BIOCHEMISTRY BIOTECHNOLOGY AN INTRODUCTION TO POLYSACCHARIDE Stephen E. Harding • Michael P. Tombs An Introduction to Gary G. Adams • Berit Smestad Paulsen POLYSACCHARIDE Kari Tvete Inngjerdingen • Hilde Barsett BIOTECHNOLOGY An Introduction to SECOND EDITION Polysaccharides and related high molecular weight glycans are hugely diverse with wide application in biotechnology and great opportunities for further exploitation. POLYSACCHARIDE An Introduction to Polysaccharide Biotechnology – a second edition of the popular original text by Tombs and Harding – introduces students, researchers, clinicians and industrialists to the properties of some of the key materials involved, how these are applied, some of the economic factors concerning their production and how they BIOTECHNOLOGY are characterised for regulatory purposes. FEATURES • Basic properties of polysaccharides and how they are very different to proteins SECOND EDITION • How these properties are affected by enzymes and how the effects of enzymes can be controlled • An introduction to some ‘patent preferred’ methodologies for polysaccharide EDITION SECOND characterisation, such as molecular weight distribution • Applications in biopharma, food and other industries • The application of marker-assisted selection to fruit ripening and preservation, the development of glycovaccines against meningitis and other serious diseases • Barsett Paulsen • Inngjerdingen • Adams Harding • Tombs • A whole chapter dedicated to a case study on bioactivity and how modern research is unlocking the way polysaccharides are at the core of many HG HG traditional medicines • Further Reading sections for follow up K23616 6000 Broken Sound Parkway, NW Suite 300, Boca Raton, FL 33487 RG-I RG-II 711 Third Avenue New York, NY 10017 2 Park Square, Milton Park Abingdon, Oxon OX14 4RN, UK www.crcpress.com An Introduction to Polysaccharide Biotechnology An Introduction to Polysaccharide Biotechnology By Stephen E. Harding, Michael P. Tombs and Gary G. Adams University of Nottingham, UK and Berit Smestad Paulsen, Kari Tvete Inngjerdingen and Hilde Barsett University of Oslo, Norway 2nd Edition CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 © 2017 by Taylor & Francis Group, LLC CRC Press is an imprint of Taylor & Francis Group, an Informa business No claim to original U.S. Government works Printed on acid-free paper International Standard Book Number-978-1-4822-4697-1 (Hardback) This book contains information obtained from authentic and highly regarded sources. Reasonable efforts have been made to publish reliable data and information, but the author and publisher cannot assume responsibility for the validity of all materials or the consequences of their use. 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CCC is a not-for-profit organization that provides licenses and registration for a variety of users. For organizations that have been granted a photocopy license by the CCC, a separate system of payment has been arranged. Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identification and explanation without intent to infringe. Library of Congress Cataloging-in-Publication Data Names: Harding, S. E. (Stephen E.) Title: An Introduction to Polysaccharide Biotechnology. Description: Second edition / Stephen E. Harding [and five others]. | Boca Raton : CRC Press, [2017] | Includes bibliographical references and index. Identifiers: LCCN 2016040404 | ISBN 9781482246971 (hbk : alk. paper) | ISBN 9781315372730 (ebk : alk. paper) Subjects: LCSH: Polysaccharides--Biotechnology. | Polysaccharides--Metabolism. | Glucosidases--Biotechnology. Classification: LCC TP248.65.P64 T66 2017 | DDC 660.6/3--dc23 LC record available at https://lccn.loc.gov/2016040404 Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com and the CRC Press Web site at http://www.crcpress.com Contents Preface to First Edition ............................................................................................................................................................................ xi Preface to Second Edition ..................................................................................................................................................................xiii Chapter 1 Polysaccharides and Their Potential for Biotechnology ....................................................................... 1 1.1 An Introduction to Polysaccharides .................................................................................................. 1 1.1.1 Glycosubstances and Glycans ........................................................................................... 2 1.1.2 Structures, Nomenclature and Abbreviations ......................................................... 3 1.1.3 Polysaccharide Polydispersity: Molecular Weight Averages ........................... 9 1.1.4 Solution Properties and Characterization Methods ..........................................10 1.1.5 Measurement of Molecular Weight Averages and Molecular Weight Distribution ...............................................................................................................10 1.1.6 Polysaccharide Conformation in Solution ...............................................................16 1.1.6.1 Overall or ‘Gross’ Conformation: Haug Triangle .............................16 1.1.6.2 Effect of Ionic Strength of the Surrounding Medium ................18 1.1.6.3 Intrinsic Viscosity ...............................................................................................18 1.1.6.4 Conformation Zones.......................................................................................19 1.1.6.5 More Specific Conformation Information: Rigid Structures ..................................................................................................20 1.1.6.6 Flexible Structures: Worm-Like Coil and Persistence Length ...........................................................................................21 1.1.6.7 Ordered Structure ............................................................................................21 1.1.7 Imaging of Polysaccharides: X-Ray Fibre Diffraction, Electron Microscopy and Atomic Force Microscopy .............................................................21 1.1.8 Polysaccharide Rheology ...................................................................................................23 1.1.9 Classification of Polysaccharides ....................................................................................25 1.1.10 The Cell Wall ...............................................................................................................................26 1.1.10.1 Cellulose .................................................................................................................26 1.1.10.2 Mannans .................................................................................................................26 1.1.10.3 Callose .....................................................................................................................26 1.1.10.4 Heteropolymers .................................................................................................26 1.1.10.5 Heteroglucans ....................................................................................................27 1.1.10.6 Heteroxylans ........................................................................................................27 1.1.10.7 Galactomannans, Glucomannans and Galactoglucomannans ..................................................................................27 1.1.10.8 Pectins .....................................................................................................................27 1.1.10.9 Glycoproteins or Extensins ..........................................................................28 1.1.10.10 Lignins .....................................................................................................................28 1.1.10.11 Alginates .................................................................................................................28 1.1.10.12 Carrageenans ......................................................................................................28 1.1.10.13 Furcellaran .............................................................................................................29 1.1.10.14 Fucans ......................................................................................................................29 1.1.10.15 Other Components