8]m ^ CD :< co THE MECHANISM OF LIFE THE MECHANISM OF LIFE DR. STEPHANE LEDUC PKOFKSSKUK A L'lVoLF, I)E MKDK.CINK D1C NANTES TRANSLATED BY w. DEANE BUTCIIKR J OK THR OF .MKUICINE " La nature a forme 1 , ct forme tons Ics jours Ics ctres les plus simples par gene-ration spontandc, LAMARCK.' LONDON WILLIAM HEINEMANN First Impression . . . March Second Impression . . . January All Rights Reserved TRANSLATOR'S PREFACE PROFESSOR LEI-MIC'S Thvorie Phtiaico-chlmlquc de la Vie et Generations Spontancefi has excited a good deal of attention, and not a little opposition, on the Continent. As recently as 1907 the Academic des Sciences excluded from its Comptes Kendus the report of these experimental researches on diffusion and osmosis, because it touched too closely on the burning question of spontaneous generation. As the author points out, Lamarck's early evolutionary hypothesis was killed by opposition and neglect, and had to be reborn in England before it obtained universal acceptance as the Darwinian Theory. Not unnaturally, therefore, he turns for an appreciation of his work to the free air and wide hori/xm of the English-speaking countries. He has entitled his book "The Mechanism of Life," since however little we may know of the origin of life, we may yet hope to get a glimpse of the machinery, and perhaps even hear the whirr of the wheels in Nature's work- shop. The subject is of entrancing interest to the biologist and the physician, quite apart from its bearing on the question of spontaneous generation. Whatever view may be entertained by the different schools of thought as to the nature and significance of life, all alike will welcome this new and important contribution to our knowledge of the mechanism by which Nature constructs the bewildering variety of her forms. There is, I think, no more wonderful and illuminating spectacle than that of an osmotic growth, a crude lump of brute inanimate matter germinating before our very eyes, putting forth bud and stem and root and branch and leaf and fruit, with no stimulus from germ or seed, without even vii viii TRANSLATOR'S PREFACE the presence of organic matter. For these mineral growths are not mere crystallizations as many suppose ; they increase by intussusception and not by accretion. They exhibit the phenomena of circulation and respiration, and a crude sort of reproduction by budding ; they have a period of vigorous youthful growth, of old age, of death and of decay. They imitate the forms, the colour, the texture, and even the microscopical structure of organic growth so closely as to deceive the very elect. When we find, moreover, that the processes of nutrition are carried on in these osmotic pro- ductions just as in living beings, that an injury to an osmotic growth is repaired by the coagulation of its internal sap, and that it is able to perform periodic movements just as an animal or a plant, we are at a loss to define any line of separation between these mineral forms and those of organic life. In the present volume the author has collected all the data necessary for a complete survey of the mechanism of life, which consists essentially of those phenomena which are exhibited at the contact of solutions of different degrees of concentration. Whatever may be the verdict as to the author's case for spontaneous generation, all will agree that the book is a most brilliant and stimulating study, founded on the personal investigation of a born experimenter. The present volume is a translation of Dr. Leducs French edition, but it is more than this, the work has been translated, revised and corrected, and in many places re- written, by the author's own hand. I am responsible only for the English form of the treatise, and can but regret that I have been able to reproduce so imperfectly the charm of the original. W. DEANE BUTCHER. EALING. PREFACE TO THE ENGLISH EDITION CV.sT par Tinitiative clu Dr. Deane Butcher que cette ouvrage est presents aux leeteurs anglais, a la race qui a dote rimmanite de tant de decouvertes originales, genial es et d\me portee tres generalc. Com me un etre vivant, unc idee exige pour naitre et se developper Ic germe et le milieu de devcloppement, II est incleniable que le peuple anglo-americain constitue un milieu particulicrcment favorable a la naissance et an developpement des idees nouvelles. Pendant notre collaboration le Dr. Deane Butcher a ete un critique judicieux et eclaire, tons les changements dans Tedition anglaise sont dus a ses observations. II s^est assimile Touvrage pour le traduire, et dans beaucoup de parties, il a mis plus de clarte et de concision qiTil n'y en avait dans le texte original. STEPHANE LEDUC. NANTES, 1911. TABLE OF CONTENTS PACK TRANSLATOR'S PREFACE , . . . vii AUTHOR'S PREFACE . . , . . . . ix INTRODUCTION . . xiii I. LIFE AND LIVING BEINGS ..... i II. SOLUTIONS . . . . . . 14 III ELECTROLYTIC SOLUTIONS . . . . .24 IV. COLLOIDS ....... 36 V. DIFFUSION AND OSMOSIS . . . . -43 VI. PERIODICITY ....... 67 VI I, COHESION AND CRYSTALLIZATION . . . .78 VIII. KARYOKINESIS . . . . . .89 IX. ENERGETICS . . . . . . -97 X. SYNTHETIC BIOLOGY . . . . . .113 XI. OSMOTIC GROWTH : A STUDY IN MORPHOGENESIS . 123 XII. THE PHENOMENA OF LIFE AND OSMOTIC PRODUCTIONS: A STUDY IN PHYSIOGENESIS . . . -147 XIII. EVOLUTION AND SPONTANEOUS GENERATION . . 160 INTRODUCTION LIFK was formerly regarded as a phenomenon entirely separated from the other phenomena of Nature, and even up to the present time Science has proved wholly unable to give a definition of Life ; evolution, nutrition, sensibility, growth, organization, none of these, not even the faculty of repro- duction, is the exclusive appanage of life. Living things are made of the same chemical elements as minerals ; a living being is the arena of the same physical forces as those which affect the inorganic world. Life is difficult to define because it differs from one living being to another ; the life of a man is not that of a polyp or of a plant, and if we find it impossible to discover the line which separates life from the other phenomena of Nature, it is in fact because no such line of demarcation exists the passage from animate to inanimate is gradual and insensible. The step between a stalagmite and a polyp is less than that between a polyp and a man, and even the trained biologist is often at a loss to determine whether a given borderland form is the result of life, or of the inanimate forces of the mineral world. A living being is a transformer of matter and energy both matter and energy being uncreateable and indestructible, i.e. invariable in quantity. A living being is only a current of matter and of energy, both of which change from moment to moment while passing through the organism. That which constitutes a living being is its form ; for a living thing is born, develops, and dies with the form and structure of its organism. This ephemeral nature of the living being, which perishes with the destruction of its form, is in xiv INTRODUCTION marked contrast to the perennial character of the matter and the energy which circulate within it. The elementary phenomenon of life is the contact between an alimentary liquid and a cell. For the essential phenomenon of life is nutrition, and in order to be assimilated all the elements of an organism must be brought into a state of solution. Hence the study of life may be best begun by the study of those physico-chemical phenomena which result from the contact of two different liquids. Biology is thus but a branch of the physico-chemistry of liquids; it includes the study of electrolytic and colloidal solutions, and of the molecular forces brought into play by solution, osmosis, diffusion, cohesion, and crystalli/ation. In this volume I have endeavoured to give as much of the science of energetics as can be treated without the use of mathematical formulae ; the conception of entropy and Garnet's law of thermodynamics are also discussed. The phenomena of catalysis and of diastatic fermentation have for the first time been brought under the general laws of energetics. This I have done by showing that catalysis is only one instance of the general law of the transformation of potential into kinetic energy, vix. by the intervention of a foreign exciting and stimulating energy which may be infinitely smaller than the energy it transforms. This conception brings life into line with other catalytic actions, and shows us a living being as a store of potential energy, to be set free by an external stimulus which may also excite sensation. In a subsequent chapter I have dealt with the rise of Synthetic Biology, whose history and methods I have described. It is only of late that the progress of physico-chemical science has enabled us to enter into this field of research, the final one in the evolution of biological science. The present work contains some of the earliest results of this synthetic biology. We shall see how it is possible by the mere diffusion of liquids to obtain forms which imitate with the greatest accuracy not only the ordinary cellular tissues, but the more complicated striated structures, such as muscle and mother-of-pearl. We shall also see how it is INTRODUCTION xv possible by simple liquid diffusion to reproduce in ordered and regular succession complicated movements like those observed in the karyokinesis of the living cell. The essential character of the living being is its Form. This is the only characteristic which it retains during the whole of its existence, with which it is born, which causes its development, and disappears with its death. The task of synthetic biology is the recognition of those physico-chemical forces and conditions which can produce forms and structures analogous to those of living beings. This is the subject of the chapter on Morphogenesis. The last chapter deals with the doctrine of Evolution. The chain of life is of necessity a continuous one, from the mineral at one end to the most complicated organism at the other. We cannot allow that it is broken at any point, or that there is a link missing between animate and inanimate nature. Hence the theory of evolution necessarily admits the physico- chemical nature of life and the fact of spontaneous generation. Only thus can the evolutionary theory become a rational one, a stimulating and fertile inspirer of research. We seek for the physico-chemical forces which produce forms and structures analogous to those of living beings, and phenomena analogous to those of life. We study the alterations in environment which modify these forms, and we seek in the past history of our planet for those natural phenomena which have brought these physico-chemical forces into play. In this way we may find the road which will, we hope, lead some day to the discovery of the origin and the evolution of life upon the earth. THE MECHANISM OF LIFE CHAPTER I LIFE AND LIVING BEINGS PRIMITIVE man distinguished but two kinds of bodies in nature, those which were motionless and those which were animated. Movement was for him the expression of life. The stream, the wind, the waves, all were alive, and each was endowed with all the attributes of life will, sentiment, and passion. Ancient Greek mythology is but the poetic expression of this primitive conception. In the evolution of the intelligence, as in that of the body, the development .ofjthc individual is but a repetitipn of the development of tlie^race. Even now children attribute life to everything that moves. For them a little bird still lives in the inside of a watch, and produces the tick-tick of the wheels. In modern times, however, we have learnt that everything in nature nioyes^ so thcOl^motionof^ itself cannot be considered as flic characteristic of [life. Heraelitus aptly compares life to a flame. Aristotle says, " Life is nutrition, growth, and decay, having for its cause a principle which has its end in itself, namely e^rsXg^g/a. This principle is itself in need of definition, and Aristotle only substitutes one unknown epithet for another. Bichat defined life as the ensemble of the .functions, jvhjch resist death. This is to define life in terms of death, but death is but the end of life, and cannot be defined without first defining life. Claude Bernard rejects jJL.defiilition of life as insufficient, and incompatible with experimental science. 2 THE MECHANISM OF LIFE Some modern physiologists regard sensibility, others irritability, as the characteristic of life, and define life as the faculty of responding, by some sort of change, to an external stimulus. As in the case of movement, we have found by more attentive observation that this faculty also is universal in nature. There is no action without reaction ; an elastic body repels the body that strikes it. Every object in nature dilates with heat, contracts with cold, and is modified by the light which it absorbs. Everything in nature responds to exterior action by a change, and hence this faculty cannot be the characteristic of life. A distinguished professor of physiology was accustomed to teach that the disproportion between action and reaction was_ the characteristic of life. " Allow a gramme weight to fall on a nerve, and the muscle will raise a weight of ten grammes. This disproportion is the characteristic of life." But there is a much greater disproportion between action and reaction when the friction of a match blows up a powder factory, or the turning of a switch lights the lamps and animates the tram- ways and the motors of a great city. The disproportion between action and reaction is therefore no characteristic of life. The essential characteristic of life is often said to be nutrition the phenomenon by which a living organism absorbs matter from its environment, subjects it to chemical metamorphosis, assimilates it, and finally ejects the destructive products of metamorphosis into the surrounding medium. But this characteristic is also common to a great number of ordinary chemical reactions, so that we cannot call it peculiar to life. Consider, for instance, a fragment of calcium chloride immersed in a solution of sodium carbonate. It absorbs the carbonic ion, incorporates it into a molecule of calcium carbonate, and ejects the chlorine ion into the surrounding medium. It may be argued that this is merely a chemical process, since the substance which determines the reaction is also modified, the chloride of calcium changing into carbonate of calcium. But every living thing is also changing its chemical LIFE AND LIVING BEINGS 3 constitution during every moment of its existence, it is this change which constitutes the process of senile involution. The substance of the child is other than that of the ovum, and the substance of the adult is not that of the child. Hence we cannot regard nutrition as the exclusive characteristic of life. Other authorities regard growth and organization as the essentials of life. But crystals also grow. It was said that the growth of a crystal differed from that of a living thing, in that the former grew by the addition of material from without the juxtaposition of bricks, as it were while the latter grew by intussusception, an introduction of fresh material into the substance of the organism. A crystal, moreover, was homo- geneous, while the tissues of a living being were differentiated such differentiation constituting the organization. At the present time, however, we recognize the existence of a great variety of purely physical productions, the so-called " osmotic growths, 1 ' which increase by a process of intussusception, and develop therefrom a marvellous complexity of organization and of form. Hence growth and organization cannot be considered as the essential characteristics of life. Since, then, we are totally unable to define the exact boundary which separates life from the physical phenomena of nature, we may fairly conclude that no such separation exists. This is in conformity with the " law of continuity, 11 the principle which asserts that all the phenomena of nature are continuous in time and space. Classes, divisions, and separa- tions are all artificial, made not by nature but by man. All the forms and phenomena of nature are united by insensible transition ; it is impossible to separate them, and in the distinction between living and non-living things we must content ourselves with relative definitions, which arc far from being precise. Life can only be defined as the sum of all phenomena exhibited by living beings, and its definition thus becomes a mere corollary to the definition of a living being. The true definition of a living being is that it is a trans- former of energy, receiving from its environment the energy 4 THE MECHANISM OF LIFE which it returns to that environment under another form. All living organisms arc transformers of energy. A living organism is also a transformer of matter. It absorbs matter from its environment, transforms it, and returns it to its environment in a different chemical condition. Living things are chemical transformers of matter. Living beings are also transformers of form. They com- mence as a very simple form, which gradually develops and becomes more complicated. The matter of which a living organism is constituted con- c5 O sists essentially of certain solutions of crystalloids and colloids. To this we may add an osmotic membrane to contain the liquids, and a solid skeleton to support and protect them. Finally, it would seem that a colloid of one of the albuminoid groups is a necessary constituent of every living being. We may say, then, that a living being is a transformer of energy and of matter, containing certain albuminoid sub- stances, with an evolutionary form, the constitution of which is essentially liquid. A living being has but a limited duration. It is born, develops, becomes organized, declines and dies. Through all the metamorphoses of form, of substance, and of energy, informing the whole course of its existence, there is a certain co-ordination, a certain harmony, which is necessary for the conservation of the individual. This harmony we call Life. Discord is disease, the total cessation of the harmony is Death. When the form is profoundly altered and the substance changed, the transformation of energy no longer follows its regular course, the organism is dead. After death the colloids which have constituted the form of the living thing pass from their liquid state as " sols " into their coagulated state as "gels." The metamorphoses of form, substance, and energy still continue, but no longer harmoniously for the conservation of the individual, but in dis-harmony for its dissolution. Finally, the form of the individual disappears, the substance and the energy of the living being is resolved and dispersed into other bodies and other phenomena. LIFE AND LIVING BEINGS 5 The results hitherto obtained from the study of life seem but inconsiderable when compared with the time and labour devoted to the question. Max Verworn exclaims, " Are we on a false track ? Do we ask our questions of Nature amiss, or do we not read her answers aright ? " Each branch of science at its commencement employs only the simpler methods of observation. It is purely descriptive. The next step is to separate the different parts of the object studied to dissect and to analyse. The science has now become analytical. The final stage is to reproduce the sub- stances, the forms, and the phenomena which have been the subject of investigation. The science has at last become synthetical. Up to the present time, biology has made use only of the first two methods, the descriptive and the analytical. The analytical method is at a grave disadvantage in all biological investigations, since it is impossible to separate and analyse the elementary phenomena of life. The function of an organ ceases when it is isolated from the organism of which it forms a part. This is the chief cause of our lack of progress in the analysis of life. It is only recently that we have been able to apply the synthetic method to the study of the phenomena of life. Now that we know that a living organism is but the arena for the transformation of energy, we may hope to reproduce the elementary phenomena of life, by calling into play a similar transformation of energy in a suitable medium. Organic chemistry has already obtained numerous victories in the same direction, and the rapid advance in the produc- tion of organic bodies by chemical synthesis may be considered the first-fruits of synthetic biology. A phenomenon is determined by a number of circumstances which we call its causes, and of which it is the result. Every phenomenon, moreover, contributes to the production of other phenomena which are called its consequences. In order there- fore to understand any phenomenon in its entirety, we must determine all its causes both qualitatively and quantitatively. Phenomena succeed one another in time as consequences 6 THE MECHANISM OF LIFE one of another, and thus form an uninterrupted chain from the infinite of the past into the infinite of the future. A living being gathers from its entourage a supply of matter and of energy, which it transforms and returns. It is part and parcel of the medium in which it lives, which acts upon it, and upon which it acts. The living being and the medium in which it exists are mutually interdependent. This medium is in its turn dependent on its entourage, and so on from medium to medium throughout the regions of infinite space. One of the great laws of the universe is the law of continuity in time and space. We must not lose sight of this law when we attempt to follow the metamorphoses of matter, of energy and of form in living beings. Evolution is but the expression of this law of continuity, this succession of phenomena following one another like the links of a chain, without discontinuity through the vast extent of time and space. The other great universal law, that of conservation, applies with equal force to living and to inanimate things. This law asserts the uncreateability and the indestructibility of matter and of energy. A given quantity of matter and of energy remains absolutely invariable through all the transformations through which it may pass. We need not here discuss the question of the possible trans- formation of matter into ether, or of ether into ponderable matter. Such a transformation, if it exists, would have but little bearing on the phenomena of life. Moreover, it also will probably be found to conform to the law of conservation of energy. In marked contrast to the permanence of matter and of energy is the ephemeral nature of form, as exhibited by living beings. Function, since it is but the resultant of form, is also ephemeral. All the faculties of life are bound up with its form, a living being is born, exists, and dies with its form. The phenomena of life may in certain cases slow down from their normal rapidity and intensity, as in hibernating LIFE AND LIVING BEINGS 7 animals, or be entirely suspended, as in seeds. This state of suspension of life, of latent life as it were, reminds us of a machine that has been stopped, but which retains its form and substance unaltered, and may be started again whenever the obstacle to its progress is removed. During the whole course of its life a living being is intimately dependent on its entourage. For example, the phenomena of life are circumscribed within very narrow limits of temperature. A living organism, consisting as it does essentially of liquid solutions, can only exist at temperatures at which such solutions remain liquid, i.e. between C. and 100 C. Certain organisms, it is true, may be frozen, but their life remains in a state of suspension so long as their substance remains solid. Since the albuminoid substances which are a necessary component of the living organism become coagulated at 44 C., the manifestations of life diminish rapidly above this temperature. The intensity of life may be said to augment gradually as the temperature rises from to 40, and then to diminish rapidly as the temperature rises above that point, becoming nearly extinct at 60 C. Another condition indispensable to life is the presence of oxygen. Life, compared by Heraclitus to a flame, is a com- bustion, an oxydation, for which the presence of oxygen at a certain pressure is indispensable. There are, it is true, certain anaerobic micro-organisms which apparently exist without oxygen,, but these in reality obtain their oxygen from the medium in which they grow. Life is also influenced by light, by mechanical pressure, by the chemical composition of its entourage, and by other conditions which we do not as yet understand. In each case the conditions which are favourable or noxious vary with the nature of the organism, some living in air, some in fresh water, and others in the sea. Formerly it was supposed that the substance of a living being was essentially different from that of the mineral world, so much so that two distinct chemistries were in existence organic chemistry, the study of substances derived from bodies which had once possessed life, and inorganic chemistry, dealing 8 THE MECHANISM OF LIFE with minerals, metalloids, and metals. We now know that a living organism is composed of exactly the same elements as those which constitute the mineral world. These are carbon, oxygen, hydrogen, nitrogen, phosphorus, calcium, iron, sulphur, chlorine, sodium, potassium, and one or two other elements in smaller quantity. It was formerly supposed that the organic combinations of these elements were found only in living organisms and could be fashioned only by vital forces. In more recent times, however, an ever increasing number of organic substances have been produced in the laboratory. Organic bodies may be divided into four principal groups. (1) Carbohydrates, including the sugars and the starches, all of which may be considered as formed of carbon and water. (2) FatSj which may be considered chemically as the ethers of glycerine, combinations of one molecule of glycerine and three molecules of a fatty acid, with elimination of water. (3) Albuminoids, substances whose molecules are complex, con- taining nitrogen and sulphur in addition to carbon, oxygen, and hydrogen. The albuminoid of the cell nucleus also contains phosphorus, and the haemoglobin of the blood contains iron. (4) Minerals or inorganic elements, such as chloride of sodium, phosphate of calcium, and carbonic acid. This group also includes water, which is the most important constituent, since it forms more than a moiety of the sub- stance of all living creatures. Wohler in 1828 accomplished the first synthesis of an organic substance, urea, one of the products of the decom- position of albumin. Since then a large number of organic substances have been prepared by the synthesis of their inorganic elements. The most recent advance in this direction is that of fimile Fischer, who has produced polypeptides having the same reactions as the peptones, by combining a number of molecules of the amides of the fatty acids. In the further synthesis of organic compounds the problems we have before us are of the same order as those already solved. There is no essential difference between organic and inorganic chemistry; living organisms are formed of the LIFE AND LIVING BEINGS 9 same elements as the mineral world, and the organic com- binations of these elements may be realized in our laboratories, just as in the laboratory of the living organism. Not only so, but a living being only borrows for a short time those mineral elements which, after having passed through the living organism, are returned once again to the mineral kingdom from which they came. All matter has life in itself or, at any rate, all matter susceptible of incorporation in a living cell. This life is potential while the element is in the mineral state, and actual while the element is passing through a living organism. Mineral matter is changed into organic matter in its passage through a vegetable organism. The carbonic acid produced by combustion and respiration is absorbed by the chlorophyll of the leaves under the stimulus of light the oxygen of the carbonic acid being returned to the air, while the carbon is utilized by the plant for the formation of sugar, starch, cellulose, and fats. Thus plants are fed in great part by their leaves, taking an important part of their nourishment from the air, while by their roots they draw from the earth the water, the phosphates, the mineral salts, and the nitrates required for the formation of their albuminoid constituents. A vegetable is a laboratory in which is carried out the process of organic synthesis by which mineral materials are changed into organic matter. The first synthetic reaction is the formation of a molecule of formic aldehyde, CH 2 O, by the combination of a molecule of water with an atom of carbon. From this formic aldehyde, or formol, we may obtain all the various carbohydrates by simple polymerization, i.e. by the association of several molecules, with or without elimi- nation of water. Thus two molecules of formol form one molecule of acetic acid, 2CH 2 O C 2 H 4 O 2 . Three molecules of formol form a molecule of lactic acid, 3CH 2 O = C 3 H 6 O S . Six molecules of formol represent glucose and levulose, 6CII 2 O = C 6 H 12 O 6 . Twelve molecules of formol minus one molecule of water form saccharose, lactose, cane sugar, and sugar of milk, l#CH 2 = C 12 II 22 O n + II 2 O 5 n times six mole- 10 THE MECHANISM OF LIFE i cules of forinol minus one molecule of water, ??(C 6 H 10 O 6 ), i form starch and cellulose. Animals derive their nourishment from vegetables either directly, or indirectly through the flesh of herbivorous animals. The mineral matter, rendered organic in its passage through a vegetable growth, is finally returned by the agency of animal organisms to the mineral world again, in the form of carbonic acid, water, urea, and nitrates. Thus vegetables may be regarded as synthetic agents, and animals and microbes as _?L_ d^ony^itimi. Here also the difference is only _ relative, for in certain cases vegetables produce carbonic acid, while some animal organisms effect synthetic combinations. Moreover, there are intermediary forms, such as fungi, which possessing no chlorophyll are nourished like animals by organic matter, and yet like vegetables are able to manu- facture organic matter from mineral salts. The work of combustion begun by the animal organism is finished by the action of micro-organisms, who complete the oxydation the rc-mincralizatioii of the chemical substances drawn originally from the inorganic world by the agency of plant life. To sum up. Vegetables ^>btai n^ Jjiej r^ jiourisl imgnt. from mineral substances, which they reduce^ dc-oxydi/c 2 and ^charge with_ so^r_energy. Anin^j3iga^sms oij_the contrary oxydi/e, and micro-organisms complete the oxydation of these sub- . ______ . ______________ -n__ ----- . ------ JL---, ------------------------------ J. ------------------ ....... _ .. - - . - - stances, returning _them to the mineral world as watej, carbonates, .nitrate^ and sulphates. Thus matter circulates eternally from the mineral to the vegetable, fiom the vegetable to the animal world, and back again. The matter which forms our structure, which is to-day part and parcel of ourselves, has formed the structure of an infinite number of living beings, and will continue to pursue its endless reincarnation after our decease. Tll!?-S!^^ ss c y c k f Hfkjs ^k? an endless cycle of energy. The combination of carbon with water carried out by the agency of chlorophyll can only take place with absorption of energy. This energy comes directly from the sun, the red and orange light radiations b^HlJ-LJ 1 ^ LIFE AND LIVING BEINGS 1 1 The arrest of vegetation during the winter months is due not so much to the lowering of temperature as to the diminution of the radiant energy received from the sun. In the same way shade is harmful to vegetation, since the radiant energy required for growth is prevented from reaching the plant. The energy radiated by the sun is accumulated and stored in the plant tissues. Later on, animals feed on the plants and utilize this energy, excreting the products of decomposition, i.e. the constituents of their food minus the energy contained in it. Thus the_ whole of the energy which animates living from the sun. To the sun also we owe all artificial heat, the energy stored up in wood and coal. We are all of us children of the sun. The radiant energy of the sun is transformed by plants into chemical energy. It is this chemical energy which feeds the vital activity of animals, who return it to the external world under the form of heat, mechanical work, and muscular contraction, light in the glow-worm, electricity in the electric eel. There is a marked difference between the forms affected by organic and inorganic substances. The forms of the mineral world are those of crystals geometrical forms, bounded by straight lines, planes, and regular angles. Livmg organisms, on the contrary, affect forms which are less regular curve( l surfaces and rounded aiigles. The physical reason for this difference in form lies in a difference of consistency, crystals being solid, whereas living organisms are liquids or semi - liquids. The liquids of nature, streams and clouds and dewdrops, affect the same rounded forms as those of living organisms. Living beings for the most part present a remarkable degree of symmetry. Some, like radiolarians and star-fish, have a stellate form. In plants the various organs often radiate from an axis, in such a manner that on turning the plant about this axis the various forms are superposed thrice, four, or more often five times in one complete revolution. It is remarkable how often this number five recurs in the 12 THE MECHANISM OF LIFE divisions and parts of a living organism. In other cases the similar parts are disposed symmetrically on either side of a median line or plane, giving a series of homologous parts which are not superposable. The most important characteristic of a living being is its form. This is implicitly admitted by naturalists, who classify animals and plants in genera and species according to the differences and analogies of their form. All living beings are composed of elementary organizations called cells. In its complete state, a cell consists of a membrane or envelope containing a mass of protoplasm, in the centre of which is a nucleus of differentiated protoplasm. This nucleus may in its turn contain a nucleolus. In some cases the cell is merely a protoplasmic mass without a visible envelope, so that a cell may be defined as essentially a mass of protoplasm provided with a nucleus. A living organism may consist merely of a single cell, which is able alone to accomplish all the functions of life. Most living beings, however, consist of a collection of in- numerable cells forming a cellular association or community. When a number of cells are thus united to constitute a single living being, the various functions of life are divided among different cellular groups. Certain cells become specialized for the accomplishment of a single function, and to each function corresponds a different form of cell. It is thus easy to recognize by their form the nerve cells, the muscle cells which perform the function of movement, and the glandular cells which perform the function of secretion. The cells of a living being arc microscopic in size, and it is remarkable that they never attain to any considerable dimensions. In order that life may be maintained in a living organism, it is necessary that a continual supply of aliment should be brought to it, and that certain other substances, the waste- products of combustion, should be eliminated. In order to be absorbed and assimilated, the alimentary substances must be presented to the living organism in a liquid or gaseous state. Thus the essential condition necessary for the LIFE AND LIVING BEINGS 13 maintenance ofMife _is__the J?Jltoet 2.? JL-liYljlS 5^lL-wlth ? | i r i^ i it_5^ '_!ML\? id. r JQi e j^i 1 !*: ui^y-T- 4?iij^i^LlMU2iiiii ( ?. i L .of life _is the coiitact_ of t\vo different liquids. This is the necessary condition which renders possible the chemical ex- changes and the transformations of energy which constitute life. It is in the study of the phenomena of liquid contact and diffusion that we may best hope to pierce the secrets of life. The physics of vital action are the physics of the phenomena which occur in liquids, and the study of the physics of a liquid must be the preface and the basis of all inquiry into the nature and origin of life. CHAPTER II SOLUTIONS WK have seen that living beings are transformers of energy and of matter, evolutionary in form and liquid in consistency ; that they are solutions of colloids and crystalloids separated by osmotic membranes to form microscopic cells, or consisting merely of a gelatinous mass of protoplasm, Avith a nucleus of slightly differentiated material. The elementary pheno- menon of life is the contact of two different solutions. This is the initial physical phenomenon from which proceed all the other phenomena of life in accordance with the ordinary chemical and physical laws. Thus the basis of biological science is the study of solution and of the phenomena which occur between two different solutions, either in immediate contact or when separated by a membrane. A solution is a homogeneous mixture of one or more solutes in a liquid solvent. Before solution the solute or dissolved substance may be solid, liquid, or gaseous. Soliites ? or substances capable of solution, may^ be divided into t wo jjjissfis substances which are cjy^ble^jof crystal- . r 5II^U?J4s au d those which are incapable^ of n^ the jcpllqids. Crystalloids may be divided again into two classes, those whose solutions arc ioni/ablc and T p ._ .. 7 -- therefore conduct electricity, chJefly jj^ and those whose solutions are non-ioni/able and are there- fore non-conductors. These latter are for the most part crystalli/able substances of organic origin, such as sugars, urea, etc. Avogadro's law asserts that under similar conditions of temperature and pressure, equal volumes of various gases SOLUTIONS 1 5 contain an equal number of molecules. Under similar conditions, the molecular weights of different substances have therefore the same ratio as the weights of equal volumes of their vapours. Hence if we fix arbitrarily the molecular weight of any one substance, the molecular weight of all other substances is thereby determined. The molecular weight of hydrogen has been arbitrarily fixed as two, and hence the molecular weight of any substance will be double its gaseous density when compared with that of hydrogen. Gramme- Molecule. A gramme-molecule is the molecular weight of a body expressed in grammes. Occasionally for brevity a gramme-molecule is spoken of as a " molecule. 1 ' Thus we may say that the molecular weight of oxygen is 16 grammes, meaning thereby that there are the same number of molecules in 16 grammes of oxygen as there are atoms in 1 gramme of hydrogen. Concentration-. The concentration of a solution is the ratio between the quantity of the solute and the quantity of the solvent. The concentration of a solution is expressed in various ways. (a) The weight of solute dissolved in 100 grammes of the solvent, (b) The weight of solute present in 100 grammes of the solution. (c) The weight of solute dissolved in a litre of the solvent, (d) The weight of solute in a litre of the solution. The most usual method is to give the concentration as the weight of solute dissolved in 100 grammes or in one litre of the solvent. Molecular Concentration. Many of the physical and biological properties of a solution are proportional, not to its mass or weight concentration, but to its molecular con- centration, i. e. to the number of gramme-molecules of tlie. S( Zk l J*L Contained in a litre of the solution. Many physical properties are quite independent of the nature of the solute, depending only on its degree of molecular concentration. Normal Solution. A iigi;imxl_solutioii is one which contains j )er Ufare. A decinormal solution contains one-tenth of a gramme-molecule of the solute per litre, and a centi normal solution one-hundredth of a gramme -molecule. A normal solution of urea, for example, 1 6 THE MECHANISM OF LIFE contains 60 grammes of urea per litre, while a normal solution of sugar contains 34 grammes of sugar per litre. The Dissolved Substance is a Gas. Van t' HofF, using the data obtained by the botanist Pfeffer, showed that the dissolved matter in a solution behaved ^exactly as if it were a gas. The analogy is complete in every respect. Like the gaseous molecules, the molecules of a solute are mobile with respect to one another. Like those of a gas, the molecules of a solute tend to spread themselves equally, and to fill the whole space at their disposal, i.e. the whole volume of the solution. The. 1 surface of the solution represents the vessel containing the gas, which confines it within definite limits and prevents further expansion. Osmotic Pressure. Like the molecules of a gas, the mole- cules of a solute exercise pressure on the boundaries of the space containing it. This osmotic pressure follows exactly the same laws jas jjaseous pressure. It has the same constants, and all the notions acquired by the study of gaseous pressure are applicable to osmotic pressure. Osmotic pressure is in fact the jgascous pressure of the molecules of the solute. When a gas dilates and increases in volume, its temperature falls, and cold is produced. Similarly, when a soluble substance is dissolved, it increases in volume, and the temperature of the liquid falls. This phenomenon is well known as a means of producing cold by a refrigerating mixture. The i_ phenomena of 'life are governed by the laws of gaseous pressure, since all these _pjienpnieii_a take place in solutions. The fLmdamental laws of biology are those of the distribution of subsj:ances_in solution, which is regulated by the laws of gaseous pressure, since all these laws are applicable also to osmotic pressure. Boyle's Law. When a gas is compressed its volume is diminished. If the pressure is doubled, the volume is reduced to one-half. The quantity V X P, that is the volume multiplied by the pressure, is constant. Gay-Lussac's Law. For a difference of temperature of a degree Centigrade all gases dilate or contract by ^f 3 of their volume at Centigrade. SOLUTIONS 1 7 Daltoii's Law. In a gaseous mixture, the total pressure is equal to the sum of the pressures which each gas would exert if it alone filled the whole of the receptacle. Pressure proportional to Molecular Concentration. The above laws are completely independent of the chemical nature of the gas, they depend only on the number of gaseous molecules in a given space, i.e. on the molecular concentration. If we double the mass of the gas in a given space, we double the number of molecules, and we also double the pressure, whatever the nature of the molecules. We may also double the pressure by compressing the molecules of a gas, or of several gases, into a space half the original size. The molecular concentration of a gas, or of a mixture of gases, is the ratio of the number of molecules to the volume they occupy. The pressure of a gas or of a mixture of gases is proportional to its molecular concentration. This is a better and a shorter way of expressing both Boyle^s law and Daltoifs law. One gramme-molecule of a gas, whatever its nature, con- densed into the volume of 1 litre, has a pressure of 22*35 atmospheres. Similarly one gramme-molecule of a solute, whatever its nature, when dissolved in a litre of water, has the same pressure, viz, 22 '35 atmospheres. Absolute Zero. According to Gay-Lussac's law, the volume of a gas diminishes by -g-j^ of its volume at C. for each degree fall of temperature. Thus if the contraction is the same for all temperatures, the volume would be reduced to zero at 273 C. This is the absolute zero of temperature. Temperatures measured from this point are called absolute temperatures, and are designated by the symbol T. If t indicates the Centigrade temperature above the freezing point of water, then the absolute temperature is equal to 2 + 273. The Gaseous Constant. Consider a mass of gas at C. under a pressure P o , with volume V . At the absolute temperature T, if the pressure be unaltered, the volume of V T this gas will be P~. Therefore the constant PV, the product "P V of the pressure by the volume, will be represented by 273 1 8 THE MECHANISM OF LIFE At the same temperature, but under another pressure P', the gas will have a different volume V'. Since, accord- ing to Boyle's law, PV is constant (P'V' = P V ), it will P V T P V still equal ~fr- - Therefore ~b~ is also constant. This f^tltJ """" """* /w I O - - "" quantity is called "the gaseous__ constant, 11 and if we represent it by the symbol R, we obtain the general formula PV = RT for all gases, or -^ = R. Suppose, for instance, we have a gramme-molecule of a gas at C. in a space of 1 litre. It has a pressure of 22'35 atmospheres at 0C., or 273 absolute temperature. PV 1 v QQ'IZ Since PV = RT, R = ^===i^Jl ) ==-0819. This number '0819 is the numerical value of the constant R for all gases, volume being measured in litres and pressure in atmospheres. Substances in sol iition behave exactly like ^ases, they follow the same laws and have the same constants. All the conceptions which have been acquired by the study of gases are applicable to solutions, and therefore to the phenomena of life. The osmotic pressure ^f_jx__sj2lutiQn_js the force with which the molecules of the^ solute, like gaseous, molecules, strive to diffuse into space, and press on the limits which confine them, the containing vessel being represented by the surfaces of the solution. Osmotic pressure is measured in exactly the samc__wa^jj^jrag^ To measure steam pressure we insert a manometer in the walls of the boiler. In the same way we may use a manometer to measure osmotic pressure. We attach the tube to the walls of the porous vessel, allow the solvent to increase in volume under the pressure of the solute, and measure the rise of the liquid in the manometer tube. Pfcffer's Apparatus. Pfeffer has designed an apparatus for the measurement of osmotic pressure. It consists of a vessel of porous porcelain, the pores of which are filled with a colloidal solution of ferrocyanide of copper. This forms a semi-permeable membrane which permits the passage of water into the vessel, but prevents the passage of sugar or of any SOLUTIONS 19 colloid. The stopper which hermetically closes the vessel is pierced for the reception of a mercury manometer. The vessel is filled with a solution of sugar and plunged in a bath of water. The volume of the solution in the interior of the vessel can vary, since water passes easily in either direction through the pores of the vessel. The boundary of the solvent has become extensible, and its volume can increase or diminish in accordance with the osmotic pressure of the solute. Under the pressure of the sugar water is sucked into the vessel like air into a bellows, the solution passes into the tube of the manometer, and raises the column of mercury until its pressure balances the osmotic pressure of the sugar molecules. Osmotic Pressure follows the Laws of Gaseous Pressure. This osmotic pressure is in fact gaseous pressure, and may be measured in millimetres of mercury in just the same way. We may thus show that osmotic pressure follows the laws of gaseous pressure as defined by Boyle, Dalton, and Gay- Lussac. The coefficient of pressure variation for change of temperature is the same for a solute as for a gas. The formula PV = RT is applicable to both. The numerical value of the constant 11 is also the same for a solute as for a gas. being "0819 for one gramme-molecule of either, when the volume is expressed in litres and the pressure in atmospheres. The formula PV = RT shows that for a given mass, with the same volume, th_Jrcs^^ absolute ten ipcra t irre. Osmotic Pressure of Sugar. A normal solution of sugar., containing 342 grammes of sugar per litre, has a pressure of 22*35 atmospheres, and it may well be asked why such an enormous pressure is not more evident. The reason will be found in the immense frictional resistance to diffusion. Frictional resistance is proportional to the area of the surfaces in contact, and this area increases rapidly with each division of the substance. When a solute is resolved into its com- ponent molecules, its surface is enormously increased, and therefore the friction between the molecules of the solute and those of the solvent. Isotonic Solutions. Two solutions which have the same 20 THE MECHANISM OF LIFE o^^tic j)r^sure are said to be isp-osmotic or isotonic. When comparing two solutions of different concentration, the solution with the higher osm otic grcssurc_ is said to be Ivy per - tqnic, and that with^ jhe jower osmotic pressure Irypotonic. Lowering of the Freezing Point. Pure water freezes at C. Haoult showed that the introduction of a non-iqnizable substance, such as .sugar or alcohol, lowers the freezing^omt of a solution in proportion to the molecular concentration o_f the solute. One gramme-molecule of the solute introduced into one litre of the solution lowers its temperature of congelation by 1'85C. Thus a normal solution of _any n on - ipnizablc substance in water freezes aj>^ 1 '85 C. The measure- ment of this lowering of the freezing point is called Cryoscopy, a method which is becoming of great utility in medicine. Cryoscopy of Blood. In order to determine the osmotic pressure of the blood at 37 C., i.e. 98*6 F., the normal temperature, we proceed as follows. On freezing the blood, we find that it congeals at "56. Its molecular concentration '56 is therefore ----- = '30, or about one-third of a gramme- 1 *oO molecule per litre. Its osmotic pressure at C. is therefore '3 x 2&'35 = 6 '7 atmospheres. The increase of pressure with temperature is the same as for a gas, viz. ^, or '00367 of its pressure at for every degree rise of temperature. The increase of pressure at 37 is therefore '00367 X 37 X 6*7= '9 atmospheres. The total osmotic pressure at 37 is therefore 6*7 + '9 = 7*6 atmospheres. Rise of Boiling Point. Water under atmospheric pressure boils at a temperature of 100 C. The addition of a solute whose solution does not conduct electricity, such as sugar, causes a rise in the boiling point proportional to the molecular concentration of that solute. Lowering of the Vapour Tension. Th^j^oi^jtoisioi^of a liquid is lowered by the addition of a solute. A liquid boils at the temperature at which its vapour tension equals thaj; of__the atmosphere. Since an aqueous solution of sugar at atmospheric pressure does not begin to boil at 100 C., it is manifest that its vapour tension is then less than that of the SOLUTIONS 2 1 atmosphere. The addition of a solute such as sugar, whose solution is not ionizable, and therefore does not conduct electricity, lowers the vapour tension of the solution in proportion to the molecular concentration of the solute. Corresponding Values. We have thus found five properties of a solution which vary proportionally, so that from the measurement of any one of them we can determine the corresponding values of all the others. These are 1. The Molecular Concentration. 2. The Osmotic Pressure. 3. The Diminution of Vapour Tension. 4. The Raising of the Boiling Point. 5. The Lowering of the Freezing Point. Cryoscopy. The usual method employed for the deter- mination of the molecular concentration and osmotic pressure of a solution is by cryoscopy the measurement of its temperature of congelation. A very sensitive thermometer is used, the scale of which extends over only 5 and is divided into hundredths of a degree. The liquid under examination is placed in a test tube, in which the bulb of the thermometer is plunged, and this is supported in a second tube with an air space all round it. The whole is then suspended to the under side of the cover of the refrigerating vessel, which may be cooled either by filling it with a freezing mixture, or by the evaporation of ether. During the whole of the operation the liquid is agitated by a mechanical stirrer. The first step is to determine the freezing point of distilled water. As the water cools the mercury gradually descends in the stem of the thermometer till it reaches a point below the zero mark at C. As soon as ice begins to form the mercury rises, at first rapidly and then more slowly, reaches a maximum, and finally descends again. This maximum reading is the true, point of congelation. The inner tube is then emptied, care being taken to leave a few small ice crystals to serve as centres of congelation for the subsequent experiment, thus avoiding supercooling of the solution. The process is then repeated with the solution under examination. The difference between 22 THE MECHANISM OF LIFE / rccx J 11 K PQJ n -t- s ..A 8 . th required " Jo \veri tig of the freezing point. " Cryoscopy is the method most used in biological research to determine molecular concentration. It has, however, some grave defects. It necessitates several cubic centimetres of the liquid under examination. It gives us the constants of the solution at the temperature of free/ing, which is far below that of life. Organic liquids are easily altered and are extremely sensible to minute differences of temperature, cryoseopy therefore gives us no information as to the con- stitution of solutions under normal conditions. It is desirable to have some other method of determining molecular con- centration and the other interdependent constants at the normal temperature of life. A much better method, were it possible, would be the direct determination of the vapour tension of the solutions under normal conditions of temperature and pressure. Molecular Lowering* of the Freezing Point. For every substance whose solution is not ionized and therefore does not conduct electricity, the lowering of the free/ing point is the same, vi/. 1'85 C. for each gramme-molecule of the solute per litre of the solution. Determination of the Molecular Concentration. In order to obtain the molecular concentration of a non-ionizable substance, we have only to determine the lowering of the freezing point. Let A be the lowering of the freezing point of any solution. Orijdividnig it by 1'85 ...(the lowering of the freezing point for a normal .solution), we obtain the lumibcr of in a litre of the solution. If n be the A number of gramme-molecules per litre, then n=- ------ . 1 *Ot) Determination of the Osmotic Pressure. The osmotic pressure P of a solution may be obtained by multiply- ing its molecular concentration n_ by 22*35 atmospheres. P = n x 22-35 = -A: x 22-35. loo Determination of Molecular Weight. The lowering of the freezing point also enables us to calculate the molecular SOLUTIONS 23 weight of any non-ionizable solute. Thus Bouchard has been able to determine by means of cryoscopy the mean molecular weight of the substances eliminated by the urine. A_wejjght x of the subs taiice is dissolved in a litre of .wateiy and _ thejowgr- ing of the free/ing point is observed. The vajue thus found divided by 1/85 gives us n, the number of gi'aninie-mplecules per_Jitre. The molecular weight M may be determined by dividing the original weight x by n. The study of osmotic pressure was begun by the Abbe Nollet ; and one of his disciples, Parrot, at an early date thus described its importance : " It is a force analogous in all respects to the mechanical forces, a force able to set matter in motion, or to act as a static force in producing pressure. It is this force which causes the circulation of heterogeneous matter in the liquids which serve as its vehicle. It is this force which produces those actions which escape our notice by their minuteness and bewilder us by their results. It is for the infinitely small particles of matter what gravitation is for heavy masses. It can displace matter in solution upwards against gravity as easily as downwards or in a horizontal direction. 1 " Thus the recognition of the fact that a substance in solution is really a gas, has at a single stroke put us in possession of the laws of osmotic pressure laws slowly and laboriously discovered by the long series of investigations on the pressure of gases. Osmotic pressure plays a most important role in the arena of life. It is found at work in all the phenomena of life. When osmotic pressure fails, life itself ceases^ CHATTER III ELECTROLYTIC SOLUTIONS Solutions wliicli conduct Electricity. The laws of solution which we have studied in the previous chapter apply only to those solutions, chiefly of organic origin, which do not conduct electricity. Solutions of electrolytes such as the ordinary salts, acids, and bases, which are ionized on solution, give values for the various constants of solution which do not accord with those required by theory. If, for instance, we take a gramme- molecule of an electrolyte such as chloride of sodium, and dissolve it in a litre of water, we find that the lowering of the free/ing point is nearly double the theoretical value of 1'85. The same holds good for the osmotic pressure, and for all the constants which are proportional to the molecular concentra- tion of the solute. The solution behaves, in each case, as if it contained more than one gramme-molecule of sodium chloride per litre. It behaves, in fact, as if it contained i times the number of molecules of solute originally introduced into it. If n be the original number of molecules, then it will apparently contain ri = m molecules. This law is universal for all electrolytic solutions; the theoretical value for their concen- tration, osmotic pressure, and all the proportional physical constants must be multiplied by this quantity, =-, which is n the ratio of the apparent number of the molecules present to the number originally introduced. A similar dissociation of the molecule is observed in the case of many gases. The vapour of chloride of ammonium, for instance, is decomposed by heat, and it may be shown experimentally that the increase of pressure on heating above ELECTROLYTIC SOLUTIONS 25 that which theory demands, is due to an increase in the number of the gaseous molecules present. Some of the vapour particles are dissociated into two or more fragments, each of which plays the part of a single molecule. Arrhenius, in 1885, advanced the hypothesis that the apparent increase in the number of molecules of an electrolytic solution was also due to dissociation. This interpretation at once threw a Hood of light on a number of phenomena hitherto obscure. Coefficient of Dissociation. We have seen that in order to obtain values which accord with experiment we have to multiply the number of gramme-molecules of the solute by the coefficient i, which is called the Coefficient of Dis- sociation. This coefficient of dissociation, i, may be found by observing the lowering of the freezing point of a normal solution, and dividing it by T85. = _~ . The coefficient of dissociation varies with the degree of concentration of the solution, rising to a maximum when the solution is sufficiently diluted. If we know /, the coefficient of dissociation for a given solute, contained in a solution of a definite concentration, we can find n' 9 the number of particles present in a solution containing n gramme-molecules of the solute per litre, since n' = in. On the other hand, if from a consideration of its free/ing point and other constants we find that an electrolytic solution appears to contain ri gramme-molecules per litre, the real number of chemical gramme-molecules in one litre / of the solution will be only =n. i Very concentrated solutions do not conform to these laws. In this they resemble gases, which as they approach their point of condensation tend less and less to conform to the laws of gaseous pressure. Electrolysis. If we take a solution of an acid, a salt, or a base, and dip into it two metallic rods, one connected to the positive and the other to the negative pole of a battery, we 26 THE MECHANISM OF LIFE find that the metals or metallic radicals of the solution are liberated at the negative pole, while the acid radicals of the salts and acids and the hydroxyl of the bases are liberated at the positive pole. The liberated substances may either be dis- charged unchanged, or they may enter into new combinations, causing a series of secondary reactions. Electrolytes. Solutions which conduct electricity are called Electrolytes, and the conducting metallic rods dipping into the solution are the Electrodes. Faraday gave the names of Ions to the atoms or atom -groups liberated at either electrode. The ions liberated at the positive electrode are the Anions, and those at the negative electrode are the Cations. The only solutions which possess any notable degree of electrical conductivity are the aqueous solutions of the various salts, acids, and bases, and in these solutions only do we meet with those phenomena of dissociation which arc evidenced by anomalies of osmotic pressure, free/ing point and the like, anomalies which show that the solution contains a greater number of molecules than that indicated by its molecular concentration. These anomalies are due to dissociation, the division of some of the molecules into fragments, each of which plays the part of a separate molecule, contributing its quota to the osmotic tension and vapour pressure of the solution, in fact to all the phenomena which are dependent on the degree of molecular concentration. The electrical conductivity of a solution is therefore proved to be dependent on its molecular dissociation. Arrheniufi' Theory of Electrolysis. In 1885, Arrhenius brought forward his theory of the transport of electricity by an electrolyte. According to this hypothesis, the electric current is carried by the ions, the positive charges by the cations, and the negative charges by the anions. In virtue of the attraction between charges of different sign, and repulsion between charges of like sign, the cations are repelled by the positive charge on the anode, and attracted by the negative charge on the cathode. Similarly the anions are repelled by the cathode and attracted by the anode. ELECTROLYTIC SOLUTIONS 27 An electrolytic solution contains three varieties of particles, positive ions or cations, negative ions or anions, and un- dissociated neutral molecules. The molecular concentration of such a solution, with the corresponding constants, depends on the total number of these particles, I.e. the sum of the ions and the undissociated neutral molecules. We may indicate an ion by placing above it the sign of its electrical charge, one + - sign for each valency. Thus Na and Cl indicate the two ions ++ of a salt solution ; Cu and S() 4 the two ions of a solution of sulphate of copper. A point is sometimes substituted for the -|- sign, and a comma for the sign. Thus Na' and (T ; Cu'-and SO 4 " My friend T)r. Lewis Jones has given a very vivid picture of the processes which go on in an electrolytic solution when an electric current is passing. He compares an electro- lytic cell to a ballroom, in which are gyrating a number of dancing couples, representing the neutral molecules, and a number of isolated ladies and gentlemen representing the anions and cations respectively. If we suppose a mirror at one end of the ballroom and a buffet at the other, the ladies will gradually accumulate around the mirror, and the gentlemen around the buffet. Moreover, the dancing couples will gradually be dissociated in order to follow this movement. Degree of Dissociation. The degree of dissociation is the fraction of the molecules in the solution which have under- gone dissociation. Let n be the total number of molecules of the solute, and n the number of dissociated molecules. Then ~ will represent the degree of dissociation. Let Jc be the n number of ions into which each molecule is split. Then a = 71 , i.e. the degree of dissociation is the ratio of the nk number of ions actually present in a solution to the number which would be present if all the molecules of the solute were dissociated. Let n be the total number of particles present in a solution 28 THE MECHANISM OF LIFE containing n molecules, each of which is composed of Jc ions. Then if a is the degree of dissociation, - = 1 + (*-!) = . n We thus obtain i the coefficient of dissociation, in terms of the degree of dissociation a and the number of ions in each molecule k. If there is no dissociation, i.e. if = (), then n' = n, and i = 1. If all the molecules are dissociated, a = 1, and i = k. Faraday's Law. Faraday found that the quantity of electricity required to liberate one gramme-molecule of any radical is 96*537 coulombs for each valency of the radical. Electrochemical Equivalent. The electrochemical equivalent of a radical is the weight liberated by one coulomb of electricity. It is equal to the molecular weight of the ion, divided by 96'537 times its valency. Electrolytic Conductivity. The conductivity of an electro- lyte is the inverse of its resistance. C= ~. For a given difference of potential the conductivity of an electrolyte is proportional to the number of ions in unit volume, the electrical charge on each ion, and the velocity of the ions. The specific conductivity A of an electrolyte is the conductivity of a cube of the solution, each face of which is one square centimetre in area. The molecular conductivity of an electrolyte is the conductivity of a solution containing one gramme-molecule of the substance placed between two parallel conducting plates, one centimetre apart. The molecular conductivity is independent of the volume occupied by the gramme-molecule of the solute, depending only on the degree of dissociation. The molecular conductivity U is equal to the product of V, the volume of the molecule, by A, its specific conductivity. U = VA. Whence A=- r , i.e. the specific ELECTROLYTIC SOLUTIONS 29 conductivity equals the molecular conductivity divided by the volume. The conductivity of an electrolyte is proportional to the number of ions in a volume of the solution containing one gram me- molecule. Let M w be the conductivity for complete dissociation and M v the molecular conductivity at the volume V. Then - = l ' =~-=a, the degree of dissociation. This M^ -iik n is OstwaWs law, which says that the degree of dissociation is equal to the ratio of conductivity when the gram me- molecule occupies a volume V, to its conductivity when the solution is so dilute that dissociation is complete. Hence the degree of dissociation may also be determined by comparing the electrical conductivities of two solutions of different degrees of concentration. SO, SO, SO, + 4- +4- + + Cu Cu Cu SO, SO, S0 4 Cu Cu Cu FIG. i. Before the passage of the current. SO 4 ++ Cu Cu Cu Cu S0 4 S0 4 SO 4 S0 4 SO. ++ ++ Cu Cu FIG. 2. After the passage of the current. Velocity of the Ions. If the electrolytic cell is divided into two segments by means of a porous diaphragm, we shall find after a time an unequal distribution of the solute on the two sides. For instance, with a solution of sulphate of copper, after the current has passed for some time there will be a diminution of concentration in the liquid on both sides of the diaphragm, but the loss will be very unequally divided. Two- thirds of the loss of concentration will be on the side of the negative electrode and only one-third on the positive side. In 1853, Hittorf gave the following ingenious explanation of this phenomenon : 30 THE MECHANISM OF LIFE Fig. 1 represents an electrolytic vessel containing a solution of sulphate of copper, the vertical line indicating a porous partition separating the vessel into two parts. Fig. 2 shows the same vessel after the passage of the current. The acid radical has travelled twice as fast as the metal. For each copper ion which has passed through the porous plate towards the cathode two acid radicals have passed through it towards the anode. Three ions have been liberated at either electrode, but in consequence of the difference of velocity with which the positive and the negative ions have travelled, the negative side of the vessel contains only one molecule of copper sulphate and has lost two-thirds of its molecular concentration, while the positive side contains two molecules of copper sulphate and has only lost one-third of its concentration. This proves clearly that the ions move in different directions with different velocities. Let u be the velocity of the anions, and v the velocity of the cations. Let n be the loss of concentration at the cathode, and 1 n the loss of concentration at the anode. Then - = --, i.e. the loss of concentration at the cathode is v 1 n to the loss of concentration at the anode as the velocity of the anions is to that of the cations. Hence by measuring the loss of concentration at the two electrodes, we have an easy means of determining the comparative velocity of different ions. In 1876, Kohlrausch compared the conductivity of the chlorides, bromides, and iodides of potassium, sodium, and ammonium respectively. He found that altering the cation did not affect the differences of conductivity between the three salts, thus showing that these differences of conductivity were dependent on the nature of the anion only, and not on the particular base with which it was combined. The difference of conductivity between an iodide and a bromide, for example, is the same whether potassium, sodium, or ammonium salts are compared. A similar experiment has been made with a scries of cations combined with various anions. The difference of conductivity of the salts in the series is the same whichever anion is used, i.e. the difference of conductivity between potas- sium chloride and sodium chloride is the same as that between ELETCROLYTJC SOLUTIONS 31 potassium bromide and sodium bromide. Hence we may con- clude that the conductivity of any salt is an ionic property. KohlrauscK's law may be expressed by the formula c = r/(?/ + t>), where c is the conductivity of the salt, d the degree of dissociation, i.e. the fraction of the electrolyte broken up into ions, and u and v the velocity of the anions and cations respectively. When all the molecules of the electrolyte arc dissociated, During the whole period of growth there is an abundant liberation of bubbles of gas, which is acurately limited to a belt around the base of the growth, and sometimes also to a cap at the summit. Since morphological differentiations of different parts is but the result of differences of evolution, i.e. of functional differences of the various parts, we may consider that osmotic growths possess the faculty of organization I ; ke living beings. An osmotic growth may be wounded, and a wound delays its growth and development like a disease or an accident in a living being. A wound in an osmotic production may also become cicatrized and covered with a membrane, when the growth will recommence exactly as in a living being. An osmotic growth is a transformer of energy. It increases in bulk, pushing aside the mother liquor, and thus doing external work. An osmotic growth has a temperature above its medium, since the chemical reaction of which it is the seat is accompanied by the production of heat. We know 154 THE MECHANISM OF LIFE but little of the transformation of energy which takes place in an osmotic production, but we may say with certainty that it is capable of transforming both chemical energy and osmotic energy into heat and mechanical motion. An osmotic production is the arena of complicated chemical phenomena which produce a veritable metabolism. It has long been known that diffusion and osmosis may determine various chemical transformations. H. St. Clair Deville has demonstrated that certain unstable salts are partially decomposed by diffusion. Thus during the diffusion of alum, the sulphate of potash is separated from the sulphate of aluminium. Similarly, when the chloride or acetate of aluminium is caused to diffuse, the acids become separated from the aluminia. This decomposition is the result of the different resistance which the medium offers to the diffusion of different ions. This difference of resistance may even cause a difference of potential between two media, similar to the differences of potential in living organisms. Frequently also a difference of hydration in the chemical substances on either side of an osmotic membrane will determine a chemical reaction, which like all other chemical reactions is accompanied by a corresponding transformation of energy. The study of these chemical metamorphoses and the transformations of energy in osmotic growths has opened up a new subject for experimental investigation in the field of organic chemistry. Coagulation. There is a most remarkable analogy between the phenomena of coagulation as seen in living beings and the phenomena which occur when the liquid in the interior of an osmotic growth comes into contact with the mother liquor. When the sap of a plant or the blood of an animal escapes into the air or water of the surrounding medium, it coagulates, i.e. it changes from a liquid to a gelatinous consistency. In the same way, when the liquid in the interior of an osmotic growth leaks out into the mother liquor it forms a gelatinous precipitate. This gelatinous precipitation is a physico- chemical phenomenon of the same nature as coagulation. It is by the study of coagulation in liquids less complex than blood that we may hope to elucidate the mechanism of the process, THE PHENOMENA OF LIFE 155 which is simply a physico-chemical phenomenon exactly analogous to gelatinous precipitation. Calcium phosphate is always prone to coagulate ; it has been called the gelatinous phosphate of lime, and we have already seen how readily tribasic calcium phosphate takes the form of beautiful trans- parent colloidal membranes which are gelatinous in texture. We may obtain colloidal precipitates exactly analogous to coagulated albumin by mixing a weak solution of chloride of calcium with potassium carbonate or tribasic phosphate. Like albumin this precipitate forms flakes, and is deposited slowly as a gelatinous colloidal mass. Like albumin also this calcic solution is coagulated by heat ; a solution of a calcic salt of a volatile acid on heating forms a precipitate which has all the appearance of albumin coagulated by heat. Finally, Arthus and Pages have shown that blood does not coagulate when deprived of its calcium salts by the addition of alkaline oxalates, fluorides, or citrates, and that the blood thus treated recovers its coagulability on the addition of a soluble salt of calcium. The coagulation of milk is also a calcium salt precipitation. Coagulation therefore would seem to be merely the colloidal precipitation of a salt of calcium. Diffusion and osmosis are the elementary phenomena of life. All vital phenomena result from the contact of two colloidal solutions, or of two liquids separated by an osmotic membrane. Hence the study of the physics of diffusion and osmosis is the very basis of synthetic biology. A living being exhibits two sorts of movements, those which are the result of stimulus from without, and those which are determined by an excitation arising from within. In the higher animals the stimulus or exciting energy coming from the entourage may be infinitely small when compared with the amount of energy transformed. Moreover, the response to an identical excitation may so vary as to give to these different responses an appearance of spontaneity. There is in reality no spontaneity, since the difference in response is governed by previous external impressions which have left their record on the machinery. There is in fact no such thing as a spontaneous action, since every action of a living i$6 THE MECHANISM OF LIFE being has as its ultimate cause a stimulus or excitation coming from without. The movements of the second category are also conditioned by an excitation, but the stimulus comes from within the organism. These movements consist principally of changes of nutrition, or movements of the circulation and respiration ; they are rhythmic in character and are probably produced by the same chemico-physical causes which determine rhythmic movements outside the living body. Just in the same wry osmotic growths present two sorts of movements, external movements and those which are connected with their nutrition. A free osmotic growth swimming in the mother liquor will alter its position and form under the influence of the slightest exterior excitation or vibration. It responds to every variation of temperature, or to a slight difference of concentration produced by adding a single drop of water, and reacts to every exterior influence by displacement or deformation. An osmotic growth also shows indications of movements which are connected with its nutrition, and these movements are rhythmic, like those of respiration or circulation in a living organism. The growth of an osmotic production shows itself not as a continuous process but periodically. The water traverses the membrane, raises the pressure, and distends the cell ; at first the cell wall resists by reason of its elasticity, it then suddenly relaxes, yielding to the osmotic pressure and bulging out at a thinner spot on the surface ; the internal pressure falls suddenly, and there is a pause in the growth. This rhythmic growth may be best observed by sowing in a solution of a tribasic alkaline phosphate, pellets composed of powdered calcium chloride moistened with glycerine, to which has been added 1 per cent, of monobasic calcium phosphate. The experiment is so arranged as to bend or incline the growing stems which shoot out from these grains. This may be done by carefully pouring above the mother liquor a layer of water, or a less concentrated solution. As the internal osmotic pressure rises, the drooping extremity of the twig will become turgescent and gradually lift itself THE PHENOMENA OF LIFE 157 up, and then suddenly fall again for several millimetres. We have frequently watched this rhythmic movement for an hour or more a slow gradual elevation of the extremity of the twig and a rapid fall recurring every four seconds or so. It may be objected that the substance of an osmotic growth is continually undergoing change, whereas a living organism transforms into its own substance the extraneous matter which it borrows from its environment. The distinction, however, is only an apparent one. The substance of a living being is also continually undergoing chemical change ; it does not remain the same for a single instant. We see an evidence of this change in the evolution of age ; the substance of the adult is not that of the infant. In some living organisms such as insects, especially the ephemeridae who have but a brief existence, this change of substance is even more rapid than that in an osmotic growth. It has been objected that osmotic productions cannot be compared with living organisms since they contain no albuminoid matter. This is to consider life as a substance, and to confound the synthesis of life with that of albumin. If albumin is ever produced by synthesis in the laboratory it will probably be dead albumin. All living organisms contain albumin; this is probably due to the fact that albuminoid matter is particularly adapted for the formation of osmotic membranes. Our osmotic productions are composed of the same elements as those which constitute living beings ; an osmotic growth obtained by sowing calcium nitrate in a solution of potassium carbonate with sodium phosphate and sulphate contains all the principal elements of a living organism, viz. carbon, oxygen, hydrogen, nitrogen, sulphur, and phosphorus. The whole of the vegetable world is produced by the osmotic growth of mineral substances, if we except the small amount of organic matter contained in the seeds. The most important problem of synthetic biology is not so much the synthesis of the albuminoids as the reduction of carbonic acid. In nature this reduction is accomplished by the radiant energy of the sun, by the agency of the catalytic action of chlorophyll. 158 THE MECHANISM OF LIFE The physico-chemical study of osmotic growth is as yet hardly begun ; we have but indicated the method, the way is open, and the problems awaiting solution are legion. Only work and ever more work and workers are required. Experiments should be made with substances which are chemically unstable like the albuminoids, substances which readily combine and dissociate again, alternately absorbing and giving up the potential energy which is the essence of life. Experiments should also be made with substances which readily unite or decompose under the influence of water, since hydration and hydrolysis appear to be the dominant mechanism in all vital reaction, as they undoubtedly are in osmotic growth, which consists of an increase of hydration on one side of an osmotic membrane and a diminution on the other side. Life is not a substance but a mechanical phenomenon ; it is a dynamic and kinetic transference of energy determined by physico-chemical reactions; and the whole trend of modern research leads to the belief that these reactions are of the same nature as those met with in the organic world. It is the grouping of physical reactions and their mode of associa- tion and succession, their harmony in fact, which constitutes life. The problem we have to solve in the synthesis of life is the proper attuning and harmonizing of these physical phenomena, as they exist in living beings, and there should be no absolute impossibility in our some day realizing this harmony in whole or in part. Albert Gaudry says : " I cannot conceive why in determin- ing the connecting links of the animal world the fact that an organic body is formed of such and such elements should be of greater importance than the manner in which these elements are grouped. Descartes regarded extension as the essential property of an organized being ; he supposed it to be inert of itself, and that it had the Deity for its motive force. To-day the hypothesis of Descartes has given way to that of Leibnitz, who regards force as the essential property of the living being, the visible and tangible matter being only of secondary importance. If we regard the living being as a force, this orce is able to aggregate matter under such and such a form, THE PHENOMENA OF LIFE 159 with such or such a structure, and such or such a chemical essence. It does not seem that the classification depending on differences of substance are any more important than those which depend on differences of form." The biological interest of osmotic productions is quite independent of the chemical nature of the substances which enter into their growth. All substances which produce osmotic membranes by the contact of their solutions exhibit phenomena analogous to those of nutrition. Osmotic morpho- genesis is a physical phenomenon resulting from the contact of the most diverse substances. It has given us our first glimpse of the manner in which a living being may be supposed to have been formed according to the ordinary physical laws of nature. We cannot at present produce osmotic growths with all the combinations found in living beings, but that is only because chemistry still lags far behind physics in the synthesis of organic forms. We are often told " not to force the analogy. " But error is equally produced by the exaggeration of unimportant differences. We have already seen that nutrition, absorption, transformation, and excitation are not the characteristics of living organisms alone ; nor is reaction to external impressions the appanage only of animate beings. To insist on the resem- blance between an osmotic production and a living being is not to force an analogy but to demonstrate a fact. Let us briefly recapitulate. An osmotic growth has an evolutionary existence ; it is nourished by osmosis and intus- susception ; it exercises a selective choice on the substances offered to it; it changes the chemical constitution of its nutriment before assimilating it. Like a living thing it ejects into its environment the waste products of its function. Moreover, it grows and develops structures like those of living organisms, and it is sensitive to many exterior changes, which influence its form and development. But these very pheno- mena nutrition, assimilation, sensibility, growth, and organization are generally asserted to be the sole character- istics of life. CHAPTER XIII EVOLUTION AND SPONTANEOUS GENERATION BY many biologists, even at the present day, the origin and evolution of living beings is considered to be outside the domain of natural phenomena, and hence beyond the reach of experimental research. The change in our views on this subject is due to a Frenchman, Jean Lamarck, who was the true originator of the scientific doctrine of evolution. At a time when the miraculous origin of every living being was regarded as an unchangeable verity, and was defended like a sacred dogma, Lamarck boldly formulated his theory of evolution, with all its attendant consequences, from spontaneous generation to the genealogy of man. In his Philosophic Zooloffique, which appeared in 1809, Lamarck put forth his claim to regard all the phenomena of life, of living beings, and of man himself as pertaining to the domain of natural phenomena. According to him, all bodies which are met with in nature, organic and inorganic alike, are subject to the same laws. Life is a physical phenomenon, and all the processes of life are due to mechanical causes, either physical or chemical. He writes : " A leur source le physique et le moral ne sont sans doute qu\me seule et mem* chose. II faut rechercher dans la consideration de Torganisation les causes memes de la vie." In the intellectual evolution of the human mind perhaps no advance has been more important than that of Lamarck the conquest of the domain of life by human intelligence. In conformity with the true scientific method, he founds his doctrine on the facts and phenomena of nature. " I confine myself," he says, " within the bounds of a simple contemplation 1 60 EVOLUTION 161 of nature. 11 It was this observation of the gradual perfecting of living organisms from the simplest to the most com- plicated that inspired Lamarck with the idea of evolution and transformation. " How," he says, " can we help searching for the cause of such wonderful results? Are we not com- pelled to admit that nature has produced successively bodies endowed with life, proceeding from the simplest to the most complex ? " The various products of nature have been divided into classes, genera, and species, simply to facilitate their study. Modern research tends to show that there is no definite line of demarcation even between the animal, vegetable, and mineral kingdoms. All our classification is artificial, and the passage from one division to another is gradual and insensible. Lamarck expresses this idea very clearly : " We must remember that classes, orders, and families, and all such nomenclature, are methods of our own invention. In nature there are no such things as classes or orders or families, but only individuals. As we become better acquainted with the productions of nature, and as the number of specimens in our collections in- creases, we see the intervals between the classes gradually fill up, and the lines of separation become effaced. 11 Lamarck also raises his voice against the supposed immutability of species. " Species have only a relative constancy, depending on the circumstances of the individuals. The individuals of a given species perpetuate themselves with- out variation only so long as there is no variation in the circumstances which influence their existence. Numberless facts prove that when an individual of a given species changes its locality, it is subjected to a number of influences which little by little alter, not only the consistency and proportions of its parts, but also its form, its faculty, and even its organiza- tion ; so that in time every part will participate in the mutations which it has undergone. 11 Lamarck also clearly affirms the fact of spontaneous generation. " I hope to prove,"he says, " that nature possesses means and faculties for the production of all the forms which we so much admire. Rudimentary animals and plants have ii 1 62 THE MECHANISM OF LIFE been formed, and are still being formed to-day, by spontaneous generation." Lamarck himself gives a resume of his doctrine in the following six propositions : 1. " All the organized bodies of our globe are veritable productions of Nature, which she has successively formed during the lapse of ages. 3. " Nature began, and still recommences day by day, with the production of the simplest organic forms. These so-called spontaneous generations are her direct work, the first sketches as it were of organization. 3. "The first sketches of an animal or a vegetable growth being begun under favourable conditions, the faculties of commencing life and of organic movement thus estab- lished have gradually developed little by little the various parts and organs, which in process of time have become diversified. 4. " The faculty of growth is inherent in every part of an organized body ; it is the primary effect of life. This faculty of growth has given rise to the various modes of multiplication and regeneration of the individual, and by its means any progress which may have been acquired in the composition and forms of the organism has been preserved. 5. " All living things which exist at the present day have been successively formed by this means, aided by a long lapse of time, by favourable conditions, and by the changes on the surface of the globe in a word, by the power which new situa- tions and new habits have of modifying the organs of a body which is endowed with life. 6. "Since all living things have undergone more or less change in their organization, the species which have been thus insensibly and successively produced can have but a relative constancy, and can be of no very great antiquity." The admirable work of Lamarck was absolutely neglected in France, where it was treated as unworthy even of consider- ation. This neglect profoundly afflicted Lamarck, who gradually sank a victim to the opposition of his contem- poraries. He left, however, one disciple, Etienne Jeoffroy St. EVOLUTION 163 Hilaire, but he too was soon reduced to silence under the weight of authority of his adversaries. Before the doctrine of evolution could live and take its proper place, it had to be reborn in England the country of liberty. This resuscitation was due to Darwin, who added to FIG. 62. Osmotic vegetation. it his illuminating doctrine of natural selection. But apart from this and a perfecting of its various details, Lamarck had already formulated the doctrine of evolution with perfect precision. Lamarck's work was still-born, whereas that of Darwin lived and grew to its full development. This was due, not to any imperfection or insufficiency in Lamarck's work, but 64 THE MECHANISM OF LIFE the milieu into which it was born. It was the environment hat stifled the offspring of Lamarck. In 1868, Ernest Haeckel speaks of the genius of Lamarck a these words : " The chief of the natural philosophers of France is Jean Lamarck, who takes his place beside Goethe nd Darwin in the history of evolution. To him belongs the nperishable glory of being the first to formulate the theory f descent, and of founding the philosophy of nature on the slid basis of. biology," and adds, " There is no country in lurope where Darwin's doctrine has had so little influence as in 'ranee " Haeckel has but done tardy justice in his discovery f and testimony to the genius of Lamarck. The spirit of opposition does not seem to have much banged in France since Lamarck's time. In 1907 the Lcademie des Sciences de Paris excluded from its Comptes benches the report of my researches on diffusion and osmosis, ecause it raised the question of spontaneous generation. The majority of scientists seem to consider that the question f spontaneous generation was definitely settled once for all hen Pasteur's experiments showed that a sterili/ed liquid, ept in a closed tube, remained sterile. Without the idea of spontaneous generation and a physical tieory of life, the doctrine of evolution is a mutilated ypothesis without unity or cohesion. On this point Lamarck Deaks most clearly : " Although it is customary when one jeaks of the members of the animal or vegetable kingdom to ill them products of nature, it appears that no definite con- option is attached to the expression. Our preconceived otions hinder us from recognising the fact that Nature herself ossesses all the faculties and all the means of producing living eings in any variety. She is able to vary, very slowly but ithout cessation, all the different races and all the different >rms of life, and to maintain the general order which we see 1 all her works." The doctrine of Lamarck is frequently misinterpreted, ^e often hear it expressed as " Function makes the organ/' or yen " Function creates the organ." This is equivalent to tying, "Life makes the living being," which is incomprehensible, EVOLUTION 165 making of function a sort of immaterial and independent entity which constructs a material organ in order to lodge within it. No such idea is to be found in all the works of Lamarck. He formulates his law in the following terms : " In every animal which is still undergoing development, the frequent and sustained use of any one organ increases its size and power, whereas the constant neglect of the use of such organ weakens and deteriorates it, so that it finally disappears." In his expression of this law Lamarck insists on the fact that organization precedes function. He affirms only that function, i.e. action and reaction, modifies the organ ; or, in other words, that organisms are modelled by the action of exterior forces acting upon them. It is in this sense only that function may be said to make an organ, but this mode of expression should be avoided, as it is apt to be misunderstood. Astronomy teaches us that our globe was detached from the sun in an incandescent state, and geology asserts that this earth has passed through a period of long ages when its temperature was incompatible with the existence of life. It was only with the cooling of the earth crust that it was possible for living beings to make their appearance. Hence they must of necessity have been produced spontaneously from terrestrial material under the influences of chemical and physical forces. This opinion imposes itself on all who reflect and judge freely. In the same way the doctrine of evolution necessitates as a corollary the doctrine of spontaneous genera- tion. The doctrine of evolution should reconstitute every link in the chain of beings from the simplest to the most compli- cated ; it cannot afford to leave out the most important of all, viz. the missing link between the inorganic and the organic kingdoms. If there is a chain, it must be continuous in all its parts, there can be no solution of continuity. Evolutionists like Lamarck and Haeckel admit spontaneous generation, not as the most probable, but as the only possible explanation of the phenomenon of life. Lamarck shows us the apparition of living things at a certain epoch of the earth's evolution, and the gradual develop- 166 THE MECHANISM OF LIFE ment of more complicated forms as the conditions changed on the surface of the globe. Darwin shows how heredity and natural selection tend to accentuate the variations which are favourable to existence. Haeckel demonstrates the parallelism between ontogenesis and philogenesis between the successive forms in the evolution of the embryo and the successive forms of the individual in the evolution of a race. These are great and admirable conquests of the human intelligence, they have FIG. 63. Marine forms of osmotic growth. demonstrated the first appearance and the progressive evolution of living beings ; it now only remains for us to explain them. The doctrine of evolution, while enforcing the fact of spontaneous generation and progressive evolution, gives us no hint as to the physical mechanism of such generation. It does not tell us by what forces, or according to what laws, the simpler forms of life have been produced, or in what manner differences of environment have acted in order to modify them. The doctrine asserts the simultaneous variations in organic forms and in the physical influences which produce them, but says EVOLUTION 167 nothing as to their mode of action. The Darwinian theory shows how acquired variations are transmitted and accentuated by natural selection, but it says nothing as to how these varia- tions may be acquired. In the same way we are in entire ignorance as to the physical mechanism of on togenetic develop- ment, the evolution of the embryo. The morphogenic action of diffusion produces osmotic growths of extreme variety. Most of these forms recall those of living things shells, fungi, corals, and algae. The analogy of function is quite as close as the resemblance of form. The study of osmosis, however, is as yet in its infancy, and osmotic productions vary with the physical conditions of chemical constitution, temperature, concentration, and the like. The study of the organizing action of osmosis on organic material has as yet been hardly attempted. Osmosis produces growths of great complexity, milch more complicated indeed than the more simple forms of living organisms. This marvellous complexity of an osmotic growth may be compared with another fact, the ontogenetic develop- ment of the ovum, a single cell which under favourable conditions of environment may evolve into a most complicated organism. These considerations lead to the belief that the beginning of life has not been the production of a simple primitive form from which all others are descended, but that a number of such primitive forms may have been produced, forms which by a rapid physical development attained a high degree of complexity. Osmotic morphogenesis shows us that the ordinary physical forces have in fact a power of organiza- tion infinitely greater than has been hitherto supposed by the boldest imagination. When we consider the ignorance in which we still remain as to the phenomena which pass before our very eyes, how can we expect to understand those which occurred in past ages, when the physical and chemical conditions were so immensely different from those which obtain in our own time ? What do we know even now of the physical and chemical phenomena which take place in the unfathomed depths of the ocean, where for aught we know even at the present time the same 1 68 THE MECHANISM OF LIFE process may be going on the genesis of life, and the emergence of living beings out of the inanimate mineral world ? " Even now," says Albert Gaudry, "polyps and oceanic animalculae are building up vast coral reefs and rocks. The oxygen and hydrogen which existed once was water, the oxygen and nitrogen which once made air, the carbon, the phosphorus, the silica and the lime which once were solid rock, now form the substance of living beings. The silica is deposited in the skeleton of a sponge or a radiolaria, the shell of a foraminifera or the carapace of a crustacean, or unites with phosphorus to form the bones of a vertebrate. A very tumult of life has succeeded to the primitive silence of inert matter. Life has invaded the earth, and we see on all sides the inanimate mineral kingdom being changed into a living world. 1 " The admission that life may have appeared on the earth under the influence of natural forces and according to physical laws arid conditions different from those of the present era throws a vivid light on the study of biogenesis, spontaneous generation, and evolution. The means of research are now indicated, and we have only to study the documents already in our possession in order to know the conditions which obtained when life first appeared on the globe. We must endeavour to reproduce these conditions and to study their effects. Since all living beings are formed of the same elements as those of the mineral world, the term "organic" 11 as applied to combinations can only be used in order to emphasize the complexity of their constitution. It was formerly believed that these organic combinations were the result of life, and could not be reproduced except by living organisms. To-day many of these organic substances are produced in the laboratory from inorganic materials. In the past history of the globe it is easy to imagine conditions which would facilitate the synthesis of organic substances without the interposition of life. At the temperature of the electric furnace, which was that of the earth at an early period of its evolution, chemical combinations are possible quite other than those obtaining under the present conditions of tempera- ture and pressure. At the higher temperature of the early EVOLUTION 169 geological era, silicides, carbides, phosphides, and nitrides were formed in stable combinations instead of the oxides, silicates, carbonates, phosphates, and nitrates of the present time. These combinations existed on the earth at a time when the conditions of temperature precluded the existence of water in a liquid state. As the temperature cooled, and the water vapour became condensed, it entered into chemical combination with the various rocks, producing organic com- pounds like acetylene, which results from the action of water on calcium carbide. H. Le'nicque has developed a theory as to the formation of various rocks under these conditions, which he communicated in 1903 to the French Society of Civil Engineers. The chemical evolution of the globe has undergone great changes as the temperature gradually fell and the constitution of its crust altered. As long as the temperature was higher than that at which water can exist, all chemical reactions must have taken place between anhydric substances, elements and salts in a state of fusion. These conditions are very different from those of the present-day chemistry, which is the chemistry of aqueous solutions. We may hope to be able to reproduce the earlier conditions by the experimental study of anhydric substances in a state of fusion. At a later period, that of the primary and secondary rocks, there was a uniform and constant temperature of about 40 C. The atmosphere was charged with water vapour, and all the conditions were present for the production of storms and tempests. The atmosphere during long ages must have been the seat of formidable and incessant electric discharges ; these discharges are the most powerful of all physical agents of chemical synthesis, and will cause nitrogen to combine directly to form various compounds nitrates, cyanides, and ammonia. Carbonic acid would also be present in abundance and would enter into combination with these nitrogenous compounds. In this way we may imagine that compounds were formed which by some process of physical synthesis subsequently gave rise to vast quantities of albuminoid matter. At that time the seas and oceans contained all those substances which have \70 THE MECHANISM OF LIFE since been fixed by the metamorphism of the primitive rocks, or deposited in the sedimentary strata. Most of the elements in our minerals were formerly in a state of solution in these primeval seas, which contained carbonates, silicates, and soluble phosphates in great abundance. As the crust gradually cooled, the terrestrial atmosphere of necessity altered in com- position, and the slow evolution of the atmosphere no doubt also exercised an influence on the development of living beings. Palaeontology teaches us that the earliest living organism appeared in the sea. The most ancient of living things, those of the primary ages, which were of greater duration than all other ages put together, were all aquatic. We find moreover that every living organism consists of liquids, solutions of crystalloids and colloids separated by osmotic membranes ; and it is significant that the ocean, that vast laboratory of life, is also a solution of crystalloids and colloids. It is evident, then, that we must look to the study of solutions if we would hope to discover the nature and origin of life. Life is an ensemble of functions and of energy-transforma- tions, an ensemble which is conditioned by the form, the structure, and the composition of the living being. Life, therefore, may be said to be conditioned by form, i.e. the external, internal, and molecular forms of the living being. All living things consist of closed cavities, which are limited by osmotic membranes, and filled with solutions of crystalloids and colloids. The study of synthetic biology is therefore the study of the physical forces and conditions which can produce cavities surrounded by osmotic membranes, which can associate and group such cavities, and differentiate and specialize their functions. Such forces are precisely those which produce osmotic growths, having the forms and exhibiting many of the functions of living beings. Of all the theories as to the origin of life, that which attributes it to osmosis and looks on the earliest living beings as products of osmotic growths is the most probable and the most satisfying to the reason. We have already seen that the seas of the primary and EVOLUTION 171 secondary ages presented in a high degree the particular conditions favourable for the production of osmotic growths. During these long ages an exuberant growth of osmotic vegetation must have been produced in these primeval seas. All the substances which were capable of producing osmotic membranes by mutual contact sprang into growth, the soluble salts of calcium, carbonates, phosphates, silicates, albuminoid matter, became organized as osmotic productions, FlG. 64. Osmotic shells and corals. were born, developed, evolved, dissociated, and died. Millions of ephemeral forms must have succeeded one another in the natural evolution of that age, when the living world was represented by matter thus organi/ed by osmosis. The experimental study of osmotic morphogeny adds its weight of evidence in the same direction. When we see under our own eyes the cells of calcium become organized, develop and grow in close imitation of the forms of life, we cannot doubt that such a transformation has often occurred in the past history of our planet, and the conviction becomes irresistible