Difference between revisions of "Pellicle"

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[[File:Brussels brett blend pellicle1.jpg|thumb|Pellicle from The Yeast Bay Brussels Brett Blend; courtesy of Dan Pixley]][[File:Lactobacillus pellicle.jpg|thumb|Lactobacillus pellicle; courtesy of Matt Humbard]] A '''pellicle''' is a subtype of biofilm that is formed on the surface of a beer.  A biofilm is a community of microorganisms that are connected to one another through the use of material that is present in the environment, such as proteins, long chains of sugars, or live (or dead) cells. It is often formed by [[Brettanomyces]], [[Pediococcus]], or [[Lactobacillus]].  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>.  
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[[File:Brussels brett blend pellicle1.jpg|thumb|Pellicle from The Yeast Bay Brussels Brett Blend; courtesy of Dan Pixley]][[File:Lactobacillus pellicle.jpg|thumb|Lactobacillus pellicle; courtesy of Matt Humbard]] A '''pellicle''' is an aggregation of cells, proteins, and long chain sugars formed on the surface of a beer.  It is often formed by [[Brettanomyces]], [[Pediococcus]], or [[Lactobacillus]].  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>.  
  
 
==Introduction==
 
==Introduction==
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(This section in progress)
 
(This section in progress)
  
As with some things in science that are not greatly explored, terminology isn't always agreed upon or fully established, and thus researching such a topic without a lot of prior knowledge can be challenging.  Pellicles are a good example of this.  It's been stated on this wiki and elsewhere that a pellicle is a subtype of biofilm.  However, the widely accepted definition of a "biofilm" in biology doesn't describe pellicles that we see in fermentation.  The widely accepted definition of a "biofilm" is an aggregate of microorganisms where the cells adhere to each other on a surface <ref>[https://books.google.com/books?id=-ZDKAQAAQBAJ&pg=PT87&lpg=PT87&dq=brock+microbiology+biofilm&source=bl&ots=PwNZ20KFqq&sig=19lvlfqYj8bE7roiA7g4oJ3Xl0E&hl=en&sa=X&ved=0CCcQ6AEwAWoVChMImZaUmN_AxwIVVFmICh1pQgk0#v=onepage&q=brock%20microbiology%20biofilm&f=false e-Study Guide for Brock Biology of Microorganisms, textbook by Michael T. Madigan.]</ref>.  Pellicles in beer do not attach to a solid surface, they appear on the "air-liquid interface" (the surface of the beer).  They are also not colonies within an adhesive <ref name="Bryan">[https://www.facebook.com/groups/MilkTheFunk/permalink/1131334646894748/?comment_id=1131986723496207&offset=0&total_comments=57&comment_tracking=%7B%22tn%22%3A%22R3%22%7D Conversation with Bryan Heit about Pellicles on MTF.  08/20/2015.]  </ref>.  To make matters even more confusing, the two widely accepted definitions of a "pellicle" in biology are the outer boundary of a protozoa cell <ref>[https://books.google.com/books?id=ACBcDjDLuNAC&pg=PA38&lpg=PA38&dq=pellicle+biology&source=bl&ots=58e43A_9h-&sig=ViakT4Ectu4DOlqoSWRCPFJ73iw&hl=en&sa=X&sqi=2&ved=0CGIQ6AEwDGoVChMIp6WkouTAxwIVgpmICh0F1wXO#v=onepage&q=pellicle%20biology&f=false Biology of Protozoa.  D.R. Khanna.  Discovery Publishing House, Jan 1, 2004.  Pg 38.]</ref>, and the protein film that forms on the surface of teeth <ref>[https://en.wikipedia.org/wiki/Dental_pellicle Wikipedia.  Dental Pellicle.  Retrieved 08/23/2015.]</ref>.
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As with some things in science that are not greatly explored, terminology isn't always agreed upon or fully established, and thus researching such a topic without a lot of prior knowledge can be challenging.  Pellicles are a good example of this.  It is sometimes stated that a pellicle is a subtype of biofilm.  However, the widely accepted definition of a "biofilm" in biology doesn't describe pellicles that we see in fermentation.  The widely accepted definition of a "biofilm" is an aggregate of microorganisms where the cells adhere to each other on a surface <ref>[https://books.google.com/books?id=-ZDKAQAAQBAJ&pg=PT87&lpg=PT87&dq=brock+microbiology+biofilm&source=bl&ots=PwNZ20KFqq&sig=19lvlfqYj8bE7roiA7g4oJ3Xl0E&hl=en&sa=X&ved=0CCcQ6AEwAWoVChMImZaUmN_AxwIVVFmICh1pQgk0#v=onepage&q=brock%20microbiology%20biofilm&f=false e-Study Guide for Brock Biology of Microorganisms, textbook by Michael T. Madigan.]</ref>.  Pellicles in beer do not attach to a solid surface, they appear on the "air-liquid interface" (the surface of the beer).  They are also not colonies within an adhesive <ref name="Bryan">[https://www.facebook.com/groups/MilkTheFunk/permalink/1131334646894748/?comment_id=1131986723496207&offset=0&total_comments=57&comment_tracking=%7B%22tn%22%3A%22R3%22%7D Conversation with Bryan Heit about Pellicles on MTF.  08/20/2015.]  </ref>.  To make matters even more confusing, the two widely accepted definitions of a "pellicle" in biology are the outer boundary of a protozoa cell <ref>[https://books.google.com/books?id=ACBcDjDLuNAC&pg=PA38&lpg=PA38&dq=pellicle+biology&source=bl&ots=58e43A_9h-&sig=ViakT4Ectu4DOlqoSWRCPFJ73iw&hl=en&sa=X&sqi=2&ved=0CGIQ6AEwDGoVChMIp6WkouTAxwIVgpmICh0F1wXO#v=onepage&q=pellicle%20biology&f=false Biology of Protozoa.  D.R. Khanna.  Discovery Publishing House, Jan 1, 2004.  Pg 38.]</ref>, and the protein film that forms on the surface of teeth <ref>[https://en.wikipedia.org/wiki/Dental_pellicle Wikipedia.  Dental Pellicle.  Retrieved 08/23/2015.]</ref>.
  
 
The usage of the term "biofilm" has been used to describe the layer of film that covers sherry known as "Flor" <ref>[http://aem.asm.org/content/76/12/4089.full Ethanol-Independent Biofilm Formation by a Flor Wine Yeast Strain of Saccharomyces cerevisiae.  Severino Zara, Michael K. Gross, Giacomo Zara, Marilena Budroni and Alan T. Bakalinsky.  2010.]</ref><ref>[http://femsle.oxfordjournals.org/content/femsle/237/2/425.full.pdf FLO11 is essential for flor formation caused by the C-terminal deletion of NRG1 in Saccharomyces cerevisiae.  Mari Ishigami, Youji Nakagawa, Masayuki Hayakawa, Yuzuru Iimura.  2004.]</ref>.  The word "pellicle" generally isn't used, although it has appeared on occasion when referring to sherry flor in the 1960's <ref>[http://www.ncbi.nlm.nih.gov/pubmed/5311988 On the pellicle formation by “flor” yeasts. Cantarelli C, Martini A.  Antonie Van Leeuwenhoek. 1969.]</ref>.  More recently, there have been studies that define a "pellicle" like we see in fermentation as a type of biofilm that forms on the air-liquid interface of a liquid (see references) <ref>[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><ref>[http://www.ncbi.nlm.nih.gov/pubmed/26047954 Identification of genes essential for pellicle formation in Acinetobacter baumannii.  Giles SK, Stroeher UH, Eijkelkamp BA, Brown MH.  BMC Microbiol. 2015 Jun 6.]</ref><ref>[http://www.ncbi.nlm.nih.gov/pubmed/26122431 Motility, Chemotaxis and Aerotaxis Contribute to Competitiveness during Bacterial Pellicle Biofilm Development.  Hölscher T, Bartels B, Lin YC, Gallegos-Monterrosa R, Price-Whelan A, Kolter R, Dietrich LE, Kovács ÁT.  J Mol Biol. 2015 Jun 26.]</ref><ref>[http://www.ncbi.nlm.nih.gov/pubmed/24035282 Modulation of curli assembly and pellicle biofilm formation by chemical and protein chaperones.  Andersson EK, Bengtsson C, Evans ML, Chorell E, Sellstedt M, Lindgren AE, Hufnagel DA, Bhattacharya M, Tessier PM, Wittung-Stafshede P, Almqvist F, Chapman MR.  Chem Biol. 2013 Oct 24.]</ref>.  Although these studies don't directly look at beer pellicles, this indicates that the definition of a pellicle as stated in this wiki may become more widely accepted in the scientific community.  
 
The usage of the term "biofilm" has been used to describe the layer of film that covers sherry known as "Flor" <ref>[http://aem.asm.org/content/76/12/4089.full Ethanol-Independent Biofilm Formation by a Flor Wine Yeast Strain of Saccharomyces cerevisiae.  Severino Zara, Michael K. Gross, Giacomo Zara, Marilena Budroni and Alan T. Bakalinsky.  2010.]</ref><ref>[http://femsle.oxfordjournals.org/content/femsle/237/2/425.full.pdf FLO11 is essential for flor formation caused by the C-terminal deletion of NRG1 in Saccharomyces cerevisiae.  Mari Ishigami, Youji Nakagawa, Masayuki Hayakawa, Yuzuru Iimura.  2004.]</ref>.  The word "pellicle" generally isn't used, although it has appeared on occasion when referring to sherry flor in the 1960's <ref>[http://www.ncbi.nlm.nih.gov/pubmed/5311988 On the pellicle formation by “flor” yeasts. Cantarelli C, Martini A.  Antonie Van Leeuwenhoek. 1969.]</ref>.  More recently, there have been studies that define a "pellicle" like we see in fermentation as a type of biofilm that forms on the air-liquid interface of a liquid (see references) <ref>[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><ref>[http://www.ncbi.nlm.nih.gov/pubmed/26047954 Identification of genes essential for pellicle formation in Acinetobacter baumannii.  Giles SK, Stroeher UH, Eijkelkamp BA, Brown MH.  BMC Microbiol. 2015 Jun 6.]</ref><ref>[http://www.ncbi.nlm.nih.gov/pubmed/26122431 Motility, Chemotaxis and Aerotaxis Contribute to Competitiveness during Bacterial Pellicle Biofilm Development.  Hölscher T, Bartels B, Lin YC, Gallegos-Monterrosa R, Price-Whelan A, Kolter R, Dietrich LE, Kovács ÁT.  J Mol Biol. 2015 Jun 26.]</ref><ref>[http://www.ncbi.nlm.nih.gov/pubmed/24035282 Modulation of curli assembly and pellicle biofilm formation by chemical and protein chaperones.  Andersson EK, Bengtsson C, Evans ML, Chorell E, Sellstedt M, Lindgren AE, Hufnagel DA, Bhattacharya M, Tessier PM, Wittung-Stafshede P, Almqvist F, Chapman MR.  Chem Biol. 2013 Oct 24.]</ref>.  Although these studies don't directly look at beer pellicles, this indicates that the definition of a pellicle as stated in this wiki may become more widely accepted in the scientific community.  

Revision as of 12:29, 4 September 2015

Pellicle from The Yeast Bay Brussels Brett Blend; courtesy of Dan Pixley
Lactobacillus pellicle; courtesy of Matt Humbard
A pellicle is an aggregation of cells, proteins, and long chain sugars formed on the surface of a beer. It is often formed by Brettanomyces, Pediococcus, or Lactobacillus. It can also be formed by Saccharomyces in rare occasions [1].

Introduction

Pellicles form when the surface of the beer is exposed to oxygen [2] One theory is that the formation of a pellicle allows the organism to access the small amount of oxygen that is present in the headspace of the fermentation vessel. Another theory, and one that may be less accurate according to Dr. Matt Humbard, is that the pellicle protects the beer from other microorganisms [1]. Popular thought is that the formation of a pellicle is not indicative of the quality of the sour beer that is being produced; it is only an indication that oxygen has entered the fermentation vessel and that the microbes are reacting to that exposure.

Scientific Terminology

(This section in progress)

As with some things in science that are not greatly explored, terminology isn't always agreed upon or fully established, and thus researching such a topic without a lot of prior knowledge can be challenging. Pellicles are a good example of this. It is sometimes stated that a pellicle is a subtype of biofilm. However, the widely accepted definition of a "biofilm" in biology doesn't describe pellicles that we see in fermentation. The widely accepted definition of a "biofilm" is an aggregate of microorganisms where the cells adhere to each other on a surface [3]. Pellicles in beer do not attach to a solid surface, they appear on the "air-liquid interface" (the surface of the beer). They are also not colonies within an adhesive [4]. To make matters even more confusing, the two widely accepted definitions of a "pellicle" in biology are the outer boundary of a protozoa cell [5], and the protein film that forms on the surface of teeth [6].

The usage of the term "biofilm" has been used to describe the layer of film that covers sherry known as "Flor" [7][8]. The word "pellicle" generally isn't used, although it has appeared on occasion when referring to sherry flor in the 1960's [9]. More recently, there have been studies that define a "pellicle" like we see in fermentation as a type of biofilm that forms on the air-liquid interface of a liquid (see references) [10][11][12][13]. Although these studies don't directly look at beer pellicles, this indicates that the definition of a pellicle as stated in this wiki may become more widely accepted in the scientific community.

The bottom line is that what is referred to as a "pellicle" in fermentation has not been widely studied, and the terminology is not yet widely established in the scientific community. Indeed, when researching "biofilms" and "pellicles", results will mostly refer to the more well established topics, which can lead to some initial confusion for brewers. Research done on traditionally defined biofilms cannot necessarily be extrapolated to the pellicle phenomenon seen in fermentation.

Images

External Resources

References

  1. 1.0 1.1 Beer Microbiology – What is a pellicle? A PhD in Beer blog. Dr. Matt Humbard. 01/30/2015. Retrieved 04/26/2015.
  2. Brewing Sour Beer at Home. The Mad Fermentationist Blog. Michael Tonsmeire. 11/06/2009. Retrieved 02/28/2015.
  3. e-Study Guide for Brock Biology of Microorganisms, textbook by Michael T. Madigan.
  4. Conversation with Bryan Heit about Pellicles on MTF. 08/20/2015.
  5. Biology of Protozoa. D.R. Khanna. Discovery Publishing House, Jan 1, 2004. Pg 38.
  6. Wikipedia. Dental Pellicle. Retrieved 08/23/2015.
  7. Ethanol-Independent Biofilm Formation by a Flor Wine Yeast Strain of Saccharomyces cerevisiae. Severino Zara, Michael K. Gross, Giacomo Zara, Marilena Budroni and Alan T. Bakalinsky. 2010.
  8. FLO11 is essential for flor formation caused by the C-terminal deletion of NRG1 in Saccharomyces cerevisiae. Mari Ishigami, Youji Nakagawa, Masayuki Hayakawa, Yuzuru Iimura. 2004.
  9. On the pellicle formation by “flor” yeasts. Cantarelli C, Martini A. Antonie Van Leeuwenhoek. 1969.
  10. Gram-negative bacteria can also form pellicles. Armitano J, Méjean V, Jourlin-Castelli C. Environ Microbiol Rep. 2014 Dec.
  11. Identification of genes essential for pellicle formation in Acinetobacter baumannii. Giles SK, Stroeher UH, Eijkelkamp BA, Brown MH. BMC Microbiol. 2015 Jun 6.
  12. Motility, Chemotaxis and Aerotaxis Contribute to Competitiveness during Bacterial Pellicle Biofilm Development. Hölscher T, Bartels B, Lin YC, Gallegos-Monterrosa R, Price-Whelan A, Kolter R, Dietrich LE, Kovács ÁT. J Mol Biol. 2015 Jun 26.
  13. Modulation of curli assembly and pellicle biofilm formation by chemical and protein chaperones. Andersson EK, Bengtsson C, Evans ML, Chorell E, Sellstedt M, Lindgren AE, Hufnagel DA, Bhattacharya M, Tessier PM, Wittung-Stafshede P, Almqvist F, Chapman MR. Chem Biol. 2013 Oct 24.