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A '''pellicle''' is an aggregation of cells, proteins, and long chain sugars (polysaccharides) formed on the surface of a beer. The "bubble" formations are caused by trapped CO2 beneath the pellicle film. Pellicles are often formed by [[Brettanomyces]], [[Pediococcus]], and [[Lactobacillus]], as well as ''Acetobacter spp'' <ref name="Perumpuli">[http://ir.kagoshima-u.ac.jp/bitstream/10232/21479/4/Diss_Perumpuli+Arachchige_Buddhika+Niroshie_RNK999_2014.pdf.pdf Pellicle of thermotolerant Acetobacter pasteurianus strains: Characterization of polysaccharide and induction patterns. Perumpuli Arachchige Buddhika Niroshie. 2014-09-30.]</ref> and other gram-negative bacteria <ref name="Armitano">[http://www.ncbi.nlm.nih.gov/pubmed/25756106 Gram-negative bacteria can also form pellicles. Armitano J, Méjean V, Jourlin-Castelli C. Environ Microbiol Rep. 2014 Dec.]</ref>. It can also be formed by [[Saccharomyces]] in rare occasions <ref name="matt">[http://phdinbeer.com/2015/01/30/beer-microbiology-what-is-a-pellicle/ Beer Microbiology – What is a pellicle? A PhD in Beer blog. Dr. Matt Humbard. 01/30/2015. Retrieved 04/26/2015.]</ref>. Pellicles Note that "pellicles" in brewing have not been scientifically defined. See [[Pellicle#Scientific_Terminology|Scientific Terminology]] for more information.