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COMPREHENSIVE BIOCHEMISTRY EDITED BY MARCEL FLORKIN Professor of Biochemistry, University of Liege (Belgium) AND ELMER H. STOTZ Professor of Biochemistry, University of Rochester, School of Medicine and Dentistry, Rochester, N.Y. (U.S.A.) VOLUME 29 PART B COMPARATIVE BIOCHEMISTRY, MOLECULAR EVOLUTION (continued) ELSEVIER SCIENTIFIC PUBLISHING COMPANY AMSTERDAM • LONDON 1975 NEW YORK CONTENTS VOLUME 2 9 B COMPARATIVE BIOCHEMISTRY, MOLECULAR EVOLUTION (continued) General Preface Preface to Section IV Errata and Corrigenda to Vol. 29A vii viii xiii Chapter HI. Biochemical Evolution in Bacteria by J. DE LEY AND K. KERSTERS I. II. III. IV. Introduction Origin of bacteria Evolution of bacteria The relative time-scale l.DNArelatedness a. The Enterobacteriaceae b. Agrobacterium 2. Evolution of metabolic processes a. Thecyy of metabolic pathway evolution b. Catabolism of carbohydrates c. Catabolism of aromatic compounds by Pseudomonads d. Energy conservation : e. Control mechanisms (() Allosteric regulation of DHAP synthase, 20 - (ii) Allosteric regulation of citrate synthase, 24 - (Hi) Different induction patterns for the same overall metabolic pathway, 25 — f. CO 2 fixation in chemo- and photo-autotrophic bacteria (i) Mechanism of CO 2 incorporation, 26 - (i/) ATP production in photoautotrophs; photosystems I and II, 27 g. Biosynthesis of tetrapyrroles h. Gram character and cell-hull structure (i) Cell-wall peptidoglycans, 32 - (ii) The peptides, 33 - (Hi) Bacterial phosphatides, 36 i. Pathways of more general occurrence (;') Biosynthesis of lysine, 38 - (ii) Patterns and biosynthesis of some lipids, 38 3. Relationship of proteins .' a. Structure and amino acid sequences of low molecular weight proteins . . . . (i) Cytochrome c, 40 - (ii) Iron-sulphur proteins, 50 - (Hi) Flavodoxin, 65 - (iv) Thioredoxin, 66 - (v) Azurin, 66 - (vi) Nuclease, 67 b.Electrophoretic analysis of cellular proteins and enzymes (() Protein profiles, 68 - (ii) Zymograms, 69 - 1 1 4 6 7 8 9 12 12 13 17 18 20 26 29 32 38 40 40 67 Acknowledgements 71 References 72 X CONTENTS Chapter IV. Biochemical Evolution in Animals by MARCEL FLORKIN Introduction I. Animal evolution A. Short survey of animal systematics and phylogeny B. Drawing phylogenetic trees from amino acid sequences C. Animal paleobiochemistry II. The genetic loss or repression of biosemes in animals 79 81 81 88 89 99 III. Amino acids and their polymers in the biochemical evolution of animals 101 A. Introduction 101 B. Evolution of protein synthesis in animals 102 1. Introduction 102 2. Lineages of proteins and peptides 103 3. Scaffolding proteins 104 4. Biosyntagms involving a proteolytic bioseme 107 a. The trypsin lineage 107 b. Mammalian pancreatic proteolytic enzymes: trypsin, chymotrypsin, elastase 108 c. Trypsinopeptides 117 d. Thrombin and plasmin in the context of the evolution of the system of blood clotting in animals 122 (i) Hemolymph coagulation in Arthropoda, 122 - (ii) Vertebrates, 123 - (in) Fibrinopeptides, 125 e. Other proteins of the trypsin family 129 f. Proteolytic enzymes not related to trypsin 130 5. The lineage: Ribonuclease-lysozyme-lactosesynthetase 130 a. Commutation in the diachronic molecular epigenesis from animal lysozyme to lactose synthetase 137 b. Rate of epigenesis 140 6. Insulin 140 7. Hypophyseal hormones : 142 8. Neurohypophyseal hormones 145 9. Myoglobin and hemoglobin 147 a. Introduction 147 b. Comparison of hemoglobin chains 148 c. Tertiary structure 148 d. Molecular diachronic epigenesis in primate hemoglobins . ' 156 e. Hemoglobin derivation from myoglobin in diachronic molecular epigenesis 157 10. Immunoglobulins 160 a. Non-self recognition in vertebrates 160 C. Extensions on the pathways of amino acid biosynthesis 166 11. Extensions on the pathway of arginine biosynthesis 166 a. Introduction 166 b. Ureogenesis 167 c. The meaning of ureogenesis 170 1 d. A terminal extension on the primary biosyntagm of arginine biosynthesis 172 CONTENTS XI 12. Extensions on the pathways of the biosynthesis of aromatic amino acids . . . a. Introduction b. Derivatives of phenylalanine and tyrosine (/) Tanning agents in Arthropoda, 174 - (ii) Iodinated thyronines, 176 - (Hi) From phenylalanine and tyrosine to phenylalkylamines, 177 c. Tryptophan derivatives (i) From tryptophan to serotonin (5-hydroxytryptamine, 5-HT) and other indolealkylamines, 177 - (ii) From tryptophan to ommochromes, 179 — D. Physiological radiations of free amino acids 13. Free amino acids as compensatory osmotic effectors in marine invertebrate tissues 14. Free amino acids as osmolar constituents of hemolymph and tissues of insects 15. Variations of the free amino acid pool of insect hemolymph during development, metamorphosis and cocoon spinning 16. Diapause 17. Free amino acids and intracellular isosmotic regulation in euryhaline invertebrates a. Introduction b. Mechanism of the participation of free amino acids in the isosmotic intracellular regulation of Crustacea 174 174 174 IV. Diversification in animals at the level of the central metabolic biosyntagm . . . . A. Introduction B. Lateral extensions on the glycolytic pathway 18. Chitin pathway a. Chitin biosynthesis b. Regressive evolution of chitin biosynthesis c. Physiological radiations of chitin d. Regressive evolution in chitinolysis 19. Trehalose biosynthesis 20. Sustained flight in bees and flies 21. Glycerol production in diapausing insects 22. Heat generation in bumblebees (Bombus vagans) 23. 2,3-Diphosphoglycerate formation in mammalian erythrocytes 24. Pathway from glucose to fructose in mammalian foetus and spermatozoa . . C. The evolution of the intracellular pathway of hydrogen transport 25. Introduction 26. Provisions for exploiting anoxic environments . 27. Cold adaptation in homeotherms 28. Brown adipose tissue (brown fat) 29. Loss of respiratory chain in trypanosomes of the brucei group 192 192 192 192 192 192 194 195 196 202 202 203 204 204 205 205 205 205 206 206 V. Some extensions on the terpenoid and steroid pathways of animals A. Introduction B. Loss of the capacity to synthesize sterols C. Terminal extensions from lanosterol 30. Steroid neurotoxins of echinoderms 31. Steroid neurotoxins of amphibians 32., Cardiac-active steroids in toad skin 33. Steroids produced as defensive secretions by water beetles 34. Insect and crustacean ecdysones 177 179 179 181 184 184 185 185 189 207 207 208 208 208 208 210 210 211 XII CONTENTS D. Lateral extensions 35. Insect juvenile hormone 36. Terpenoid coactones used by insects E. Physiological radiations 37. A physiological radiation of ecdysone 38. Physiological radiations of carotenoids and their derivatives 213 213 213 215 215 215 Acknowledgements 219 References 220 Chapter V. Ideas and Experiments in the Field of Prebiological Chemical Evolution by MARCEL FLORKIN 1. Heterogeny (spontaneous generation) 2. Agenesis or the origin of life 3. The primitive atmosphere 4. Synthesis of monomers and of polymers 5. The proteinoid theory of the origin of life 6. The gene hypothesis of theorigin of life. Nucleic acids before proteins 7. The thermodynamics of the irreversible processes and the origin of life 8. Epistemological aspects 231 235 242 242 244 252 254 256 References 259 Subject Index 261