INTERNATIONAL UNION OF BIOCHEMISTRY SYMPOSIUM SERIES Vol. 1. The Origin of Life on the Earth—A, I. OPARIN et al. (Editors) Vol. 2. Enzyme Chemistry: Proceedings of the International Symposium in Tokyo- Kyoto PROCEEDINGS OF THE FOURTH INTERNATIONAL CONGRESS ON BIOCHEMISTRY VIENNA, 1958 Vol. 3. (I) Carbohydrate Chemistry of Substances of Biological Interest Vol. 4. (II) Biochemistry of Wood Vol. 5. (Ill) Biochemistry of the Central Nervous System Vol. 6. (IV) Biochemistry of Steroids Vol. 7. (V) Biochemistry of Antibiotics Vol. 8. (VI) Biochemistry of Morphogenesis Vol. 9. (VII) Biochemistry of Viruses Vol. 10. (VIII) Proteins Vol. 11. (IX) Physical Chemistry of High Polymers of Biological Interest Vol. 12. (X) Blood Clotting Factors Vol. 13. (XI) Vitamin Metabolism Vol. 14. (XII) Biochemistry of Insects Vol. 15. (ΧΙΠ) Colloquia Vol. 16. (XIV) Transactions of the Plenary Sessions Vol. 17. (XV) Biochemistry Vol. 18. Biochemistry of Lipids—G. POPJÁK (Editor) Vol. 19. Haematin Enzymes (Parts 1 and 2)—J. E. FALK, R. LEMBERG and R. K. MORTON Vol. 20. Report of the Commission on Enzymes, 1961 (I.U.B.) PROCEEDINGS OF THE HFTH INTERNAΉONAL CONGRESS ON BIOCHEMISTRY MOSCOW, 1961 (Provisional titles) Vol. 21. (I) Biological Structure and Function at the Molecular Level Vol. 22. (II) Functional Biochemistry of Cell Structures Vol. 23. (Ill) Evolutionary Biochemistry Vol. 24. (IV) Molecular Basis of Enzyme Action and Prohibition Vol. 25. (V) Intracellular Respiration: Phosphorylating and Non-Phosphorylating Systems Vol. 26. (VI) Mechanism of Photosynthesis Vol. 27. (VII) Biosynthesis of Lipids Vol. 28. (VIII) Biochemical Principles of the Food Industry Vol. 29. (IX) Transactions of the Plenary Sessions Vol. 30. (X) Abstracts of Papers and Indexes to the Volumes of the Proceedings The building of the Australian Academy of Science, Canberra, where the Symposium was held. H A E M A T IN E N Z Y M ES A SYMPOSIUM OF THE INTERNATIONAL UNION OF BIOCHEMISTRY ORGANIZED BY THE AUSTRALIAN ACADEMY OF SCIENCE CANBERRA 1959 Edited by J. E. FALK, R. LEMBERG and R. K. MORTON PART 1 (Pages 1 to 362) SYMPOSIUM PUBLICATONS DIVISION P E R G A M ON P R E SS OXFORD · LONDON · N EW YORK · PARIS 1961 PERGAMON PRESS LTD. Headington Hill Hall, Oxford 4 & 5 Fitzroy Square, London W,l PERGAMON PRESS INC. 122 East 55th Street, New York 22, N. Y, 1404 New York Avenue N.W., Washington 5 D.C, Statler Center 640, 900 Wilshire Boulevard, Los Angeles 17, California PERGAMON PRESS S.A.R.L. 24 Rue des Eco les, Paris PERGAMON PRESS G.m.b.H. Kaiserstrasse 75, Frankfurt am Main Copyright © 1961 PERGAMON PRESS LTD. LIBRARY OF CONGRESS CARD NUMBER 60-53463 Set in Monotype Times 10ll2pt and Printed in Great Britain by the Pitman Press, Bath PREFACE Tffls volume contains the papers and discussion material presented at the Symposium on Haematin Enzymes. It was held at Canberra between 31st August and 4th September, 1959, and was organized by the Australian Academy of Science for the International Union of Biochemistry. The Symposium was arranged for the Academy by a Conmiittee com­ prising A. H. Ennor, J. Ε. Falk, R. Lemberg and R. K. Morton (Convener). The Committee is grateful to several organizations, cited in the address by Dr. R. Lemberg, President of the Symposium, for financial and other assistance. The titles and addresses of participants are given on pp. xvii-xx. For convenience, titles have been omitted from the scientific communications. It is with profound regret that we record the untimely death in October, 1960, of Professor Enzo Boeri, one of the distinguished participants in the Symposium. He made many notable contributions to our knowledge of haematin enzymes and he will be remembered with admiration, respect and affection by all who were privileged to know him. R. K. MORTON PARTICIPANTS DR. C. A. ApPLEBY Biochemistry Section, Division of Plant Industry, C.S.I.R.O., Canberra, Australia. MR. J. McD. ARMSTRONG DepartmentofAgriculturalChemistry, Waite Agricultural Research Institute, University of Adelaide, Adelaide, Australia. MR. J. BARRETI Institute of Medical Research, Royal North Shore Hospital, Sydney, Australia. DR. N. K. BoARDMAN Biochemistry Section, Division of Plant Industry, C.S.I.R.O., Canberra, Australia. PROFESSOR E. BOERI· Institute of Human Physiology, University ofFerrara, Ferrara, Italy. DR. W. D. BONNER Johnson Research Foundation, University of Pennsylvania, Philadelphia, U.S.A. MME. PROFESSOR P. CHAIX Laboratory of Biological Chemistry, Univer sity ofParis, Paris, France. PROFESSOR B. CHANCE Johnson Foundation for Medical Physics, University of Pennsylvania, Philadelphia, U.S.A. DR. P. S. CLEZY Institute of Medical Research, Royal North Shore Hospital, Sydney, Australia. DR. E. CUTOLO Italian Serum Research Institute, Naples, Italy. PROFESSOR F. DICKENS Courtauld Institute of Biochemistry, Middle sex Hospital Medical School, University of London, London, England. PROFESSOR D. L. DRABKIN DepartmentofBiochemistry,GraduateSchool of Medicine, University of Pennsylvania, Philadelphia, U.S.A. PROFESSOR F. P. DWYER The John Curtin School ofMedical Research, Australian National University, Canberra, Australia. • Died. 1960. H.E.-VOL. 1-8 xvii xviii PARTICIPANTS PROFESSOR F. EGAMI Department of Biophysics and Biochemistry, Faculty of Science, University of Tokyo, Tokyo, Japan. DR. R. W. ESTABROOK Johnson Foundation for Medical Physics, University of Pennsylvania, Philadelphia, U.S.A. DR. J. E. FALK Biochemistry Section, Division of Plant Industry, C.S.I.R.O., Canberra, Australia. PROFESSOR P. GEORGE John Harrison Laboratory of Chemistry, University of Pennsylvania, Philadelphia, U.S.A. DR. R. W. HENDERSON Department of Biochemistry, University of Melbourne, Melbourne, Australia. DR. T. HORIO Graduate Department of Biochemistry, Brandeis University, Waltham, U.S.A. PROFESSOR M. D. KAMEN Graduate Department of Biochemistry, Brandeis University, Waltham, U.S.A. PROFESSOR K. KAZIRO Biochemical Laboratory, Nippon Medical School, Tokyo, Japan. MR. J. W. LEGGE Department of Biochemistry, University of Melbourne, Melbourne, Australia. DR. R. LEMBERG, Institute of Medical Research, Royal North Prof. a. D. (Heidelberg) Shore Hospital, Sydney, Australia. MR. W. H. LOCKWOOD Institute of Medical Research, Royal North Shore Hospital, Sydney, Australia. DR. G. D. LUDWIG Hospital of the University of Pennsylvania, Philadelphia, U.S.A. DR. E. MARGOLIASH Laboratory for the Study of Hereditary and Metabolic Disorders, College of Medicine, University ofUtah, Salt Lake City, U.S.A. DR. D. B. MORELL Institute of Medical Research, Royal North ShoreHospital,Sydney,Australia. DR. M. MORRISON Department of Biochemistry, School of Medicine and Dentistry, University of Rochester, Rochester, U.S.A. PARTICIPANTS PROFESSOR R. K. MORTON Department ofAgricultural Chemistry, Waite Agricultural Research Institute, University of Adelaide, Adelaide, Australia. DR. F. J. Moss School of Biological Sciences, University of New South Wales, Sydney, Australia. PROFESSOR J. B. NEILANDS Department of Biochemistry, University of California, Berkeley, U.S.A. DR. R. A. NEVE Department of Biochemistry, University of California, Berkeley, U.S.A. DR. A. P. NYGAARD Nutrition Institute, Blindern, Oslo, Norway. DR. Y. OGURA Department of Biophysics and Biochemistry, Faculty of Science, University of Tokyo, Tokyo, Japan. DR. J. E. O'HAGAN Red Cross Blood Transfusion Service, Brisbane, Australia. DR. L. E. ORGEL University Chemical Laboratory, University ofCambridge, Cambridge, England. DR. S. PALEUS Biochemistry Department, Nobel Institute of Medicine, Stockholm, Sweden. DR. D. D. PERRIN Department of Medical Chemistry, John CurtinSchoolofMedicalResearch,Australian National University, Canberra, Australia. DR. J. N. PHILLIPS Biochemistry Section, Division of Plant Industry, C.S.I.R.O., Canberra, Australia. DR. J. POSTGATE Microbiological Research Establishment, Porton, Wiltshire, England. Assoc. PROFESSOR Department of Biochemistry, University of W. A. RAWLINSON Melbourne, Melbourne, Australia. PROFESSOR E. C. SLATER Laboratory of Physiological Chemistry, University of Amsterdam, Amsterdam, The Netherlands. DR. L. SMITH Department of Biochemistry, Dartmouth Medical School, Hanover, New Hampshire, U.S.A. DR. C. F. STRIITMATIER Department ofBiologicalChemistry, Harvard Medical School, Boston, U.S.A. DR. A. TISSIERES The Biological Laboratories, Harvard Univer sity, Cambridge, U.S.A. DR. P. TRUDINGER Biochemistry Section, Division of Plant Industry, C.S.I.R.O., Canberra, Australia. PROFESSOR S. F. VELICK School of Medicine, Washington University, St. Louis, Missouri, U.S.A. PROFESSOR W. W. WAINIO Bureau of Biological Research and Depart mentofPhysiologyandBiochemistry,Rutgers, The State University, New Brunswick, New Jersey, U.S.A. PROFESSOR J. H. WANG Department of Chemistry, Yale University, New Haven, U.S.A. DR. R. J. P. WILLIAMS Inorganic Chemistry Laboratory, and Wad ham College, Oxford, England. DR. M. E. WINFIELD Division of Physical Chemistry, Indus trial Chemical Laboratories, C.S.I.R.O., Melbourne, Australia. PRESroENHAL ADDRESS I DECLARE open the Symposium on Haematin Enzymes of the International Union of Biochemistry and welcome all who attend it. It is a great pleasure for me to welcome the many distinguished scientists from overseas who have come to Austraha to discuss with us the problems which interest us all. I hope that the stimulation which you may receive will repay you for your long and strenuous journeys to our distant shores, that you may carry back happy memories of this week spent in Austraha—^and that you will return! Your visit will certainly be a stunulus to Austrahan science. I am particularly happy to receive you in this home of the Austrahan Academy of Science.* When we discussed plans for its erection in the Council of the Academy only a few years ago, I Uttle dreamt that I should have the honour of opening the first International Conference in it. Our thanks are due to the International Union of Biochemistry, not only for accepting our invitation to hold the Symposium, but also for financial support; to the Austrahan Commonwealth Government; to the Wellcome Trust; and to the various Academies and national bodies, particularly to the National Science Foundation of the United States and to the Royal Society. The support of these various organizations made this meeting possible. We decided that we wanted an intimate symposium in which all participants could be rehed upon to make valuable contributions. We wanted to discuss our problems critically, but with some degree of leisure. This is a Symposium and even if we cannot do as the Greeks did, having a meal, wine and dancers in this hall, the comfortable lounge chahrs are the nearest approach to it possible in these hurried times. There you may recline for a httle snooze if the richness of the intellectual feast endangers your mental digestion. Our Symposium has a pecuhar note in that it calls together scientists of different branches, from quantum mechanics to microbiology, and asks them to direct the spothghts of their knowledge on to a comparatively narrow field, but a field of great biological importance and chemical interest. After reading the prepubhshed papers I am convinced that we were right in assuming that the difiiculties of finding sufficient common ground for our various denomin­ ations are no longer insuperable. Still, we shall have to exert some patience and forbearance. I ask the theorist not to be impatient with the experimenter, if he asks questions which reveal his lack of knowledge of theory, but to answer them in * See frontispiece (Editors), xxi