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THE DYNAMICS OF LIVING MATTER IW JACQUES LOEB PROFESSOR OF l'HYSIOLOGY IN THb UNIVERSITY OF CALIFORNIA NdD l!!ork THE COLUMBIA UNIVERSITY PRESS THE MACMILLAN COMPAI\'Y. AGENTS LONDON: MACMILLAN & CO., LTD, AI/rigltts reS(!f't)ed I!;;olumbia mLnib't.it~ llliological .sni,. ED1TED BY HENRY FAIRFIELD OSBORN EDMUND B. WILSON r. FROM THE GREEKS TO DARWIN By Henry Fan'field Osborn II. AMPHIOXUS AND THE ANCESTRY OF THE VERTEBRATES By Arthur Willey Ill. FISHES, LIViNG AND FOSSIL. An Introductory Study By Bashford Dean IV. THE CELL IN DEVELOPMENT AND INHERITANCE By Edmund B Wilson V. THE FOUNDATIONS OF ZOOLOGY By W. K. Brooks VI. THE PROTOZOA By Gary N. Calkins VII. REGENERATION By T. H. Morgan VlI1. THE: DYNAMICS OF LIVlNG MATTER By Jacques Loeb IX. STRUCTURE AND HABITS OF ANTS. Un preparation) By W. M. Wheeler X. BEHAV10R OF THE LOWER ORGANISMS. (II! preparation) By H. S. Jennings COPYRIGHT, 1906, By THE MAC)ULLAN COMPANY. 06 Set up and electrotyped. Published MaTch, 19 . NorlDo(lb~rtft J. B. Cushing 6.. Co._Herwick & Smith Co. Norwood, Mass., U.B.A. ~n ANNE LEONARD LOEB PREFACE THIS book owes its origin to a series of eight lectures delivered upon the invitation of Professor E. B. Wilson and Professor H. F. Osborn at Columbia University in the spring of 1902. The aim of the lectures was to give a presentation of my researches on the dynamics of living matter and the views to which they had led me. In preparing the book I have tried to give a somewhat more com- plete survey of the field of experimental biology than was possible in the lectures, without, however, trying to alter their character. In the introductory lecture use was made of my address at the Inter- national Congress at St. Louis. To Dr. S. S. Maxwell, who has undertaken the main burden of reading tbe proof and preparing the index, and to Professor J. B. MacCallum, who also assisted me in the reading of the proof, my sincere thanks are due. BERKELEY. CALIFORNIA, January I, 1906. "ii CONTENTS LECTURE I INTRODUCTORY REMARKS • LEcrVRE II CONCERNING THE GENERAL CHEMISTRY OF LIFE PHENOMENA I. Historical Remarks 2. Reversible Enzyme < Action - Lipase -Action - Reversible Enzyme Action in the Carbohydrate Group - The General Occurrence of Protein-splitting Enzymes 3. Respiration as a Catalytic Process • 13 (a) The Oxidases • 13 Cob) Further Remarks on the Significance of Oxygen in Life Phenomena. 16 (c) Death in Lack of Oxygen and the Protective Action of Oxygen 18 (d) Changes of Structure in Lack of O"ygen 19 4. The Production of C02 through Enzymes 5. Concerning the Theory of Enzyme Action 24 (a) Stereochemical Attempt!; . 24 (0) The Theory of Intermediary Reactions 26 LECfVRE III THE GENERAL PHYSICAL CoNSTITUTION OF LIVING MA'ITER I. The Limits of Divisibility of Living Matter 29 2. Foam Structures and Emulsions 31 3. The Colloidal Character of Living Matter 33 4. The Formation of Surface Films and Traube's Membranes of Precipitation- Overton's and Meyer's Work on Narcotics and the Nature of Surface Films. 38 5. Osmotic. Pressure and the Exchange of Liquids hetween the Cells and the Sur~ rounding Liquid . 41 6. Further Limitations of Traube's Theoty of Semipenneability . 45 7. The Antagonistic Effects of Salts 46 LECfVRE lV ON SOME PHYsICAL MANIFEsTATIONS OF LIFE 1. Hypotheses of Muscular Contraction 53 2. Quincke's Theory of Protoplasmic Motion 55 3. Concerning tbe Theory of Cell Division. 58 4- The Origin of Radiant Energy in Living Organisms 66 5. Electrical Phenomena ill Livjng Organisms 68 ;x x CONTENTS LECTURE V THE R6cE OF ELECTROLYTES IN THE FORMATION ANI) PRESERVATION OF L1VING MATTER 1. On the Specific Difference between the Nutritive Solutions for Plants and Animals -l)rotective Solutions and Nutrient Solutions 71 2. Concerning a Theory of Irritability and the Role of Na, K, and Ca for Animal Ufe - Rhythmkal Contractions in Skeletal Muscle, in Medusa:, and the Ven- tricle of the Heart - Contact Irritability in Muscle - Analogies to the Role of Salts in Coagulation of Milk-Significance for the Understanding of Functional Nervous Diseases - The Action of Purgatives 78 3. The Reaction of Living Matter and the Role of Bicarbonates in the Preservation ~~. ~ 4. Electrical Stimulation 98 LECTURE VI THE EFFECTS OF HEAT AND RADIANT ENERGY UPON LIVING MAITER I. Effects of Heat-Upper Temperature Limit of Life-Influence of Reaction- velocity upon Biological Processes - M:>wer Temperature Limit - Other Biolugical Effects of Heat 106 2. General Effects of Radiant Energy upon Living Matter- Oxemical Action of Light upon Organisms . lIZ LECTURE VII HELIOTROPISM l. The Heliotropism of Sessile Organisms. JI7 2. Heliotropism of Free-moving Animals . 124 3. The Control of the Precision and Sense of Heliotropic Reactions in Animals. 130 4. The Reaction of Animals to Sudden Changes in the Intensity of Light 135 LECTURE VIII FuRTHER FACTS CONCERNING TROPISMS AND RELATED PHENOMENA. I. General Theory of Tropisms . 2. Galvanotropism 3. Geotropism 4. Chemotropism and Related Phenomena , 5. Stereotropism . 6. Concluding Remarks concerning Tropismlike Reactions. LECTURE IX FERTILIZATION I. The SpecifiC Chara-eter of the Fertilizing Power of the SpennatozoGn-Hybrid Fertil~. 161 2. Artificial Parthenogenesis and the Theory of Fertilization 164 CONTENTS xi LECTURE X HEREDITY I. The Hereditary Effects of the Spermatozoon and Egg-The Prevailing Influence of the Egg in the Early Stages of Development - Merogony-Toxicity of the Blood of Forms not closely Related-Mendel's Experiments. 179 2. The Determination of Sex and the Secondary Sexual Characters 186 3. Egg Structure and Heredity . 191 LECTURE XI ON THE DYNAMICS OF REGENERATIVE PROCESSlts I. Sachs's Hypothesis of the Formation of Organs 199 2. Heteromorphosis and Regeneration in Tubularia • 201 3. Regeneration i.n an Actinian (Cerialtthus memhranactw) 207 4. Regeneration and Heteromorphosis in Planarians . 210 5. On the Influence of the Central Nervous System upon Regeneration and on Phe- nomena of Correlation in Regeneration. 213 6. The Effect of Some External Conditions upon Regeneration and the Transforma~ tion of Organs 217 7. The Role of Re'lersible }>rocesses in Phenomena of Regeneration - The Distribu· t10D of the Power of Regeneration in the Animal Kingdom • 218 LECTURE XII CONCLUDING REMARKS 223 INDEX LECTURE I INTRODUCTORY REMARKS IN these lectures we shall consider living organisms as chemical machines, consisting essentially of colloidal material, which possess the peculiarities of automatically developing, preserving, and reproducing themselves. The fact that the machines which can be created by man do not possess the power of automatic development, self-preservation, and reproduction constitutes for the present a fundamental difference between living machines and artificial machines. We must, however, admit that nothing contradicts the possibility that the artificial produc- tion of living matter may one day be accomplished. It is the purpose of these lectures to state to what extent we are able to control the phe- nomena of development, self-preservation, and reproduction. Living organisms may be called chemical machines, inasmuch as the energy for their work and functions is derived from chemical pro- cesses, and inasmuch as the material from which the living machines are built must be formed through chemical processes. It is therefore only natural that the dynamics of living matter should begin with an analysis of the specific character of the chemical processes in organisms. It is neither our intention nor is it possible for us to give an exhaustive analysis, and we shall only go far enough to satisfy ourselves that no variables are found in the chemical dynamics of living malter which cannot be found also in the chemistry of inanimate nature. The material of which living organisms consist is essentially col- loidal in its character. Graham introduced the discrimination between colloidal and crystalloidal substances: the latter diffuse easily, the former only wtth difficulty, or not at all, through animal membranes. The colloidal substances may be in solution or fine suspension, or they may appear in a jellylike or coagulated or precipitated form. In the former case where they are liquid we speak of sols, in the latter of gels. The structures which we find in living matter originate mostly through a gelation or coagulation of liquid colloids. We shall see in these lectures that liquefactions and gelations or coagulations may possibly play a great TIlle in various physical manifestations of life; but as the physics of colloids.is still in its beginning, we must not be surprised that it is as yet impossible to carry its application to life phenomena very far.

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