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FOREWORD R. KEITH CANNAiNT Chairman, Division of Medical Sciences National Academy of Sciences-National Research Council "The subject of this book is the red blood of vertebrates." With this sentence Lawrence J. Henderson introduced his classical monograph entitled "Blood: A Study in General Physiology." He then proceeded to justify his choice of subtitle by giving the following description of his purpose: "We shall study this substance as a physico-chemical system and as a tissue, seeking in its properties the exemplification of the general properties of protoplasm. In its physiological function and relation with other parts of the body we shall look for an illustration of organic integration and adaptation. We shall also study it compara- tively, from species to species, in rest and activity, in health and disease. So far as is possible these studies will be quantitative and mathematical." It was Claude Bernard who gave to general physiology its first charter. He cc.nceived it as the science of "the elementary condition of the phenomena of life" and its goal the description of this elementary condition in the language of physical science. To Bernard we also owe the concept of blood as the "milieu interieur" of the body. It is a happy circumstance of history, therefore, that a succession of general physiologists- Bohr, Krogh, Haldane, Barcroft, Van Slyke, Hen- derson, and others too numerous to name have found in blood the biological system most accessible to controlled investigation and most amenable to exacting physico-chemical description. Prom their world we have inherited those quantitative and mathematical descriptions of the respiratory, osmotic, and ionic functions of the blood that so elegantly illustrate the harmonious organization and integration of components that we have learned to regard as the special characteristics of living systems in general. The subject of the Conference of which this book is the record is also "the red blood of vertebrates." It is true that attention was deliberately re- stricted to a single component of blood. This, however, may be defended as a logical and appropriate refinement of approach not only because hemoglobin is the predominant constituent of the system but also because its own unique . v
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V1 FOREWORD properties so largely determine the characteristics of some of the major physiological functions of blood. The purpose of the Conference was to . ~ refine the physico chemical characterization of the various forms of hemo- g]obin as a contribution to a better understanding of the "elementary con- dition" of blood as the internal environment of the body. In this sense the Conference is offered as an exercise in general physiology. Ten years ago a Symposium on Hemoglobin was held in Cambridge, England, as a memorial to Joseph Barcroft. Its proceedings were duly pub- lished in a notable volume. It is our hope that the papers and discussions recorded in the book which is now in your hands have recaptured something of the spirit and the zest of the earlier meeting. In the intervening years much progress has been made. Experimental tools that were only beginning to attract attention ten years ago are now the standard equipment of the protein chemist. Concepts which were, at that time, novel and hypothetical are now the accepted foundation for current ideas of the structure and interactions of hemoglobin. New experimental devices have opened up new avenues of investigation and new knowledge has posed new questions. The center of interest continues to shift. In the last fees years the discovery that the hemoglobin of a single species may exist in a variety of related natural forms has stimulated wide interest in the chemical nature of these "abnormal" molecules and in their genetic and clinical signifi- cance. This new interest is reflected in the pages that follow. In one of his reports on his work on the respiratory function of the blood, Barcroft reflected wistfully that the introduction of his refined differential manometric methods for the analysis of the blood gases had served only "to make more certain the uncertainty of our position." It may be true that hemo- globin has been more precisely characterized from a physico-chemical point of view than any other protein. However, as we dwell with satisfaction on the deepening invasion of the territory of biology by the physical sciences, it may be well to remind ourselves that action and reaction are equal and opposite and to recall the admonition of A. V. Hill that "physics and chem- istry will only dominate biology by becoming biology."