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 High-Performance
Liquid
Chromatography
of
Peptides and Proteins:
Separation, Analysis,
and
Conformation
Editors

Colin T. Mant
R esearch Associate
D epartm ent o f B iochem istry
U niversity o f A lberta
Edm onton, A lberta, Canada

Robert S. Hodges
Professor
D epartm ent o f B iochem istry
U niversity o f A lberta
Edm onton, A lberta, C anada

CRC Press
Boca Raton London New York W ashington, D.C.
 L ibrary of Congress Cataloging-ln-Publlcation D ata

High-performance liquid chromatography o f peptides and proteins:

separation, analysis, and conformation / editors, Colin T. Mant,
Robert S. H odges,
p. cm.
Includes bibliographical references and index.
ISBN 0-8493-6549-X
1. High performance liquid chromatography. 2. Proteins— ^Analysis.
3. Peptides— ^Analysis. I. Mant, Colin T. II. H odges, Robert S.
QP519.9.H53H5355 1991
574.19'245— dc20 91-17549
CIP

This book contains information obtained from authentic and highly regarded sources. Reprinted material is
quoted with permission, and sources are indicated. A wide variety of references are listed. Reasonable efforts
have been made to publish reliable data and information, but the author and the publisher cannot assume
responsibility for the validity of all materials or for the consequences of their use.

Neither this book nor any part may be reproduced or transmitted in any form or by any means, electronic or
mechanical, including photocopying, microfilming, and recording, or by any information storage or retrieval
system, without prior permission in writing from the publisher.

All rights reserved. Authorization to photocopy items for internal or personal use, or the personal or internal
use of specific clients, may be granted by CRC Press LLC, provided that $.50 per page photocopied is paid
directly to Copyright Clearance Center, 222 Rosewood Drive, Danvers, MA 01923 USA. The fee code for
users of the Transactional Reporting Service is ISBN 0-8493-6549-X/99/$0.00+$.50. The fee is subject to
change without notice. For organizations that have been granted a photocopy license by the CCC, a separate
system of payment has been arranged.

The consent of CRC Press LLC does not extend to copying for general distribution, for promotion, for creating
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copying.

Direct all inquiries to CRC Press LLC, 2000 N.W. Corporate Blvd., Boca Raton, Florida 33431.

IVademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used
only for identification and explanation, without intent to infnnge.
Visit the CRC W eb site a t w w w .crcpress.com

© 1991 by CRC Press LLC

No claim to original U.S. Government works

International Standard Book Number 0-8493-6549-X
Library of Congress Card Number 91-17549
14 15
 PREFACE

On the face of it, preparing what the editors wished to be a comprehensive and practical
publication encompassing high-performance liquid chromatography (HPLC) of peptides and
proteins appeared to be a somewhat daunting ambition. There is certainly no scarcity of material
on the topic at hand (from sources including published papers, books, industrial communica­
tions, and even word-of-mouth), a reflection of the explosive growth in the application of HPLC
to peptide/protein separations during the past decade. However, to paraphrase a comment in one
of the enclosed articles (“the approach to their separation must be tailored to the separation
goals”)»we felt that our approach to the book had to be tailored to our perceived goal of a practical
teaching manual for both die novice HPLC user as well as more experienced chromatographers.
Thus, in addition to being a hands-on teaching manual and textbook, we also wanted the book
to represent the perfect reference guide and training tool for personnel in well-established HPLC
laboratories.
With these aims in mind, we requested our contributors, scientists of international renown,
to help us prepare a series of precise, easy-to-read articles, emphasizing the practical approach
to HPLC with minimal theory, although with clear expression of the underlying principles to
peptide and protein separations. We felt that by having a large number of contributors (117)
producing a substantial number (82) of short articles (containing clear and simplistic figures and
only key references), with the concomitant advantage of expressing different viewpoints and
approaches, we could produce a publication representing the flavor of the HPLC field as a whole,
instead of just a confined vision. Although coordinating this considerable number of contribu­
tors and articles necessarily took time and put some stress on us as editors, it was undoubtedly
worth it. All of the major modes of microbore, ultrafast, and analytical HPLC (size-exclusion,
ion-exchange, reversed-phase, hydrophobic interaction, affinity, and immunoaffinity chroma­
tography) are discussed, together with a presentation of preparative HPLC, including displace­
ment techniques. Problem-solving approaches to the separation of various classes of biologi­
cally active peptides and proteins (e.g., peptide hormones, viral proteins, hydrophobic integral
membrane proteins, receptor proteins and enzymes) are thoroughly explored. The importance
of peptide standards for monitoring column and instrument performance and for optimizing
separation conditions is heavily emphasized. In order to maximize the practical potential of
HPLC, several articles focus on the choice of the correct detection method (electrochemical,
UV, fluorescence) as well as the need for a proper knowledge of approaches to column and
instrument maintenance and trouble-shooting. A section of predictive approaches, an area of
ever-growing interest, deals with computer simulation of both peptide separations and peptide
structure. Also included is the value of HPLC for protein conformation/folding studies and
protein structure determination and verification by peptide mapping. Aids to HPLC users
include descriptions of trace enrichment techniques, batch extraction of peptides, and sample
preparation methods. Finally, various complementary techniques to HPLC are described,
including amino acid analysis, preparation of proteins for microsequence analysis, sequencing
techniques, and capillary zone electrophoresis.
Good intentions are all very well, but to produce the kind of publication we envisaged
required the support and cooperation of all our contributors. One of the prime rewards of this
whole endeavor has been our making the acquaintance of a host of scientists we may not
otherwise have met, considering our diverse interests. Our sincere thanks to one and all.

Colin Mant
Bob Hodges
University of Alberta
Edmonton, Canada
 ACKNOWLEDGMENTS

We are grateful for the continuing support of the Medical Research Council of Canada and
the Alberta Heritage Foundation for Medical Research for providing the funding and equipment
necessary for us to maintain international competitiveness in the HPLC field. We also thank the
co-directors (Dr. Kay and Dr. Smillie) of the MRC Group for their support and understanding
of the time taken to complete this worthwhile project.
Our thanks to secretaries Vicki Luxton and Dawn Lockwood for putting up with an enormous
typing burden.
Finally, our thanks to wives Janice Mant and Phyllis Hodges for their encouragement and
patience with the whole venture. Like us, they believe it was worth it. For this reason, and many
others, we dedicate this book to them.
 THE EDITORS

Dr. Colin T. M antj Ph.D., is a Research Associate in the laboratory of Dr. Robert S. Hodges
at the University of Alberta, Edmonton, Alberta, Canada.
Dr. Mant attended the University of Manchester Institute of Science and Technology
(UMIST) in Manchester, England and received a B.Sc. (Hons.) degree in biochemistry from the
Victoria University of Manchester in 1975. He carried out his postgraduate work at the
University of Kent at Canterbury, England, under the supervision of Dr. Ronald B. Cain. This
work, involving research into the microbial catabolism of herbicide-related synthetic pyrimidi­
nes and carried out in collaboration with I.C.I. (Plant Protection) Ltd. in Berkshire, England, led
to Dr. Mant receiving his Ph.D. in Biochemistry in 1982.
Dr. Mant joined the laboratory of Dr. Hodges in 1982 as a Medical Research Council of
Canada postdoctoral fellow (1982 to 1987) and Research Associate (1987 to present) in the
renowned Medical Research Council of Canada Group in Protein Structure and Function. He has
more than 45 publications in the area of HPLC methodology.
Apart from the day-to-day management of the HPLC projects in Dr. Hodges’ group, now an
important part of the total research carried out in the editors’ laboratory, Dr. Mant also acts as
key collaborator with Dr. Hodges on many of the other research interests of the group,
particularly on projects involving protein design, computer simulation, and muscle regulation.
He is also a local organizer for the 13th American Peptide Symposium, to be held in Edmonton
in 1993.

Dr. R obert S. Hodges, Ph.D., is Professor of Biochemistry and a member of the Medical
Research Council of Canada Group in Protein Structure and Function at the University of
Alberta, Edmonton, Alberta, Canada.
Dr. Hodges graduated in 1965 from the University of Saskatchewan, Saskatoon, Sas­
katchewan, Canada with a B.Sc. degree in honors biochemistry. After graduation, he worked for
2 years as Research Scientific Service Officer, Department of National Defence, Defence
Research Board, Government of Canada, in 1971, he obtained his Ph.D. degree in biochemistry
from the University of Alberta. This work was supervised by Dr. L. B. Smillie and involved the
amino acid sequence determination of the first two-stranded a-helical coiled-coil (tropomyosin).
This research led to the hypothesis that a hydrophobic 3— 4 repeat was responsible for the
formation and stabilization of this unique structure. After graduation, he worked as a Medical
Research Council of Canada postdoctoral fellow (1971 to 1973) and Research Associate (1973
to 1974) in the laboratory of Professor Bruce Merrifield, Nobel Prize winner in Chemistry in
1984, at the Rockefeller University, New York. The research involved structure-function
studies of the enzyme ribonucléase A through the chemical synthesis of protein fragments using
the Merrifield solid-phase method.
In 1974, Dr. Hodges was appointed Assistant Professor of Biochemistry and became one of
the five founding members of the Medical Research Council of Canada Group In Protein
Structure and Function. The deficiencies of traditional chromatographic methods for purifica­
tion of synthetic peptides, proteins, and protein fragments were evident to the group and Dr.
Hodges purchased their first HPLC in 1979 to maintain international competitiveness. He
became an Associate Professor in 1977 and Professor in 1984.
The demand on Dr. Hodges’ research laboratory for synthetic peptides and peptide-
conjugates by the Canadian research community resulted in the formation, in 1985, of a cost-
recovery service facility, the Alberta Peptide Institute, of which he remains Director.
In 1986, Dr. Hodges formed S.P.I. Synthetic Peptides Incorporated, a University of Alberta
spin-off research and development company. The objectives of S.P.I. under his supervision as
President, are to conunercialize peptide-based products (peptide pharmaceuticals, peptide
diagnostics, and synthetic vaccines). The company also markets HPLC peptide standards,
HPLC software (ProDigest LC) and peptide/protein structure prediction software. S.P.I. offers
a custom synthesis and HPLC methodology development service for the pharmaceutical and
biotechnology industry.
In 1990, the Government of Canada began a new program to promote excellence in Canadian
science. The Networks of Centres of Excellence (involving universities and industrial partners
across Canada). Dr. Hodges is a participant in two of these programs. First, the Protein
Engineering Network: 3D-Structure, Function and Design. The second, the Bacterial Diseases
Network: Molecular Strategies for the Study and Control of Bacterial Pathogens of Humans,
Animals, Fish and Plants.
Dr. Hodges in 1986 received special recognition for contributions to biotechnology in “A
Tribute to Biotechnology” sponsored by the Government of Alberta. In 1988, he joined the
Editorial Board of the journal Peptide Research, In 1990, he became a member of the program
committee for the 12th American Peptide Symposium and will chair the 13th American Peptide
Symposium in Edmonton, in 1993. In 1991, he was elected a Fellow of the Royal Society of
Canada.
It is noteworthy that HPLC methodology has played and will continue to play a critical role
in the success of biotechnology. Dr. Hodges has published more than 50 HPLC methodological
papers from over 170 research papers. His current major research interests include the de novo
design of peptides and proteins with novel structural and biological properties, immunogenicity/
antigenicity of peptides and proteins, development of synthetic vaccines to viral and bacterial
pathogens and understanding muscle regulation. Peptide synthesis and HPLC are the major
methodologies utilized to achieve his goals and the importance of HPLC is reflected by the dozen
HPLC instruments utilized by his group.
 CONTRIBUTORS

Andrew J . Aipert Ron Biehler

President Senior Applications Chemist
PolyLC, Inc. Department of Applications
Columbia, Maryland Beckman Instruments, Inc.
Palo Alto, California
Firoz D. Antia
Research Assistant Jam es A. Black
Department of Chemical Engineering Facility Manager
Yale University Alberta Peptide Institute
New Haven, Connecticut University of Alberta
Edmonton, Alberta, Canada
Vivian Arrizon-Lopez
Senior Application Chemist Michael M. B rasseur
Research and Applications Department Manager
Beckman Instruments, Inc. Department of Peptide Chemistry
Palo Alto, California Magainin Sciences, Inc.
Plymouth Meeting, Pennsylvania
John E. Battersby
Senior Research Associate T. W. L om e Burke
Department of Analytical Chemistry Technologist
Genentech, Inc. Department of Biochemistry
San Francisco, California University of Alberta
Edmonton, Alberta, Canada
Wayne J. Becktel
Assistant Professor Klaus B üítner
Department of Biochemistry Post-Doctoral Fellow
The Ohio State University Torrey Pines Institute for Molecular
Columbus, Ohio Studies
San Diego, California
Suzanne Benjannet
Research Assistant Paul J. Cachia
Department of Molecular Senior Research Scientist and Manager
Neuroendocrinology Peptide Synthesis and Services Division
Clinical Research Institute of Synthetic Peptides, Inc.
Montreal and
Montreal, Quebec, Canada Research Associate
Department of Biochemistry
Hugh P. J. Bennett University of Alberta
Associate Professor Edmonton, Alberta, Canada
Department of Medicine
McGill University Michael R. C arpenter
and Technologist
Department of Endocrine Laboratory Department of Biochemistry
Royal Victoria Hospital University of Alberta
Montreal, Quebec, Canada Edmonton, Alberta, Canada
Rosanne C. Chloupek John W . Dolan
Senior Research Associate President
Department of Analytical Chemistry LC Resources Inc.
Genentech, Inc. Lafayette, California
San Francisco, California and
Editor
Michel C hrétien LC Troubleshooting
Scientific Director and Chief Executive LC/GC Magazine
Officer Eugene, Oregon
Laboratory of Molecular
Neuroendocrinology Jing Dong
Clinical Research Institute of Montreal Research Scholar
Montreal, Quebec, Canada Hormone Receptor Laboratory
University of Louisville
Joel C. C olburn Louisville, Kentucky
Product Manager
Department of Research and Development Stefan Einarsson
Applied Biosystems, Inc. Research Associate
San Jose, California Icelandic Fisheries
Laboratories
J. M yron C raw ford Reykjavik, Iceland
Associate in Research
Department of Molecular Biophysics and Heinz E ngelhardt
Biochemistry Professor
Yale University Department of Angewandte Physikalische
New Haven, Connecticut Chemie
Universität des Saarlandes
Judy Cummings Saarbrücken, Federal Republic of
Applications Specialist Germany
Research and Applications Department
Beckman Instruments, Inc. Ulrich Esser
Palo Alto, California Hartmann & Braun
Frankfurt, Germany
Raym ond D. DeAngells
Associate in Research II P etr Folk
Department of Molecular Biophysics and Research Associate
Biochemistry Department of General
Yale University Physiology
New Haven, Connecticut Charles University
Prague, Czechoslovakia
C. Dewaele
Chief HPLC Assistant Helene Hagestam Freiser
Laboratory of Organic Chemistry Senior Research Scientist
University of Gent SynChrom, Inc.
Gent, Belgium Lafayette, Indiana
R obert Galyean Milton T. W. H earn
Research Assistant Professor of Biochemistry
The Clayton Foundation Laboratories for Director, Centre for Bioprocess
Peptide Biology Technology
The Salk Institute Monash University
La Jolla, California Clayton, Victoria, Australia

Heinz Goetz R ichard A. H enry

Application Chemist President
Hewlett-Packard Keystone Scientific, Inc.
Waldbronn, West Germany Bellefonte, Pennsylvania

K aren M. Gooding R obert S. Hodges

Analytical Director and President Professor
SynChrom, Inc. Department of Biochemistry
Lafayette, Indiana University of Alberta
Edmonton, Alberta, Canada
Paul D. Grossm an
Scientist C arl A. Hoeger
Department of Research and Development Senior Research Associate
Applied Biosystems, Inc. The Clayton Foundation Laboratories for
Foster City, California Peptide Biology
The Salk Institute
Joachim R. G rün La Jolla, California
Chemist
Max-Planck Institute for Molecular Anita L. Hong
Genetics Department of Custom Peptide Synthesis
Berlin, Germany Applied Biosystems, Inc.
Foster City, California
William S. Hancock
Staff Scientist Csaba H orváth
Department of Analytical Professor and Chairman
Chemistry Department of Chemical Engineering
Genentech Yale University
San Francisco, California New Haven, Connecticut

G oran Hansson R ichard A. Houghten

Graduate Student Director
Department of Analytical and Marine Torrey Pines Institute for Molecular
Chemistry Studies
University of Göteborg San Diego, California
Göteborg, Sweden
Victor J. H ruby
Jon H arbaugh Regents Professor
Product Planning Manager Department of Chemistry
Department of Product and
Planning Department of Biochemistry
Beckman Instruments, Inc. University of Arizona
Palo Alto, California Tucson, Arizona
Salman M. Hyder Andrew Kawasaki
Research Scientist Research Associate
Department of Pharmacology Department of Chemistry
University of Texas Medical School University of Arizona
Houston, Texas Tucson, Arizona

Niggi Iberg Cyril M, Kay

Research Associate Professor
Department of Surgical Research, Department of Biochemistry
Children’s Hospital University of Alberta
Harvard Medical School Edmonton, Alberta, Canada
Boston, Massachusetts
Djohan Kesum a
R ichard H. Ingraham Chemist
Senior Scientist Department of Custom Peptide Synthesis
Department of Medicinal Chemistry Applied Biosystems
Boehringer Ingelheim Pharmaceutical, Inc. Foster City, California
Ridgefield, Connecticut
Michael K lagsbrun
Toshiaki Isobe Associate Professor
Instructor in Biochemistry Department of Biological Chemistry and
Department of Chemistry, Faculty of Surgery
Science Harvard Medical School
Tokyo Metropolitan University Boston, Massachussets
Setagaya-ku, Tokyo, Japan
and I. K oornneef
Instructor Technician
Department of Physiology Hubrecht Laboratory
Saitama Medical School Netherlands Institute for Developmental
Iruma-gun, Saitama, Japan Biology
Utrecht, The Netherlands
D juro Josic Present status
Research Scientist Department of Biochemistry
Institut feur Molekularbiologie University of Nÿmegen
Freie Universitaet Berlin Nymegen, The Netherlands
Berlin, Germany
H erm an K rap f
K rishna K. Kalghatgi Product Group Manager
Research Scientist Separation Products Division
Department of Chemical Engineering Pharmacia/LKB
Yale University Biotechnology AB
New Haven, Connecticut Uppsala, Sweden

B arry L. K arger Michael K unitani

Professor and Director Senior Scientist and Director
Chemistry Department and Barnett Department of Analytical
Institute Chemistry
Northeastern University Cetus Corp., Inc.
Boston, Massachusetts Emeryville, California
Henk H. L auer M ary B. LoPresti
Lauer Labs Associate in Research
Emmen, The Netherlands Department of Molecular Biophysics and
Biochemistry
Claude Lazure Yale University
Associate Director New Haven, Connecticut
Laboratory of Molecular
Neuroendocrinology Colin T. M ant
Clinical Research Institute of Montreal Research Associate
Montreal, Quebec, Canada Department of Biochemistry
University of Alberta
Kok K. Lee Edmonton, Alberta, Canada
Postdoctoral Fellow
Synthetic Peptides, Inc. H eidrun M atern
Edmonton, Alberta, Canada Ph.D.
Department of Internal Medicine III
M aryann Lee Aachen University of Technology
Staff Researcher II Aachen, West Germany
Department of Pharmaceutical Analysis I
Syntex Research Siegfried M atern
Palo Alto, California Professor of Medicine, Director
Department of Internal Medicine III
Albert Light Aachen University of Technology
Professor Aachen, West Germany
Department of Chemistry
Purdue University R ichard A. McClintock
West Lafayette, Indiana Research Assistant
The Clayton Foundation Laboratories for
Susanne Linde Peptide Biology
Staff Scientist The Salk Institute
Department for Protein Chemistry La Jolla, California
Hagedom Research Laboratory
Gentofte, Denmark Stephen M oring
Research and Development Chemist
M argaret A. LIndorfer Department of Research and
Senior Research Associate Development
Institute of Molecular Biology Applied Biosystems, Inc.
University of Oregon Foster City, California
Eugene, Oregon
R obert L, M oritz
K aren L ockhart Senior Research Officer
Supervisor Joint Protein Structure Laboratory
Department of Pharmaceutical Analysis 1 Ludwig Institute for Cancer Research
Syntex Research (Melbourne Branch)
Palo Alto, California and
The Walter and Eliza Hall Institute for
D. C. Lommen Medical Research
LC Resources, Inc. P. O. Royal Melbourne Hospital
Lafayette, California Parkville, Victoria, Australia
Michael N attriss J. M. R obert P ark er
Instrument Technologist Research Associate
Department of Biochemistry Department of Biochemistry
MRC Group in Protein Structure and University of Alberta
Function Edmonton, Alberta, Canada
Edmonton, Alberta, Canada
M atthew V. Piserchio
Sai M. Ngal Keystone Scientific, Inc.
Graduate Student Bellefonte, Pennsylvania
Department of Biochemistry
University of Alberta T erry M. Phillips
Edmonton, Alberta, Canada Director
Department of Immunochemistry
George Washington University
Cecilia Nguyen
Medical Center
Chemist II
Washington, D.C.
Department of Pharmaceutical
Analysis I
F ran k W. Putnam
Syntex Research
Distinguished Professor Emeritus
Palo Alto, California
Department of Biology
Indiana University
R. Nieuwland Bloomington, Indiana
Graduate Student
Hubrecht Laboratory R ichard R einhardt
Netherlands Institute for Developmental Senior Scientist
Biology Max-Planck-Institute for Molecular
Utrecht, The Netherlands Genetics
Present status Berlin, Germany
Research Associate
Department of Haematology W erner R eutter
Academic Hospital Utrecht Professor
Utrecht, The Netherlands Institut feur Molekularbiologie
Freie Universitaet Berlin
M ark P« Nowlan Berlin, Germany
Senior Research Scientist
SynChrom, Inc. Jean E. Rivier
Lafayette, Indiana Professor
The Clayton Foundation Laboratories
for Peptide Biology
K erry D. Nugent
The Salk Institute
President
La Jolla, California
Michrom Bioresources
Pleasanton, California
Jam es A. Rochemont
Research Associate
John M. O stresh Department of Biochemical
Director of Research Neuroendocrinology
Multiple Peptide Systems Clinical Research Institute of Montreal
San Diego, California Montreal, Quebec, Canada
Timothy D. Schlabach K athryn L. Stone
Product Manager Lab Manager
Spectra-Physics Department of Molecular Biophysics and
San Jose, California Biochemistry
Yale University
Nabil G. Seidah New Haven, Connecticut
Director
Department of Biochemical Nobuhiro Takahashi
Neuroendocrinology Chief Scientist
Clinical Research Institute of Montreal Department of Life Science
Montreal, Quebec, Canada Toa Nenryo Kogyo K. K.
Iruma-gun, Saitama, Japan
Paul Semchuk
Technologist I Geza Toth
Department of Biochemistry Research Associate
University of Alberta Department of Chemistry
Edmonton, Alberta, Canada University of Arizona
Tucson, Arizona
William Shalongo
Department of Biochemistry Klaus K. Unger
University of Iowa Professor
Iowa City, Iowa Institut für Anorganische and Analytische
Chemie
R ichard J. Simpson Johannes Gutenberg-Universität
Head Mainz, Federal Republic of Germany
Joint Protein Structure Laboratory
Ludwig Institute for Cancer Research Jennifer E. Van Eyk
(Melbourne Branch) Ph.D. candidate
and Department of Biochemistry
The Walter and Eliza Hall institute for University of Alberta
Medical Research Edmonton, Alberta, Canada
P. O. Royal Melbourne Hospital
Parkville, Victoria, Australia A. J. M. van den Eijnden-van Raaij
Doctor
Lawrence B. Smillie Hubrecht Laboratory
Professor Netherlands Institute for Developmental
Department of Biochemistry Biology
University of Alberta Utrecht, The Netherlands
Edmonton, Alberta, Canada
E. J. J. van Zoelen
Lloyd R. Snyder Doctor
Vice President Hubrecht Laboratory
LC Resources, Inc. Netherlands Institute for Developmental
Orinda, California Biology
Utrecht, The Netherlands
E arle Stellwagen Present status
Professor Professor
Department of Biochemistry Department of Cell Biology
University of Iowa University of Nymegen
Iowa City, Iowa Nymegen, The Netherlands
M. Verzele K .J . Wilson
Professor Vice President, General Manager
Laboratory of Organic Chemistry Applied Biosystems
University of Gent Foster City, California
Gent, Belgium
Jam es L. W ittliff
C. Timothy W ehr Professor
Manager, Chemistry R & D Hormone Receptor Laboratory
High Performance Electrophoresis University of Louisville
Bio-Rad Laboratories Louisville, Kentucky
Richmond, California
Shiaw-Lin W u
Benny S, W elinder Scientist
Head Department of Medicinal and Analytical
Department for Protein Chemistry Chemistry
Hagedom Research Laboratory Genentech, Inc.
Gentofte, Denmark San Francisco, California

G jalt W. Welling Paul Zavitsanos

Senior Research Scientist Application Engineer
Department of Medical Microbiology Hewlett-Packard
Rijksuniversiteit Groningen Montreal, Canada
Groningen, The Netherlands
Nian E. Zhou
Sytske W elling-W ester Postdoctoral Fellow
Senior Research Scientist Department of Biochemistry
Department of Medical Microbiology University of Alberta
Rijksuniversiteit Groningen Edmonton, Alberta, Canada
Groningen, The Netherlands
Lynne R. ZIeske
Ronald D. Wiehle Scientist
Research Associate Department of Research and Development
Hormone Receptor Laboratory Applied Biosystems, Inc.
University of Louisville Foster City, California
Louisville, Kentucky
G. J. T. Zw artkruis
Kenneth R. Williams Research Associate
Professor (Adj.) of Research Hubrecht Laboratory
Department of Molecular Biophysics and Netherlands Institute for Developmental
Biochemistry Biology
Yale University Utrecht, The Netherlands
New Haven, Connecticut
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