See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/284725490 Biochemistry of beer fermentation Book · January 2015 CITATIONS READS 5 3,183 2 authors: Eduardo José Pires Tomáš Brányik University of Minho University of Chemistry and Technology, Pra… 13 PUBLICATIONS 111 CITATIONS 79 PUBLICATIONS 1,215 CITATIONS SEE PROFILE SEE PROFILE Some of the authors of this publication are also working on these related projects: Biosorption of heavy metals View project Yeast biotechnology View project All content following this page was uploaded by Tomáš Brányik on 27 November 2015. The user has requested enhancement of the downloaded file. SpringerBriefs in Biochemistry and Molecular Biology More information about this series at http://www.springer.com/series/10196 Eduardo Pires · Tomáš Brányik Biochemistry of Beer Fermentation 13 Eduardo Pires CEB—Centre of Biological Engineering University of Minho Braga Portugal Tomáš Brányik Department of Biotechnology Institute of Chemical Technology Prague 6 Czech Republic ISSN 2211-9353 ISSN 2211-9361 (electronic) ISBN 978-3-319-15188-5 ISBN 978-3-319-15189-2 (eBook) DOI 10.1007/978-3-319-15189-2 Library of Congress Control Number: 2014960345 Springer Cham Heidelberg New York Dordrecht London © The Author(s) 2015 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made. Printed on acid-free paper Springer International Publishing AG Switzerland is part of Springer Science+Business Media (www.springer.com) Contents 1 An Overview of the Brewing Process . . . . . . . . . . . . . . . . . . . . . . . . . . . A Brief History of Brewing. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Ingredients. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Water . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Malted Barley and Adjuncts. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Malting. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hops . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Yeast . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wort Production . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Milling. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mashing. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wort Boiling. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fermentation and Maturation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 1 2 2 3 4 4 5 5 5 6 7 7 8 2 The Brewing Yeast. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Yeast Flocculation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carbohydrate Transport and Metabolism . . . . . . . . . . . . . . . . . . . . . . . . . . Main Glucose Repression Pathway. . . . . . . . . . . . . . . . . . . . . . . . . . . . Glucose-Sensing System—Ras/cAMP/PKA Pathway. . . . . . . . . . . . . . The Impact of the Glucose-Sensing System on Fermentation. . . . . . . . Transport of α-Glucosides. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nitrogen Metabolism. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Target of Rapamycin (Tor) Pathway. . . . . . . . . . . . . . . . . . . . . . . . . . . . Nitrogen Catabolite Repression (NCR). . . . . . . . . . . . . . . . . . . . . . . . . General Amino Acid Control (GAAC). . . . . . . . . . . . . . . . . . . . . . . . . . Transport and Control of Nitrogen Sources. . . . . . . . . . . . . . . . . . . . . . Alcoholic Fermentation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11 13 15 16 18 20 21 22 23 27 29 30 33 36 v vi Contents 3 By-products of Beer Fermentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pleasant By-products. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Higher Alcohols. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Transamination. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Decarboxylation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Reduction to Higher Alcohols . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Regulation of Higher Alcohols. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Anabolic Pathway. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Esters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Biosynthesis of Acetate Esters. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Biosynthesis of Ethyl Esters. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ester Regulation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Esters in Beer Aging. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Unpleasant By-products . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vicinal Diketones (VDKs). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Yeast Response to Fermentation Parameters. . . . . . . . . . . . . . . . . . . . . . . . Yeast Strain. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Temperature. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrostatic Pressure. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wort Composition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sugars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Free Amino Nitrogen (FANs). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oxygen and Unsaturated Fatty Acids (UFAs) . . . . . . . . . . . . . . . . . . . . References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . View publication stats 47 47 48 48 49 50 51 52 52 54 55 57 57 58 59 59 61 61 62 63 64 64 65 67 68
