Handbook of Biochemistry and Molecular Biology Handbook of Biochemistry and Molecular Biology 3rd Edition I Proteins -Volume EDITOR Gerald D. Fasn1an, Ph. D. Rosenfield Professor of Biochemistry Graduate Department of Biochemistry Brandeis University Waltham, Massachusetts Boca Raton London New York CRC Press is an imprint of the Taylor & Francis Group, an informa business First published 1976 by CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 Reissued 2018 by CRC Press © 1976 by CRC Press, Inc. © 1970, 1968 by The Chemical Rubber Co. CRC Press is an imprint of Taylor & Francis Group, an Informa business No claim to original U.S. Government works 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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Fasman, Ph. D. Rosenfield Professor of Biochemistry Graduate Department of Biochemistry Brandeis University Waltham, Massachusetts The following is a list of the four major sections of the Handbook, each consisting of one or more volumes Ptoteins - Amino Acids, Peptides, Polypeptides, and Proteins Nucleic, Acids — Purines, Pyrimidines, Nucleotides, Oligonucleotides, tRNA, DNA, RNA Lipids, Carbohydrates, Steroids Physical and Chemical Data, Miscellaneous - Ion Exchange, Chromatog raphy, Buffers, Miscellaneous, e.g., Vitamins ADVISORY BOARD Gerald D. Fasman Editor Herbert A. Sober (deceased) Consulting Editor MEMBERS Bruce Ames Victor Ginsburg Professor, Department of Biochemistry Chief, Biochemistry Section, National University of California Institute of Arthritis, Metabolism and Berkeley, California 94720 Digestive Diseases Department of Health, Education, and Sherman Beychok Welfare Professor, Department of Biological National Institutes of Health Sciences Bethesda, Maryland 20014 Columbia University New York, New York 10027 Walter Gratzer MRC Neurobiology Unit Waldo E. Cohn Department of Biophysics Senior Biochemist, Biology Division Kings College Oak Ridge National Laboratory University of London Oak Ridge, Tennessee 37830 London England Harold Edelhoch National Institute Arthritis, Metabolism Lawrence Grossman and Digestive Diseases Department of Health, Education, and Professor, Department of Biochemical and Welfare Biophysical Sciences National Institutes of Health School of Hygiene and Public Health The Johns Hopkins University Bethesda, Maryland 20014 Baltimore, Maryland 21205 John Edsall Professor Emeritus, Biological Laboratories Frank Gurd Harvard University Professor, Department of Chemistry Cambridge, Massachusetts 02138 Indiana University Bloomington, Indiana 47401 Gary Felsenfeld Chief, Physical Chemistry Laboratory Laboratory of Molecular Biology William Harrington National Institute of Arthritis, Professor, Department of Biology Metabolism, and Digestive Diseases The Johns Hopkins University National Institutes Of Health Baltimore, Maryland 21218 Bethesda, Maryland 20014 William P. Jencks Edmond H. Fischer Professor, Graduate Department of Professor, Department of Biochemistry Biochemistry University of Washington Brandeis University Seattle, Washington 98195 Waltham, Massachusetts 02154 ADVISORY BOARD (continued) 0. L. Kline Julius Marmur Executive Officer Professor, Department of Biochemistry American Institute of Nutrition and Genetics 9650 Rockville Pike Albert Einstein College of Medicine Bethesda, Maryland 20014 New York, New York 10461 1. M. Klotz Alton Meister Professor, Department of Chemistry Professor, Department of Biochemistry Northwestern University Cornell University Medical College Evanston, Illinois 60201 New York, New York 10021 Robert Langridge Kivie Moldave Professor, Department of Biochemistry Princeton University Professor, Department of Biochemistry Princeton, New Jersey 08540 California College of Medicine University of California Irvine, California 92664 Philip Leder Chief, Laboratory of Molecular Genetics National Institute of Child Health D. C. Phillips and Human Development Professor, Laboratory of Molecular National Institutes of Health Biophysics Bethesda, Maryland 20014 Department of Zoology Oxford University I. Robert Lehman Oxford Professor, Department Biochemistry England School of Medicine Stanford University William D. Phillips Stanford, California 94305 The Lord Rank Research Centre Ranks Hove, McDougall Ltd. Lawrence Levine Lincoln Road, High Wycombe Professor, Graduate Department of Bucks Biochemistry England Brandeis University Waltham, Massachusetts 02154 G. N. Ramachandran John Lowenstein Professor, Molecular Biophysics Unit Professor, Graduate Department of Indian Institute of Science Biochemistry Bangalore Brandeis University India Waltham, Massachusetts 02154 Michael Sela Emanuel Margoliash Professor, Department of Chemical Professor, Department of Biological Immunology Sciences The Weizmann Institute of Science Northwestern University Rehovot Evanston, Illinois 60201 Israel ADVISORY BOARD (continued) Waclaw Szybalski Ignacio Tinoco, Jr. Professor, McArdle Laboratory for Professor, Department of Chemistry Cancer Research University of California The University of Wisconsin Berkeley, California 94720 Madison, Wisconsin, 53706 Bert L. Valiee Serge N. Timasheff Professor, Biophysics Research Professor, Graduate Department of Laboratory Biochemistry Peter Bent Brigham Hosoital Brandeis University Harvard Medical School Waltham, Massachusetts 02154 Boston, Massachusetts 02115 PREFACE The rapid pace at which new data is currently accumulated in science presents one of the significant problems of today — the problem of rapid retrieval of information. The fields of biochemistry and molecular biology are two areas in which the information explosion is manifest. Such data is of interest in the disciplines of medicine, modern biology, genetics, immunology, biophysics, etc., to name but a few related areas. It was this need which first prompted CRC Press, with Dr. Herbert A. Sober as Editor, to publish the first two editions of a modern Handbook of Biochemistry, which made available unique, in depth compilations of critically evaluated data to graduate students, post-doctoral fellows, and research workers in selected areas of biochemistry. This third edition of the Handbook demonstrates the wealth of new information which has become available since 1970. The title has been changed to include molecular biology; as the fields of biochemistry and molecular biology exist today, it becomes more difficult to differentiate between them. As a result of this philosophy, this edition has been greatly expanded. Also, previous data has been revised and obsolete material has been eliminated. As before, however, all areas of interest have not been covered in this edition. Elementary data, readily available elsewhere, has not been included. We have attempted to stress the areas of today’s principal research frontiers and consequently certain areas of important biochemical interest are relatively neglected, but hopefully not totally ignored. This third edition is over double the size of the second edition. Tables used from the second edition without change are so marked, but their number is small. Most of the tables from the second edition have been extensively revised, and over half of the data is new material. In addition, a far more extensive index has been compiled to facilitate the use of the Handbook. To make more facile use of the Handbook because of the increased size, it has been divided into four sections. Each section will have one or more volumes. The four sections are titled: Proteins — Amino Acids, Peptides, Polypeptides, and Proteins Nucleic Acids — Purines, Pyrimidines, Nucleotides, Oligonucleotides, tRNA, DNA, RNA Lipids, Carbohydrates, Steroids Physical and Chemical Data, Miscellaneous - Ion Exchange, Chromatography, Buffers, Miscellaneous, e.g., Vitamins By means of this division of the data, we can continuously update the Handbook by publishing new data as they become available. The Editor wishes to thank the numerous contributors, Dr. Herbert A. Sober, who assisted the Editor generously, and the Advisory Board for their counsel and cooperation. Without their efforts this edition would not have been possible. Special acknowledgments are due to the editorial staff of CRC Press, Inc., particularly Ms. Susan Cubar Benovich, Ms. Sandy Pearlman, and Mrs. Gayle Tavens, for their perspicacity and invaluable assistance in the editing of the manuscript. The editor alone, however, is responsible for the scope and the organization of the tables. We invite comments and criticisms regarding format and selection of subject matter, as well as specific suggestions for new data (and their sources) which might be included in subsequent editions. We hope that errors and omissions in the data that appear in the Handbook will be brought to the attention of the Editor and the publisher. Gerald D. Fasman Editor August 1975