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Chemical Specificity of Nucleic Acids and Mechanism of Their Enzymatic Degradation 1

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Chemical Specificity of Nucleic Acids and Mechanism of Their Enzymatic Degradation 1 EXPERIENTIA Vol. VI - Fasc. 6 Pag. 201-240 15. VI. 1950 Chemical Specificity of Nucleic Acids and Mechanism of their Enzymatic Degradation 1 By ERWIN CHARGAFF 2, New York, N.Y. I. Introduction abounds more than the chemistry of the cell. Since an The last few years have witnessed an enormous re- ounze of proof still weighs more than a pound of vival in interest for the chemical and biological proper- prediction, the important genetical functions, ascribed • ties of nucleic acids, which are components essential for --probably quite rightly--to the nucleic acids by the life of all cells. This is not particularly surprising, many workers, will not be discussed here. Terms such as the chemistry of nucleic acids represents one of the as "template" or "matrix" or "reduplication" will not remaining major unsolved problems in biochemistry. be found in this lecture. It is not easy to say what provided the impulse for this rather sudden rebirth. Was it the fundamental work of II. Identity and Diversity iu High Molecular E. HAMMARSTEN3 on the highly polymerized desoxy- Cell Constituents ribonucleic acid of calf thymus ? Or did it come from The determination of the constitution of a compli- the biological side, for instance the experiments of cated compound, composed of many molecules of a BRACHE'ra and CASPERSSON5 ? Or was it the very im- number of organic substances,• evidently requires the portant research of AVERY ~ and his collaborators on exact knowledge of the nature and proportion of all the transformation of pneumococcal types that started constituents. This is true for nucleic acids as much as the avalanche ? for proteins or polysaccharides. It is, furthermore, It is, of course, completely senseless to formulate clear that the value of such constitutional determin- a hierarchy of cellular constituents and to single out ations will depend upon the development of suitable certain compounds as more important than others. methods of hydrolysis. Otherwise, substances repre- The economy of the living cell probably knows no senting an association of many chemical individuals conspicuous waste; proteins and nucleic acids, lipids can be described in a qualitative fashion only; precise and polysaccharides, all have the same importance. decisions as to structure remain impossible. When our But one observation may be offered. It is impossible to laboratory, more than four years ago, embarked upon write the history of the cell without considering its the study of nucleic acids, we became aware of this geography; and we cannot do this without attention to difficulty immediately. what may be called the chronology of the cell, i. e. the The state of the nucleic acid problem at that time sequence in which the cellular constituents are laid found its classical expression in LEVENE'S monograph 1. down and in which they develop from each other. If (A number of shorter reviews, indicative of the develop- this is done, nucleic acids will be found pretty much at ment of our conceptions concerning the chemistry of the beginning. An attempt to say more leads directly nucleic acids, should also be mentioned2.) The old into empty speculations in which almost no field tetranucleotide hypothesis-it should never have been 1 This article is based on a series of lectures given before the called a theory--was still dominant; and this was Chemical Societies of Ziirich and Basle (June ~29th and 30th, 1949), characteristic of the enormous sway that the organic the Soci~tfi de chimie biologique at Paris, and the Universities of chemistry of small molecules held over biochemistry. Uppsala, Stockholm, and Milan. Department of Biochemistry, College of Physicians and Sur- I should like to illustrate what t mean by one example. geons, Columbia University, New York. The author wishes to thank If in the investigation of a disaccharide consisting of the John Simo~ Guggenheim 3Iemorial Fo~ndatio~ for making pos- two different hexoses we isolate 0.8 mole of one sugar sible his ½tay in Europe. The experimental work has been supported by a research grant from the United States Public Health Service. and 0.7 mole of the other, this will be sufficient for the 3 E.HAMMARSTEN, Biochem. Z. 144, :183 (1924). 4 J. BRACHET in Nucleic Acid, Symposia See. Exp. Biol. No. i 1 P.A. LEVENE and L.W.BAss, Nucleic Acids (Chemical Catalog (Cambridge University Press, 1947), p. 207. Cp. J. BRACltET, in Co., New York, 1931). Nucleic Acids and Nucleoproteins, Cold Spring Harbor Syrup. Quant. H. BRED~RECK, Fortschritte der Chemic organischer Naturstoffe Biol. 12, 18. (Cold Spring Harbor, N.Y., 1947). 1, 1~21 (1938). - F.G. FISCHER, Naturwissenseh. 30, 377 (19.12). - s T.CAsPERSSON, in Nucleic Acid, Syrup. Soc. Exp. Biol., No, 1 R. S.TIPsON, Adv. Carbohydrate Chenl. 1, 193 (1945). - J. M. GUL- (Cambridge University Press, 1947), p. 127. LAND, G. R. BARKER, and D. O. JoRDAn, Ann. Rev. Bioehem. 14, * O. T. AVERY, C. M. MAcLEoD, and M. MeCARTY, J. Exp. Med. 175 (1945). - E.CHARGAFF and E.VlscltER, Ann. Rev. Biocbem. 17, 79, 137 (1944). ~01 (1948). - F. SCHLENK, Adv. Enzymol. 9, ,155 (1949). 202 E. CHARGAFF" Chemical Specificity of Nucleic Acids and Mechanism of their Enzymatic Degradation [EXPERtF.NTIA VOL. VI]6~ recognition of the composition of the substance, pro- primitive in creating in a cell, such as the tubercle vided its molecular weight is known. The deviation of bacillus, a host of novel compounds, new fatty acids, the analytical results horn simple, integral proportions alcohols, etc., that are nowhere else encountered. is without importance in that case, But this will not There, the recognition of chemical peculiarities is hold for high-molecular compounds in which variations relatively easy. But in the case of the proteins and in the proportions of their several components often nucleic acids, I believe, nature has acted most subtly; will provide the sole indication of the occurrence of and the task facing us is much more difficult. There is different compounds. nothing more dangerous in the natural sciences than In attempting to formulate the problem with some to look for harmony, order, regularity, before the exaggeration one could say: The validity of the identi- proper level is reached. The harmony of cellular life fication of a substance by the methods of classical may well appear chaotic to us. The disgust for the organic chemistry ends with the mixed melting point. amorphous, the ostensibly anomalous--an interesting When we deal with the extremely complex compounds problem in the psychology of science--has produced of cellular origin, such as nucleic acids, proteins, or many theories that shrank gradually to hypotheses polysaccharides, a chemical comparison aiming at the and then vanished. determination of identity or difference must be based We must realize that minute changes in the nucleic on the nature and the proportions of their constituents, acid, e. g. the disappearance of one guanine molecule on the sequence in which these constituents are ar- out of a hundred, could produce far-reaching changes ranged in the molecule, and on the type and the po- in the geometry of the conjugated nucleoprotein; and sition of the linkages that hold them together. The it is not impossible that rearrangements of this type smaller the number of components of such a high- are among the causes of the occurrence of mutations 1. molecular compound is, the greater is the difficulty of The molecular weight of the pentose nucleic acids, a decision. The occurrence of a very large number of especially of those from animal tissue cells, is not yet different proteins was recognized early; no one to my known; and the problem of their preparation and knowledge ever attempted to postulate a protein as a homogeneity still is in a very sad state. But that the compound composed of equimolar proportions of 18 desoxypentose nucleic acids, prepared under as mild or 20 different amino acids. In addition, immunological conditions as possible and with the avoidance of investigations contributed very much to the recognition enzymatic degradation, represent fibrous structures of of the multiplicity of proteins. A decision between high molecular weight, has often been demonstrated. identity and difference becomes much more difficult No agreement has as yet been achieved on the order of when, as is the case with the nucleic acids, only few magnitude of the molecular weight, since the inter- primary components are encountered. And when we pretation of physical measurements of largely asym- finally come to high polymers, consisting of one metric m~lecules still presents very great difficulties. component only, e. g. glycogen or starch, the charac- But regardless of whether the desoxyribonucleic acid terization of the chemical specificity of such a com- of calf thymus is considered as consisting of elementary pound becomes a very complicated and laborious task, units of about 35,000 which tend to associate to larger While, therefore, the formulation of the tetranucleo- structures ~ or whether it is regarded as a true macro- tide conception appeared explainable on historical molecule of a molecular weight around 820,0008, the grounds, it lacked an adequate experimental basis, fact remains that the desoxypentose nucleic acids are especially as regards "thymonucleic acid". Although high-molecular substances which in size resemble, or only two nucleic acids, the desoxyribose nucleic acid even surpass, the proteins. It is quite possible that of calf thymus and the ribose nucleic acid of yeast, had there exists a critical range of molecular weights above been examined analytically in some detail, all con- which two different cells witl prove unable to synthe- clusions derived from the study of these substances size completely identical substances. The enormous were immediately extended to the entire realm of number of diverse proteins may be cited as an example.
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