Saccharides are widely distributed in all living life forms and glycobiology is the study of their structure, biosynthesis, biology, and evolution. It is a rapidly growing field, relevant to many areas of basic research, biomedicine, and biotechnology. The field includes the chemistry of
carbohydrates, the enzymology of glycan formation and degradation, the recognition of glycans by specific proteins, roles of glycans in complex biological systems, and their analysis and manipulation. Essentials of Glycobiology is the field's most authoritative textbook, offering an overview
suitable for advanced undergraduates and graduate students in molecular and cellular biology and biomedicine. In an entirely revised and updated fourth edition, the book retains a broad focus and applicability to fields ranging from biology and medicine to chemistry and materials science, with
excellent artwork and chapters written by an international selection of leading investigators.

Foreword
Preface
Praise for Previous Editions of Essentials of Glycobiology
Contributors
Editors of Earlier Editions
Consultant Reviewers
Classic Books and Monographs
Abbreviations

GENERAL PRINCIPLES
Historical Background and Overview
Ajit Varki and Stuart Kornfeld
Monosaccharide Diversity
Peter H. Seeberger
Oligosaccharides and Polysaccharides
Carlito B. Lebrilla, Jian Liu, Goran Widmalm, and James H. Prestegard
Cellular Organization of Glycosylation
Karen J. Colley, Ajit Varki, Robert S. Haltiwanger, and Taroh Kinoshita
Glycosylation Precursors
Hudson H. Freeze, Michael Boyce, Natasha E. Zachara, Gerald W. Hart, and Ronald L. Schnaar
Glycosyltransferases and Glycan-Processing Enzymes
James M. Rini, Kelley W. Moreman, Benjamin G. Davis, and Jeffrey D. Esko
Biological Functions of Glycans
Pascal Gagneux, Thierry Hennet, and Ajit Varki
A Genomic View of Glycobiology
Nicolas Terrapon, Bernard Henrissat, Kiyoko F. Aoki-Kinoshita, Avadhesha Surolia, and Pamela Stanley

STRUCTURE AND BIOSYNTHESIS
N-Glycans
Pamela Stanley, Kelley W. Moremen, Nathan E. Lewis, Naoyuki Taniguchi, and Markus Aebi
O-GalNAc Glycans
Inka Brockhausen, Hans H. Wandall, Kelly G. Ten Hagen, and Pamela Stanley
Glycosphingolipids
Ronald L. Schnaar, Roger Sandhoff, Michael Tiemeyer, and Taroh Kinoshita
Glycosylphosphatidylinositol Anchors
Sneha Sudha Komath, Morihisa Fujita, Gerald W. Hart, Michael A.J. Ferguson, and Taroh Kinoshita
Other Classes of Eukaryotic Glycans
Robert S. Haltiwanger, Lance Wells, Hudson H. Freeze, Hamed Jafar-Nejad, Tetsuya Okajima, and Pamela Stanley
Structures Common to Different Glycans
Pamela Stanley, Manfred Wuhrer, Gordon Lauc, Sean R. Stowell, and Richard D. Cummings
Sialic Acids and Other Nonulosonic Acids
Amanda Lewis, Xi Chen, Ronald L. Schnaar, and Ajit Varki
Hyaluronan
Melanie Simpson, Liliana Schaefer, Vincent Hascall, and Jeffrey D. Esko
Proteoglycans and Sulfated Glycosaminoglycans
Catherine L.R. Merry, Ulf Lindahl, John Couchman, and Jeffrey D. Esko
Nucleocytoplasmic Glycosylation
Christopher M. West, Chad Slawson, Natasha E. Zachara, and Gerald W. Hart
The O-GlcNAc Modification
Natasha E. Zachara, Yoshihiro Akimoto, Michael Boyce, and Gerald W. Hart

GLYCANS IN EVOLUTION AND DEVELOPMENT
Evolution of Glycan Diversity
Pascal Gagneux, Vladislav Pannin, Thierry Hennet, Markus Aebi, and Ajit Varki
Eubacteria
Chris Whitfield, Christine M. Szymanski, Amanda Lewis, and Markus Aebi
Archaea
Benjamin H. Meyer, Sonja-Verena Albers, Jerry Eichler, and Markus Aebi
Fungi
Francoise H. Routier, Tamara L. Doering, Richard D. Cummings, and Markus Aebi
Viridiplantae and Algae
Malcolm A. O'Neill, Alan G. Darvill, Marilynn E. Etzler, Debra Mohnen, Serge Perez, Jenny C. Mortimer, and Markus Pauly
Nematoda
Iain B.H. Wilson, Katharina Paschinger, Richard D. Cummings, and Markus Aebi
Arthropoda
Kelly G. Ten Hagen, Hiroshi Nakato, Michael Tiemeyer, and Jeffrey D. Esko
Deuterostomes
Michael Pierce, Iain B. H. Wilson, Katharina Paschinger, and Pamela Stanley

GLYCAN-BINDING PROTEINS
Discovery and Classification of Glycan-Binding Proteins
Maureen E. Taylor, Kurt Drickamer, Anne Imberty, Yvette van Kooyk, Ronald L. Schnaar, Marilynn E. Etzler, and Ajit Varki
Principles of Glycan Recognition
Richard D. Cummings, Ronald L. Schnaar, Jeffrey D. Esko, Robert J. Woods, Kurt Drickamer, and Maureen E. Taylor
Structural Biology of Glycan Recognition
Jesus Angulo, Jochen Zimmer, Anne Imberty, and James H. Prestegard
R-Type Lectins
Richard D. Cummings, Ronald L. Schnaar, and Yasuhiro Ozeki
L-Type Lectins
Richard D. Cummings, Marilynn E. Etzler, T.N.C. Ramya, Koichi Kato, Gabriel A. Rabinovich, and Avadhesha Surolia
P-Type Lectins
Nancy Dahms, Thomas Braulke, and Ajit Varki
C-Type Lectins
Richard D. Cummings, Elise Chiffoleau, Yvette van Kyook, and Rodger P. McEver
I-Type Lectins
Takashi Angata, Stephan von Gunten, Ronald L. Schnaar, and Ajit Varki
Galectins
Richard D. Cummings, Fu-Tong Liu, Gabriel A. Rabinovich, Sean R. Stowell, and Gerardo R. Vasta
Microbial Lectins: Hemagglutinins, Adhesins, and Toxins
Amanda L. Lewis, Jennifer J. Kohler, and Markus Aebi
Proteins That Bind Sulfated Glycosaminoglycans
Ding Xu, James H. Prestegard, Robert J. Linhardt, and Jeffrey D. Esko

GLYCANS IN PHYSIOLOGY AND DISEASE
Glycans in Glycoprotein Quality Control
Tadashi Suzuki, Richard D. Cummings, Markus Aebi, and Armando Parodi
Free Glycans as Bioactive Molecules
Antonio Molina, Malcolm A. O'Neill, Alan G. Darvill, Marilynn E. Etzler, Debra Mohnen, Michael G. Hahn, and Jeffrey D. Esko
Glycans in Systemic Physiology
Robert Sackstein, Sean R. Stowell, Karin M. Hoffmeister, Hudson H. Freeze, and Ajit Varki
Bacterial and Viral Infections
Amanda L. Lewis, Christine M. Szymanski, Ronald L. Schnaar, and Markus Aebi
Parasitic Infections
Richard D. Cummings, Cornelis H. Hokke, and Stuart M. Haslam
Genetic Disorders of Glycan Degradation
Hudson H. Freeze, Richard Steet, Tadashi Suzuki, Taroh Kinoshita, and Ronald L. Schnaar
Congenital Disorders of Glycosylation
Dirk J. Lefeber, Hudson H. Freeze, Rich Steet, and Taroh Kinoshita
Glycans in Acquired Human Diseases
Robert Sackstein, Karin M. Hoffmeister, Sean R. Stowell, Taroh Kinoshita, Ajit Varki, and Hudson H. Freeze
Glycosylation Changes in Cancer
Susan L. Bellis, Celso A. Reis, Ajit Varki, Reiji Kannagi, and Pamela Stanley

METHODS AND APPLICATIONS
Glycan-Recognizing Probes as Tools
Richard D. Cummings, Marilyn Etzler, Michael G. Hahn, Alan Darvill, Kamil Godula, Robert J. Woods, and Lara K. Mahal
Glycosylation Mutants of Cultured Mammalian Cells
Jeffrey D. Esko, Hans H. Wandall, and Pamela Stanley
Structural Analysis of Glycans
Stuart M Haslam, Daron I. Freedberg, Barbara Mulloy, Anne Dell, Pamela Stanley, and James H. Prestegard
Glycomics and Glycoproteomics
Pauline M. Rudd, Niclas G. Karlsson, Kay-Hooi Khoo, Morten Thaysen-Andersen, Lance Wells, and Nicolle H. Packer
Glycoinformatics
Kiyoko F. Aoki-Kinoshita, Matthew P. Campbell, Frederique Lisacek, Sriram Neelamegham, William S. York, and Nicolle H. Packer
Chemical Synthesis of Glycans and Glycoconjugates
Peter H. Seeberger and Hermen S. Overkleeft
Chemoenzymatic Synthesis of Glycans and Glycoconjugates
Hermen S. Overkleeft and Peter H. Seeberger
Chemical Tools for Inhibiting Glycosylation
David J. Vocadlo, Todd L. Lowary, Carolyn R. Bertozzi, Ronald L. Schnaar, and Jeffrey D. Esko
Glycosylation Engineering
Henrik Clausen, Hans H. Wandall, Matthew P. DeLisa, Pamela Stanley, and Ronald L. Schnaar
Glycans in Biotechnology and the Pharmaceutical Industry
Peter H. Seeberger, Daron I. Freedberg, and Richard D. Cummings
Glycans in Nanotechnology
Martina Delbianco, Benjamin G. Davis, and Peter H. Seeberger
Glycans in Bioenergy and Materials Science
Malcolm A. O'Neill, Robert J. Moon, William S. York, Alan G. Darvill, Kamil Godula, Breeanna Urbanowicz, and Debra Mohnen
Future Directions in Glycosciences
Gerald W. Hart and Ajit Varki

Glossary
Study Guide
Index