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update to Rpoiness
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. When a beta-glucosidase enzyme was added to help break down the EPS produced by the bacteria, the lysozyme 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 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>