the Exopolysaccharide Gene Gtf. Vanessa Pittet and Kendra Morrow and Barry Ziola. 2011. DOI: https://doi.org/10.1094/ASBCJ-2011-0124-01.]</ref>. Along these same lines, [https://www.academia.edu/27927065/Lysozyme_resistance_of_the_ropy_strain_Pediococcus_parvulus_IOEB_8801_is_correlated_with_beta_glucan_accumulation_around_the_cell Coulon et al. (2012)] reported that an EPS producing strain of ''Pediococcus parvulus'' was more resistant to the antimicrobial enzyme [https://en.wikipedia.org/wiki/Lysozyme lysozyme], which is often added to wine to kill lactic acid bacteria. The beta-glucan EPS formed a "coat" around the cells of the bacteria, thus protecting it from the lysozyme enzyme. When a beta-glucosidase glucanase enzyme was added to help break down the EPS produced by the bacteria, the beta-glucan "coat" disappeared from the cell walls and the lysozyme was once again effective at killing this strain of ''Pediococcus'' <ref>[https://www.academia.edu/27927065/Lysozyme_resistance_of_the_ropy_strain_Pediococcus_parvulus_IOEB_8801_is_correlated_with_beta_glucan_accumulation_around_the_cell Coulon, Joana et al. “Lysozyme Resistance of the Ropy Strain Pediococcus Parvulus IOEB 8801 Is Correlated with Beta-Glucan Accumulation around the Cell.” International Journal of Food Microbiology 159.1 (2012): 25–29. Web.]</ref>.
It has been observed that ''Lactobacillus'' species can produce EPS (''Lactococcus lactis'', ''Lactobacillus delbrueckii'', ''Lactobacillus casei'', and ''Lactobacillus helveticus'') <ref name="ESP"></ref>. Some ''Oenococcus oeni'' strains can also produce EPS <ref>[https://pubmed.ncbi.nlm.nih.gov/19659698/ Ciezack G, Hazo L, Chambat G, Heyraud A, Lonvaud-Funel A, Dols-Lafargue M. Evidence for exopolysaccharide production by Oenococcus oeni strains isolated from non-ropy wines. J Appl Microbiol. 2010 Feb;108(2):499-509. doi: 10.1111/j.1365-2672.2009.04449.x. Epub 2009 Jun 30. PMID: 19659698.</ref>. Some species of yeast can also produce EPS, including ''Candida'', ''Cryptococcus'', ''Debaryomyces'', ''Lipomyces'', ''Pichia'' <ref>[https://www.facebook.com/groups/MilkTheFunk/permalink/2465980936763439/ Zach Taggart. Milk the Funk Facebook group post on EPS from yeast. 01/16/2019.]</ref>, ''Pseudozyma'', ''Rhodotorula'' and ''Sporobolomyces'' <ref name="Gientka_2015">[https://www.researchgate.net/publication/283498621_Exopolysaccharides_from_yeast_insight_into_optimal_conditions_for_biosynthesis_chemical_composition_and_functional_properties_-_review?fbclid=IwAR1X6Y0rnquoF6SD-eH9m6EWpLIefgZJFUJK51NJYBooJWngxEVS2aR3PKE Exopolysaccharides from yeast: insight into optimal conditions for biosynthesis, chemical composition and functional properties - review. Iwona Gientka, Stanisław Błażejak, Stanisław Błażejak, Lidia Stasiak, Lidia Stasiak, Anna Chlebowska-Śmigiel, Anna Chlebowska-Śmigiel. 2015.]</ref>.
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