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==Proteins in grain==
==Proteins in grain==
The mature [[barley]] grain contains a spectrum of proteins that differ in function, location, structure, and other physical and chemical characteristics. The level of protein is a critical aspect of the quality of [[malt]] and [[beer]], and it is influenced by [[grain]] variety, soil conditions, crop rotation, fertilization, and weather conditions.<ref name=steiner/><ref name=picariello/><ref name=mahalingam/> Generally, the protein content in barley grain represents approximately 8–16% of its total mass.<ref name=mashing>Evans E. [[Library|''Mashing.'']]  American Society of Brewing Chemists and Master Brewers Association of the Americas; 2021.</ref><ref name=silva>Silva F, Nogueira LC, Goncalves C, Ferreira AA, Ferreira IMPLVO, Teixeira N. [https://www.sciencedirect.com/science/article/abs/pii/S0308814607006085 Electrophoretic and HPLC methods for comparative study of the protein fractions of malts, worts and beers produced from Scarlett and Prestige barley (''Hordeum vulgare'' L.) varieties.] ''Food Chem.'' 2008;106(2):820–829.</ref><ref name=mahalingam>Mahalingam R. [https://bmcgenomics.biomedcentral.com/articles/10.1186/s12864-016-3408-5 Shotgun proteomics of the barley seed proteome.] ''BMC Genomics.'' 2017;18(44).</ref> Brewers prefer lower protein levels as long as there is plenty of soluble nitrogen for good yeast nutrition and beer foam potential.<ref name=crit>Bamforth CW, Fox GP. [https://www.brewingscience.de/index.php?tpl=table_of_contents&year=2020&edition=0009%2F0010&article=92781 Critical aspects of starch in brewing.] ''BrewingScience.'' 2020;73(9/10):126–139.</ref> This is because excessive protein content decreases the relative amount of [[carbohydrates]] (mainly [[starch]]) and also has other negative influences on the [[brewing]] process.<ref name=yu/> The barley used for [[malting]] should generally fall at the lower end of this range, with about 9–11% protein.<ref name=fix/><ref>Yu W, Tao K, Gidley MJ, Fox GP, Gilbert RG. [https://www.researchgate.net/publication/328971139_Molecular_brewing_Molecular_structural_effects_involved_in_barley_malting_and_mashing Molecular brewing: Molecular structural effects involved in barley malting and mashing.] ''Carbohydr Polym.'' 2019;206:583–592.</ref> Six-row barley tends to have more protein content and thus less starch than two-row.<ref name=mahalingam/> Hordeins (storage proteins) are the most abundant proteins found in a barley grain, and they form a matrix around the [[starch]] granules, increasing grain hardness.<ref name=celus>Celus I, Brijs K, Delcour JA. [https://www.sciencedirect.com/science/article/abs/pii/S0733521006000762 The effects of malting and mashing on barley protein extractability.] ''J Cereal Sci.'' 2006;44(2):203–211.</ref><ref name=iimure/> These storage proteins are created specifically to be broken down and used as a source of amino acids for building other proteins during seed germination, facilitating the plant's growth. A large variety of other proteins are present in barley, with over a thousand unique proteins identified.<ref name=kerr>Kerr ED, Fox GP, Schulz BL. [https://www.sciencedirect.com/science/article/pii/B9780081005965228692 Grass to glass: Better beer through proteomics.] In: Cifuentes A, ed. ''Comprehensive Foodomics.'' Elsevier; 2020:407–416.</ref> Notable among these are [[lipid transfer protein]]s (LTP) and [[protein Z]], which are the two major proteins in finished beer.
The mature [[barley]] grain contains a spectrum of proteins that differ in function, location, structure, and other physical and chemical characteristics. The level of protein is a critical aspect of the quality of [[malt]] and [[beer]], and it is influenced by [[grain]] variety, soil conditions, crop rotation, fertilization, and weather conditions.<ref name=steiner/><ref name=picariello/><ref name=mahalingam/> Generally, the protein content in barley grain represents approximately 8–16% of its total mass.<ref name=mashing>Evans E. [[Library|''Mashing.'']]  American Society of Brewing Chemists and Master Brewers Association of the Americas; 2021.</ref><ref name=silva>Silva F, Nogueira LC, Goncalves C, Ferreira AA, Ferreira IMPLVO, Teixeira N. [https://www.sciencedirect.com/science/article/abs/pii/S0308814607006085 Electrophoretic and HPLC methods for comparative study of the protein fractions of malts, worts and beers produced from Scarlett and Prestige barley (''Hordeum vulgare'' L.) varieties.] ''Food Chem.'' 2008;106(2):820–829.</ref><ref name=mahalingam>Mahalingam R. [https://bmcgenomics.biomedcentral.com/articles/10.1186/s12864-016-3408-5 Shotgun proteomics of the barley seed proteome.] ''BMC Genomics.'' 2017;18(44).</ref> Brewers prefer lower protein levels as long as there is plenty of soluble nitrogen for good yeast nutrition and beer foam potential.<ref name=crit>Bamforth CW, Fox GP. [https://www.brewingscience.de/index.php?tpl=table_of_contents&year=2020&edition=0009%2F0010&article=92781 Critical aspects of starch in brewing.] ''BrewingScience.'' 2020;73(9/10):126–139.</ref> This is because excessive protein content decreases the relative amount of [[carbohydrates]] (mainly [[starch]]) and also has other negative influences on the [[brewing]] process.<ref name=yu/> The barley used for [[malting]] should generally fall at the lower end of this range, with about 9–11% protein.<ref name=fix/><ref>Yu W, Tao K, Gidley MJ, Fox GP, Gilbert RG. [https://www.researchgate.net/publication/328971139_Molecular_brewing_Molecular_structural_effects_involved_in_barley_malting_and_mashing Molecular brewing: Molecular structural effects involved in barley malting and mashing.] ''Carbohydr Polym.'' 2019;206:583–592.</ref> Six-row barley tends to have more protein content and thus less starch than two-row.<ref name=mahalingam/> Hordeins (storage proteins) are the most abundant proteins found in a barley grain, and they form a matrix around the [[starch]] granules, increasing grain hardness.<ref name=celus>Celus I, Brijs K, Delcour JA. [https://www.sciencedirect.com/science/article/abs/pii/S0733521006000762 The effects of malting and mashing on barley protein extractability.] ''J Cereal Sci.'' 2006;44(2):203–211.</ref><ref name=iimure/> These storage proteins are created specifically to be broken down and used as a source of amino acids for building other proteins during seed germination, facilitating the plant's growth.<ref name=mashing/> A large variety of other proteins are present in barley, with over a thousand unique proteins identified.<ref name=kerr>Kerr ED, Fox GP, Schulz BL. [https://www.sciencedirect.com/science/article/pii/B9780081005965228692 Grass to glass: Better beer through proteomics.] In: Cifuentes A, ed. ''Comprehensive Foodomics.'' Elsevier; 2020:407–416.</ref> Notable among these are [[lipid transfer protein]]s (LTP) and [[protein Z]], which are the two major proteins in finished beer.


Although there are some differences in protein structure, the overall protein profile of beers produced from other cereal grains used for brewing tends to be similar, including [[wheat]] and [[oats]].<ref name=picariello/><ref name=klose>Klose C, Thiele F, Arendt EK. [https://www.tandfonline.com/doi/abs/10.1094/ASBCJ-2010-0312-01 Changes in the protein profile of oats and barley during brewing and fermentation.] ''J Am Soc Brew Chem.'' 2010;68(2):119–124.</ref>
Although there are some differences in protein structure, the overall protein profile of beers produced from other cereal grains used for brewing tends to be similar, including [[wheat]] and [[oats]].<ref name=picariello/><ref name=klose>Klose C, Thiele F, Arendt EK. [https://www.tandfonline.com/doi/abs/10.1094/ASBCJ-2010-0312-01 Changes in the protein profile of oats and barley during brewing and fermentation.] ''J Am Soc Brew Chem.'' 2010;68(2):119–124.</ref>