Ill %\\t J. £- ^iii ICtbrarg . .V ^orilj (Earnlma JSfate College QH54I H3 . *fc% i— j 11481 This book may be kept out TWO \ ONLY, and is subject to a fine of t. CENTS a day thereafter. It is due on t day indicated below: i 57 SE )fc7 A/ 5 to»>5| J ' 0 Apr54 1 ^a1** | MAY 3Q 20ct5g3 Mflfl" 1 8 1963 0t- 1964 iii -^^■■^■i^k FEB I : MAR 23 — ., .-c THE FITNESS OF THE ENVIRONMENT THE MACMILLAN COMPANY NEW YORK • BOSTON • CHICAGO DALLAS • SAN FRANCISCO MACMILLAN & CO., Limited LONDON • BOMBAY • CALCUTTA MELBOURNE THE MACMILLAN CO. OF CANADA, Ltd. TORONTO THE FITNESS OF THE ENVIRONMENT AN INQUIRY INTO THE BIOLOGICAL SIGNIFICANCE OF THE PROPERTIES OF MATTER BY LAWRENCE J. HENDERSON ASSISTANT PROFESSOR OF BIOLOGICAL CHEMISTRY IN HARVARD UNIVERSITY IN PAET DELIVERED AS LECTURES IN THE LOWELL INSTITUTE, FEBRUARY, 1913 THE MACMILLAN COMPANY 1924 AU rights reserved PRINTED IN THE UNITED STATES OF AMERICA , Copyright, 1913, Bt THE MACMILLAN COMPANY. Set up and electrotyped. Published February, 1913. Nortoooti ^reas J. 8. Cushing Co. — Berwick 1 VII. The Ultimate Problem VIII. The Method of Solution \ . C7 xiii xiv CONTENTS CHAPTER III PASS WATER 72 I. Thermal Properties 80 A. Specific Heat 80 B. Latent Heat 92 C. Thermal Conductivity 106 D. Expansion before Freezing . . . .106 II. The Action of Water upon Other Substances 110 A. Water as a Solvent Ill B. Ionization 118 C. Surface Tension 126 CHAPTER IV CARBONIC ACID 133 I. Solubility 136 II. Acidity 140 CHAPTER V THE OCEAN 164 I. The Regulation of Physico-chemical Conditions 164 II. The Circulation of Water 180 III. The Ocean as Environment .... 183 CHAPTER VI THE CHEMISTRY OF THE THREE ELEMENTS . 191 I. Organic Chemistry 191 A. Valence 196 B. Hydrocarbons 197 CONTENTS , v lA. K C. Compounds of Carbon, Hydrogen, and Oxygen MM D. Other Organic Compounds . t |qq £. The Characteristics of Organic Substances . MM F. The Sugars ^ G. Hydrolysis ^ II. Inorganic Chemistry .... Mr III. Thermochemistry .... W;j CHAPTER VII THE ARGUMENT 249 I. Analysis of the Evidence . . . $50 II. The Exhaustiveness of the Treatment . |fl III. Summary ag» CHAPTER VIII LIFE AND THE COSMOS 274 I. The Significance of Fitness . 274 II. Vitalism ^Sl» A. The Vitalism of Bergson 293 B. Vitalism and Teleology 298 HI. Cosmic Evolution 30 1 A. The Periodic System 303 B. Teleology 305 CHAPTER I FITNESS PURPOSE AND ORDER IDEAS of purpose and order are among the first concepts regarding their en- vironment which appear, as vague antici- pations of philosophy and science, in the minds of men. In truth, when the manifold phenomena and experiences of daily life stored in the memory are critically scruti- nized, purpose and order seem naturally to suggest themselves as explanations of the universe. Day and night, the changing but recurring seasons, the fertilizing sunshine and rain, the flight of birds, the powers of the human hand, and all the beau tits and mysteries of nature cannot fail of such in- terpretation by the simple and untrained mind. Alike anthropology and the history of primitive civilizations bear witness to this natural tendency of thought. Such ideas pre- B 1 2 THE FITNESS OF THE ENVIRONMENT cede exact knowledge and civilization, and arise spontaneously among savage peoples. They are the solvent of the chaos, as which the outer world first presents itself to our eyes and hands, and they are the fabric of all theologies. As civilization has progressed, these early hypotheses have received endless criticism, and their definition has been continually sharpened. Meantime natural science has sought and provided ever more accurate accounts of the phenomena which first sug- gested them to man, and of countless other forms and transformations of matter and energy, and the discovery of laws of nature has steadily changed once quite mysterious order and purpose into the plainest of neces- sary results. Upon the advent of modern science order speedily began to receive its true account when, after only a half century of progress, dynamics through Newton provided a formu- lation of the laws which govern the most striking of all the orderly phenomena of nature.1 Since Newton's day the explana- 1 '* Die Newton'schen Principien sind genligend, um ohne Hinzuziehung eines neuen Princips jeden praktisch vorkom- menden mechanischen Fall, ob derselbe nun der Statik oder der Dynamik angehort, zu durchschauen. Wenn sich hierbei Schwierigkeiten ergeben, so sind dieselben immer nur mathe- FITNESS 3 tion of natural order as the automatic result of natural law has not ceased, and at length has become so nearly complete that the ap- pearance of order under any circumstances is now taken as proof of the existence of a law. The fate of the hypothesis of purpose in nature has been less simple, because the dis- covery of law, or even of the possibility of law, underlying adaptation and fitness was more difficult. Until the middle of the nine- teenth century the countless adaptations of organisms to the environment and the mani- fest fitness of nature for the activities of living things seemed to many biologists only explicable as the result of some directing force.1 Even skeptics were nearly or quite mathischer (formeller) und keineswegs mehr principieller Natur." — Mach, "Die Mechanik in Ihrer Entwickelung Historisch-Kritisch Dargestellt." Leipzig, 1897, 3d ed., p. 257. " Dann hat er auch die Aufstellung der heute angenom- men Principien der Mechanik zu einem Abschluss gebracht." — Mach, ibid. p. 181. 1 See for example that remarkable series of works, the Bridgewater Treatises "On the Power, Wisdom, and Goodness of God, as manifested in the Creation ; illustrating such work by all reasonable arguments, as for instance the variety and formation of God's creatures in the animal, vegetable, and mineral kingdoms ; the effect of digestion, and thereby of conversion ; the construction of the hand of man, and an infinite variety of other arguments ; as also by discoveries ancient and modern, in arts, sciences, and the whole extent of literature." — Whewell, "Astronomy and General Phys- 4 THE FITNESS OF THE ENVIRONMENT unable, however strong their desire, to ac- count for the facts with a plausible theory. The dogma of final causes had led a thou- sand times to the truth by teaching the investigator that the true description of an organ or physiological process was to be found in its utility to the organism as a whole. Such considerations were far too numerous and too patent for science to shirk some explanation, and the only weighty explana- tion at hand seemed the teleological one. n FITNESS With a suddenness which to many seemed catastrophic Darwin's hypothesis of natural selection changed the whole aspect of the problem. Law appeared as the basis of purpose just as it had appeared as the basis of order, and adaptations became, in the judg- ment of most men, the necessary results of an automatic process. To-day, after a half century, there is no longer room for doubt that the fitness of organic beings for their life in the world has been won in whole or in part ics Considered with Reference to Natural Theology." Lon- don, 1834, 4th ed., p. ix. To this series such men as Whewell and Sir Charles Bell contributed. FITNESS 5 by an almost infinite series of adaptations of life to its environment, whereby, through a corresponding series of transformations, pres- ent complexity has grown out of former sim- plicity.1 The great and fruitful ideas which Darwin brought to the attention of the whole world have long since been incorporated into hu- man thought. Not the least important among them is the new scientific concept of fitness, as it emerges from the discussion of natural selection. Before Darwin, this concept possessed all the vagueness of an idea which, though in part founded on observation, was not to be explained with the help of existing scientific theories. But although Darwin's fitness involves that which fits and that which is fitted, or more correctly a reciprocal rela- tionship, it has been the habit of biologists since Darwin to consider only the adaptations of the living organism to the environment.2 1 The ideas which are associated with the names of de Vries, as well as the very different hypotheses of Driesch, Bergson, and others are, of course, concerned with the manner, not with the fact of adaptation and organic evolution. 2 Far different was the earlier point of view. An examina- tion of Whewell's Bridgewatcr Treatise at once reveals im- portant, if often fallacious, discussions of environmental fit- ness; e.g. "It has been shown in the preceding chapters that a great number of quantities and laws appear to have been selected in the construction of the universe ; and that by the 6 THE FITNESS OF THE ENVIRONMENT For them, in fact, the environment, in its past, present, and future, has been an inde- pendent variable, and it has not entered into any of the modern speculations to consider if by chance the material universe also may be subjected to laws which are in the largest sense important in organic evolution. Yet fitness there must be, in environment as well as in the organism. How, for example, could man adapt his civilization to water power if no water power existed within his reach ? adjustment to each other of the magnitudes and laws thus selected, the constitution of the world is what we find it, and is fitted for the support of vegetables and animals in a manner in which it could not have been, if the properties and quan- tities of the elements had been different from what they are. We shall here recapitulate the principal of the laws and magnitudes to which this conclusion has been shown to apply. 1. The Length of the Year, which depends on the force of the attraction of the sun, and its distance from the earth. 2. The Length of the Day. 3. The Mass of the Earth, which depends on its magni- tude and density. 4. The Magnitude of the Ocean. 5. The Magnitude of the Atmosphere. 6. The Law and Rate of the Conducting Power of the Earth. 7. The Law and Rate of the Radiating Power of the Earth. 8. The Law and Rate of the Expansion of Water by Heat. 9. The Law and Rate of the Expansion of Water by Cold, below 40 degrees. FITNESS 7 At first sight it may well seem that inquiry into such a problem must end unsuccessfully in vague and unprofitable guesses. Indeed the past has brought forth no lack of such vain attempts, usually guided by a devotion to the doctrine of design in the service of the- ology. Yet other sciences have grown since 1859, and physical and chemical data in abundance are now at hand to aid in a recon- sideration of the environment's fitness, if 10. The Law and Quantity of the Expansion of Water by Freezing. 11. The Quantity of Latent Heat absorbed in Thawing. 12. The Quantity of Latent Heat absorbed in Evapora- tion. 13. The Law and Rate of Evaporation with regard to Heat. 14. The Law and Rate of the Expansion of Air by Heat. 15. The Quantity of Heat absorbed in the Expansion of Air by Heat. 16. The Law and Rate of the Passage of Aqueous Vapor through Air. 17. The Laws of Electricity; its relations to Air and Moisture. 18. The Fluidity, Density, and Elasticity of the Air, by means of which its vibrations produce Sound. 19. The Fluidity, Density, and Elasticity of the Ether, by means of which its vibrations produce Light." — Whe- well, "Astronomy and General Physics Considered with Reference to Natural Theology." London, 1834, 4th ed., pp. 141-143. It is hard to understand how such ideas could have fallen into oblivion. 8 THE FITNESS OF THE ENVIRONMENT such exist. Clearly it is well to seek among these data for a more precise formulation of the problem, which may then perchance lead to some more ambitious quest, or at least to new understanding of the old failure. Ill THE ENVIRONMENT The world of our senses is a world of matter and energy, space and time. After centuries of philosophical and scientific study, these, the very logical elements of science, are no doubt still without a final description. None the less is there sound foundation for the belief that our preliminary accounts of all four possess completeness in some respects and for certain purposes. Nor are we to-day less confident of the finality of some of our ideas regarding the nature of life and the vital processes, as they exist in this world. But both of these conclusions call for further consideration. A MATTER Many facts contribute to the belief, uni- versal among chemists, that the known ele- ments constitute by far the greatest part of FITNESS 9 a system of materials out of which the uni- verse is formed, within which all chemical changes (except certain phenomena of radium and a few other anomalies, including perhaps unknown changes in the interiors of the celestial bodies) take place. It is certain that nearly all the chemical transformations upon the earth consist of rearrangements of the atoms of the known elements. A century and a half of scientific chemistry guarantee that conclusion with a security rarely attained in descriptive science. And the testimony of the spectroscope is equally conclusive that the visible stars, like the sun itself, are made up almost or quite exclusively of the same chemical ele- ments. Such facts, so familiar that thev re- quire no comment or explanation, might sufficiently justify the acceptance of the chem- ist's known elements as the only important matter in the universe. But even more weighty evidence is at hand ; I mean the so-called periodic classification of the ele- ments. It has long been evident that simple rela- tionships exist in some cases between the atomic weights of similar elements. For ex- ample, the atomic weights of bromine, stron- tium, and selenium are approximately equal 10 THE FITNESS OF THE ENVIRONMENT to the means of the atomic weights of chlorine and iodine, of calcium and barium, and of sulphur and tellurium respectively. More general relationships between the atomic weights and properties of the elements were first pointed out by Newlands in 1864 and were extended by Mendeleeff and Lothar Meyer a little later. Out of these studies has arisen the law that the properties of the elements are periodic functions of their atomic weights. The essential characteristics of this law are best illustrated by a consideration of the relative volumes occupied by atoms of the various elementary substances, the so-called atomic volumes, which may be expressed by dividing atomic weights by specific gravities. The facts are graphically represented upon the accompanying diagram, where atomic weights are plotted horizontally, atomic vol- umes vertically. Beginning with lithium the volumes fall to boron and carbon, then rise irregularly to sodium. A second fall leads to aluminium, a second rise to potassium, and then the rises and falls of the curve are repeated until, among the elements of higher atomic weight, gaps break the continuity of the relationship. On the whole curve similar elements occupy FITNESS 11 -8 -8 .© -s -8 -3 -a -O 3 s 8 % 12 THE FITNESS OF THE ENVIRONMENT similar positions. Thus the alkali metals, lithium, sodium, potassium, rubidium, and caesium, occupy the crests of the waves; the halogens, fluorine, chlorine, bromine, and iodine, fall about midway between crests and troughs, and a little further study dis- closes a host of other corresponding relation- ships. Similar periodic variations may be shown to occur in other physical properties of the elements ; — the melting points, the boiling points, the magnetic characteristics, etc. Even more striking are the periodic varia- tions in chemical properties, including the general characteristics which first led to the idea of rational classification, and more spe- cific qualities like the combining powers for hydrogen, oxygen, and other elements. The clearest proof of the value of the periodic classification has been the predic- tion of "new' elements, and accurate fore- knowledge of their properties. Thus when Mendeleeff first described the system, the element germanium, discovered by Winkler in 1886, was unknown; but from the proper- ties of the elements surrounding a gap in the system the Russian chemist was able to predict its properties with almost incredible exactness, as the following table shows. FITNESS 13 Atomic weight . . . Specific gravity . . . Atomic volume . . . Formula of oxide . . Specific gravity of oxide Formula of chloride Boiling point of chloride Specific gravity of chloride Formula of fluoride . . Formula of ethyl compound Specific gravity of ethyl compound Prediction 72.0 5.5 13 Ge02 4.7 GeCU Less than 100° 1.9 GeFl4 Ge(C2H5)4 0.96 Observation 72.3 5.469 13.2 Ge02 4.703 GeCU 86° 1.9 GeFU Ge(C2Hfi)4 Lower than water Finally it is to be especially noted that, upon arranging the known elements in a table rationally constructed upon the basis of the above recorded facts, comparatively few spaces within the range of known atomic weights remain to be filled. The conclusion is obvious that very few elements now un- known are possible unless they possess very high atomic weights. But the apparent transmutation of radium into helium is a pretty clear indication that elements of very high atomic weight may be unstable. If they have existed in number and large quan- tity, they probably have long since ceased so to exist, except perhaps in the interior of celestial bodies, and they are not likely elsewhere to complicate natural phenomena by their unknown properties. 14 THE FITNESS OF THE ENVIRONMENT H H S H W H K H O a n < H 00 CO .* *- no 00 Ph © i—i Ph © i— i t— 1 O© J3°- © 1— I > Fe C 55.9 59 Ru R 101.7 103 t- CO t— 1 © 1-1 OS © b- t-> © 1— 1 O* M o fl9 PQ os l-H > I— t <->*0 co 53 *o i> rt-. • * • • ««! © o © CO © >— i .- © u n i> > CO W5 53© 1-1 • £2 • «5 Lj co" ^ co m «5 o* © > © © K^ «* >^J <*> 1-1 i-H 00 «5 °s s • H3 • (U "^ © © > © i-H CC co Q4 H co o^ d os coS 1— 1 Ce 140.2 • Th 232.5 c3 o fl2 i—i — i 1— 1 »— ( © I— 1 i—i O^ © ><© GO " i-( La 138.9 Yb 173.0 n © • a* •n*. 60 5 1— 1 1— 4 D i— 1 m© © *2 N^ K®. 0% 1—1 HH © w © «8 UO CO *> 00 CQoo • rjf O* 3^ CO o* »— t «3§ Na 23.05 «5 " OS CO Ag 107.9 o2 • 2*: <3 © • O o «© © <1 OS CO • " 00 • *2 • • • FITNESS 15 The behavior of radium and the classifica- tion itself suggest one further idea: the hy- pothesis that the elements are genetically related, that they have been evolved by some unknown process according to unknown laws. Certain it is that the properties of matter are no chance phenomena, and that transmuta- tion has ceased to be merely a philosopher's dream. All of these familiar facts of chemical science fully justify us in dealing with matter as a known factor in the study of life conditions in the universe. For, whatever mav be the fate of present theories, our present practical knowledge of the behavior of matter cannot fail us in the future. B ENERGY Contemporary with the work of Darwin and of Mendeleeff were the feats of Maver, Joule, Helmholtz, Kelvin, and Clausius, whereby ideas of energy assumed their mod- ern aspect. In the revolution wrought by these men imponderables and fluids vanished from this domain, and energy became that which, not being matter, is conserved. The new principles of the 'fifties have held their own until to-day. Meantime they have 16 THE FITNESS OF THE ENVIRONMENT made of thermodynamics (the department of science which is especially concerned with the laws of energy transformation) a sub- ject which few who cultivate the physical sciences may disregard. Countless develop- ments and achievements of thermodynamics give very real ground for the belief that we may speculate about the transformations of energy in the universe with the same assur- ance that we have in discussing chemical changes. Our reasons for confidence in the truth of current general notions of energy, and in their adequacy to account for any phenomena so far as energy is concerned, wherever life exists in the universe, are manifold, and not unlike those which have been reviewed in dis- cussing the elements. Centuries of search have revealed, in addi- tion to that most obvious form which is studied in dynamics, a very small number of varieties or manifestations of energy, such as heat, electricity, magnetism, optical energy, and chemical energy. Such manifestations of energy are by no means confined to the earth or to the solar system. Indeed Newton first worked out the general laws of dynamics and erected them into a complete science with the aid, not of terrestrial, but of astro- FITNESS 17 nomical phenomena.1 And recent most re- markable studies of the stars have enabled astronomers to account for obscure events in far distant parts of the universe by the application of the principles of dynamics. Similarly light, heat, and chemical energy, as we know them, are unquestionably universal. No doubt the manifestations of energy within the sun and stars, like the accompany- ing material phenomena there, can to-day only be surmised. For aught we know, these places may, as has been guessed, be the birthplace of elements and the seat of mani- festations of energy quite different from 1 "What the Occasion of Sir Isaac Newton's leaving the Cartesian Philosophy, and of discovering his amazing Theory of Gravity was, I have heard him long ago, soon after my first Acquaintance with him, which was 1694, thus relate, and of which Dr. Pemberton gives the like Account, and somewhat more fully, in the Preface to his Explication of his Philos- ophy : It was this. An Inclination came into Sir Isaacs Mind to try, whether the same Power did not keep the Moon in her Orbit, notwithstanding her projectile Velocity, which he knew always tended to go along a strait Line the Tangent of that Orbit, which makes Stones and all heavy Bodies with us fall downward, and which we call Gravity? Taking this Postulatum, which had been thought of before, that such Power might decrease, in a duplicate Proportion of the Dis- tances from the Earth's Center." — "Memoirs of the Life of Mr. William Whiston by Himself." London, 1749, Vol. I, pp. 35-38. Quoted by Ball. "An Essay on Newton's Prin- cipia." London, 1893, p. 8. G 18 THE FITNESS OF THE ENVIRONMENT what we have ever observed. But, however interesting and important such processes may be, it is not to be supposed that they are of direct moment in physiological processes. These conditions are far beyond the limits of our present investigation. Accordingly, everything that observation has taught con- firms the belief that energy, like matter, is in general well known to us. Its manifesta- tions are few, and they are universal. But just as the generalizations of science yield further assurance regarding matter, so they do not fail to confirm our conclusions in the study of energy. The law of the conserva- tion of energy and the law of the degradation of energy, otherwise known as the first and second laws of thermodynamics, clearly in- dicate that the manifestations of energy are not accidental nor independent of one an- other. They are orderly, and they obey laws. Energy is one and indestructible. Such are the apparently irrefragable con- clusions of the brief half century of creative development, from the time when Young first used the word "energy" and Bolton and Watt first employed the idea of measuring energy in horse power, through the period of Carnot's brilliant intuition regarding the re- lation between heat and work, to the epoch FITNESS 1 0 of the foundation of thermodynamics.1 To- day we know that just so much heat, neither more nor less, may be obtained by the com- plete conversion of a unit of electrical energy or by a given chemical process. We know, moreover, that not every conceivable change from one form of energy to another is possible. On the whole, energy can flow in but one direction; perpetual motion is impossible; and useful energy is steadily becoming de- graded, dissipated, and useless. Such laws are fully worthy of a place be- side the periodic law, and they justify equal confidence in the adequacy of our current descriptions of matter and of energy for the purposes of biology. C SPACE AND TIME Since Kant revolutionized modern phi- losophy, the whole world has steadily realized that between matter and energy on the one hand, and space and time on the other, there is a real and highly significant difference.2 1 An excellent account of this period may be found in Merz's "History of European Thought in the Nineteenth Century," Vol. II, Chap. VII, "On the Physical View of Nature." 2 For a brief statement of Kant's argument see Royce, "The Spirit of Modern Philosophy," pp. 121-125. 20 THE FITNESS OF THE ENVIRONMENT But, however important such distinctions may be for the philosopher, the man of science in his practical task is obliged to put them aside and to make the best of whatever evi- dence experience, observation, and experi- ment may supply. Out of such studies space and time have emerged, usefully defined by mathematical criticism as substantial parts of the edifice of science.1 There is no small difficulty in the exposi- tion of modern critical results regarding space and time, but fortunately there is little need of considering them on the present occasion. For in spite of all assaults of phi- losophers and mathematicians space remains for practical purposes more certainly than ever the Euclidian space of the ancients, only it has become somewhat richer in char- acteristics. And time is now and forever that which flows equably, wholly independ- ent of all else, though almost all else is dependent upon time. It is Euclidian space in which the earth moves and describes its ellipse, parallel rays of light never do meet in our practical experience, and our crystals 1 The works of Poincare, "La Science et l'Hypothese," "La Valeur de la Science," and "Science et Methode," published by Flammarion, may be consulted for a popular statement of such mathematical studies. FITNESS 21 are in form the figures of Euclidian geometry. Our time flows ever in proportion to the swings of a pendulum, the propagation of light, and the progress of a chemical change. Time and space are thus bound to matter and energy by experience, and for practical purposes we accept all four as science at present knows them.1 We cannot doubt that knowledge of them will increase and ideas of them change. But we can scarcely think that our present ideas are inadequate for our present purposes, or that, for life, matter will ever be other than the elements of the periodic classification, energy that set of quantities to which we apply the laws of thermodynamics, and time and space the concepts which were familiar to Galileo and Euclid. IV THE ORGANISM Thus the growth of physical science has provided the speculative biologist with a very accurate and extensive description of the physico-chemical structure of the ma- terial universe and with a well-founded con- 1 In the present work we need have no concern for the so-called principle of relativity. 22 THE FITNESS OF THE ENVIRONMENT fidence in his right to make use of the descrip- tion in investigating the relationship between life and the environment. The biologist studies living organisms as inhabitants of this world, and by holding fast to physics and chemistry he has created modern physiology, a science which unites many, indeed nearly all, of the departments of physics and chemistry in the task of de- scribing the processes of life. That task has proved an arduous one, even in comparison with the other enterprises of science, and it must be confessed that few of the departments of physiology wear an aspect of finality which has long been famil- iar in such sciences as mechanics and crystal- lography, for example. Yet, as time has passed, and the nature of the material basis of life and the conspicuous features of the mechanism which the organism presents for study have become more familiar, assurance has steadily grown of the possibility of decid- ing upon fundamental and essential char- acteristics of the life process. No doubt opinions have fluctuated, and in different periods of the history of science particular phenomena of living organisms have been examined, criticized, and then well-nigh for- gotten. But gradually ideas, ever more and more precise, have arisen and been accepted. FITNESS 23 Until very recent times, however, the main interest has centered upon morpho- logical problems and upon the processes of growth and development. The ancient con- troversies regarding types and homologous parts, the question of spontaneous genera- tion and the whole science of embryology, and inquiries into the nature of fermentation and the role of microorganisms are examples of the older tendencies. Such interests have, it need hardly be said, lost none of their im- portance, but they scarcely touch the physico- chemical problem of the nature of living things. Yet there is in these subjects one point of view, a favorite of Cuvier's, now, though still familiar, less often emphasized, which states a most important characteristic of life in terms of matter and energy, space and time.1 Living things preserve, or tend 1 "La vie est done un tourbillon plus ou moins rapide, plus ou moins complique, dont la direction est constante, et qui entrafne toujours des molecules de memes sortes, mais ou les molecules individuelles entrent et d'ou elles sortent continuellement, de maniere que la forme du corps vivant lui est plus essentielle que la matiere." ("Regne animal," p. 13, etc.) "II vient sans cesse des elements du dehors en dedans : il s'en echappe du dedans en dehors : toutes les par- ties sont dans un tourbillon continue!, qui est une condition essentielle du phenomene, et que nous ne pouvons suspendre longtemps sans l'arreter pour jamais. Les branches les plus simples de l'histoire naturclle participent deja a cette compli- ft C State College 24 THE FITNESS OF THE ENVIRONMENT to preserve, an ideal form, while through them flows a steady stream of energy and mat- ter which is ever changing, yet momentarily molded by life; organized, in short. This idea, to which we must later return, could not possess in the early nineteenth century the significance and value which now attach to it. It needed the explanation which the study of metabolism has at length provided. METABOLISM Metabolism is the term applied to the in- flow and outflow of matter and energy and their intermediarv transformations within the organism. Its serious investigation began with Lavoisier, the principal founder of mod- cation et a ce mouvement perpetuel, qui rendent si difficile l'applieation des sciences generates. " (" Rapport,'5 p. 150, etc.) " Dans les corps vivans chaque partie a sa composition propre et distincte ; aucune de leurs molecules ne reste en place ; toutes entrent et sortent successivement : la vie est un tour- billon continuel, dont la direction, toute compliquee qu'elle, est, demeure constante, ainsi que l'espece des molecules qui y sont entrainees, mais non les molecules individuelles elles- memes. . . . Ainsi la forme de ces corps leur est plus essentielle que leur matiere," etc. (Ibid. p. 200.) — " Eloges historiques," Vol. I, p. 200. Quoted by Merz, "A History of European Thought in the Nineteenth Century," Vol. I, p. 129. Edinburgh and London, 1898. FITNESS 25 ern chemistry, who by ingenious experiments discovered that the essential feature of the chemical process in the animal is combus- tion or oxidation, and that the amount of oxy- gen required by such combustion is not much less than that needed to burn substances which resemble the foods in the air. The problems which thus arose have been studied by a host of later investigators, notably by Liebig and Voit, and gradually a vast array of facts concerning the turnover of matter and en- ergy in the body have been accumulated. Among other achievements is the proof that the principle of the conservation of energy applies to the living organism. These have been chemical investigations, carried out by chemists, and for that reason, until quite recently, they have not received their due in general biology. Meantime, as knowledge of the balance sheet of the body, the total metabolism so- called, has been perfected, more and more interest has developed in the changes which attend the passage of matter and energy in their various stages through the organism. Such problems at once demand a physico- chemical description of protoplasm as a nec- essary basis for their solution. The same demand has also arisen in other quarters. 26 THE FITNESS OF THE ENVIRONMENT Thus the microscope, with all its brilliant contributions to knowledge of the form and more gross structural elements of the cell, hardly at all contributes to knowledge of its physico-chemical organization as a mechan- ism. Out of such needs a preliminary, if very imperfect, rational description of proto- plasm has arisen, and gradually the physical and chemical laws governing protoplasm, its form, composition, and stability, its con- stituent parts and their mode of action, and the physical and chemical changes within it are being discovered.1 The idea of dur- able form in matter and energy that change can now be applied to the cell with greater advantage, in that descriptions of the form and of the change are now at hand, though as yet all too imperfect. Another profoundly important contribu- tion of the science of metabolism to our knowledge of the characteristics of life is the discovery of the cycle of matter through plants and animals.2 The plant takes up carbonic acid and water and a few other simple substances from air and soil, and 1 This subject is extensively treated by Hober, "Physik- alische Chemie der Zelle und der Gewebe." Leipzig, 1911, 3d ed. 2 This was originally clearly stated by Lavoisier. FITNESS 27 transforms them into oxygen, which renews the air, and sugar, starch, and other sub- stances, which are the food of the animal.1 These products the animal burns, thereby forming once more carbonic acid and water, which return to the plant and so pass through the cycle again and again. The changes in energy which accompany this process are quite different from the chemical changes. Starch and sugar and oxygen, formed in the leaf of the plant, are compounded of carbonic acid, water, and sunshine. This sunshine, or solar energy, wh'en changed into the chemical energy of the carbohydrate, is pre- served and transmitted to the animal.2 In his body it is set free as muscular force and heat, and then dissipated. Accordingly, when carbonic acid and water are combined to form sugar and oxygen in the leaf, it is al- ways a new store of solar energy which they bear, and while matter goes round and round, energy is being constantly degraded and lost. The one process is cyclic, the other moves steadily in one direction from 1 Our knowledge of photosynthesis is largely based upon the classical work of N. T. de Saussure, "Recherches Chi- miques sur la Vegetation." Paris, 18(U. 2 Only after the establishment of the principle of the conservation of energy was it possible to gain a clear con- ception of the energetics of metabolism. 28 THE FITNESS OF THE ENVIRONMENT sunshine to the waste heat of the animal body.1 B ORGANIC CHEMISTRY Independent alike of general biology and of the science of metabolism there has grown up still another department of natural sci- ence, organic chemistry, which contributes very materially to the description and com- prehension of living things. During a large part of the nineteenth century the efforts of chemists were mainly directed to the cultiva- tion of this subject, which seeks to describe the molecular constitution of all the com- pounds of carbon, including nearly all the individual substances which make up animals and plants. Gradually, as organic chemis- try has progressed, very complete descrip- tions of the atomic groupings within the molecules of fats,2 carbohydrates,3 and pro- 1 Here, as in so many other cases, it is not the conservation of matter and energy, but the conservation of matter and the degradation of energy which are important. For an exten- sive development of this important difference see B. Brunhes, "La Degradation de l'Energie." Paris, 1909. 2 Chevreul, "Recherches Chimiques sur les Corps Gras." Paris, 1823. 3 E. Fischer, " Untersuchungen liber Kohlenhydrate und Fermente." Berlin, 1909. FITNESS 29 teins,1 the chief of such things, and most of the other biologically important sub- stances have been obtained, and we are at length in possession of exceedingly clear and reliable ideas as to the chemical constitu- tion of living matter. In fact, the nature and laws of the chemical composition of pro- toplasm are actually more certain than the nature and laws of its physical structure.2 In this manner, by slow degrees, the de- scription of living things has progressed, and gradually the characteristics of life have become less obscure and their aspects more simple. It cannot be denied that many traits like consciousness and inheritance are, at least for the present, beyond the scope of description in terms of matter and energy, and the fundamental riddle shares this de- tachment.3 But the physico-chemical basis 1 E. Fischer, " Untersuchungen iiber Aminosauren, Poly- peptide und Proteine." Berlin, 1906. 2 Substantial progress in the latter field is nearly all of very recent date, almost wholly since the sudden rise of physical chemistry. 3 "But now, having confessed that Life as a principle of activity is unknown and unknowable — that while its phe- nomena are accessible to thought the implied noumenon is inaccessible — that only the manifestations come within the range of our intelligence while that which is manifested lies beyond it ; we may resume the conclusions reached in the preceding chapters. Our surface knowledge continues to be 30 THE FITNESS OF THE ENVIRONMENT of life is firmly established in the world of our senses. On the whole the composition of living matter, its physical structure, the changes of matter and energy which consti- tute the metabolic process, together with the totality of such changes, which make up the fundamental economic process of that largest community which consists of all living beings, are all clearly defined. C THE CHARACTERISTICS OF LIFE Under the circumstances it is certainlv no t/ rash enterprise to seek a definition of some of the essential characteristics of life. Al- a knowledge of its kind, after recognizing the truth that it is only a surface knowledge. "For the conclusions we lately reached and the definition emerging from them, concern the order existing among the actions which living things exhibit ; and this order remains the same whether we know or do not know the nature of that from which the actions originate. We found a distinguish- ing trait of Life to be that its changes display a correspond- ence with coexistences and sequences in the environment; and this remains a distinguishing trait, though the thing which changes remains inscrutable. The statement that the continuous adjustment of internal relations to external relations constitute Life as cognizable by us, is not invali- dated by the admission that the reality in which these rela- tions inhere is incognizable." — Herbert Spencer, "The Principles of Biology." New York and London, 1909, Vol. I. Revised and enlarged edition, pp. 122-123. FITNESS 31 though it is probably far beyond our present power to make a complete study of the prob- lem, I feel sure that a brief analysis will justify certain very definite conclusions. Life as we know it is a physico-chemical mechan- ism, and it is probably inconceivable that it should be otherwise.1 As such, it possesses, and, we may well conclude, must ever pos- sess, a high degree of complexity, — physi- cally, chemically, and physiologically; that is to say, structurally and functionally. We cannot imagine life which is no more complex than a sphere, or salt, or the fall of rain, and, as we know it, it is in fact a very great deal more complex than such simple things. Next, living things, still more the community of living things, are durable. But complexity and durability of mechanism are only pos- sible if internal and external conditions are stable. Hence, automatic regulations of the environment and the possibility of regulation of conditions within the organism are essen- tial to life. It is not possible to specify a large number of conditions which must be regulated, but certain it is from our present experience that at least rough regulation of 1 I mean, of course, for the purposes of physical and chem- ical study. With such qualifications the statement is prob- ably no longer open to objection from any quarter. 32 THE FITNESS OF THE ENVIRONMENT temperature, pressure, and chemical consti- tution of environment and organism are really essential to life, and that there is great advantage in many other regulations and in finer regulations. Finally, a living being must be active, hence its metabolism must be fed with matter and energy, and accordingly there must always be exchange of matter and energy with the environment. Returning to the concept of the organism as a durable form through which flow matter and energy, it is now possible to make these ideas more vivid. The complex structure of the living being is relatively stable, alike in the chemical composition of its individual constituent molecules, in their proportions and amounts, in their aggregation into the invisible structural elements of protoplasm, in the visible parts of the cell, in the organs and tissues, and finally in toto, as a man or a tree. Similarly stable are the physical con- ditions within this structure: temperature, pressure, alkalinity, and osmotic pressure. Finally, that which surrounds it, the imme- diate environment, possesses also a high de- gree of stability, or if the organism be very complex, it may be that it has an efficient pro- tection against change of environment; a skin which insulates, for instance. But in this case FITNESS 33 it has also acquired an environment, a milieu interieur for its cells, — like the blood and lymph, — which serves the same purpose as stability of the external environment, and ex- ercises the further function of supplying food. It is through this structure, in the process of metabolism, that matter and energy flow. Entering in various forms and quantities, they are temporarily shaped exactly to the form and condition of the organism; they conform to the characteristics of the king- dom, class, order, family, genus, species, and variety to which it belongs, and they assume even the characteristics of the individual itself.1 Then they depart through the va- rious channels of excretion. When these ideas are reduced to their very simplest forms, it appears that life must be highly complex in structure and function; that the conditions of the environment must be regulated, and that there must be very exact regulation of conditions, both structural and functional within the organism, and finally, that, while life is active, there must be ex- change of both matter and energy with the environment. Complexity, regulation, and food are essential to life as we know it, and 1 Science is, of course, still at a loss for an adequate gen- eral explanation of such processes. 3-t THE FITNESS OF THE ENVIRONMENT in truth we cannot otherwise conceive of life, or indeed of any other durable mechanism. For my part I do not doubt that these pos- tulates are quite as true of the world of our senses as are the fundamental laws of matter and energy, space and time. Obviously these few conclusions can make no claim to completeness. Fully to describe life, the discovery of many other fundamen- tal characteristics is necessary, including such as are related to inheritance, variation, evo- lution, consciousness, and a host of other things. But in the formation and logical development of such ideas there is danger of fallacy at every step, and since the present list will suffice for the present purpose, further considerations of this sort are best dispensed with. This subject should not be put aside, however, without clear emphasis that the postulates which have been adopted above are extremely meager. The only motives for abandoning further search are the economy and the security which are thus insured, and the very great difficulty of extending the list. Any one who is familiar with similar efforts to elucidate the essential character- istics of life, such as that of Wallace,1 cannot, I fear, fail to perceive the extreme limitations 1 A. R. Wallace, "Man's Place in the Universe." New FITNESS 35 which are imposed upon inquiry by assuming complexity, regulation, and metabolism ex- clusively. Perhaps in reality these postu- lates are only two. Metabolism might with- out difficulty be included under regulation, but the consideration of such purely logical questions is beside the present purpose. However, these are probably the character- istics of the organism which are best fitted for discussion in relation to the physico-chemical phenomena of matter and energy, and it is barely possible that no others bear the same simple relations to the outside world. York, 1903, Chaps. X and XI, especially the following statement : — "The physical conditions on the surface of our earth which appear to be necessary for the development and main- tenance of living organisms may be dealt with under the fol- lowing headings : — " 1. Regularity of heat supply, resulting in a limited range of temperature. "2. A sufficient amount of solar light and heat. " 3. Water in great abundance, and universally distributed. "4. An atmosphere of sufficient density, and consisting of the gases which are essential for vegetable and animal life. These are Oxygen, Carbonic-acid gas, Aqueous vapor, Ni- trogen, and Ammonia. These must all be present in suitable proportions. "5. Alternations of day and night." It must be remembered, however, that such conclusions depend upon reasoning from analogy, a dangerous proceed- ing. 36 THE FITNESS OF THE ENVIRONMENT THE PROBLEM We may now return to the problem of the fitness of the environment. So long as ideas of the nature of living things remain vague and ill-defined, it is clearly impossible, as a rule, to distinguish between an adaptation of the organism to the environment and a ease of fitness of the environment for life, in the very most general sense. No doubt there are clear instances of both phenomena which require no close analysis for their interpreta- tion. Thus the hand is surely an instance of adaptation, and the anomalous expansion of water on cooling near its freezing point an instance of environmental fitness. But how much weight is to be assigned to adapta- tion and how much to fitness in discussing the relations between marine organisms and the ocean ? Evidently to answer such questions we must possess clear and precise ideas and definitions of living things. Life must by ar- bitrary process of logic be changed from the varying thing which it is into an independ- ent variable or an invariant, shorn of many of its most interesting qualities to be sure, but no longer inviting fallacy through our FITNESS 37 inability to perceive clearly the questions involved. Such is the purpose, and the justification, for setting up the postulates of complexity, regulation, and metabolism as inherent in that mechanism which is called the living organism. With them, at length, we face the problem which awaits us. To what ex- tent do the characteristics of matter and energy and the cosmic processes favor the existence of mechanisms which must be com- plex, highly regulated, and provided with suitable matter and energy as food ? If it shall appear that the fitness of the environment to fulfill these demands of life is great, we may then ask whether it is so great that we can- not reasonably assume it to be accidental, and finally we may inquire what manner of law is capable of explaining such fitness of the very nature of things. CHAPTER II THE ENVIRONMENT ASTRONOMY AN examination of the relationship be- tween life and the environment, in which by means of the simplifying postulates pre- viously developed life is arbitrarily taken as an invariant, should, if it is to be quite general, rest upon a physico-chemical description of the whole universe. We require to know the form and structure of stars and of interstellar space, of nebulae and of solar systems, and the conditions and changes which accompany such aggregations of matter. Evidently this requirement can be but imperfectly fulfilled, and yet one need not be too apologetic in venturing the attempt. In the end, to be sure, we shall found our argument upon the safe basis of terrestrial phenomena, but mean- time it will be an advantage to consider con- ditions far and wide. S8 THE ENVIRONMENT 39 The science of cosmography is probably the earliest of all the natural sciences, and cosmological speculation appears to accom- pany it from the outset. Long before the dawn of history the Chaldeans possessed much accurate information about the stars, and the zodiac was known to the Egyptians not less than fifteen centuries before our era. Always pursued with great interest, such studies received their first provisional systematic for- mulation at the hands of Hipparchus in the second century B.C. He, the greatest of the astronomers of antiquity, succeeded in bring- ing the apparent movements of the sun, moon, and planets into an arbitrary scheme which was nearly perfect for the sun, though less so for the other movable celestial bodies. He also measured and catalogued the positions of a large number of fixed stars. Upon this secure foundation of quantitative observa- tions modern astronomy has built. At the beginning of the modern period Copernicus, Tycho Brahe, Kepler, Galileo, and Newton reduced the phenomena of the solar system to law. At a later day speculations based upon their results and upon growing knowl- edge of physics and chemistry led Thomas Wright, Kant, and finally Laplace to a ra- tional, if somewhat imperfect, cosmological 40 THE FITNESS OF THE ENVIRONMENT theory of the solar system. Finally, ever more accurate observations and the marvel- ous fertility of spectroscopical investigations have brought the stars within our reach. The whole universe now appears to be not unlike our part of it, both chemically and physically. The same forms of matter, the same material aggregates, the same manifes- tations of energy, and similar movements are everywhere present. The stars are no longer changeless, but violently active bodies ; they are no longer permanent, but evolving sys- tems; they are born, they grow, age, and die; and throughout their evolution they obey laws, which, though as yet imperfectly known, appear to be common to all. Mean- time the study of nebulae, comets, and meteor- ites has kept pace with other departments of the science, and our interpretation of the re- sults of stellar astronomy1 constantly gains from ever increasing knowledge of the physical and chemical processes in the sun. The universe which thus gradually has been revealed to the astronomer is made up of a relatively small number of types of material 1 A description of such facts from the physico-chemical point of view may be found in Arrhenius's "Lehrbuch der kosmichen Physik." Leipzig, 1903. A brief popular account f some of the facts in the same author's "Worlds in the Mak- ing," translated by Borns. New York and London, 1908. THE ENVIRONMENT 41 aggregation. These include luminous dense bodies like the sun and stars ; non-luminous dense bodies like the earth, the moon, the planets, and invisible partners of certain stars ; nebulae, comets, and meteorites. The larger of these bodies are separated by vast extents of space which contain only rare meteorites, perhaps minute traces of gaseous material, and cosmic dust. There can be little doubt that other types of bodies do not commonly occur in that portion of the universe which is open to astronomical investigation. Both the enormous collections of astronomical data which are now at hand and the beginnings of clear knowledge of cosmic processes justify this belief. Of what may lie beyond the visible stars we can, of course, know nothing. The nature of the stars is revealed to us chiefly by study of their spectra, according to which they have been roughly classified, by Vogel * for example, quite simply into three principal types. I. White stars in which there is marked evidence of the presence of hydrogen, or, in some instances, helium. The stars of this class undoubtedly are extremely hot, the helium stars probably especially so. Their atmospheres seem to be very dense and to 1 See Arrhenius's "Lehrbuch," pp. 23-27. 42 THE FITNESS OF THE ENVIRONMENT consist of hydrogen or helium or a mixture of the two gases. There is evidence that some of these stars possess very high rotational velocities. II. Yellow stars, including the sun, whose spectra indicate the presence of hydrogen and numerous metals, — sodium, iron, calcium, magnesium, etc. The lines which show the presence of hydrogen in the stars of this type vary in intensity. The current belief is that those stars which appear to possess more hydrogen are the hottest. The stars of this type are less hot than those of type I. III. Reddish stars whose spectra show little or no sign of the presence of hydrogen, but indicate that of chemical compounds, in- cluding hydrocarbons. The presence in these spectra of the lines of sodium, iron, calcium, and magnesium is clearly established. Stars of this type are evidently the coolest of lumi- nous dense bodies. This classification is, of course, provisional and unsatisfactory, and probably sometimes results in bringing together relatively unlike stars and in separating such as are very much akin. Moreover, subdivisions in the classifica- tion are necessary and hard to make. Other better but more complex classifications appear to exist, but they suffer only in less degree THE ENVIRONMENT [.', from like defects. In short all the known facts can best be explained by the assump- tion that the stars represent different stag - of development of suns.1 In that case this 1 "Bei Durchmusterung der Specktra der verschiedenen Sterne kann man sich nicht des Gedankens erwehren, dasa die verschiedenen Sterngruppen verschiedenen Entwick- lungsstadien entsprechen. Die jiingsten mid wiirmsten aller Sterne waren (nach der allgemeinen Ansiclit, vgl. weiter unten Kap. Kosmogonie) diejenigen der ersten Gruppe. Das kontinuierliche Licht, welches von dem eigentlichen Sternkorper ausstrahlt, riihrt haupstachlich von Kondensa- tionen, wolkenartigen Bildnngen in der Atmosphiire der Sterne, zum geringeren Teil von den stark verdiehteten Metalldampfen im Inneren des Sterns her. In den hoheren Schichten dieser Atmosphiire finden sich die leichten Gase, Wasserstoff oder Helium oder alle beide, weiter unten Metal 1- dampfe. Bei den Sternen erster Klasse ist die Atmosphiire der leichten Gase so dick und heiss, dass die fur uns sicht- baren Kondensationen beinahe alle in diesen oberen Schichten vor sich gehen. Wir sehen deshalb keine oder nur schwache Metalllinien, dagegen sehr starke Wasserstoff- oder Helium- linien. Bisweilen ist die Menge und Temperatur der Leichten Gase geniigend, um helle Umkehrungen dieser Linien zu verursachen. Bei dem zweiten Spektraltypus ist die Ab- kiihlung weiter fortgeschritten, so dass Kondensationen nicht nur in den hochsten Schichten der Atmosphiire, sondern auch innerhalb der Metallatmosphiire vorkommen. Man sieht dann die dunklen Metalllinien scharf hervortreten. Das Zuriicktreten des violetten Endes des Spcktrums und einige schwache Bander im roten Teil deuten auf niedrigere Tem- peratur hin. Bei den rotliehen Sternen treten tiefe Temper- atur andeutende Erscheinungen noch mehr hervor. Die bei denselben gewohnlich vorkommende Verttnderlichtkeit liisst auf das Vorkommen von kalteren und wiirmercn Perioden 44 THE FITNESS OF THE ENVIRONMENT development or evolution must be a contin- uous process through which every sun slowly passes, and on the whole all suns must be much alike. Certainly we have the best of evidence to justify the assumption that most stars, including the sun, have very much the same chemical composition, and that differ- ences in spectra are due to the slowly progress- ing physico-chemical changes which have accompanied the process of cooling. Needless to say, the chemical composition of the sun itself is far better known than that of the stars. Particularly prominent among his constituent elements are those above mentioned: hydrogen, sodium, calcium, mag- schliessen, wie solche in geringerem Maasstab bei unserer Sonne durch die Fleckenperiode sich kundgeben. Zuletzt wird die Leuchtkraft der Sterne sehr schwach und das Licht ausgepragtrot, der relativ niedrigen Temperatur entsprech- end. Nach diesem Stadium kommt dasjenige, worin die dunklen ultraroten Strahlen allein herrschen, der Stern ist in einen nichtleuchtenenden Himmelskorper Ubergegangen (vgl. weiter unten Kap. Kosmogonie). Im Grossen und Ganzen zeigen die Sterne dieselbe chemi- sche zusammensetzung wie die Sonne. Die hervorragende Rolle des Wasserstoffs und Heliums, sowei des Eisens, Na- triums, Calciums und Magnesiums, macht sich iiberall be- merkbar. Es ist dann kein Zweifel, dass unsere Sonne mit den Fixsternen sehr nahe verwandt ist, und zwar ist sie als ein Fixstern der ersten Abteilung in der zweiten Klasse an- zusehen." — Arrhenius, "Kosmische Physik." Leipzig, 1903, p. 27. THE ENVIRONMENT 45 nesium, and iron. Also some others, which because of their low density are concentrated near the surface, are known to occur. The elements which have not yet been discovered are those like the metalloids which do not un- der ordinary circumstances give well-marked spectra, and those like gold, platinum, and mercury, whose higher specific gravities may be supposed to cause their accumulation in the interior. Carbon is certainly present, and almost certainly oxygen as well. Indeed it is only reasonable to conclude, as Kirchhoff originally suggested,1 that all the elements 1 "Diese Vorstellung von der Beschaffenheit der Sonne stimmt mit der von Laplace begrlindeten Hypothese liber die Bildung unseres Planetensystems uberein. Wenn die Masse, die jetzt in den einzelnen Korpern dieses Systems verdichtet ist, in fruheren Zeiten einen zusammenhangenden Nebel von ungeheurer Ausdehnung bildete, durch dcssen Zusammenziehung Sonne, Planeten und Monde entstanden sind, so mussten alle diese Korper bei ihrer Bildung im wesentlichen von ahnlicher chemischer Zusainmensetzung sein. Die Geologic hat gelehrt, dass die Erde einst in glii- hend fliissigem Zustande sich befundcn hat ; man muss anneh- men, dass auch die anderen Korper unseres Systems einmal in einem solchen gewesen sind. Die Abkiihlung, die infolge der Ausstrahlung der Warme bei alien eingetreten ist, hat aber bei ihnen sehr verschiedene grade erlangt ; und wiihrend der Mond kalter als die Erdegeworden ist, ist die Temperatur des Sonnenkorpers noch nicht unter die Weissgllihhitze gesunken. Die irdische Atmosphare, die jetzt nur wenige Elemente enthalt, musste, als die Erde noch gliihte, eine viel mannigfaltigere Zusammensetzung haben ; alle in der Gliih- 46 THE FITNESS OF THE ENVIRONMENT are present in the sun, and very often, at least, in the stars. Exceptions may arise, but probably they will hardly suffice to invalidate the rule. The large, dark, dense bodies which are directly known to us are the planets and their satellites. There are, however, many in- dications that the heavens are occupied by great numbers of "dead" suns, incrusted and therefore no longer luminous. Such appear to be the only conclusions which can be drawn from a study of the energetics of solar evolu- tion, for sooner or later a sun must cool from loss of energy until at length a crust forms, and, barring catastrophe, it must then endure for- ever. Moreover, as we have seen, the varying aspects of the stars seem to disclose suns in all stages of such a process. More nearly direct is the evidence furnished by study of variable stars of the Algol type. Algol itself Q8 Persei) is a star of second magnitude with a period of 2 days, 20 hours, 48 minutes, 53.8 seconds. During each period hitze fliichtigen Stoffe mussten in ihr vorkommen. Eine entsprechende Beschaffenheit muss heute noch die Atmosphare der Sonne besitzen." — G. Kirchhoff, — " Untersuchungen iiber das Sonnenspektrum und die Spektren der Chemischen Elemente." Abhandlungen der Koniglichen Akademie der Wissenschaften zu Berlin, 1861. Zweite, durch einen An- hang vermehrte Ausgabe. Berlin, 1862. THE ENVIRONMENT 47 for about 2j days it shines with constant in- tensity; thereupon it begins to decline and in approximately 4^ hours sinks to its minimum of brightness; then it becomes gradually brighter until after 4^ hours more it has re- attained its full brilliancy. This behavior is explained by the supposition that Algol is accompanied by a dark star and that their movements are such that a partial eclipse occurs every 69 hours. Pickering has suc- ceeded in calculating, upon the assumption that the dark star as a whole intercepts the rays of Algol, the approximate sizes, veloci- ties, and orbits of these two stars, one of which is quite invisible. Many similar phe- nomena lead to similar conclusions regarding other variable stars. It is apparent that such dark bodies, whether extinct suns or planets, represent another stage in celestial evolution. Their past his- tories may be various, for there is still room for much doubt as to the manner of formation and origin of planets, but at any rate all are probably derived from luminous stars or planets through the process of cooling, with its accompanying crust formation. Like their earlier forms, they must therefore be made up of matter as we know it, since when a heavenly body puts on a crust, it does not change the 48 THE FITNESS OF THE ENVIRONMENT matter of which it is composed. Finally terrestrial chemistry completes the evidence regarding the composition of such astronom- ical bodies; geophysics that regarding their state. The number of extinct suns is prob- ably very great ; Arrhenius thinks it not un- likely that they may be one hundred times more numerous than the luminous stars.1 It is more difficult to gain a clear idea of the nebulae, for such aggregations of matter are very diverse in appearance, and none lie near enough to the earth for us to study them as we study the solar system. However, in- vestigation of new stars, of the spiral forms of many nebulae, of the so-called star rifts which appear to be due to the movement of a large body through a nebula, sweeping up smaller bodies and leaving a channel behind, and a variety of considerations dependent upon the modern development of the sciences of physics and chemistry, all contribute to a growing belief that nebulae may often, and sometimes at least do certainly arise from col- lisions between dense bodies. Further, the nature of the processes by which stars may be formed out of nebulae becomes constantly better understood, and while there is small ground to regard our present science of nebulae 1 " Worlds in the Making," p. 151. THE ENVIRONMENT 49 as final, there is none at all for the belief that anything essentially inexplicable either physico-chemically or genetically will be dis- covered in their organization. Putting aside all contentious matters, it is abundantly clear that nebulae, however they may vary among themselves, are made up of vast extents of gaseous material and dust which are exceedingly rare and at very low temperature, and that they may contain all kinds of foci of condensation, from stars to meteorites, in great variety of forms and conditions. On the whole the common-sense judgment that the solar system may be taken as a fair sample of the universe, and that its probable evolution is in the main typical of cosmic evolution in general seems to be well founded. Any other hypothesis does violence to a host of facts, and to the larger generalizations of modern science. n POSSIBLE ENVIRONMENTS If now we seek to make the best of existing astronomical knowledge, as hastily sketched, in the study of our biological problem, cer- tain considerations at once present themselves* 50 THE FITNESS OF THE ENVIRONMENT Obviously it is not everywhere in such a uni- verse that life can exist. The visible stars, like the sun, certainly cannot support life. Throughout such bodies durable and complex arrangements of matter are impossible, for if formed, they must be at once dissipated by catastrophes far greater than any which can occur upon the earth's crust. The enormous intensity of heat, even in the most superficial parts of suns, must effectually preclude any state of matter but the gaseous, and thus prevent the existence there of anything of the nature of a mechanism. Such bodies, apart only from continuous variation from center to periphery, in the proportions of the elements, in density, and in the nature of the chemical unions between the elements, must be essentially homogeneous. They can scarcely possess relatively as much structure as the earth's atmosphere. In truth the sun itself seems to be the one and only durable solar mechanism. Not less evident is the impossibility of active life in interstellar space or in nebulae. Dor- mant life (panspermia) may indeed be possible universally, except only in the neighborhood of suns. But if life is to be fed, if there is to be active metabolism, including exchange of matter with the environment, something more THE ENVIRONMENT 51 nourishing than the rare molecules of a nebula, or the still rarer particles of interstellar space, must be provided. We may safely conclude, therefore, on the basis of our reliable knowledge of the universe, that active life can exist probably only upon a dense, crusted body1; for, of course, the in- terior of the earth is no better suited to life than is the interior of the sun. We have perhaps taken a long road to arrive at so familiar an idea. But our task involves the consideration of every conceivable form of life, not merely that relatively anthropo- morphic kind which we commonly think of when speculating loosely regarding life in other worlds. It is indeed possible that the common-sense judgment of the universe which declares our solar system to be on the whole, in its funda- mental characteristics, typical of every such system may turn out to be in some respects unjustified. For the present, however, so long as we use it only as an indication of the direction in which we are to turn our atten- tion, there is certainly no risk whatever in fol- lowing this hypothesis in the later discussion. 1 For an interesting discussion of the necessary conditions of existence see P. Lowell, "Mars as the Abode of Life." The Macmillan Company, New York, 1908. 52 THE FITNESS OF THE ENVIRONMENT III GEOPHYSICS Let us, accordingly, now examine such phenomena as are likely to occur upon the surfaces of bodies which in the course of cosmic evolution have acquired a solid crust. In faithfully carrying out such a plan, the sciences of geology and meteorology must be brought under contribution, and climatic conditions must receive especial attention. Not, to be sure, that our globe in every respect can fairly be taken as meteorologically typical of all incrusted bodies. On the contrary, there are a large number of phenomena which are unquestionably of highest significance in fa- voring the existence of life on this particular planet which appear to be accidental and prob- ably somewhat uncommon.1 Such are the 1 These also have been favorite subjects in the works on natural theology. The Bridgewater Treatises of Whewell and of Prout are replete with illustrations, those of Whewell often moderately expounded, while Prout's are, as a rule, most curious and antiquated. "Lastly, who will venture to assert that the distribution of sea and of land, as they now exist, though apparently so disproportionate, is not actually necessary as the world is at present constituted ? What would be the result, for instance, if the Pacific or the Atlantic oceans were to be con- verted into continents ? Would not the climates of the exist- THE ENVIRONMENT 53 size of the sun taken in relation with its dis- tance from the earth ; the size of the earth, which enables it to retain its present atmos- phere; the eccentricity of its orbit and the inclination of the ecliptic; the relative amounts of land and sea, and a host of other factors. Together these probably make of the earth, in comparison with other bodies, an extremely favorable abode for the living organism. Yet it cannot be denied that in detailed chemical constitution the earth is certainly more or less typical of all similar bodies. Moreover the earth's crust and its atmosphere, being formed in accordance with ing continents, as formerly observed, be completely changed by such an addition to the land, and the whole of their fertile regions be reduced to arid deserts ? Now, this distribution of sea and of land, so wonderfully adapted as it appears to be to the present state of things, depends of course in a great measure upon the absolute quantity of water in the world. While, on the other hand, the relative gravity of water, as com- pared with that of the earth, keeps the ocean within its destined limits, notwithstanding its incessant motion. Thus Laplace has shown that the world would have been con- stantly liable to have been deluged from the slightest causes. had the mean density of the ocean exceeded that of the earth ! Hence the adjustment of the quantity of water and of its density, as compared with that of the earth, afford some of the most marked and beautiful instances of design." — Prout, The Bridgewater Treatises, Treatise VIII, "Chemistry, Meteorology, and the Function of Digestion." London, 1834, pp. 186-187. 54 THE FITNESS OF THE ENVIRONMENT general laws, are likewise typical or at all events were so at the time of their origin. Neither can the change in crust and atmos- phere which time has wrought be wholly unique, though here a possible exception may again arise in the action of life itself. Since we do not at present positively know of the existence of life elsewhere and certainly have no detailed knowledge of its nature, we can- not feel sure that the conversion of atmos- pheric carbonic acid into oxygen and coal is either a universal or a common occurrence. In details of the geological process indeed there may well be marked differences. Probably the greatest variation will occur in the relative duration of conditions like those which we now enjoy on the earth, the length or brevity of the period from the full establishment of the circulation of water by evaporation, cloud formation, rainfall, with the flow of lakes and streams, until its extinction by cold. Thus there is more liability of error in an analyis of the general characteristics of those spon- taneous changes which must occur upon the surface of a body after the formation of a crust than there is in the attempt to dis- cover the general characteristics of stellar evolution. But here again our knowledge is not based upon terrestrial phenomena alone. THE ENVIRONMENT 55 With greater or less completeness and accu- racy the atmospheres of the moon, of Mars, and of other planets have been studied and accounted for. IV THE ATMOSPHERE Even at the earliest period in the evolu- tion of a typical star there appears to he a progressive variation in chemical composi- tion from center to periphery. Theoreti- cally it seems inevitable that the heaviest elements should be concentrated in the interior and that those of lowest atomic weight should be present in greatest amount near the sur- face. Actually, as above stated, spectro- scopic investigation fully confirms this view. Thus the spectra of typical hot stars show that hydrogen is an invariable constituent of their superficial parts. Indeed the uni- versal presence of hydrogen under such cir- cumstances is undoubtedly one of the most clearly established facts of stellar astronomy. As stars cool and become red the spectral changes quite as unmistakably point to the presence of carbon. Accordingly we possess the best of evidence and the best of reasons for the belief that large quantities of hy- 56 THE FITNESS OF THE ENVIRONMENT drogen and carbon must exist at or near the surface when a crust forms upon a cool- ing star. The nature of the chemical combinations into which these elements at first enter is perhaps open to some question. But as the temperature falls in the cooling of a sun or planet the affinities of carbon and hydrogen for oxygen increase, so that carbonic acid and water must normally result. For oxygen is almost certainly present in the sun; it is found in meteorites, and the vast store of it in the earth's atmosphere and crust (roughly one half of their total mass) justifies the be- lief that it is everywhere one of the commonest of elements. Hence an atmosphere contain- ing water and carbonic acid appears to be a normal envelope of a new crust upon a cooling body. Even were not these substances at first present in such an atmosphere, volcanoes must soon belch them, forth in enormous quantities to relieve the pressure which inevi- table chemical processes set up. It is clear that no one can give an exhaus- tive description of the formation of the earth's atmosphere and the changes which underlie vulcanism, so long as the theoretical considera- tions involved remain often more obscure than the facts. However, be the process what it THE ENVIRONMENT 57 may, it is at least automatic, and must repeal itself in other similar circumstances. There is, moreover, direct evidence in sup- port of the above conclusions. Spectro- scopic investigation has proved the presence of water vapor in the atmospheres of Mars, Venus, Jupiter, and Saturn, and nobody has suggested what the snowcaps of Mars may be unless they are real snow (hoarfrost) or, im- probably, carbonic acid. Lowell and Arrhe- nius agree in considering them snowcaps.1 In the earth's atmosphere carbonic acid has been very largely converted into oxygen and vegetable matter, which later has been turned into enormous quantities of coal. It is, in fact, possible, in accordance with the sugges- tion of Koene, that all the oxygen of the atmosphere has been thus formed from car- bon dioxide, and that therefore coal, peal, and other similar substances within the earth are chemically equivalent to the oxygen now free. If a typical atmosphere must contain water and carbon dioxide, its evolution must ob- viously be in part conditioned by the presence of these substances. Hence terrestrial mete< >r- ology, no less than terrestrial geology, must be in greater or less degree a special case of a 1 Arrhenius, " Kosmische Physik," p. 173; Lowell, " Mars as the Abode of Life," p. 81. 58 THE FITNESS OF THE ENVIRONMENT general process, and meteorological condi- tions on the earth cannot be perfectly unique. A number of circumstances, however, cause far greater variations in meteorological pro- cesses than in most other phenomena which have yet been discussed. For instance, on small astronomical bodies with weak gravi- tational attractions atmospheres cannot long endure. Like the moon these bodies must gradually lose nearly all their gases to space. Such loss has almost certainly occurred from the earth itself, and probably accounts for the absence of hydrogen and helium from the air. These gases, being very light, are not attracted with sufficient force to the earth, and gradually rise to the upper level of the atmosphere and fly away. Again, in the ab- sence of a near-by sun which steadily provides energy to balance loss by radiation, the period during which water and carbonic acid can remain in an atmosphere must be relatively short. Graduallv, but in a time whollv in- considerable in comparison with the duration of the terrestrial atmosphere, the gases sur- rounding bodies so placed must condense and then solidify. Finally, a body which con- stantly turns one face to a sun must slowly condense its whole atmosphere upon its dark, THE ENVIRONMENT 59 cold surface, and so no less certainly be de- prived of a gaseous envelope and of oceans. Accordingly it appears safe to say that really durable atmospheric conditions depend upon sufficient size of the planet, the presence of a sun, and rotation.1 No doubt a host of other factors which exist in the case of the earth are only less important. In any event, all such phenomena, though varied by chance, are of automatic origin, and whatever may be the peculiarities of our solar system there is no reason to suppose that like conditions are not of frequent occurrence. Throughout space there must be thousands of planets which, like the earth and Mars, are enveloped in an atmosphere that endures through count- less centuries, and that contains great quan- tities of water and carbon dioxide. All such atmospheres must in greater or less degree manifest general meteorological phenomena. There must be winds and clouds, rain and snow and ice, the formation of oceans and ocean currents, streams and lakes, all in- terrelated by complex cyclic processes which endure. Tides, too, and magnetic and elec- trical phenomena cannot be absent, while 1 A full discussion of all such problems will be found in the "Lehrbuch" of Arrhenius and in S. Giinther's "Handbuch der Geophysik." Two volumes, Stuttgart, 1897-1898. 60 THE FITNESS OF THE ENVIRONMENT the action of water through long ages must accomplish its gigantic work of disintegration and sedimentation. Soil must be formed, and water must penetrate it. In short a possible abode of life not unlike the earth apparently must be a frequent occurrence in space. GENERAL COSMOGRAPHICAL CONCLUSIONS Such, in the present state of knowledge, are the general cosmographical views which naturally suggest themselves to one who considers the possibility of life throughout the universe. The solar system appears to be, in its most general traits, a fair sample of the whole ; the sun is a typical star ; the planets are certainly members of a large class of similar bodies. These various types of material aggregation are a good deal alike wherever they occur. They are formed of the same matter, probably in very much the same proportions. They are actuated by the same manifestations of the same energy, and their evolutionary histories are similar. One and all are likely to possess, for a longer or shorter time, climates which make life possible. On the other hand, it is already obvious that THE ENVIRONMENT 61 the solar system, and especially the eartli among planets, are very favorable for life, partly through apparently accidental circum- stances. Putting aside, therefore, the biological fitness of the special climates of the earth both as a familiar fact, and as possibly in no small de- gree accidental, we may more advantageously give our attention at once to other phenomena which appear to be of a far more general character, — the occurrence of large quanti- ties of water and carbon dioxide in the atmos- phere, and the fundamental meteorological processes which their presence involves. Ni- trogen and various other substances auto- matically find a place beside water and car- bonic acid, but it will be convenient to pass them by or to postpone the consideration until other aspects of the subject have been more fully developed. VI THE PRIMARY CONSTITUENTS OF THE ENVIRONMENT Of course a consideration of the prop- erties of water and carbonic acid might be approached from a study of terrestrial processes exclusively. But since the assump- tion that such phenomena are common occur- 62 THE FITNESS OF THE ENVIRONMENT rences throughout the universe, and that they are the normal result of cosmic evolution, does not in any way modify the subsequent course of the inquiry, there appears to be no loss of logical security from its introduction. Mean- while the evidence that the special phenomena under discussion are probably the nowise ex- ceptional outcome of the operation of general laws, and not merely sporadic, cannot fail to lend weight to whatever conclusions may ultimately be reached. Obviously it is in the physical and chemical attributes of these two compounds and their constituent elements that we find very many of the conditions which make life possible upon the earth. They are material, provided and mobilized automatically, out of which living things undoubtedly can be formed. Moreover if we limit our study to the physico- chemical properties of water and carbonic acid, and to the compounds of carbon, hydro- gen, and oxygen, we shall greatly simplify our problem. It cannot be denied that this re- striction, no less than the earlier decision to restrict the postulated characteristics of life to complexity, regulation, and metabolism, is sure to limit the inquiry, often perhaps in a very unwelcome manner. On the other hand, the gain in economy and security is once THE ENVIRONMENT 63 more of great importance, and at present is perhaps essential to clear thinking. Such a procedure manifestly influences not at all the validity of any conclusions which may be reached ; only it must not be forgotten that the conclusions apply directly to our limited field of inquiry alone. VII THE ULTIMATE PROBLEM Such is the outcome of a preliminary glance at the many departments of science which are necessarily involved in the ques- tion of fitness of the environment. Living things permit themselves to be simplified into mechanisms which are complex, regulated, and provided with a metabolism ; the environ- ment, by a series of eliminations, is reduced to water and carbonic acid. These are sim- plifications counseled solely by expediency. Neither logical process is necessary ; each in- volves a disregard for many circumstances which might be of weight in the present in- quiry. But in the end there stands out a perfectly simple problem which is undoubt- edly soluble. That problem may be stated as follows : In wrhat degree are the physical, chemical, and general meteorological char- 64 THE FITNESS OF THE ENVIRONMENT acteristics of water and carbon dioxide and of the compounds of carbon, hydrogen, and oxygen favorable to a mechanism which must be physically, chemically, and physiologi- cally complex, which must be itself well regu- lated in a well-regulated environment, and which must carry on an active exchange of matter and energy with that environment? The first step in seeking a solution must be to review the data of physics and chemistry which describe the properties of water and carbonic acid, having due regard to their meteorological significance. Such data of the highest accuracy exist in great profusion, for almost every conceivable property of these substances has been studied with patient care. Next, the properties of the compounds of carbon, hydrogen, and oxygen must be con- sidered, and some of the characteristics of the chemical reactions into which they enter must be discussed. For this examination the unparalleled development of the science of organic chemistry provides ample material. All of these things must be scrutinized quan- titatively as well as qualitatively, and here again there is no lack of necessary information. Immediately one advantage of the method here proposed becomes evident. We can deal with the familiar abstractions of physical THE ENVIRONMENT 65 science, — specific heat, coefficient of expan- sion, solubility, heat of reaction, etc., — and thereby we shall gain all the advantages of the most exact sciences. No qualifications, no doubtful or contentious matter, no imper- fect descriptions need enter. In this manner it will be easy to estimate the absolute biological fitness in certain re- spects of water and carbonic acid, and at once a host of automatic results of their proper- ties will become evident. Many of these re- sults, such as the nearly constant temperature of the ocean, the ample rainfall, the freezing of water upon the surface, the great variety of carbon compounds, are familiar subjects of speculation, though since Darwin little in- terest has been manifested in them ; others, only recently brought to light by the growth of physical science, are nearly or quite un- known in this connection. All deserve to receive more serious attention from biologists than is at present vouchsafed them, for they constitute a part of the very foundation of general biology, and they cause many of the phenomena with which man is concerned in his struggle for mastery of the environment. Yet the mere exposition of such facts and relationships cannot suffice in a discussion of the fitness of the environment. In the first p 66 THE FITNESS OF THE ENVIRONMENT place these are in the main familiar ideas, and if they were altogether conclusive to prove the existence of really significant fitness, if they could be regarded as alone adequate to establish the necessity of putting fitness by the side of adaptation as a coordinate factor in causing the marvels of life, it is hard to be- lieve that they would have been so long neg- lected. In the second place there is nothing comparative about such information. Water is indeed a wonderful substance which fills its place in nature most satisfactorily, but would not another substance do as well ? Is not ammonia, for example, a possible substi- tute ? And are there not many other chemical bodies which might, in a very different world, serve equally useful purposes ? Perhaps, too, the great variety of carbon compounds which are known to the chemist are known only be- cause the vital processes furnish an abundance of material with which to experiment. Is it not possible, therefore, that another element, silicon, for instance, may enter into even greater varieties of compounds ? It is such questions, ever present in the minds of men of science, yet never carefully scrutinized to see if an answer be possible, which, I suspect, have long deflected attention from this subject. Clearly, therefore, it will be necessary to THE ENVIRONMENT 67 compare the proper! ios of water and carbonic acid and of the carbon compounds with those of other substances. It will be necessary to find out whether these substances are not only fit but fittest, — and this no doubt is a task of a very different sort. It may even seem at first sight an impossible one, but I hope to show that this is not the case, and that in spite of the incompleteness of our physical and chemical knowledge, it may be pressed to a satisfactory issue. A few remarks may now indicate the general line of thought we shall pursue, and then the actual study must pro- vide the proof. VIII THE METHOD OF SOLUTION The very constant temperature of the ocean is a most important factor in the economy of nature. It constitutes, for example, a vital regulation of the environment of a large pro- portion of all the living organisms of the world, and it has many other important "functions.' This constancy of temperature is in large part due to the magnitude of the specific heat of water. Other things being equal, the greater the specific heat of water, the more constant must be the temperature of the ocean. If, 68 THE FITNESS OF THE ENVIRONMENT then, the specific heat of water, as is actually the case, be nearly or quite a maximum among all specific heats, it follows that the fitness of water in this respect is nearly maximal. Again the ocean contains an astonishing variety of substances in solution, and they are present often in large quantities. In this manner a very great supply of food in very great variety is offered marine organisms. Of course such richness of the environment is an exceedinglv favorable circumstance for the organism, and it is due principally to the ability of water to dissolve a multitude of things in large quantities. It is not to be supposed that the substances present in sea water are all of use to every organism. This need not be the case at all ; but a variety of supplies which may be adapted to special re- quirements as they arise, here iodine, there cop- per, for instance, is a very genuine advantage. Further, the vast utility of the solvent action of water in blood, lymph, and all the body fluids is too patent to call for comment. If, now, it can be shown that the efficiency of water is nearly or quite a maximum, as it really is, among all known solvents, then it must be evident that in another respect the fitness of water is nearly or quite maximal. Again the amount of energy that is re- THE ENVIRONMENT 69 quired to tear apart molecules of water, and to liberate hydrogen and oxygen, is very great indeed, and when hydrogen and oxygen re- combine to form water, this energy must reappear, — under ordinary circumstances as heat. This fact, too, is very favorable for the organism, because almost all compounds which contain hydrogen yield a great deal of energy when they are burned ; they are, in short, great reservoirs of energy which can be tapped in the process of metabolism. If, therefore, the heat of combustion of hydrogen be nearly or quite a maximum, as it is, among all sub- stances, it is clear that water is again, in an- other respect, most wonderfully fitted for life. Finally, if it be true, and such is the case, that very few of the substances which share the fitness of water in one of these character- istics also share or approach its fitness in either of the others, and that none possesses all these qualifications in a degree that merits consideration, it must, I conceive, be admitted that so far as the investigation has proceeded water is the only possible fit substance. A criticism may here be made ; are there not other substances which possess other groups of qualifications which water lacks ? And that is a difficulty which is even harder to meet. But in the first place it is evident 70 THE FITNESS OF THE ENVIRONMENT that there are not an infinity of important physical properties ; in fact there are very few. And in the second place it is evident, both from centuries of experience in physical science and from the postulates above adopted regarding life, which undoubtedly do in the main describe its physico-chemical character- istics, that very few properties indeed are of importance in the least comparable with those which I have mentioned. Finally, it is in the highest degree probable that we are acquainted with most of the truly essential physical properties, and know them as biologically important, when they are so ; and I think we shall find it possible to consider them all, and thus to make the argument complete. Meanwhile it should be noted that there are two different ways of illustrating the fitness of a physical property. Properly employed, both are free from fallacy, and it will be de- sirable for us to employ both. Thus it may be shown, as in the case of the temperature of the ocean, that a particular property of water, its high specific heat, automatically produces a maximum of something which is favorable to life. Or again, as in the case of the regula- tion of the temperature of the human body by the process of perspiration, it may be shown that a particular property of water, its high THE ENVIRONMENT 71 heat of vaporization, has been utilized through adaptation of the organism to secure very high efficiency in a physiological process. Such is the method which must be followed in order to decide the question of the fitness of the environment. The physico-chemical char- acteristics of water, carbonic acid, and the carbon compounds are to be taken up one by one, and their absolute and relative magnitudes considered. The possible utility of such proper- ties, both automatically and through process of organic adaptation, must then be estimated, bearing in mind the fundamental characteris- tics of the living organism which have been arbitrarily postulated. Finally the various favorable qualities of water, carbonic acid, and the carbon compounds must be grouped together in order to see if they constitute a unique ensemble of fitness, among all possible chemical substances, for a living organism which must be complex, regulated, and en- gaged in active metabolism. At length the problem of fitness appears in a simple form. The road to a solution is open, and we may now proceed to an untram- meled discussion of unexceptionable data and well-known laws of physics, chemistry, meteor- ology, and physiology. Without further hypo- thetical difficulties, these must lead to the goal. CHAPTER III WATER GENERAL CONSIDERATIONS IT was assuredly not chance that led Thales to found philosophy and science with the assertion that water is the origin of all things. Whether his belief was most influenced by the wetness of animal tissues and fluids, or by early poetic cosmogonies, or by the ever pres- ent importance of the sea to the Ionians,1 however vague his conception of water may, indeed must, have been, he at least expressed a conclusion which proceeded from experience and serious reflection. Later, when positive knowledge had already grown to be a sub- stantial basis for speculation, both meteor- ological and chemical views contributed to the decision of Empedocles and Aristotle to include water among the elements.2 And it is especially worthy of note that of earth, air, 1 Windelband, "Ilandbuck der Altertumswissenschaft," V. 1. 139. Nordlingen, 1888. 2 Windelband, I.e.; S. Giintlier, " Geschichte der Natur- wissenschaften." Reclam, Leipzig, Vol. I, p. 19. 72 WATER 7 ; fire, and water the last is the only one which happens to be an individual chemical com- pound. From that day to this the unique position of water has never been shaken. It remains the most familiar and the most im- portant of all things. Within a comparatively recent time, to be sure, it has definitely lost its claim to be a true element, in the modern sense, but meanwhile almost every great development of science has but contributed to make its importance more clear. In physics, in chemistry, in geology, in meteorology, and in biology nothing else threatens its preeminence. The physicist has perforce chosen it to define his standards of density, of heat capacity, etc., and as a means to obtain fixed points in thermometry. The chemist has often been almost exclusively concerned with reactions which take place in aqueous solution, and the unique chemical properties of water are of fundamental sig- nificance in most of the departments of his science. In geology neptunism has at length won a certain though incomplete truimph over plutonism, and the action of water now appears to be far the most momentous factor in geolog- ical evolution.1 The meteorologist perceives 1 "Of all geological agencies water is the most obvious and apparently the greatest, though its efficiency is conditioned 74 THE FITNESS OF THE ENVIRONMENT that the incomparable mobility of water, which depends upon its peculiar physical prop- erties and upon its existence in vast quanti- ties in all three states of solid, liquid, and gas, is the chief factor among the properties of matter to determine the nature of the phenom- upon the presence of the atmosphere, upon the relief of the land, and upon the radiant energy of the sun. Through the agency of rainfall, of surface streams, of underground waters, and of wave action, the hydrosphere is constantly modifying the surface of the lithosphere, while at the same time it is bearing into the various basins the wash of the land and depositing it in stratified beds. It thereby becomes the great agency for the degradation of the land and the building up of the basin bottoms. It works upon the land partly by dissolving soluble portions of the rock substance, and partly by mechanical action. The solution of the soluble part usu- ally loosens the insoluble, and renders it an easy prey of the surface waters. These transport the loosened material to the valleys and at length to the great basins, meanwhile roll- ing and grinding it and thus reducing it to rounder forms and a finer state, until at length it reaches the still waters or the low gradients of the basins and comes to rest. The hydrosphere is, therefore, both destructive and constructive in its action. As the beds of sediment which it lays down follow one another in orderly succession, each later one lying above each earlier one, they form a time record. And as relics of the life of each age become more or less imbedded in these sediments, they furnish the means of following the history of life from age to age. The historical record of geology is, therefore, very largely dependent upon the fact that the waters have thus buried in systematic order the successive life of the ages." — Chamberlin and Salisbury, "Geology." New York, 1904, Vol. I, p. 8. WATER 75 ena which he studies;1 and the physiologist has found that water is invariably the prin- 14'0f all the terrestrial agents by which the surface of the earth is geologically modified, by far the most important is water. We have already seen, when following livpogene changes, how large a share is taken by water in the phenom- ena of volcanoes and in other subterranean processes. Re- turning to the surface of the earth and watching the opera- tions of the atmosphere, we soon learn how important a part of these is sustained by the aqueous vapor that pervades the atmosphere. "The substance which we term water exists on the earth in three well-known forms: (1) gaseous, as invisible vapor; (2) liquid, as water; and (3) solid, as ice. The gaseous form has already been noticed as one of the characteristic ingredients of the atmosphere. Vast quantities of vapor are continually rising from the surface of the seas, rivers, lakes, snow fields, and glaciers of the world. This vapor remains invisible until the air containing it is cooled down below its dewpoint, or point of saturation, — a result which follows upon the union or collision of two aerial currents of different temperatures, or the rise of the air into the upper cold regions of the atmosphere, where it is chilled by expansion, by radiation, or by contact with cold mountains. Condensa- tion appears only to take place on free surfaces, and the formation of cloud and mist is explained by condensation upon the fine microscopic dust of which the atmosphere is full. At first minute particles of water vapor appear, which either remain in the liquid condition, or, if the temperature is sufficiently low, are frozen into ice. As these changes take place over considerable spaces of the sky, they give rise to the phenomena of clouds. Further condensation augments the size of the cloud particles, and at last they fall to the sur- face of the earth, if still liquid, as rain ; if solid, as snow or hail; if partly solid and partly liquid, as sleet. As the vapor is largely raised from the ocean surface, so iu great 76 THE FITNESS OF THE ENVIRONMENT cipal constituent of active living organisms.1 ' Water is ingested in greater amounts than all measure it falls back again directly into the ocean. A con- siderable proportion, however, descends upon the land, and it is this part of the condensed vapor which we have now to follow. Upon the higher elevations it falls as snow, and gathers there into snow fields, which, by means of glaciers, send their drainage towards the valleys and plains. Else- where it falls chiefly as rain, some of which sinks underground to gush forth again in springs, while the rest pours down the slopes of the land, swelling the brooks and torrents which, fed both by springs and rains, gather into broader and yet broader rivers that bear the accumulated drainage of the land out to sea. Thence once more the vapor rises, con- densing into clouds and rain to feed the innumerable water channels by which the land is furrowed from mountain top to seashore. "In this vast system of circulation, ceaselessly renewed, there is not a drop of water that is not busy with its allotted task of changing the face of the earth. When the vapor ascends into the air, it is, comparatively speaking, chemi- cally pure. But when, after being condensed into visible form, and working its way over or under the surface of the land, it once more enters the sea, it is no longer pure, but more or less loaded with material taken by it out of the air, rocks, or soils through which it has traveled. Day by day the process is advancing. So far as we can tell, it has never ceased since the first shower of rain fell upon the earth. We may well believe, therefore, that it must have worked marvels upon the surface of our planet in past time, and that it may effect transformation in the future." — Geikie, "Textbook of Geology." London, 1903, 4th ed., Vol. I, pp. 447, 448. 1 Thus water makes up from 70 to 85 per cent of fishes, about 87 per cent of oysters, 85 per cent of apples, 78 per cent of potatoes, 95 per cent of the edible portion of lettuce, etc. WATER 77 other substances combined, and it is no less the chief excretion. It is the vehicle of the principal foods and excretory products, for most of these are dissolved as they enter or leave the body.1 Indeed, as clearer ideas of the physico-chemical organization of proto- plasm have developed it has become evident that the organism itself is essentially an aqueous solution in which are spread out col- loidal substances of vast complexity.2 As a result of these conditions there is hardly a physiological process in which water is not of fundamental importance. All of these circumstances, which completely justify the interest in water which Thales and Aristotle, and nearly all later students of nature have manifested, depend in great part upon the quantity of water which is present out- side the earth's crust, and upon its often unique physical and chemical properties. 1 Properly speaking, the entrance of the foods into the body is across the wall of the intestine; at this point the foods have all undergone digestion and are almost exclu- sively in solution. In like manner excretion takes place across the renal epithelium, or the epithelium of the lungs, or across that of the sweat glands; these too are traversed only by substances in solution. 2 "Der Organismus, Pflanze wie Tier, ist ein Geftss voll wasseriger Losung, in dem sich als disperse Phase verschieden- artige Kolloide befinden." — Bechhold, "Die Kolloide in Biologie und Medizin." Dresden, 1912. 78 THE FITNESS OF THE ENVIRONMENT Such properties are our present concern. Doubtless if it were not for the enormous quantity of water which exists upon our planet, all its physical properties would be of little avail to bring about its universal im- portance in nature. This, however, as has been above explained, appears to be neither an accidental nor an uncommon phenomenon. Of the total extent of the earth's surface the oceans make up about three fourths, and they contain an amount of water sufficient, if the earth were a perfect sphere, to cover the whole area to a depth of between two and three miles. This corresponds to about 0.2 per cent of the volume of the globe. The occur- rence of water is, moreover, not less important and hardly less general upon the land. In addition to lakes and streams, water is al- most everywhere present in large quantities in the soil, retained there mainly by capillary action, and often at greater depths. The atmosphere also contains an abundance of water as aqueous vapor and as clouds. Now i the very occurrence of water upon the earth, and especially its permanent presence, is due in no small degree to its chemical stability in the existing physical and chemical con- ditions. This stability is of great moment in the various inorganic and organic processes WATER 79 in which water plays so large a part. In the first place the chemical reactions in which it is concerned during the process of geological evolution, though they are no doubt in the total of great magnitude, are both slow and far from violent. Long since any very active changes of this sort, so far as the superficial part of the crust is concerned, have run their course. In the second place water is really, at the temperature of the earth and in com- parison with most other chemical substances, an extremely inert body, for the union of hydro- gen with oxygen is so firm that it is not readily dissolved. Thus water exists as a singularly inert con- stituent of the atmosphere, as a liquid nearly inactive in chemical processes on the surface and in the soil, and everywhere as a mild sol- vent which does not easily attack the sub- stances which in great variety dissolve in it. The chemical changes which do follow upon solution are not such as to produce substan- tial chemical transformations, and most sub- stances can pass through water unscathed. The nature of water, then, is a great factor in the chemical stability, which, no less than the physical stability of the environment, is es- sential to the living mechanism. But it may be questioned if such stability would not 80 THE FITNESS OF THE ENVIRONMENT necessarily be ultimately attained in greater or less degree with almost any other substance, as a result of the general tendency of chemical processes to reach a condition of equilibrium, and it will therefore be well to turn to more secure fields of inquiry. THERMAL PROPERTIES The most familiar among such are certain characteristics of water which have been long known, and which, as the Bridgewater Trea- tises and other works on natural theology testify, were formerly favorite subjects of metaphysical speculation, — the thermal prop- erties. These characteristics of water were recognized at an early stage in the develop- ment of modern science, and in many cases their special importance in meteorology and in other departments of the sciences of nature is almost self-evident. A SPECIFIC HEAT First among these is the heat capacity or, as it is more commonly termed, the specific heat of water. This quantity has the value of WATER 81 1.000 for the interval between 0° and 1° centigrade, a number which is due to the choice of water in defining the calorie or fundamental unit of heat. The caloric, small calorie, or gram calorie is that quantity of heat which is required to raise the temperature of one gram of water through 1° centigrade, and it varies slightly with the temperature, having the relative values 1.000 for the in- terval from 0° to 1°, 0.998 for the interval from 4° to 5°, 0.992 for the interval from 15° to 16°, and its mean value for the interval from 0° to 100° is 1.004. The heat capacity of ^ water is then 1.000, in that 1.000 calorie is re- quired to raise the temperature of 1.000 gram of water through 1.000 degree centigrade. The approximate specific heats of a variety of important substances are as follows: — [liquid .... 1.00 Water I solid . . . .0.50 [gas .... 0.3-0.5 Lead 0.03 Iron 0.10 Quartz 0.19 Salt 0.21 Marble 0.22 Glass 0.20 Sugar 0.30 Ammonia, liquid . . .1.23 Chloroform O.ii Hydrogen 3.4 Alcohol 0.5-0.7 Hexane 0.50 It is unnecessary to enter upon an elaborate analysis of the data concerning specific heats, for the magnitude of the specific heat of a substance is dependent upon its chemical nature, as was first made clear b}T Dulong 82 THE FITNESS OF THE ENVIRONMENT and Petit in 1819. The law which bears their name consists of the statement that in the case of elementary substances the product of specific heat and atomic weight is a constant, — roughly, 6.4. Certainly this so-called law is a mere approximation, and some elements, notably carbon, silicon, and boron, at the ordinary temperature, depart widely from its requirements, but in the main the approxi- mation holds good. Later the researches of Neumann, Gamier, Cannizaro, and especially of Kopp made possible an extension of the law to compounds. It is evident that the law of Dulong and Petit amounts to the statement that for all elementary substances the quantity of heat which is required to change the temperature of every atom, regardless of its nature, is a constant. A brief discussion will serve to make this plain. According to the law the specific heat of an element varies inversely as its atomic weight, diminishing as the atomic weight increases, so that the product of the two quantities remains constant. But of course the number of atoms per gram of sub- stance also varies inversely as the atomic weight. Hence the specific heats of elemen- tary substances and the number of atoms per gram are always roughly proportional, which WATER 83 can only be the case if all atoms, no matter of what element, require a constant amount of heat to raise their temperatures one degree. That is to say, in all elementary substances the heat capacity of the atom is constant, and independent of the nature of the element (with the qualifications above noted). The study of compounds has shown that this same generalization is also true of them. This means that in all substances the heat capacity of every atom is nearly constant and is independent of its nature and of that of the compound in which it finds itself. Accordingly the law of Dulong and Petit may be formulated as follows ; — the specific heat of a substance multiplied by the average of the atomic weights of all the constitutent atoms in the molecule is often equal to about 6.4, and is always not very different from this number. This conclusion may be tested with the data above recorded. Substance Molecular Weight Number of Atoms Average Atomic Weight Sfbczfio 1 Ii:at Specific Heat X Atomic Weight Water Ammonia Quartz . Salt . . Sugar . . Hexane . 18 17 60 58 342 86 3 4 3 2 45 20 6 4 20 29 8 4 1.0 1.2 0.2 0.2 0.3 0.5 6.0 4.8 4.0 5.8 2.4 2.0 84 THE FITNESS OF THE ENVIRONMENT It must be confessed that such data are not a brilliant confirmation of the law. A series of numbers which vary from 2.0 to 6.0 is something quite different from constancy, and every one of these numbers is less than 6.4. It is, however, certain that these quanti- ties are uniformly of the same order of magni- tude, and this is all that is of importance for our present purpose. For accordingly they prove that unless the average atomic weight of a substance be very low its specific heat cannot be very high. Of course only com- pounds which are largely made up of hydrogen can possess very low average atomic weights, and among such those will be lowest in this respect which contain a relatively small number of atoms of another element of low atomic weight, like carbon, nitrogen, oxygen, etc. Of such substances the hydrocarbons make up the only numerous group, and for the most part their specific heats appear to be, like that of elementary carbon itself, considerably lower even than would be predicted by the rule. So it is that the conclusion is warranted that water shares the characteristic of very high specific heat with a very small number of substances, among which hydrogen and am- monia are probably the only important chemi- cal individuals. From this conclusion another WATER 85 follows directly; namely, that water possesses certain nearly unique qualifications which are largely responsible for making the earth habitable, or at least very favorable as a habitation for living organisms. It need hardly be pointed out that this importance of the high heat capacity of water is a very well-known fact. Even in the early decades of the nineteenth century, when natural theology and argument from design were the subject of lively controversy, es- pecially in England, such subjects were very familiar, and an excellent temperate dis- cussion from the theologian's side will be found in Whewell's Bridgewater Treatise.1 At that time, before a clear formulation of the concept of adaptation existed, it was of course impos- sible to disentangle such natural fitness from the results of the organic evolutionary process. In the more modern period since the publi- cation of "The Origin of Species," the late Professor J. P. Cooke of Harvard has dwelt upon this and other properties of water and sought to show that, lying wholly apart from the new ideas, such phenomena remain 1 Chapter IX of this work deals with "The Laws of Heat with Respect to Water." Although the ideas are somewhat vague, the importance of the capacity of water to absorb heat is clearly brought out. 86 THE FITNESS OF THE ENVIRONMENT unexplained and inexplicable by our present laws of natural science. He, too, endeavored to employ such facts as theological arguments but, in spite of many sound contentions, with less success in a more skeptical age.1 The most obvious effect of the high specific heat of water is the tendency of the ocean and of all/lakes and streams to maintain a nearly constant temperature. This phenomenon is of course not due to the high specific heat of water alone, being also dependent upon evaporation, freezing, and a variety of cir- cumstances which automatically mix and stir water. But in the long run the effect of high specific heat is of primary importance. It will be convenient to postpone considera- tion of the regulation and importance of the constant temperature of the ocean until the other properties of water which contribute thereto have been discussed. A second effect of the high specific heat of 1 "Assume that the variations preserved by natural selec- tion are all accidental, a point on which naturalists greatly differ, still what is the result ? An adaptation to the environ- ment. According to the theory, then, the conditions of the environment are a determining cause ; and unless we believe that all nature was the result of a fortuitous concourse of atoms, we can find in these conditions abundant opportunities where intelligent causation can act." — Josiah Parsons Cooke, "The Credentials of Science." New York, 1888, p. 251. WATER 87 water is the moderation of both summer and winter temperatures of the earth. It is not easy to estimate the total magnitude of this effect, but the manner in which it comes about is well illustrated by the differences between seaboard and inland climates or between the climate of a large part of the United States, which is a continental climate, and that of Western Europe, which is essentially an in- sular climate. In the most extreme form such moderation of climate is to be observed on the high seas and upon small islands. There are found the smallest known differences between the mean temperatures of different months of the year and of different hours of the day, and the least tendency to violent changes of temperature. The calculation of Zenker regarding normal temperatures may Latitude Continental Climate Marine Climate Difference Degrees Degrees Degrees Degrees 0 34.6 26.1 -8.5 10 33.5 25.3 -8.2 20 30.0 22.7 -7.3 30 24.1 18.8 -5.3 40 15.7 13.4 -2.3 50 5.0 7.1 2.1 60 -7.7 0.3 8.0 70 -19.0 -5.2 13.8 80 -24.9 -8.2 16.7 90 -26.1 * -8.7 17.4 88 THE FITNESS OF THE ENVIRONMENT be cited as a good illustration of the nature of the case.1 It is unnecessary to discuss the effects upon living organisms of the equable tem- perature of the ocean and of the moderation of climate, for obviously we are here confronted by a true instance of regulation of the environ- ment. The high heat capacity of water operates in still another manner to regulate tempera- ture upon the land and at the same time to increase the mobility of the environment of marine organisms. For directly or indirectly it is involved in the formation and duration of ocean currents, especially the movement of water in the depths from the polar to the tropical seas, and it determines the amount of heat carried by such currents. A similar and even more important "function' is the direct promotion of winds, with the resulting distribution of aqueous vapor throughout the atmosphere, a primary factor in the dissemina- tion of water by means of the rainfall. Here the essential thing is the existence of a vast warm reservoir in the tropics and of two similar cold reservoirs at the poles. Under 1 A discussion of Zenker's work will be found in Hanns "Handbook of Climatology," translated by Ward, pp. 210- 215. WATER 89 these circumstances the circulation of winds, bearing away water vapor from the tropical oceans, is inevitable, and the process is intensified by the high specific heat of water. The living organism itself is directly favored by this same property of its principal constit- uent, because a given quantity of heat pro- duces as little change as possible in the tem- perature of its body. Man is an excellent case in point. An adult weighing 75 kilo- grams (165 pounds) when at rest produces daily about 2400 great calories, which is an amount of heat actually sufficient to raise the temperature of his body more than 32° centigrade. But if the heat capacity of his body corresponded to that of most substances, the same quantity of heat wTould be sufficient to raise his temperature between 100° and 150°. In these conditions the elimination of heat would become a matter of far greater difficulty, and the accurate regulation of the temperature of the interior portion of his body, especially during periods of great muscular activity, well-nigh impossible. Extreme con- stancy of the body temperature is, of course, a matter of vital importance, at least for all highly organized beings, and il is hardly conceivable that it should be otherwise. In 90 THE FITNESS OF THE ENVIRONMENT the first place marked influence of change of temperature upon chemical reaction is almost universal, and as a rule an increase of 10° centigrade in temperature will more than double the rate of a chemical change.1 Sec- ondly all living organisms contain both chemi- cal substances and physico-chemical struc- tures or systems which begin to be altered, and usually irreversibly altered, at a tempera- ture which is very little above that of the human body.2 It is perhaps imaginable that 1 If the velocity of a chemical reaction be represented by a coefficient, k, the increase in its magnitude with rising tem- perature is unlike that of ordinary physical coefficients, and in many cases amounts to a two or threefold rise for a tem- perature increase of 10° centigrade. The well-known data concerning the transformation of dibromsuccinnic acid into brommaleic acid and hydrobromic acid in aqueous solution illustrate a typical case. t k 15° 0.00000967 40° 0.0000863 50° 0.000249 60.2° 0.000654 70.1° 0.00169 80° 0.0046 89.4° 0.0156 101° 0.0318 2 This is attested not only by the low temperature at which many proteins coagulate, but also by the action of temperatures between 50° and 60° to inactivate enzymes, and WATER 91 conditions might be otherwise in beings of a very different kind, but to-day every chemist well knows that if he is to control a chemical process, almost the first desideratum is rigid regulation of the temperature at which the process takes place.1 It is therefore incontestable that the un- usually high specific heat of water tends automatically and in most marked degree to regulate the temperature of the whole envi- ronment, of both air and water, land and sea, and that of the living organism itself. Like- wise the same property favors the circulation of water by facilitating the production of winds, besides contributing to the formation of ocean currents. Here is a striking instance of natural fitness, which in like degree is un- attainable with any other substance except ammonia. to produce alterations in many of the complex substances that are involved in the phenomena of immunity and other similar things. 1 Almost the most conspicuous change in the equipment of modern chemical laboratories, as a result of the growth of physical chemistry, is the introduction everywhere of thermo- stats. 92 THE FITNESS OF THE ENVIRONMENT B LATENT HEAT Very different from specific heat in their relationship to the chemical constitution of a substance, but not unlike it in biological importance, are the so-called latent heats of melting and of evaporation. The latent heat of melting is expressed as the number of calories which are required to convert one gram of solid at the freezing point into one gram of liquid at the same tempera- ture. For water its value is approximately 80, which indicates that the same quantity of heat must be employed to melt ice as to raise the temperature of the resulting ice- water to 80° centigrade. The latent heat of evaporation is similarly defined as the number of calories required to change one gram of liquid into vapor. Its magnitude depends upon the temperature at which the process takes place. The latent heat of evaporation of water is approximately 536. There is required, accordingly, as much heat to boil away one gram of water as to raise the temperature of 536 grams through 1° centigrade. There are a number of important effects of WATER 93 the high latent heals of fusion and evapora- tion of water upon the meteorological pro- cesses. When, for example, a body of water becomes cooled to its freezing point, the further abstraction of heal cannol lower il> temperature below that point, which, to be sure, is somewdiat variable in the case of sail water. And so long as water and ice exi>t in contact, the system constitutes a ther- mostat, a very accurate one if the water be fresh, which changes only in respect to the quantities of ice and water as heat is added or removed.1 Heating serves merely to melt the ice, cooling to freeze the water. Accord- ingly, as long as the earth shall remain habit- able the cooling of its oceans and seas will remain rigidly limited by their freezing point. However inclement the atmosphere, the ocean can always support life until the final extinc- tion of water by cold. It is worthy of note that the freezing point of water, though to man with his carefully regulated body tem- perature apparently low, is in reality very high indeed compared with that of any like substances, — perhaps 100° centigrade above the average. 1 In fact, there is no better means of obtaining a constant temperature in the chemical laboratory than by mixing pure ice with pure water. 94 THE FITNESS OF THE ENVIRONMENT Table of Melting Points Water Hydride of antimony . Hydride of arsenic . , Hydrobromic acid . , Hydrochloric acid Hydrofluoric acid . , Hydriodic acid . . , Methane Carbon dioxide . . , Hydride of phosphorus Hydrogen sulphide . , Sulphurous oxide . . , Ammonia .... Nitric oxide . . . Degrees H20 0 SWI3 -91.5 AsH3 -113.5 HBr -87 HC1 -112.5 HF -92.3 HI -50 CKU -185.8 C02 - 57 PH3 -132.5 H2S -85.6 S02 -72.7 NH3 -75 NO -167 This is, no doubt, one of the most important facts with which we are concerned, for while a very large number of chemical processes take place quite freely at 0°, the conditions are very different at the freezing point of am- monia, for instance. At that temperature the velocity of most chemical processes is but a fraction of one per cent of their velocity at 0°, and a large part of the chemical activity which is familiar to us ceases. The result of the unusually high freezing point of water and of the phenomenon of latent heat is felt, however, not merely in the avoidance of an excessive fall in the tempera- ture of lakes and seas. As above explained, WATER 95 whenever the ocean comes in contact with climates of very low temperature it tends to moderate them, the more effectively the greater the disparity between the temperature of the air and that of the water, and here latent heat is quite as important a factor, though indirectly, as specific heat. It remains to point out that the latent heat of melting of water is nearly the greatest which has yet been discovered, being ex- ceeded, in fact, by that of ammonia alone. Table of Latent Heats of Melting Substance Formula Melting Point Latent Heat of Fusion Pb 326° 5.4 Calories Br -7.3 16.2 Cd 321 13.7 Iron Fe 23-33 Ga 13 19.1 I 11.7 K 58 15.7 Cu Na 96.5 43 31.7 Ni 464 Pd 36.3 Phosphorus .... P Pt 27.35 1779 4.7 27.18 Hg S Ag 115 999 2.82 9.37 21.07 Bi 266.8 12.64 Zn 415 28.13 Tin ! Sn 233 14.25 96 THE FITNESS OF THE ENVIRONMENT Melting Latent Heat Substance Formula Point of Fusion Water H20 0° 80 Calories Ammonia .... NH3 -75 108 Antimony chloride . SbCla 73 13.4 Antimony bromide . SbBr3 94 9.7 Lead chloride . . PbCU 485 20.9 Calcium chloride . CaCl2-6H20 28.5 40.7 Potassium nitrate . KX03 339 47.4 Sodium nitrate . . NaNOj 310.5 63 Phosphoric acid H3PO4 18 25.7 Nitric acid . . . HNO3 -47° 9.5 Sulphuric acid . . H2S04 10.3 24. Sulphuric oxide . . S03 76.7 Ethylene bromide . C2H4Br2 8 13 Formic acid . . . H.COOH -7.5 57.4 Chloral hydrate C2H3Cls02 46 33.2 Dimethyl oxalate . C204(CH3)2 49.5 42.6 Acetic acid . . . CH3COOH 43.7 Glycerine .... C3H8U3 13 42.5 Stearic acid . Ci8Hr,602 64 47.6 Benzene .... CeHe 5.3 30.1 Nitrobenzene . . C6H5N02 -9.21 22.3 Di-chlorbenzene CeH4Cl2 52.5 29.9 p-Toluidine . . . C7H9N 35.8 Phenol . . . . C6H5OH 25.4 24.9 Menthol . . . . CioII2oO 42 18.9 Phenylhydrazine . CH6.NH.NHa 24.5 Phenylacetic acid . C6H5.CH2.COOH 75 25.4 Naphthaline . . . CioHg 80 35.7 Accordingly, the processes above described possess nearly the highest possible efficiency. A very large amount of heat must be ab- stracted from a body of water before it can be solidified; after a given amount of cool- WATER 97 ing a very large quantity of water must re- main liquid ; a body of water at 0° centi- grade can warm up a very large amount of colder air with the formation of a very small quantity of ice. Thus the permanency of the ocean, and the moderating effect of water upon cold climates are very nearly maximal. These are also facts, directly dependent upon the physico-chemical nature of water, which are remarkably favorable to the organism. Still more important is the latent heat of evaporation of water. Wherever water is in contact with the air, evaporation must take place until, if the system be of small dimen- sions, equilibrium is established between aque- ous vapor and the liquid; in short until the air is saturated with water. Unlike freezing, which occurs only at one particular tempera- ture, this process goes on continuously through- out all ranges of temperature at which liquid water can exist, and even upon ice at low tem- peratures. It is always accompanied by the conversion of heat, in the' amount measured by the latent heat of evaporation, into other forms of energy ; the heat becomes latent. And since air in contact with water is rarely saturated with aqueous vapor, owing to the constant movement of the atmosphere, the process of evaporation, with the accompany- 98 THE FITNESS OF THE ENVIRONMENT ing conversion of heat into latent heat, is a continuous process. The phenomenon is a variable one, however, for while at high tem- perature, both because of the greater supply of heat and because of the greater amount of water vapor that the air can hold, the process is very important and active, at low tem- perature it is far less considerable. This in itself is no doubt a benefit because it tends especially to restrict the upward march of temperature when the temperature is high, but is of minor importance when the tempera- ture is low. In view of the other favorable qualities of water it is perhaps not surprising to find that its latent heat of evaporation is by far the highest known. So great, in truth, is this quantity and so important the process that the latent heat of evaporation is one of the most important regulatory factors at present known to meteorologists. When the sun shines upon a body of water, only a small part of the energy which the water receives contributes to the elevation of its temperature. Thus Fitzgerald has con- cluded from his studies of Lough Derg in Ireland during clear hot summer weather l 1 See Hann, "Handbook of Climatology," translated by Ward, p. 131. WATER 99 Table of Latent Heats of Evaporation Temper- Sub tance Formula ature op VAPOR- IZATION Latent Heat 0? Vaporization Water H20 100° 536 Calories Ammonia NH3 295 Bromine Br2 61.5 43.7 Chlorine Cl2 -22 67.4 Iodine I2 174 23.9 Hydrofluoric acid . . HF 360 Oxygen o2 -188 58 Nitrogen N2 49.8 Phosphorus . . . . P 287 130.4 Mercury Hg 350 62 Sulphur S2 316 362 Nitrous oxide . . . N20 100.6 Nitric acid .... HNO3 115 Sulphurous oxide . . S02 0 91.2 Sulphuric oxide . . . S03 18 147.5 Sulphuric acid . . . H2SO4 326 122.1 Thionylchloride . . SOCl2 82 54.5 Arsenic chloride . . AsCl3 69.7* Phosphorus trichloride PCI3 67.2* Stannic chloride . . SnCU 46.8* Silicon chloride . . . SiCL. 37.3 Carbon dioxide . . . C02 72.2 Carbon disulphide . . CSa 0 90 Carbon tetrachloride . CCI, 0 52 Cyanogen (CN)2 0 103 Hydrocyanic acid . . HCN 20 211* Methyl alcohol . . . CH3OII 0° 289.2 Ethyl alcohol . . . r.IUHI 0 ^:;«J.5 Amyl alcohol CbHuOH 131 120 Cetyl alcohol . . . QeHuOB 58.5 Hexane CHM 68 79.4 Methyl chloride . . CH3C1 0 !»<;.9 * Total heat of vaporization. 100 THE FITNESS OF THE ENVIRONMENT Temper- Substance Formula ature of Vapor- ization Latent Heat of Vaporization Ethyl bromide . . . C2H5Br 38.2 60.4 Calories Amyl iodide . C5HtlI 47.5 Aldehyde . . CH3.CHO 136.4 Chloroform . CHCl, 0 67 Ether . . . (C2H5)20 34.9 90.4 Acetone . . CH3.CO.CH3 56.6 125.3 Formic acid . HCOOH 103.7 Acetic acid . . CH3COOH 118 84.9 Acetic anhydride (CH3CO)20 137 66.1 Dichlor-acetic acid CHCloCOOH 138.4 79.1 Valerianic acid . CsHio02 103.5 Ethyl acetate C4H7O0 105.8 Acetyl chloride CH3COCI 78.9 Acetonitrite . . CH3CN 81.5 170.6 Ethyl amine . C2H5NH2 146.2 Benzene . Cell 6 0 109 Toluene . . C6X15CH3 111 83.5 Nitro-benzene C6H5N02 151.5 79.1 Aniline . . C6H5NH2 93.3 Acetophenone C6H5COCH3 203.7 77.2 Benzonitrite . C6H5CN 191 87.7 Piperidine C5HnN 105.8 88.9 Pyridine . . C5H5N 115.5 101.4 that in the morning the surface temperature rises about 0.6° per hour. This, however, appears to account for but a small fraction of the solar heat which the lake had taken up ; the rest must have been expended in evaporation. Another element of great im- portance is the transparency of water. As a result the rays of the sun are not absorbed WATER 101 by the mere surface alone, but a considerable layer of water near the surface receives the heat. At the equator the evaporation of the ocean appears to be about 2.3 meters per year,1 which involves more than 1,000,000,000,000,- 000 calories of latent heat per square kilo- meter. The amount of heat which is em- ployed in evaporating water from 100 square kilometers of the tropical ocean is accordingly vastly more than all the energy employed in the metabolism of the total population of the United States, and it amounts to more than 100,000,000 horse power. This is equiv- alent to more than one horse power per square meter day and night throughout the year. To a greater or less extent all over the earth this same process goes on, and as a result the water vapor in the air probably averages between 15 and 20 kilograms per square meter of the earth's surface, an ample supply for the formation of rain. The effect of this enormous evaporation to moderate the tem- perature of the tropics is very considerable; but the heat which thus disappears is not lost. Rendered latent at the place of evap- oration, it is turned back into actual heat at 1 This and other similar facts will also be found in the work of Hann. 102 THE FITNESS OF THE ENVIRONMENT the point of condensation, and thus serves to warm another and cooler locality. This process, so vast that all the water power of the globe may be regarded as its secondary by-product, possesses, in respect to its ten- dency to moderate and equalize the temper- ature of ocean, of lakes, and of the climates of all the earth, a maximal value. No other liquid could, during the evaporation of a given quantity of material, bind so much heat; no other vapor could yield so much heat upon condensation. Quite as important to man as this great power of meteorological regulation is the corresponding physiological activity, evap- oration of water from the skin and lungs. In an animal like man, whose metabolism is very intense, heat is a most prominent ex- cretory product, which has constantly to be eliminated in great amounts, and to this end only three important means are available: conduction, radiation, and the evaporation of water. The relative usefulness of these three methods varies with the temperature of the environment. At a low temperature there is little evaporation of water, but at body temperature or above there can be no loss of heat at all by conduction and radiation, and the whole burden is therefore thrown upon WATER 1 03 evaporation. The manner in which evapora- tion becomes important in temperature reg- ulation is well illustrated by the following cal- culation from a chart of Rubner's.1 The experiments upon which the chart is based were made upon the dog, an animal which lacks man's apparatus of sweat glands. The values of the table are only approximate. Temperature Heat Loss by Evaporation Degrees Per Cent 9 16 11 19 13 22 15 25 17 27 19 30 21 32 23 32 27 36 29 42 31 58 33 64 35 79 In plants evaporation is even more impor- tant than in animals. Evidently such adap- tation of the physiological processes to the conditions of the environment is enormously i favored by the high latent heat of evapora- tion.2 1 Lusk, "The Science of Nutrition," p. 99. 2 A full discussion of this subject will be found in the work of Lusk (see above), Chap. Ill, especially pp. 98-100. 104 THE FITNESS OF THE ENVIRONMENT There is still another beneficial result of this property : the great variation in the vapor tension of water which accompanies variation in temperature. Vapor tension measures the amount of vapor which is present in the atmosphere when it is in contact with a liquid and after it has become saturated with the liquid's vapor. Now, according to a well- known law, the rate of increase of vapor ten- sion, or in other words the amount of vapor which the air can hold, is greater the greater the latent heat of vaporization.1 Hence, degree by degree there is more variation in the vapor tension of water than there could be if the latent heat were lower. Such great variability in the quantity of water which the air can hold is in meteorology the most important characteristic of aqueous vapor. The relationship between vapor tension and temperature (centigrade) is shown in the accompanying table. 1 Near the freezing point an increase of 10° in temperature doubles the amount of water which the air can hold. The increase is proportional to the latent heat of vaporization according to the formulae 8.30.5 log ^ = fj(^) p0 1.99 \ l0li / where W stands for the latent heat of vaporization, p for vapor tension, and T for temperature. 2 Arrhenius, "Kosmische Physik," p. 612. 2 WATER Temperature Vapor Tension Degrees Millimeters 0 4.58 10 9.18 20 17.41 30 31.55 40 54.07 50 92.17 80 355.47 90 526.00 100 700.00 105 These variations are what make possible both the evaporation of water and its precipitation as rain and as dew in the meteorological cycle. And therefore the high latent heat of vapori- zation of water is in still another manner a most favorable circumstance in its effect upon the organisms. To sum up, this property appears to possess a threefold importance. First, it operates powerfully to equalize and to moderate the temperature of the earth; secondly, it makes possible very effective regulation of the tem- perature of the living organism; and thirdly, it favors the meteorological cycle. All of these effects are true maxima, for no other substance can in this respect compare with water.1 1 This conclusion mav be contrasted with that of Whcwell (Bridgewater Treatise, p. 142). He includes the expansion of water by heat, the expansion of water by cold below 40°, the expansion of water in freezing, the latent heats of melting and 106 THE FITNESS OF THE ENVIRONMENT c THERMAL CONDUCTIVITY The heat conduction of water is also a maxi- mum among ordinary liquids, and, though very low compared with good conductors like metals, must favor the equalization of tempera- ture within the living cells whose structure hin- ders the establishment of convection currents. Table of Heat Conductivities Water 0.0125 Rubber 0.0004 Alcohol 0.00048 Tin 0.15 Ether 0.00034 Lead 0.08 Benzene 0.00033 Iron 0.16 Glycerine 0.00066 Copper 0.72 Crown glass .... 0.0016 Silver 1.10 D EXPANSION BEFORE FREEZING A final thermal property of water remains to be considered; namely, its anomalous ex- pansion when cooled at temperatures near the freezing point. The facts are illustrated by the accompanying table. evaporation, and the rate of evaporation of water. It will be seen that eighty years ago it was already possible to make out a strong case for the fitness of water ; but it should not be forgotten that at that time ideas were in some respects still very vague, and comparative data few. WATER 107 Temperature Density of Water Expansion 1 to .: l c- . ,■ Per cent 0 0.99987 0.01:; 1 0.!)!)!)!):; 0.007 2 0.99997 0.003 3 0.99999 0.001 4 1.00000 0.000 5 0.99999 0.001 6 0.99997 0.003 7 0.99993 0.007 8 0.99988 0.012 9 0.99981 0.019 10 0.99973 0.027 20 0.99824 0.176 30 0.99567 0.44 40 0.99233 0.77 50 0.98813 1.19 100 0.95934 4.07 This unique property of water is the most familiar instance of striking natural fitness of the environment, although its importance has perhaps been overestimated.1 If, how- ever, water, like all other common substances, 1 It scarcely merits the curious rhapsody of Prout. for instance: "The above anomalous properties of the expan- sion of water and its consequences have always struck us as presenting the most remarkable instances of design in the whole order of nature — an instance of something done ex- pressly, and almost (could we indeed conceive such a thing of the Deity), at second thought, to accomplish a particular object." — Prout, Bridgewater Treatise, "Chemical Meteor- ology and the Function of Digestion." London, 1834, pp. 249-250. 108 THE FITNESS OF THE ENVIRONMENT steadily contracted on cooling, so that its point of maximum density fell at the freezing point, it is impossible to say how great would be the disadvantage for living organisms. Certain it is that life upon the earth would be thereby very greatly restricted. For this prop- erty, together with the by no means unique phe- nomenon of expansion upon solidification,1 is very largely responsible for the permanence in liquid state of many bodies of water in cold climates. In salt water the anomalous contraction disappears, and the lack of paleo- crystic ice is due to the density of ice and to the great mass of the ocean and the movement of its waters.2 There is an old experiment of Rumford's which well illustrates what conditions must have been had the contraction of water been normal and ice denser than water.3 He found that in a vessel filled with water, which contains ice confined at the bottom, it is possible to heat and even boil the superficial portion of the water without melting the ice. And so it would be with lakes, streams, and oceans were it not for the anomaly and the 1 The density of ice at the melting point is 0.91674. 2 A full discussion of this subject will be found in S. G unther's "Handbuch der Geophysik." 3 See Whewell's Bridgewater Treatise. WATER 109 buoyancy of ice. The coldest water would continually sink to the bottom and there freeze. The ice, once formed, could not be melted, because the warmer water would stay at the surface. Year after year the ice would increase in winter and persist through the summer, until eventually all or much of tlir body of water, according to the locality, would be turned to ice. As it is, the tempera- ture of the bottom of a body of fresh water cannot be below the point of maximum den- sity ; on cooling further the water rises ; and ice forms only on the surface. In this way the liquid water below is effectually pro- tected from further cooling, and the body of water persists. In the spring the first warm weather melts the ice, and at the earliest possible moment all ice vanishes. Such are the important thermal properties of water, and in briefest outline their unique fitness for the living mechanism. No other known substance could be substituted for water as the material out of which oceans, lakes, and rivers are formed, and as the sub- stance which passes through the meteorolog- ical cycle, without radical sacrifice of some of the most vital features of existing condi- tions. Ammonia in these respects is the only substance now known which approaches the 110 THE FITNESS OF THE ENVIRONMENT fitness of water. But not only is it almost inconceivable that ammonia should ever occur in sufficiently vast quantities upon a planet's surface, but it is evident as well that am- monia wholly lacks the qualification of anoma- lous expansion, and also in some of the most important of the other thermal properties falls far short of water ; while in latent heat of melting and in specific heat its advantage over water is inconsiderable. It is obvious that upon a body like the earth the state of the oceans and the meteor- ological phenomena are of the utmost impor- tance to all living things. Unless these be favorable, human experience and reflection alike agree that life could not widely exist. It seems, therefore, almost safe to say, on the basis of its thermal properties alone, that water is the one fit substance for its place in the process of universal evolution, when we regard that process biocentrically. II THE ACTION OF WATER UPON OTHER SUB- STANCES Although the thermal properties of water make up the classical subject-matter for dis- cussions of the fitness of the natural en- WATER 1 1 1 vironment, other no less important physical properties exist. Such especially arc those characteristics of liquid water which in no small measure determine the nature of the resulting physico-chemical systems when other substances, whether soluble or insoluble, crys- talline or colloidal, are brought into contact with it ; I mean the solvent power, the dielec- tric constant, together with the related ioniz- ing power, and the surface tension. WATER AS A SOLVENT As a solvent there is literally nothing to compare with water. In truth its qualifi- cations are on this point so unique and ob- vious that nobody seems to have taken the trouble to gather together the evidence, and, accordingly, beyond the bare assertion, a brief statement of the facts is not easy.1 In the first place the solubility in water of acids, bases, and salts, the most familiar classes of inorganic substances, is almost universal. 1 Nearly the whole science of chemistry has been built around water and aqueous solutions. A reference to anv text- book will at once reveal the truth of this statement. At first sight such a condition appears to be a matter of chance, but, as one becomes more familiar with the true character of the science, realization of a rational justification for the historical fact steadily grows. 112 THE FITNESS OF THE ENVIRONMENT Relatively few of these bodies are highly insoluble; very many are exceedingly soluble in water. Apart from their electrolytic dis- sociation and hydrolysis, which will be later discussed, the chemical changes wrought upon such dissolved substances in solution are commonly very unimportant. For chemical inertness, depending upon great stability, is a most significant characteristic of water, and undoubtedly a highly advantageous one as well. On the whole the best evidence for the efficiency of water as a solvent of inorganic sub- stances is to be found in the data of geology. Of all geological agents water appears to have been by far the most active within the periods of which investigation is made pos- sible by the geological record.1 Rainfall, the movement of surface streams and of water beneath the ground, and wave action, all contribute to the work of disintegration, sedimentation, etc., partly by dissolution of soluble material, partly by mechanical action. But mechanical action is itself much in- creased by the loosening which earlier dis- solution has caused. In this manner the great solvent power of water throughout its meteor- ological cycle largely contributes to the mobil- 1 Geikie, "Textbook of Geology," pp. 447-597. WATER 113 ization of many materials which could not otherwise be brought to the organisms which need them. It has been calculated by Murray1 that the total yearly run off of all the rivers of the earth is about 6500 cubic miles, carrying nearly 5,000,000,000 tons of dissolved mineral matter and prodigious quantities of sediment. The average composition of such water has been estimated as follows : — Parte per Million Potassium as K20 2 40 Sodium as Na20 7 10 Lithium as Li20 0 20 Calcium as CaO 43 20 Magnesium as MgO 14.70 Manganese as M113O4 1 20 Iron as FeO 2 80 Aluminium as AI2O3 310 Silicon as Si02 16.40 Carbonic acid as C02 46 Phosphorus as P2O5 , q.SO Nitric acid as N205 380 Sulphuric acid as S03 8 Chlorine as CI 3 70 Ammonia as NH3 0.07 Total mineral matter 152.97 It is, of course, almost exclusively to these constant accessions that the ocean owes its salinity, which in the course of time has reached well-nigh inconceivable magnitude. The common salt alone in the oceans of all 1 Russell, ''Rivers of North America," p. 80. 1 114 THE FITNESS OF THE ENVIRONMENT the earth amounts to not less than 35,000,000,- 000,000,000 tons. Quite as significant of the solvent power of water is the variety of elements whose presence in sea water can be demonstrated, thus proving that the total store of them is in any case enormous. They include hydrogen, oxygen, nitrogen, carbon, chlorine, sodium, magnesium, sulphur, phos- phorus, which are easily demonstrated; further, arsenic, csesium, gold, lithium, ru- bidium, barium, lead, boron, fluorine, iron, iodine, bromine, potassium, cobalt, copper, manganese, nickel, silver, silicon, zinc, alumin- ium, calcium, and strontium.1 Equally striking is the evidence in regard to the first stages of this geological process. Under the action of water, aided, to be sure, in many cases by dissolved carbonic acid, every species of rock suffers slow destruction. All substances yield in situ to the solvent work of water, and the dissolved parts may all be found in the great final reservoir, the ocean. It has been proved that nearly every one of the substances which are thus set in motion upon the face of the earth are placed under contribution by life, for bio- chemical analysis reveals them as constitu- ents of living organisms, absorbed either 1 Arrhenius, "Kosmische Physik." WATER 115 on their way down from the mountain tops to the ocean or by the marine flora and fauna. Not less valuable to the community of living things than the dissolution of the rocks, is the disintegration and transport of solid material, largely dependent thereon, which among its many results includes the prelim- inary steps of soil formation. By these familiar and enduring geological means chemi- cal substances are mobilized in the greatest variety of forms and conditions, and thus rendered available for the living organism. It is clearly evident from the chemist's long experience with solvents that no other fluid could permanently carry on this process with such acceptable regularity and efficiency. For no other chemically inert solvent can compare with water in the number of things which it can dissolve, nor in the amounts of them which it can hold in solution ; and of course any chemically active solvent must sooner or later exhaust itself by its chemical action, when the cycle must cease. Here, then, is a fitness of water which is open to no doubt. Let us turn for further proof to the organ- ism itself, taking blood serum as a source of information. The composition of this sub- 116 THE FITNESS OF THE ENVIRONMENT stance (in the case of the cow) is roughly as follows : ' — Parts per 1000 Water 913.6 Protein 72.5 Sugar 1.05 Cholesterine 1.24 Lecithine 1.68 Fat 0.93 Organic phosphoric acid 0.01 Na20 4.31 K20 0.26 CaO 0.12 MgO 0.45 CI 3.69 P205 0.08 Most of these substances are in solution, and unquestionably a host of others are pres- ent with them, in small and varying amounts. Among these may be mentioned iodine, bromine, iron, sulphates, urea, ammonia, ben- zoic acid, amino-acids, etc. But indeed all substances found in urine (see below) also occur in blood. It cannot be doubted that if the vehicle of the blood were other than water, the dissolved substances would be greatly restricted in variety and in quantity, nor that such restriction must needs be ac- companied by a corresponding restriction of the life processes. 1 A full discussion of the following data may be found in such works as those of Hammarsten and Abderhalden on physiological chemistry. WATER 1 1 7 The composition of the urine provides another excellent illustration of the utility of the solvent power of water. In the course of its complex chemical processes a higher organism produces a host of end products which must be removed, and also finds itself accidentally in possession of a great variety of other useless substances which require excretion. The solvent power of water is one of the great factors in facilitating this task. Human urine has been reported to contain in solution the following substances: urea, carbamic acid, creatinine, creatine, uric acid, xanthine, guanine, hypoxanthine, adenine, paraxanthine, heteroxanthine, epi- sarkine, epiguanine, oxalic acid, allantoine, hippuric acid, phenaceturic acid, benzoic acid, phenolsulphuric acid, skatoxylsulphuric acid, paraoxyphenylacetic acid, homogentisic acid, urobiline, urochrome, uroerythrine, glucose, levulose, lactose, numerous compounds of glycuronic acid, glycine, alanine, leucine, tyro- sine, and other amino-acids, various enzymes, putrescine, cadavarine, and countless other organic substances, chlorides, bromides, and iodides, phosphates and sulphates, potassium, sodium, ammonia, calcium, magnesium, iron, carbonic acid, nitrogen, argon, etc. Here again it is sure that such variety could 118 THE FITNESS OF THE ENVIRONMENT not be attained with another solvent. It is no exaggeration to say that except atmos- pheric oxygen and carbonic acid, nearly all the food of living organisms is water borne, and all material in its passage into the body, through the body, and out of the body nearly always employs the same vehicle. Cer- tainly no other form of transport would be so efficient and so economical. B IONIZATION If, therefore, aqueous solutions are, ap- parently of necessity, the very basis of the life processes, the state of substances when in this condition, and also when in contact with water, is of vital importance. Here two prop- erties of water, the dielectric constant and the surface tension, exert a cardinal influence. Among the phenomena of solution those which are related to electrolytic dissociation, as suggested by the hypothesis of Arrhenius, have deservedly received a great deal of at- tention since the secure establishment of the new science of physical chemistry in the eighties of the last century. In the course of time the belief that in aqueous solution the molecules of all acids, bases, and salts are more or less split into particles which bear WATER 1 1 g electrical charges has been universally ac- cepted. These so-called ions arc the sourer of nearly all the electrical phenomena of solu- tion, whether in batteries, in the manifesta- tions of animal electricity, or in simple con- duction through an aqueous solution. But the more familiar electrochemical processes are by no means the only results of ioniza- tion. An infinite number of chemical inter- actions between dissociated bodies follow inevitably. These changes are not, to be sure, decisive and irreversible, but balanced actions, which, however, vastly increase the variety of substances that exist in water. Let us consider, for example, a system which has been made by dissolving in water the simple salts sodium chloride, NaCl, potas- sium bromide, KBr, and lithium iodide, Lil. According to the ionization hypothesis, more than half of the molecules of every one of these salts will at once dissociate into ions as follows : — NaCl = Na + Cl KBr = K + Br LiI = Li + I These reactions are balanced, and it is confidently believed that the ions are con- 120 THE FITNESS OF THE ENVIRONMENT stantly recombining to form molecules and the molecules constantly dissociating once more to form ions. At the same time, noth- ing hinders the union of sodium ions with bromine ions, or of any other pair of positive and negative ions. Accordingly, the solu- tion at once contains not only the three origi- nal salts and the six different varieties of ions, but also the following new molecules: — sodium bromide, NaBr, sodium iodide, Nal, potassium chloride, KC1, potassium iodide, KI, lithium chloride, LiCl, and lithium bromide, LiBr. All nine varieties of molecules and all six of ions are concerned in a complicated system of chemical reactions which are now well understood, the state of equilibrium depending upon known conditions. For in- stance, if the solution be a moderately dilute one and the original substances be present in chemically equivalent quantities, about 90 per cent of the material will be in the ionic state, each variety of ions making up about 15 per cent of the total, and about 10 per cent will be in the form of molecules, each variety constituting about 1.1 per cent. There can be no doubt that ionization plays a great part in determining the char- acteristics of solutions of acids, bases, and salts, and in bringing about the reactions which WATER 121 occur among them, and between them and other substances. Such, then, is the process to which are due most of the electrical phenomena and many of the chemical phenomena of solutions, and it is certain that the extent and variety of ionization in water far surpass what is possi- ble in any other solvent. One reason for this is most simple. The ionizing substances are so very much more often soluble in water than in any other solvent, and when soluble are in general so much more highly soluble, that the opportunity for ionization in water is quite unparalleled. Further, ionization in solution unquestionably depends upon the dielectric constant of the solvent, in accord- ance with the principle first stated by Nernst that the greater the dielectric capacity of tin' solvent, the greater is the degree of electro- lytic dissociation of substances dissolved in it. when the conditions are otherwise the same.1 1 "The following consideration will make this principle clearer: The positively and negatively charged ions would unite to form electrically neutral molecules because <>f the electrostatic attraction which exists between them ii' it were not for the action of another and opposing force the nature <»f which is as yet unknown. The equilibrium between these two forces gives rise to the equilibrium between the ions and the undissociated molecules, or determines the degree of dis- sociation. When the dielectric constant i> increased, the 122 THE FITNESS OF THE ENVIRONMENT This is the case because the tendency of the electrically charged ions to reunite and form electrically neutral molecules must be less the greater the dielectric constant of the sol- vent. Now the dielectric constant of water is nearly the highest at present known, and therefore ionization in water is on that account also more extensive than in almost any other solvent. Finally, for reasons that are not yet under- stood, the process of ionization in other sol- vents than water is a much more complex affair, and there can be no doubt that such complexity limits the phenomena which are dependent upon ionization.1 electrostatic attraction between the ions is alone weakened, and hence the degree of dissociation is increased." — Le Blanc, "A Textbook of Electro-Chemistry." New York, 1907, p. 147. 1 "It would be natural to expect that the conceptions which have been found serviceable in the case of solutions in water could be applied directly to solutions in other solvents, keeping in mind that, according to the individual nature of any given solvent, the degree of dissociation, the migration velocity of the ions, and consequently the conductivity of a solution of a given concentration would be different. It is a noteworthy fact, however, that the behavior of non-aque- ous is much more complicated than that of aqueous solutions. This is shown especially by the investigation of the conduc- tivity of solutions of various substances in liquid sulfur dioxide made by Walden and Centnerszwer. Neither the law of the independent migration of the ions, nor the law that by increas- WATER 123 Physiologically, as researches of the last twenty years clearly prove, the action of ions is of fundamental significance. The brilliant investigations of J. Loeb, and the long series of studies by various other physiologists of the influence of electrolytes upon colloids form perhaps the most telling evidence for this belief.1 At all events there is no ques- tion that the simple equilibria between acids and bases and salts are of extreme importance in physiological processes. They lie at the very basis of the structure of all protoplasm, ing dilution the conductance approaches a maximum value, nor, finally, the dilution law, was found to hold. Molecular weight determinations carried out at the same time by the boiling-point method gave normal values for non-electrolytes, and abnormally large values for electrolytes, whereas abnor- mally small values would be expected. This indicates that association has taken place, to a considerable extent, which in all probability takes place not only between molecules of dissolved substance, but also between these molecules and those of the solvent. Considering these circumstances, it is very fortunate for the advance of the sciences of chemistry and electro-chemistry that such complications arc generally, although not always, absent in the case of aqueous solutions. It is due to this fact that it has been possible to deduce simple laws from a study of such solutions." — Le Blanc, "A Text- book of Electro-Chemistry." New York, 1907, pp. H2-1 \:\. 1 Full discussion of such subjects will be found in the "Dynamics of Living Matter," by Loeb, and in his contribu- tion to Oppenheimer's "Handbuch der Biochemie," as well as in Hober's " Physikalische Chemie der Zelle und der Gewebe." 124 THE FITNESS OF THE ENVIRONMENT and the sure and definite relations between such bodies provide, as it were, a secure foun- dation for the more complex organic structures. More obvious is the value of ions as sources of electricity. If the older electro-physiology of the third quarter of the nineteenth cen- tury has proved in some respects a sterile field, there can yet be no doubt that more subtly, and quite apart from the nervous im- pulse and the peculiar phenomena of electrical fishes, electrical phenomena are everywhere involved in the most intimate of the physio- logical processes. Even without further discussion of a sub- ject that must soon lead into difficult and highly technical considerations, I feel sure that the existence of another important fitness of water is patent. For ions are evidently a real contribution to the richness of the environment. They enhance the Variety of chemical substances and of chemical reactions ; they constitute a group of singularly mobile ( chemical agents; they provide electricity; and, finally, aqueous solutions are by far the best source of ions. It must be pointed out before leaving this subject that the dielectric constant, hence the ionizing power, is somehow related to various other properties of the solvent. In WATER 125 the accompanying table KD stands for the dielectric constant, IIV for the latent heat of vaporization, and KH for the absolute conductivity for heat. It is to be observed that on the whole all three quantities decrease simultaneously. These properties are also related to the critical pressure, to the van der Waals constant a, and to the molecular volume at the boiling point. Solvent Water, H20 Methyl alcohol, CH3OH . . . Ethyl alcohol, C2H50H . . . Formic acid, H • COOH . . . Acetic acid, CH3 ■ COOH . . Ammonia, NH3 Methylamine, CH3 • NH2 . . Sulphurous oxide, SO2 . . . Acetone, CH3 • CO • CH3 . . . Ethylacetate, CH3 • CO • O • C2H5 Benzene, CeHe Toluene, CaHs • CH3 .... Ether, (QHs^O Chloroform, CHCI3 .... Tetrachlormethane, CCI4 . . Stannic chloride, SnCl4 . . . KD Hv 81.7 536.5 32.5 267.5 21.7 205 57.0 103.7 6.5 89.8 16 329 <10.5 14 92.5 20.7 125.3 5.85 86.7 2.26 93.5 2.31 83.6 4.3G 84.5 4.95 58.5 2.18 46.35 3.2 30.53 K 17 0.154 0.0495 0.0423 0.0648 0.0472 0.0348 0.0333 0.0307 0.0303 0.0288 0.0252 Such evidence clearly suggests that some of the manifold fitnesses of water proceed from a single cause or group of causes. For the present, however, these relationships are 126 THE FITNESS OF THE ENVIRONMENT obscure, and in any case there seems to be no immediate hope of bringing them into connection with the science of biology. C SURFACE TENSION Of all common liquids, except mercury, water has the greatest surface tension. Table of Surface Tensions Water 75 Carbonic acid 1.8 Ammonia 41.8 Mercury 436 Benzene 28.8 Methyl alcohol 23 Ethyl alcohol 22 Ether 16.5 Glycerine 65 Acetone 23 Formic acid 37.1 Acetic acid 23.5 Chloroform 26 This fact is of enormous moment in biology, most obviously perhaps in its influence upon the soil. For surface tension and density determine the height to which a liquid will rise in a capillary system, and thus it comes about that the principal factor in bringing water within reach of plants is the exceptional surface tension of water. The nature of the case is clearly explained by Hilgard. "The WATER 127 liquid water held in the pores of the soil, in the form of surface films representing the curved surface seen in capillary tubes, and therefore tending to cause the water to move upwards, as well as in all other directions, until uniformity of tension is established, is of vastly higher importance to plant growth than hygroscopic moisture. It not only serves normally as the vehicle of all plant food ab- sorbed during the growth of the usual crops, but also, as a rule, to sustain the enormous evaporation by which the plant maintains, during the heat of the day, a temperature sufficiently low to permit of the proper op- eration of the processes of assimilation and building of cell tissue." * The rise of water in capillary systems re- sembling soil, under the action of surface tension, may be as much as ten feet. In soil itself the highest rise under the usual circum- stances is unquestionably as much as four or five feet; but if the surface tension of water were like that of most liquids it could be, under similar conditions, but two or three feet. There seems to be little doubt that the rise of fluids in tall plants is also in large part due to the action of surface tension, and accordingly it must be much favored by the 1 Hilgard, "Soils." New York, 1907, p. 201. 128 THE FITNESS OF THE ENVIRONMENT magnitude of that quantity in the case of water. Finally, surface tension is of great impor- tance, indeed in simple cases is the one effective agent, in the phenomenon called adsorption.1 On the basis of thermodynamical considera- tions first developed by Willard Gibbs, it is easy to show that whenever the dissolution of a substance changes the surface tension of a solvent, the distribution of the dis- solved substance will not be strictly homo- geneous. If the solution has a lower surface tension than the solvent, the surface of the solution will become more concentrated than the interior; or if the surface tension of the solution be greater than that of the solvent, the surface of the solution will become less concentrated than the interior. This result, quite insignificant in simple solutions, becomes a matter of much moment when, as in the case of suspensions of fine particles like ani- mal charcoal, in emulsions, jellies, or any other system of like disperse heterogeneity of physical constitution, there occurs very great increase of surface area. Then it is 1 A familiar example of adsorption is the use of bone- black to decolorize sirup in the process of sugar refining. The colored matters are almost completely removed from solution, and cling to the surface of the charcoal. WATER log that adsorption becomes a factor of the great- est weight; for, other things being equal, the total force of surface tension in a system is proportional to the area of surface. Under these circumstances dissolved substances are no longer distributed with any approach to equality or regularity in the system, but they collect at the surface in very great quantities, and in the most irregular manner. Now of all known physical structures there is none which rivals protoplasm in its fine complexity, and adsorption is therefore un- questionably a prominent agent in deciding its physico-chemical constitution. Moreover, adsorption influences and complicates almost every process of chemical physiology, for no product of life is without its colloids, i.e. sub- stances which are finely divided and there- fore endowed with great surface areas. In truth colloids are probably quite essential to fine complexity, and so to every conceivable form of life.1 The evidence for this universal importance 1 "Eines aber mochte ich beliaupten, welches auch immer die stoffliche Zusammensetzung jener Lebewesen (living organisms in another world) sein mag : es miissen Kolloide sein. . . . Welcher andere Zustand, ausser dem Kolloiden, konnte derart veranderliche, derart plastische Formen bilden und ware doch im stande, diese Formen, wenn notig, unver- anderlich zu wahren." — Bechhold, I.e., p. 194. 130 THE FITNESS OF THE ENVIRONMENT of adsorption in biology is not to be briefly presented, but it may be found in almost endless profusion in such works as those of Freundlich and Bechhold.1 It must not be supposed that the phenomena of adsorption in biology are simple and exactly understood. What is certain is that they are universal, and that surface tension lies at the root of the matter. This is because all living things are colloidal, and I am in- clined to think that most physiologists will admit that life without colloids is probably unthinkable, even in a world very differently constituted from our own. Colloidal struc- tures are, in fact, the first and greatest factors in physical complexity of organization, and the principal force, unless it be in exceptional cases an electrical charge due to ions, which operates upon the colloidal structures is sur- face tension. This, then, is another striking fitness of water above all other things. Such are the facts which I have been able to discover regarding the fitness of water for the organism. The following properties ap- pear to be extraordinarily, often uniquely, 1 Freundlich, "Kapillarchemie." Leipzig, 1909. Bech- hold, "Die Kolloide in Biologie und Medizin." Dresden, 1911. WATER 131 suited to a mechanism which must be complex, durable, and dependent upon a constant metabolism: heat capacity, heat conductiv- ity, expansion on cooling near the freezing point, density of ice, heat of fusion, heat of vaporization, vapor tension, freezing point, solvent power, dielectric constant and ionizing power, and surface tension.1 In no case do the advantages which these properties confer seem to be trivial ; com- monly they are of the greatest moment; and I cannot doubt, even after allowances have been made for the probability of occa- sional fallacies in the development of an argu- ment which, though simple, is beset with many pitfalls, that they are decisive. Water, of its very nature, as it occurs automatically in the process of cosmic evolution, is fit, with a fitness no less marvelous and varied than that fitness of the organism which has been won by the process of adaptation in the course of organic evolution. If doubts remain, let a search be made for 1 Contrasting this statement with that of Whewell in his Bridgewater Treatise, it is evident that while the progress of science has provided much novel information, and elim- inated many false views, the present situation differs from the earlier one only in the better definition of the issue, and in our modern freedom from metaphysical and theological associations. 132 THE FITNESS OF THE ENVIRONMENT any other substance which, however slightly, can claim to rival water as the milieu l of simple organisms, as the milieu interieur of all living things, or in any other of the count- less physiological functions which it performs either automatically or as a result of adapta- tion. In truth Darwinian fitness is a perfectly reciprocal relationship. In the world of modern science a fit organism inhabits a fit environment. 1 Here and elsewhere the word " milieu " has been employed when it is desired to express only that part of the conception of environment which is involved in the literal meaning of the word, leaving to environment the added significance of that which provides food. CHAPTER IV CARBONIC ACID TWO chemical individuals stand alone in importance for the great biological cycle upon the earth. The one is water, the other carbon dioxide. The one, for reasons which we have just reviewed, is the most familiar of all the varieties of matter. The other rarely is seen except by chance, and with- out scientific research never could have been known for what it is in value to living things. Yet these two simple substances are the com- mon source of every one of the complicated substances which are produced by living beings, and they are the common end prod- ucts of the wearing away of all the constitu- ents of protoplasm, and of the destruction of those materials which yield energy to the body.1 1 A man weighing 60-70 kilograms excretes daily the fol- lowing quantities of material : — Water 2500-3500 grams Carbon dioxide .... 750-900 grams All other substances . . 00-1-25 grams 133 134 THE FITNESS OF THE ENVIRONMENT Once perhaps the atmosphere of the earth consisted chiefly of water and carbon dioxide; but cooling has caused the condensation of most of the water, and geological processes, more recently aided by the action of vegetation with coal and peat formation, have removed nearly all of the carbon dioxide. The latter transformations have resulted in the substitu- tion of oxygen in the atmosphere. However, the interior of the earth continues to deliver through volcanoes large amounts of carbon dioxide, and thus the original source of atmos- pheric carbon dioxide persists as a diminish- ing supply. To-day carbon dioxide makes up only a little more than 0.03 per cent by volume of the whole atmosphere, approximately 4.6 kilograms per square meter of the earth's surface, or about 2,300,000,000,000 metric Water is, therefore, three fourths, carbon dioxide one fifth, of the total, and all other substances amount to but 2 to 3 per cent. In like manner the materials ingested by an ordinary green plant are proportioned, water making up more than nine tenths, and carbon dioxide amounting to fully five times the sum of all other substances combined. Needless to say, a large part of the water which enters and leaves plants and animals has had no real share in their organization. It is merely the bearer of dissolved substances or the means, through evaporation, of lowering the tempera- ture of their bodies. CARBONIC ACID 135 tons for the whole earth.1 In the oeean the amount of carbonic acid is about 0.1 gram per liter, or approximately 0.01 per cenl by weight. Here, however, much (lie larger part is in chemical union with bases, chiefly in the form of bicarbonates.- There can be no doubt that the physical properties of carbon dioxide are less important to the living organism than are those of water. But indeed the characteristics of water so largely determine the nature of the environ- 1 The composition of dry air is as follows : — Per Cubic Meter Nitrogen . . Oxygen . . Argon . . . Carbon dioxide Nitrogen . . Oxygen . . Argon . . . Carbon dioxide 2 See below in the discussion of the alkalinity of the ocean. 136 THE FITNESS OF THE ENVIRONMENT ment and the conditions within the organism alike, that this is necessarily the case. How- ever, the less conspicuous substance is not without its physical fitnesses, and we must now turn to them. SOLUBILITY The most obvious of the properties of car- bonic acid is its all-pervasiveness. Originally formed in vast quantities by the cosmic pro- cess, and accumulated in the atmosphere, the store has been steadily replenished there by vulcanism. It is probable that of the enor- mous quantities now deposited as limestone in the earth's crust, a quantity sufficient to yield an atmospheric pressure greater ten- fold than the present atmospheric pressure, only a fraction was at any one time actually gaseous. For it happens that the presence of carbonic acid in the atmosphere insures the occurrence of greater, or at least nearly equal, quantities in the ocean and in all the natural waters of the earth. This is due to the solu- bility of carbon dioxide, to the magnitude of its absorption coefficient in water. The absorption coefficient is the volume of gas absorbed by one liter of liquid when CARBONIC ACID 137 the gas pressure^ is one atmosphere. Accord- ing to the law of Henry, however, the absorp- tion of the gas is always proportional to its Table of Absorption Coefficients at 0° Oxygen 0.01!) Hydrogen 0.0*1 Nitrogen 0.0*1 Carbon monoxide 0.035 Carbon dioxide 1.797 Sulphurous oxide 79.79 Ammonia 1299 partial pressure in the atmosphere, and there- fore the absorption coefficient measures the ratio, after equilibrium has been attained, between the concentration of a substance in solution and in the gaseous state, no matter what that concentration may be. The absorption coefficient is not constant under all circumstances, and varies especially with the temperature. Table of Absorption Coefficients of C02 Temperature Absorption Coefficient 0° 1.797 10° 1.185 20° 0.901 37.29° 0.563 100° 0.2 44 It will be seen from the tables that, differ- ing from most gases, carbon dioxide has an absorption coefficient nearly equal to one, and 138 THE FITNESS OF THE ENVIRONMENT that for the ordinary temperature of the earth's waters, where they are in contact with car- bonic acid gas, it is very close indeed to 1.0. Hence, when water is in contact with air, and equilibrium has been established, the amount of free carbonic acid in the water is almost exactly equal to the amount in the air. Unlike oxygen, hydrogen, and nitrogen, carbonic acid enters water freely ; unlike sulphurous oxide and ammonia, it escapes freely from water. Thus the waters can never wash carbonic acid out of the air, nor the air keep it from the waters. It is the one substance which thus, in considerable quantities relative to its total amount, everywhere accompanies water.1 In earth, air, fire, and water alike these two sub- stances are always associated. Accordingly, if water be the first primary constituent of the environment, carbonic acid is inevitably the second, — because of its solubility possessing an equal mobility with water, because of the reservoir of the at- mosphere never to be depleted by chemical 1 "Carbonic acid being more soluble than the other gases, is contained in rain water in proportions between 30 and 40 times greater than in the atmosphere." — Geikie, "Text- book of Geology." 4th ed., Vol. I, p. 449, 1903. It must not be forgotten that carbonic acid in subterranean water, by which so much geological change is accomplished, originates, not in the air, but from organic matter in the soil. CARBONIC ACID 189 action in the oceans, lakes, and streams. In truth, so close is the association between these two substances that it is scarcely con-eel logi- cally to separate them at all; together they make up the real environment and I hey never part company. Carbonic acid thus possesses the first great qualification of a food: its occurrence is universal and its mobility a maximum. This is due to the fact that its absorption coefficient is on the average ap- proximately one, the most favorable value. Needless to say the absorption coefficient of carbonic acid is also of great importance in many physiological processes, chiefly per- haps in excretion. In the course of a day a man of average size produces, as a result of his active metabolism, nearly two pounds of carbon dioxide. All this must be rapidly re- moved from the body. It is difficult to im- agine by what elaborate chemical and physical devices the body could rid itself of such enor- mous quantities of material were it not for the fact that, in the blood, the acid can cir- culate partly free ' and, in the lungs, by a pro- cess which under ordinary circumstances has all the appearances of a simple physical phe- 1 Of the total carbonic acid of the blood 5-10 per cent exists as the free acid, partly in the plasma, partly in the cor- puscles. 140 THE FITNESS OF THE ENVIRONMENT nomenon,1 can escape into air which is charged with but little of the gas.2 Were carbon dioxide not gaseous, its excretion would be the greatest of physiological tasks ; were it not freely soluble, a host of the most universal existing physiological processes would be im- possible. II ACIDITY The only other property of this substance which calls for consideration at this point is its acid nature and strength. Very few min- erals are freely soluble in pure water, and nearly all yield to the process of weathering far more readily because of the carbonic acid which all natural waters contain.3 The latter 1 The determination of the exact nature of the process by which carbonic acid is excreted across the lung membrane is one of the standing difficulties of physiology, but we need not here enter upon its consideration. 2 Expired air contains on the average more than 4 per cent by volume of carbonic acid. 3 "A few minerals (halite, for example) are readily soluble in water without chemical change, and without the aid of any intermediate element; hence the copious brine-springs of salt regions. In the great majority of cases, however, solu- tion is effected through the medium of carbonic acid or other reagent. Limestone is soluble to the extent of about 1 part in 1000 of water saturated with carbonic acid. The solution and removal of lime from the mortar of a bridge or vault, CARBONIC ACID m constituent is probably a necessary adjuvant in the most far-reaching geological phenom- ena. Indeed, it is the united net ion of water and carbonic acid, aided in lesser degree by nitric acid, which has been formed in the atmos- phere by electrical action, and by acid prod- ucts of vegetation, which sets free the in- organic constituents of the earth's crust and turns them into the stream of metabolism. But apart from the solvent action of car- bonic acid, there is another group of phenomena which depend upon its acid character. These must now be explained. They are the neu- trality or faint alkalinity of the ocean, and of protoplasm. According to the modern theory of solution, water itself, like the dissolved electrolvtes, is dissociated into ions, though only to a very slight degree.1 The reaction is expressed as follows : — H20=H+OH and the deposit of the material so removed in stalactites and stalagmites, likewise the rapid effacement of marble epi- taphs in our church yards, are instances of this solution. . . . Among the sulphates, gypsum is the most important example of solution. It is dissolved in the proportion of aboul 1 part in 400 parts of water. Even silica is abstracted from rocks by natural waters." — Geikie, "Geology," pp. 451-452. 1 For a discussion of this subject the textbook of M.llor, "Chemical Statics and Dynamics," p. 405, may be consulted. 142 THE FITNESS OF THE ENVIRONMENT If the water be pure, the concentrations of hydrogen and hydroxyl ions are necessarily the same, for water is electrically neutral. A variety of independent methods of estimation have shown that at 25° (centigrade) this concentration amounts almost precisely to 0.0000001N, in the ordinary units.1 This corresponds to 0.0000001 gram of ionized hydrogen and 0.0000017 gram of ionized hydroxyl in 1000 grams of water. Further, the theory of solution explains acidity in water by the occurrence of hydrogen ions, formed from dissolved electrolytes, in excess of hydroxyl ions ; and alkalinity by a similar excess of hydroxyl over hydrogen ions. Neu- trality is then the condition when, as in pure water, the two concentrations are equal. In short, expressing the concentration of ionized 1 Concentrations are expressed in terms of chemical equivalents, gram-molecules, or moles. N (normal) is the symbol for this unit. The values of the concentration of ionized hydrogen at neutrality as estimated by different investigators are as follows : — 6. X10"7 Kohlrausch 1884 1.0 XKT7 Ostwald 1893 1.1 X 10 7 Arrhenius, Shields 1893 1.2 X10"7 Wijs 1893 0.9 X10"7 Kanolt 1907 1.02 X 10"7 Heydweiller 1909 1.02 X10-7 Lunden 1907 CARBONIC ACID 148 * + hydrogen by (H) and of ionized hydroxyl by (OH) : - If (H) = 0.0000001 N = (OH) the solution is neutral; if (H) > 0.0000001 N > (OH) the solution is acid; if (H) < 0.0000001N < (OH) the solution is alkaline. It remains to point out that implicit in these definitions is the well-founded hypothesis that in water the concentrations of hydrogen and hydroxyl ions vary inversely, so that with constant temperature under all circum- stances their product is constant : ! — (H) X (OH) = K Substituting in this equation the value 0.0000001 of the concentrations at neutral- ity, we obtain the value K - 0.00000000000001 u ra\ 0.00000000000001 whence (H) = (OH) 1 This corresponds with the requirements of the Mass- Law. 144 THE FITNESS OF THE ENVIRONMENT Whenever a weak acid is present in aqueous solution in company with such bases sa sodium, potassium, calcium, magnesium, etc., which are invariable constituents of the ocean, blood, protoplasm, etc., provided the acid be in excess, it is a simple matter to determine the reaction, which can best be measured by + the values of (H) and (OH), following the considerations above. Now there is a certain characteristic prop- erty of an acid, its ionization constant, k, which measures its tendency to dissociate in aqueous solution, thereby to produce hydro- gen ions, and hence to increase the intensity of acidity. Strong acids have ionization con- stants which are of the order of magnitude of 1.0, weak acids of the order of magnitude of 0.0001, the weakest acids, 0.00000001, or less. Table of Ionization Constants HC1, HN03, etc 1. H3P04 0.011 H3As04 0-005 HNO2 0.0005 H2C03 0.0000003 NaH2P04 0.0000002 H2S 0.000000091 H3BO3 0.0000000007 Na2HP04 0.00000000000036 It has been discovered that in the general case above discussed the concentration of CARBONIC ACID 145 ionized hydrogen is always almosl exactly proportional to the ratio of free acid to Bait, and is equal, in very close approximation, to the product of this ratio by the ionization constant of the acid. That is to say, repre- senting free acid by HA and sail by BA, + whence, if Jc = (H) HA BA From this relationship, therefore, follows the conclusion, fully established by experiment, that whenever in such a solution the excess of acid, HA, is chemically equivalent to the quantity of salt, BA, the hydrogen ion concen- tration is almost exactly equal to the ioniza- tion constant of the acid. But the ionization constant of carbonic acid (first hydrogen atom) is 0.0000003. Hence in a solution containing exactly equivalent quantities of free earl ionic acid and, for example, sodium bicarbonate, the hydrogen ion concentration must be 0.0000003 N. Further, since HA _ (H) BA k 146 THE FITNESS OF THE ENVIRONMENT if the amount of acid be ten times the amount of salt (ll^j = loY the hydrogen ion concentra- tion must be 0.000003 N, and if the reverse be the case ( =f = — ) the value must be VBA 10/ 0.00000003 N. The range of variation of concentration of hydrogen ions in the usual solutions of the chemical laboratory considerably surpasses the limits 1.0 N and 0.00000000000001 N. In comparison with such enormous differences those between 0.000003 N and 0.00000003 N are J_ . 1 100 * 100,000,000,000,000, Hence ordinarily it is quite accurate enough to speak of any solution containing both free carbonic acid and a bicarbonate, when the disparity between the concentrations of the two substances is not very great, as of constant neutral reaction. For, obviously, the neutral point, which at a temperature of 25° amounts to a concentration of hydrogen and hydroxyl ions 0.0000001 N, falls well within the narrow range of reaction of such solutions, being characterized by a ratio of carbonic acid to bicarbonate of about 1:3. Thus carbonic acid, like the almost equally weak acids sulphuretted hydrogen and phos- almost negligible (-J- : nOO.OOo) CARBONIC ACID 1 fl phoric acid (after its first hydrogen baa been neutralized by base), has the remarkable property of preserving a neutral reaction whenever it exists in solution will) its sails, provided there be an execs, of acid. All acids whose strength is even a little cither greater or less than carbonic acid lack the property.1 This characteristic of carbonic acid is of the utmost significance, first by regulating one of the most fundamental of physico- chemical conditions, and secondly bv niv- serving throughout nature the characteristic chemical inactivity of water, which disappears whenever the reaction becomes either appre- ciably acid or appreciably alkaline. Almost the only case of important geological action due to acidity or alkalinity of water is the action of fresh water, containing carbonic acid itself, to weather the rocks. This process is, however, self-limited, for the dissolved material forms bicarbonates, and thus at once pro- vides permanently inactive balanced solutions. It is impossible to understand the efficiency with which neutrality is preserved by carbonic acid, without the actual discussion of a par- ticular case. Let us therefore consider a solu- 1 Henderson, "The Relation between the Strength! of Acids and their Capacity to Preserve Neutrality,*' American Journal of Physiology, XXI, 173, 1908. H. C. State College 148 THE FITNESS OF THE ENVIRONMENT tion of 1 kilogram of carbon dioxide in 100 liters of water, to which sodium hydrate is being added. At the beginning of the experi- ment the hydrogen ion concentration will be approximately 0.0001 N, almost exactly one thousand times that at neutrality, and the hydroxyl ion concentration 0.0000000001 N, one one-thousandth that at neutrality. If, now, the sodium hydrate be added to the solution in successive portions, the change of reaction will be as indicated in the following table: — Amount of (H) (OH) Intensity compared with a Neutral Solution of NaOH Acidity Alkalinity Grams 0 0.0001 N 0.0000000001 N 1000 0.001 50 0.0000052 0.000000002 52 0.02 100 0.0000024- 0.000000004 24 0.04 150 0.0000015 0.000000006 15 0.06 200 0.0000011 0.000000009 11 0.09 250 0.0000008 0.000000012 8 0.12 300 0.0000006 0.000000017 8 0.17 350 0.0000005 0.000000020 5 0.20 400 0.0000004 0.000000025 4 0.25 450 0.0000003 0.000000033 3 0.33 500 0.00000025 0.00000004 2.5 0.4 550 0.00000020 0.00000005 2.0 0.5 600 0.00000015 0.00000007 1.5 0.7 650 0.00000012 0.00000008 1.2 0.8 700 0.00000009 0.00000011 0.9 1.1 750 0.00000006 0.00000017 0.6 1.7 800 0.00000004 0.00000025 0.4 2.5 850 0.00000002 0.0000005 0.2 5.0 CARBONIC ACID 149 From the tabic4 it appears that the first 50 grams of alkali reduce the hydrogen ion concentration to but 50 times that of neutral solutions, and 200 grams of alkali have made it only about 10 times that of pure water, in spite of the fact that there are more than 750 grams of free carbonic acid still present in the solution. So much acidity can at once be obtained by dissolving merely 0.004 gram of hydrochloric acid in 100 liters of water. Thereafter neither acidity nor alkalinity sur- passes this intensity until 450 grams more of sodium hydrate have been added to the solu- tion. Yet in pure water 0.005 gram of sodium hydrate would make the reaction more alka- line than that. Such is the case when the equilibrium is homogeneous, i.e. in an isolated solution. But when, in similar cases, an atmosphere containing carbon dioxide is present, the con- ditions are still more striking. Suppose, for example, a solution of 100 liters containing 1 kilogram of sodium bicarbonate in equilib- rium with an atmosphere containing 1 gram of carbon dioxide per liter. Let hydrochloric acid be added in successive small portions to the solution. Further let the solution be constantly stirred and shaken, and let the experiment be conducted slowly, so that there 150 THE FITNESS OF THE ENVIRONMENT shall always be equilibrium between the car- bonic acid in the solution and in the atmos- phere. Further, let the temperature be such that the absorption coefficient of carbon dioxide shall be 1.000. Then the successive states of the solution will be approximately as recorded in the following table: — HCl H2CO3: Added NaHCOj Grams 0 2.27:11.9 10 2.27: 11.5 50 2.27 : 10.0 100 2.27: 8.2 150 2.27: 6.3 200 2.27: 4.4 250 2.27: 2.6 300 2.27:0.68 310 2.27 : 0.31 318 00 320 330 (H) 000000057 000000059 000000068 000000083 000000108 000000154 00000026 0000010 0000022 00026 00045 0027 N Relative (OH) Acidity 0.000000176 N 0.57 0.000000170 0.59 0.000000147 0.68 0.000000120 0.83 0.000000093 1.08 0.000000065 1.54 0.000000039 2.6 0.000000010 10 0.0000000045 22 0.00000000039 260 0.00000000022 450 0.000000000037 2700 Relative Alka- linity 1.76 1.70 1.47 1.20 0.93 0.65 0.39 0.10 0.045 0.0039 0.0022 0.00037 From the beginning of the experiment until almost 250 grams of hydrochloric acid have been added, neither alkalinity nor acidity is double in intensity the values wThich obtain in a perfectly neutral solution. This amounts to a constancy of reaction which, until a few years ago, was scarcely known to the chemist at all. Such close approach to neutrality CARBONIC ACID 151 can be attained with pure water only after elaborate and very difficult purification, yet in the presence of carbonic acid it is the natural condition. Of course the case above dis- cussed is an extreme one. In nature the con- centrations of dissolved substances arc less, the mixing less efficient, and the variations of reaction a little greater. There is also, in nature, likely to be a considerable excess of bicarbonate over free carbonic acid present. The cause of the greater constancy of re- action in the case of the heterogeneous equi- librium is simple enough. At the beginning of the experiment the free carbonic acid of the solution is exactly in equilibrium with that of the atmosphere. Accordingly, when hydro- chloric acid is poured in and reacts with so- dium bicarbonate to form sodium chloride and more carbonic acid, NaHC03 + HC1 - NaCl + H20 + C02 every bit of the latter escapes to the atmos- phere, and the total amount of acid is just what it was before. But a certain amount of carbonic acid has been replaced by the equiv- alent amount of hvdrochloric acid. This latter substance, however, is wholly in union with sodium, as its salt. Thus the addition of the strong acid diminishes the amount of 152 THE FITNESS OF THE ENVIRONMENT alkaline salt (sodium bicarbonate), but does not increase the amount of free acid. Not until the bicarbonate is entirely decomposed (318 grams HC1) does the hydrochloric acid begin to exert its own action as an acid, and then 2 grams cause about as much rise in acidity as 318 grams have previously caused, or nineteen times the rise effected by the first 300 grams, or about two hundred times the rise caused by one hundred times the quantity of hydrochloric acid at the beginning of the experiment. These statements all rest upon facts which have been accurately established by experi- ment and are brought forward in company with the theoretical treatment, based upon the Mass Law, only for the sake of complete- ness of statement and because brief exposition is otherwise scarcely possible. The extraor- dinary capacity of carbonic acid to preserve neutrality in aqueous solution, which is ex- plained by its strength and solubility in water, is a well-established experimental fact, and no other known substance shares this power.1 Hydrogen sulphide might perhaps be thought of as an exception. But while its solubility 1 Henderson, "The Theory of Neutrality Regulation in the Animal Organism," American Journal of Physiology, XXI, 427, 1908. CARBONIC ACID \S$ in water is not favorable for the best resnlu its instability is a fatal obstacle. Such are the physico-chemical facts re- garding neutrality regulation in heterogeneous systems by means of carbonic acid and bicar- bonates, and, though the exposition is difficult, it has seemed necessary to make them clear. For there is, I believe, except in celestial me- chanics, no other case of such accuraev in a natural regulation of the environment. More- over, the chemist has discovered no means of rivaling the efficiency and delicacy of adjust- ment of the process. Finallv, acidity and alkalinity surpass all other conditions, even temperature and concentration of reacting substances, in the influence which they exert upon many chemical processes.1 Almost wholly through this mechanism the oceans are always nearly neutral. Chiefly with its aid protoplasm and blood possess an unvarying reaction. Quite recently the con- centration of hydrogen ions in the ocean has been very carefully studied by Palitzsch,1 1 Of all catalytic agents these ions are by far the moat important. In their influence upon the stability of collo* systems they are also unapproached by other Bubstani 2"Etant donne* que l'eau n seulemenl elk V i entoure de ses flots, mais qu'elle traverse leura branchiei el impregne en partie Lea corp dea invertebrfe, il seinhle . 154 THE FITNESS OF THE ENVIRONMENT who finds very great constancy, the extreme variations (with the exception of the Black Sea) being 0.000000011 N to 0.0000000045 N. Allowing for the change of the ionization constant of water with the temperature, and the fact that in such systems the hydrogen ionization is nearly independent of the tem- perature, these values roughly correspond to hydroxyl ion concentrations 0.000002 N and 0.000005 N respectively at the lower temper- atures and slightly higher values at the higher temperatures of sea water. This is a sufficient excess of hydroxyl ions properly to be termed faint alkalinity, though it amounts to but about 0.00005 gram per liter, or 0.000005 per cent. It is in part due to the fact that the quantity of carbonic acid in the air is now very small, while in the ocean the concentra- tion of bicarbonates is great. Indeed, the ocean has unquestionably grown alkaline; justifie de la placer dans la meme categorie que les autres- liquides physiologiques. Des determinations exposes dans ce qui precede il ressort que, de meme que ces liquides, l'eau de mer est douee d'une grande capacite de regler sa concentra- tion en ions hydrogene, bien que cette capacite soit moins prononcee que celle constatee par example dans le sang." — S. Palitzsch, "Sur le mesurage et la grandeur de la concen- tration en ions hydrogene de l'eau salee," Comptes-rendus des travaux du Laboratoire de Carlsberg, lOme Volume, Ire Livraison, 1911, p. 93. CARBONIC ACID 155 for its origin must have been acid from the presence of carbonic acid unbalanced by base. The present ratio between bicarbonates and carbonic acid in sea water has not been accu- rately estimated, but it is perhaps 50 : 1 or 100:1. The conditions are complicated by flora and fauna, and such influences have not yet been determined. Turning to the acid-base equilibrium of blood and protoplasm, we encounter a subject which is better understood, though not more significant in the present discussion. The alkalinity of the blood is one of the familiar subjects of physiological investigation, and its clinical importance has long been clear. Not until the introduction of the ionization hypothesis, however, was it possible to explain the conditions. The outcome of these studies has been to assign to the equilibrium between carbonic acid and bicarbonates a first place in the regulation of the reaction of blood;1 and since such substances are invariably constitu- ents of all protoplasm, to make evident the universal biological importance of this equi- 1 Friedenthal, "Archiv fur Physiologic, Verhandlungen der Physiologischen Gesellschaft Berlin," May 8, 1908. Hen- derson, American Journal of Physiology, XV, 457, 190G, "Ergebnisse der Physiologie," VIII, 454-345, 1909 (the last a review of the equilibrium between acids and bases in the organism). 156 THE FITNESS OF THE ENVIRONMENT librium. The process is complicated by the intervention of all the other acids and bases of the body. Of these, however, only phos- phates, and in lesser degree proteins, are im- portant. Thus it is certain that in the one universal chemical equilibrium of protoplasm which has thus far been defined and quantita- tively described the carbonates take a prin- cipal part. It is not possible to explain the significance of carbonic acid in this physiological process as chiefly an adaptation ; for natural selection can have nothing to do with the occurrence of carbonic acid in the living organism, or, pre- sumably, with the nature of the original living things upon the earth.1 The presence of carbonic acid is inevitable, and whatever the first forms of terrestrial life may have been, certain it is that carbonic acid was one of the constituent substances. From that day to this it has steadily fulfilled the function of regulating the reaction of protoplasm, and of body tissues and fluids. The recent studies of Hasselbach and Lunds- gaard2 indicate that the hydrogen ion concen- tration of normal blood at body temperature 1 It was this obvious fact which originally led me to a re- consideration of fitness. 2 " Biochemische Zeitschrift, Vol. 38, p. 77, 1912. CARBONIC ACID 157 is about 0.000000044 N. This value is sub- ject to constant slight variations, diminish- ing as the blood passes through the lungs, increasing during the greater circulation; but variations of this kind arc certainly very slight in the healthy organism. The value corresponds to a concentration of bicarbonatcs about ten times as great as that of free car- bonic acid. Together the acid and its salts make up the larger part of all the carbonic acid, and a very considerable fraction of all the dissolved molecules of the blood. In disease, especially diabetic coma, the hydro- gen ion concentration may rise to 0.0000001 N, or perhaps higher ; when acid is injected into the blood the value may be greater still, but death speedily ensues, and it is certainly im- possible during life that there should be any considerable permanent variation. Increase in acidity of the blood can occur only in association with marked diminution in the concentration of bicarbonates, which may fall to less than one third of their normal amount, greatly impoverishing the blood in respect to carbonic acid, and interfering with its excretion. This is due to the fact that the amount of free carbonic acid in the blood is under the independent control of the respira- tory center, and when acids decompose bicar- 158 THE FITNESS OF THE ENVIRONMENT bonates the carbonic acid which is liberated escapes through the lungs into the air. This action is analogous to the simple heterogeneous equilibrium explained above, and calls for no special discussion. The importance of this physiological equi librium is to be sought in part in one of the primary characteristics of the metabolic process, — the chiefly oxidative formation of acid. In the main the foodstuffs are neutral substances, but their principal end products, except water, are almost exclusively acid compounds, — carbonic acid, phosphoric acid, sulphuric acid, uric acid. Moreover, there is a well marked tendency, at least in man and the vertebrates, under very many pathological conditions, to form other acids, such as lactic acid, /3-oxy butyric acid, acetoacetic acid, etc. This tendency toward acidity of reaction and the accumulation of acid in the body is one of the inevitable characteristics of metabo- lism ; the constant resistance of the organism one of the fundamental regulatory processes. Now it comes about through the carbonate equilibrium that the stronger acids, as soon as they are formed, and wherever they are formed, normally find an ample supply of bicarbonates at their disposal, and accordingly react as follows : — CARBONIC ACID 159 HA + NaHC03 = NaA + II2C03 This reaction corresponds exactly to the simple case first discussed, and so causes an inconsiderable change in the concentration of ionized hydrogen. The free carbonic acid then passes out through the lungs, and the salt is excreted in the urine. Other processes are involved, including a device for final re- tention of a part of the alkali which has neutral- ized acid,1 but in the whole complex function nothing is more important than the simple reaction written above. The hydrogen ion concentration exerts a marked influence upon the rate of progress of chemical reactions. Thus, for example, the so-called inversion of cane sugar by a proc- ess of hydrolytic cleavage into glucose and fructose, C12H22OH + H20 = C6H1206 + C6H1206, is commonly accomplished by warming a solu- tion of sugar to which a little acid has been added. It was shown by the classical investi- gation of Wilhelmi that the velocity of this process depends upon the strength of the acid, or, according to the modern view, upon 1 Henderson, Journal of Biological Chemistry, IX, 403, 1911. 160 THE FITNESS OF THE ENVIRONMENT the concentration of hydrogen ions. Indeed, these ions are the effective agents in the process. Reactions of this type, in which carbohy- drates, fats, proteins, and other substances take part, make up a very large, if not the largest fraction of all the processes of metabo- lism, and there can be no doubt that for their regulation very accurate adjustment of acidity and alkalinity is essential. In the body, to be sure, such reactions are under the control of enzymes, but the concentration of hydrogen and hydroxyl ions is not less, rather more important for that reason. Beside retaining their direct influence upon the reaction, the ions also exert an influence upon the enzymes themselves. Moreover, not only enzymes, but almost all colloidal bodies, especially such unstable structures as the colloids of proto- plasm, are profoundly affected by changes of reaction, and for the preservation of their stability they also are absolutely dependent upon constancy of acidity and alkalinity. Finally, it is to be noted that glucose, which is the principal source of energy in physiolog- ical processes, is very unstable in even faintly alkaline solutions, and that its stability varies in most marked degree with the slightest change in the concentration of hydroxyl CARBONIC ACID 1G1 ions.1 Further, there is also reason to be- lieve that the concentration of hydrogen and hydroxyl ions may sometimes be concerned in adjustments of great magnitude, which involve the distribution of water between the tissues and the body fluids, or at least be- tween red blood corpuscles and the plasma. - Recently, for example, a theory has been sug- gested which seeks to account for certain forms of oedema as the result of local or general increase in the acidity of the organism.3 In any case, whatever the fate of these latter 1 Henderson, Journal of Biological Chemistry, X, 3, 1911. 2 The changing distribution of material between red blood corpuscles and plasma as the tension of carbonic acid changes has, since its discovery by Zuntz, been investigated by a series of physiologists. (See Spiro and Henderson, Biochemische Zeitschrift, 15, 114, 1908.) In each complete cycle of the circulation there occurs a complete cycle of changes between the constituents of the corpuscles and of the plasma. As the blood passes through the tissues and receives carbonic acid the volume of the red corpuscles increases from tin- entrance of water coming from the plasma; chlorine simul- taneously passes in the same direction; and various other changes occur. In the lungs, with the escape of carbonic acid, the process is reversed. There seems t<» be DO doubt that this process is largely dependent upon changes in re- action accompanying changes in the concentration of carbonic acid. 3 M. H. Fischer, "(Edema." New York, 1910. It is impos- sible to judge of the correctness of many of the views set forth in this work. At present they appear to be in part ill- founded. M 162 THE FITNESS OF THE ENVIRONMENT views may be, the common physiological processes, and the structural conditions which depend for their integrity upon constancy of the concentration of hydrogen and hydroxyl ions are certainly manifold. y The principal conditions and processes, both inorganic and organic, which rest upon the acid nature of carbonic acid and its char- acteristic distribution between the atmos- phere and aqueous solutions have now been indicated. In their origin at least they are nowise due to the agency of organic evolution. Yet directly, because of the nature of carbon dioxide as a gas, because of its solubility in water, and on account of the precise degree of its weakness as an acid, they possess the highest possible efficiency. This conclusion might be established with rigorous accuracy by means of a mathematical analysis, but the above discussion is sufficient for the present purpose.1 In this manner carbonic acid shows itself in its physico-chemical traits variously fitted for the organic mechanism. Less various, to be sure, and less obvious than those of water, such fitnesses as it does possess are quite as 1 Henderson, American Journal of Physiology, XXI, 173, 1908. CARBONIC ACID 163 genuine. But they arc dependent upon water; secondary in their nature; resting upon solu- bility and ionization; upon interactions be- tween the two substances. So they lead us to a consideration of the most intricate of all mutual relations, to a purely chemical study of the compounds of carbon, hydrogen, and oxygen, into which water and carbonic acid can be transformed. CHAPTER V THE OCEAN AT every stage of our inquiry we have seen that the unique properties of water and carbonic acid contribute vitally important characteristics to the ocean. Such conclusions accord with the ordinary experiences of life, and they gain in significance from the un- doubted fact that organic beings first existed, and for a very long time existed only in the waters. On this account it will be well to pause before attacking the problems of organic chemistry, and, in somewhat greater detail, to examine the ocean in its relation to the inhabitants of the globe. Thus we shall be able more clearly to perceive the manner in which, in one most important instance, the properties of water and carbonic acid operate to fit the world for life. THE REGULATION OF PHYSICO-CHEMICAL CONDITIONS The most striking of all the ocean's qualities is its constancy. No doubt since its origin 164 THE OCEAN 1G5 it has grown colder and more saline, and has changed its reaction from faintly acid to faintly alkaline. But a million years are little in such great slow processes, and no living thing has ever experienced appreciable change in any one of them. Modern research in oceanography has de- tected surprisingly little variation in the tem- perature of the ocean.1 The temperature of surface water depends upon the climatic character of the locality, but it is subject to far less variation than the temperature of the atmosphere above it, and is higher than the latter. The accompanying tables 2 indicate the nature of some of the variations in the temperature of sea water. Annual Ranges of Temperatures of Ocean Water and of the alr over land 0° 10° 20° 30° 40° 50° 2.3 2.4 3.6 5.9 7.5 5.6 — 3.3 7.2 10.2 14.0 25.4 1 Nearly all the facts contained in the present chapter have been drawn from the following works: S. Giinther, "Hand- buch der Geophysik " ; Arrhenius, "Kosmische Physik " ; and Hann's "Climatology," translated by Ward. 2 See Hann's "Climatology," translated by Ward, p. 135. 166 THE FITNESS OF THE ENVIRONMENT Water and Air Temperatures at Lesina Winter Spring Summer Autumn Year Range Surface water . Water at 10 meters 9.2° 13.5 13.9 14.8° 15.0 14.7 24.4° 22.0 20.3 17.9° 19.5 18.4 1G.6° 17.5 1G.8 15.2° 8.5 6.4 According to Schott the variation of surface temperature of the open ocean over the whole globe is never less than 1° nor more than 15°. Such variations are slight on the equator, larger in the region of the trade winds, less again in the northern and southern seas.1 The surface temperature extends only a short distance into the water, thus constituting a warm surface layer. It has indeed been known since the time of Aristotle that the depths of the sea are cold. In the open ocean the temperature increases steadily as the depth increases, but more rapidly near the surface, more slowly at greater depths. The bottom water at great depth varies from a temperature of 2° in the tropics to about — 2° in the polar waters. Such low temperature of the depths of the tropical oceans is almost certainly due to the inflow of cold water from high latitudes. 1 Arrhenius, I.e. THE OCEAN 167 In the Atlantic the temperature varies ap- proximately as follows with increasing depth: — Depth in Meters Temperature Centigrade 0 19° 500 16 1000 9 1500 4 2000 3 2500 2.5 3000 2 In the Mediterranean, where no cold cur- rent flows in at the bottom of the ocean, the temperature sinks rapidly to 11° at a short distance below the surface, and thereafter re- mains constant. This is due to the fact that in winter the surface water is cooled to that temperature and sinks, remaining then pro- tected from the summer heat by the warmer layer of less density above. The slight range of ocean temperatures, whether with changing seasons, with chang- ing depth, or with changing latitude, depends primarily upon the latent heat of water, especially its heat of vaporization, and upon the very high freezing point, as already ex- plained in the discussion of these physical properties. Far more constant than the temperature is the alkalinity of sea water. It has been stated above that the extreme variation in concent ra- 168 THE FITNESS OF THE ENVIRONMENT tion of ionized hydrogen is from 0.00000001 IN to 0.0000000045 N. In his studies during a voyage of five months in the summer season of 1910 of the Danish steamship Thor, Palitzsch made the following observations: l surface water from the Skager-Raek, from the south- ern portions of the North Sea, and from the west of the Baltic ranged from about 0.000000010 N to about 0.000000009 N ; in the North Sea the surface water varied from 0.0000000083 N to 0.0000000074 N; at the lati- tude of Murray Firth, twenty miles from the coast, the values were between 0.0000000071 N and 0.0000000066 N. In the Atlantic the surface water corresponded in the most north- ern portions to that of the North Sea, while in the Bay of Biscay and along the coast of Portugal the values were 0.0000000056 N, corresponding to slightly increased alkalinity, especially if account be taken of the rising temperature. In general the waters of the Mediterranean corresponded to those of the coast of Portugal. But from the Sea of Mar- mora, the Bosphorus, and the Black Sea samples were obtained which gave the value 0.0000000045 N. In general, as the depth increased, the Palitzsch, " Comptes-rendus des travaux du Laboratoire dc Carlsberg" lOme Volume, p. 85, 1911. THE OCEAN 169 hydrogen ion concentration increased, and the alkalinity accordingly diminished. Depth in + (H) x 100,000,000 Meters Mediterranean Atlantic North Sea Black Sea 0 0.59 0.G0 0.74 0.46 10 0.56 25 0.65 50 0.59 0.66 0.71 75 0.93 85 1.03 100 0.62 0.74 0.81 1.38 200 0.65 0.83 2.1 300 2.8 400 0.65 0.91 0.93 3.0 600 0.98 700 1.05 800 0.68 0.98 1000 0.72 0.98 1200 0.72 1.05 1500 0.76 1.13 2000 0.81 1.13 2500 0.85 3200 0.85 The only variation from the truly remark- able constancy of reaction of the ocean, so far as we know, is in the case of the Black Sea. But this sea, at depths below 180 meters, con- tains sulphurous acid, which undoubtedly accounts for the slight diminution of alkalin- ity recorded in the table. This last obser- 170 THE FITNESS OF THE ENVIRONMENT vation also provides a striking example of the efficiency with which the reaction of sea water is maintained. In spite of the unusual circumstances here the variation is inconsider- able. The only known important factor which operates to establish and to preserve the re- action of sea water is the carbonate equilib- rium. There is one consideration which must be especially noted before passing on. The most obvious effect of slight changes of temperature, and of slight changes of alkalinity as well, is upon the velocity of chemical reactions. In this respect the effect of hydroxyl ions is likely to be in proportion to their concentra- tion,1 and the effect of temperature is usually such that a change of about ten degrees doubles the velocity of the reaction.2 Hence ordinary chemical reactions will progress about eight times as fast in the hottest as in the coldest ocean waters, and about seven times as fast in the most alkaline as in the least alkaline parts of the ocean. But in the case of any organism inhabiting a particular locality such changes in reaction velocity will be scarcely 1 The chief actions of hydroxyl ions are catalytic, and, as in the case of the catalysis of esterification, the effect is pro- portional to the concentration of the hydroxyl ions. 2 This observation is due to van't Hoff. THE OCEAN 171 appreciable. Accordingly, the regulation is physiologically adequate. The concentration of sea water is another nearly constant characteristic, though gener- ally speaking the salinity is somewhat greater on the high seas than near the coasts, where fresh water is constantly diluting the salt water, and there are some other causes which produce slight variations. The average salt content is about 3.45 per cent. The quan- tities of the more important constituents, calculated from Dittmar's data,1 are as fol- lows : — Sodium, Na . Magnesium, Mg Calcium, Ca . Potassium, K Chlorine, CI . Sulphate, SO< Carbonate, CO3 Bromine, Br . Per Cent 1.049 0.130 0.041 0.038 1.89G 0.263 0.007 0.006 Relative Amount 30.59 3.79 1.20 1.11 55.27 7.66 0.21 0.19 Most of the other numerous constituents are present in very small quantities. For in- stance, in each metric ton of sea water there 1 Dittmar, Report of Voyage of the Challenger, 1884, p. 203. The original data were calculated upon the erroneous assumption that the various salts exist in solution independ- 172 THE FITNESS OF THE ENVIRONMENT are dissolved about 0.019 gram of silver and about 0.006 gram of gold, amounts which correspond to 0.0000019 per cent and 0.0000006 per cent, respectively. It has been calculated that 166,000,000 years have been required for the streams to carry into the sea the sodium chloride which is now present. The calcium carbonate of the rivers wTould however suffice to supply the present amount of that substance in 500,- 000 years. Accordingly, the present store of the latter substance represents but a very small fraction of what has passed through the ocean, and as a result of the intervention of life has finally been deposited as sedimentary limestone. Since the formation of the ocean, if present conditions correspond with the past, water must have carried to the dwellers of ently of one another. Thus the composition of sea water is stated as follows : — Sodium chloride, NaCl Magnesium chloride, MgCU . Magnesium sulphate, MgS04 Calcium sulphate, CaS04 Potassium sulphate, K2S04 . Calcium carbonate, CaC03 . Magnesium bromide, MgBr2 Relative Amount 77.758 10.878 4.737 3.600 2.465 0.345 0.217 THE OCEAN 173 the sea not less than 300,000,000,000,000,000 tons of calcium carbonate, which they have temporarily utilized as structural material. Whether this estimate be correct or not, the process is certainly the cause of the most considerable change wrought by life upon the face of the earth. The relative quantities of the several saline constituents of the ocean are hardly at all subject to variation. Chlorine, for example, makes up never less than 55.21 per cent and never more than 55.34 per cent of all the dis- solved inorganic substances, so that the total salinity may be readily estimated with consid- erable accuracy by titration of the chlorides. Such constancy is due to the elaborate mixing of the waters resulting from ocean currents. There can be no doubt, however, that the relative amounts of the different acids and bases have slowly but steadily changed during the progress of geological evolution. Many substances, like calcium carbonate, have been steadily removed, a few, like sodium chloride, have steadily accumulated without loss. The total salinity of the ocean, as stated above, is subject to slight variation. Along the North American coast, in the polar cur- rent, upon the coast of Norway, and toward the south of South America, the concentration 174 THE FITNESS OF THE ENVIRONMENT of the Atlantic is 3.2 per cent to 3.3 per cent. Areas where the concentration ranges from 3.3 per cent to 3.4 per cent are still more extensive. The greater part of the North Atlantic ranges in concentration from 3.5 per cent to 3.6 per cent. In general there is a region of maximum concentration between the Equator and the Pole. The Mediterranean, the Red Sea, and other similar bodies of water possess a somewhat higher salt concentration, dependent upon excessive evaporation and the absence of great currents ; but only in exceptional cases and small isolated bodies of water does the concentration rise above 4.1 per cent. The salinity of ocean water varies also with the depth. In seas where there is a great influx of fresh water the surface is less con- centrated than the depths; in seas where there is much evaporation the surface is more concentrated than the depths. In the latter case the higher temperature of the surface causes expansion of the more concentrated water, and enables it to remain above. When these two influences of dilution and evapora- tion are combined, they may bring about a yearly variation of salinity. Such variations of the environment are important to animal life, slight though they may be. Thus the THE OCEAN 175 herring in their migrations keep to a water whose concentration ranges from 3.2 per cent to 3.3 per cent. From the constancy of the relative pro- portion of the salts in sea water it follows that every such constituent is subject to no greater variations than the sum of all. Interesting recent experiments have shown this fact to be of vital consequence to living organisms. Thus a host of experiments of Loeb and his pupils, and of others, have demonstrated re- markable toxicity in the action of pure salts, physiologically antagonistic actions of vari- ous pairs of salts, and peculiar advantages of media containing a variety of salts in definite relative amounts.1 Of all such balanced solu- tions sea water is by far the best, a condition which is almost certainly due to the processes of organic evolution. Herbst 2 has shown that the development of the fertilized eggs of sea urchins can only take place in the presence of the chlorides, sulphates, and car- bonates of sodium, potassium, calcium, and magnesium, and in a faintly alkaline reaction. Every one of these substances is essential, 1 See the article by Loeb in Oppenheimer's "Handbuch der Biochemie." 2 Herbst, Archiv.fiir Ejitwickelungsmechanik, 5, 050, 1897; 7, 486, 1898; 11, 617, 1901; 17, 300, 1904. 176 THE FITNESS OF THE ENVIRONMENT except that in a measure potassium may be replaced by rubidium and caesium, and chlo- rine by bromine. Moreover the relative con- centrations are of the highest importance. Thus it has become clear that the remark- able relative and absolute constancy of the chemical composition of sea water is biolog- ically far more important than formerly could be surmised. This characteristic of the ocean undoubtedly fits it for living things as they exist. It is further to be noted that the salinity of sea water is proportional to its osmotic pres- sure. This important property also is there- fore nearly constant. When a solution is inclosed in a membrane, a bladder, for example, and the latter is im- mersed in water, both water and dissolved substance pass through the wall of the mem- brane. Ordinarily, however, the water will move much more rapidly than the dissolved substance, hence the volume of the solution will increase, and hydrostatic pressure will be established. If a well-supported mem- brane of cupric ferrocyanide be substituted for the bladder, the process will be modified, in that water alone, not the dissolved sub- stance, can pass through the membrane, which is accordingly termed semipermeable. THE OCEAN 177 Under these circumstances the pressure, called osmotic pressure by van't Hoff, may be very great. According to the theories of van't Hoff and Arrhenius this pressure is, in the case of dilute solutions, proportional to the total number of particles (molecules plus ions) which are present in solution. In its magni- tude and the laws governing its variation such pressure corresponds exactly to gaseous pres- sure. In fact the theory of solution consists primarily in the extension of the laws of Boyle and Gay-Lussac, of the hypothesis of Avo- gadro, and of the manifold theoretical develop- ments which have been based upon them, to solutions. The great force of osmotic pres- sure always comes into action when solutions are in contact with permeable or semiper- meable membranes. It must therefore al- ways be reckoned with in physiology. The biological importance of the constancy of the osmotic pressure of sea water is strikingly exemplified by the precision with which every higher vertebrate preserves constant the os- motic pressure of its own body fluids, all at about seven or eight atmospheres. It may readily be shown that the osmotic pressure of a solution is proportional to the depression of its freezing point, and accord- 178 THE FITNESS OF THE ENVIRONMENT ingly osmotic pressure is commonly estimated with the help of this relationship. In the following table the facts regarding blood sera of certain animals are collected.1 Depression of the Freezing Point of Blood Serum Degree Man 0.526 Cow 0.585 Horse 0.564 Pig 0.615 Rabbit 0.592 Dog 0.571 Cat 0.638 Sheep 0.619 In man the freezing point depression of the blood is, under ordinary circumstances, practi- cally constant, and there can be no doubt that such is the case for all highest organisms. Accordingly the differences in the above table may be taken to indicate slight constant dif- ferences between the different species. The marine animals, except a few of the vertebrates, are adjusted in their osmotic pressures to the water which surrounds them. As in the following table so in another lo- cality where the freezing point of the water was —1.9° the bodv fluids of the animal were again found to agree with it. It is therefore evident that constancy of osmotic 1 Hober, " Physikalische Chemie der Zelle und der Gewebe." 1 a < M u •< - - - E « THE OCEAN 179 pressure is for marine animals a matter of real moment. Depressions of the Freezing Point Degrees Calentratc, Alcyonium palmafum. 2.196 Echinodcrm, Astcropcctcn aurantiacus 2.312 Echinodcrm, Flolotluiriu lubulosa 2.315 Worm, Sipuneuhu nudus 2.31 Crustacean, Maja squinada 2.3G Crustacean, Homarus vulgaris 2.29 Cephalopod, Octopus macropus 2.24 Selachian, Torpedo marmorata 2.26 Selachian, Mustelus vulgaris 2.36 Selachian, Trygon violacea 2.44 Teleost, Charax puntazzo 1.04 Teleost, Cerna gigas 1.035 Teleost, Crenilabrus pavo 0.74-0.76 Teleost, Box salpa 0.82-0.88 Reptile, Thalassochelys caretta 0.61 Sea Water 2.3 The great importance of osmotic pressure is also attested by many of the facts of phys- iology. The study of this subject has in- deed from its origin always been closely associated with the biological sciences, and it was in great part biological experiments and wholly experiments of biologists which were employed by van't Hoff in his development, on the basis of osmotic phenomena, of the theory and laws of dilute solutions. Absorption, secretion, excretion, and the movement of substances across membranes, 180 THE FITNESS OF THE ENVIRONMENT — to say nothing of the establishment of liquid currents within the body, — are all related to osmotic pressure. The forces in- volved in such processes are large, and osmotic phenomena assume a special importance wherever colloidal systems occur. It appears to be certain that osmotic pressure is now se- curely established as one of the fundamental factors in the physico-chemical organization of the living mechanism, and one of the constant conditions, like concentration of the several constituents, alkalinity, temperature, etc., whose preservation is of vital importance. n THE CIRCULATION OF WATER There are a number of causes which bring about ocean currents. In the tropics high temperature causes a far greater evaporation of water than can be offset by rainfall and the flow of rivers ; near the poles this relation is reversed. Hence water must steadily flow from high to low latitudes, there to evaporate and complete the cycle in the atmosphere and on the land. In polar regions the cold water sinks and penetrates along the bottom of the sea in great deep currents to the tropics. THE OCEAN 181 The surface currents of the ocean have a different origin, for they depend upon winds, especially trade winds, etc. Such continuous action of moving air upon water has been theoretically explained by Ilelmholtz and Zopperitz. Needless to say, in addition to these principal causes there are a great variety of lesser factors which assist in the formation and preservation of ocean currents. It is impossible here to undertake an analy- sis of the phenomenon, but certain it is that into the processes that constantly stir the ocean, beside the rotation of the earth, the eccentricity of its orbit, and the inclination of its axis, the thermal properties of water enter as fundamentally important factors. The magnitude and the extent of the movements which result from such influences are very considerable. The principal surface currents are oval in form, one in the North Pacific between 10° and 50° north latitude, one in the North Atlantic between 10° and 30° north latitude, one in the South Pacific between 5° and 45° south latitude, one in the South Atlantic between 0° and 40° south latitude, and one in the Indian Ocean between 0° and 40° south latitude. The greatest of these are the Pacific currents. In the far south is an Antarctic current flowing from 182 THE FITXESS OF THE ENVIRONMENT west to east ; in the north a current flows from east to west, from the Siberian coast to Northeast Greenland and thence along the east coast ; another flows from Baffin's Bay along the east coast of North America. Of all ocean currents, the Gulf Stream, a branch of the northern equatorial current, has been most carefully studied. Its maxi- mum velocity is 220 kilometers per day, greater therefore than that of the Rhine at Coblentz ; the mean about 134 kilometers a day. In the Straits of Yucatan the Gulf Stream carries 0.2 cubic kilometer (200,000,- 000 tons) per second. If all this water were to be cooled to the temperature of the polar ocean this would be equivalent to the trans- port of about 5,000,000,000,000,000 gram calories per second. The magnitude of this quantity, of course, depends upon the specific heat of water. In this manner vast quantities of water, carrying enormous stores of heat, are constantly in motion all over the globe. The result is that homogeneity of the ocean which has been discussed above, — constancy of concentra- tion, of composition, of temperature, of alka- linity, and of osmotic pressure. THE OCEAN 183 III THE OCEAN AS ENVIRONMENT There are, in accordance with our funda- mental postulates of the characteristics of life, two principal requirements of the living organism which an environment must ful- fill, — such a supply of matter and energy for food as may be suitable to a complex mechanism, and stability of conditions. After a general review of the chief character- istics of the ocean, it is therefore necessary to examine them more particularly in relation to such requirements. In so doing it must not be forgotten, however, that these characteristics of the ocean which we have just discussed are only in part due to the unique physical proper- ties of water which have been alreadv dis- cussed in Chapter III, and to those of carbonic acid which have been discussed in Chapter IV. In part they depend upon the mere magnitude of the sea, on the stability of the solar system and the consequent antiquity of the geological and meteorological processes, and upon a great variety of astronomical and geophysical conditions. However, we shall only a little extend the scope of our inquiry if we now consider water not only as an indi- 184 THE FITNESS OF THE ENVIRONMENT vidual chemical substance, that is to say ab- stractly, but also naturally, as automatic processes of cosmic and geological evolution have fashioned it into the principal constit- uent of the face of the globe. Primarily, at any rate, the outcome of such processes is dependent upon the inherent properties of water and upon the quantity of it which is present on the surface of the earth, and the subject is too important to be passed com- pletely by. Perhaps the first desideratum in an en- vironment as a source of food is mobility. Any organism which, like the lilies of the field, need not toil for its nourishment, is in most favorable conditions, and such conditions are the principal cause of the enormous wealth of vegetation upon the earth. Now the ocean, apart from the flora and fauna which inhabit it, is perfectly homogeneous ; hence its mobility brings to an organism all that it has to offer, and even sweeps along organic nourishment as well. In the ocean not only plants but many animals may remain motionless and, like the oyster, await the food that will surely be borne to them ; or they may float freely, relying on the mixing of the water to bring them into contact with their food. After mobility, richness and variety of THE OCEAN 185 environment are important. It is certainly impossible to imagine a medium more rich and varied in elementary constituents than sea water, unless it be sea water with still other sub- stances added to it. But, in the first place, there are few absent elements which might be added, and, in the second place, the addition of other substances would be likely to cause the escape of bodies which are present. At all events, the ocean is certainly more favor- able in these three respects than if it were anything else that could occur in the course of nature. Almost ideally mobile, rich, and varied, the sea is an almost perfect source of supply for a complex mechanism. To be sure there are great difficulties in extracting its constituents from sea water, and the efficiency of physiological processes is a factor essential to their utilization, but at least there stand materials ready for the mechanism which can employ them. The predominance of water, no doubt, forces that substance upon living beings as their chief constituent; in view of the fitness of water for the purpose that is in itself a favor- able circumstance. Otherwise the organism is left free to choose from all the common elements, and from some of the rare ones, what may be most suitable to its purposes at every 186 THE FITNESS OF THE ENVIRONMENT stage of the infinitely varied process of organic evolution. As a result we find here and there in the marine flora and fauna almost every element which the sea affords concentrated and put to use. After what has gone before it will not be necessary further to discuss the second great qualification of an environment — stability of conditions in the ocean. The principal physical conditions and chemical compounds therein are constant ; that is the whole case. But it is a case which cannot be bettered. Certainly nowhere else where life is possible, probably in no other place in the universe except another ocean, are so many conditions so stable and so enduring. The regulatory devices of our modern labo- ratories have not yet succeeded in rivaling the ocean. Singly, certain conditions, for example, temperature, alkalinity, and concen- tration, may be more accurately regulated by man, though on a small scale only ; but the regulation of all such properties together is not yet possible. The only known improve- ment upon the ocean is the body of a higher warm-blooded animal. Here, however, the processes of organic evolution have begun with the ocean, and in several respects merely perfected existing arrangements. THE OCEAN 187 This statement is far from fanciful. Not only do the body fluids of the lower forms of marine life correspond exactly with sea water in their composition, but there are at least strong indications that the fluids of the highest animals are really descended from sea water, from the sea water of an earlier epoch, to be sure, and they are not changed beyond recog- nition by the transformations of evolution.1 A comparison of the relative amounts of various saline constituents in sea water and in mammalian blood (roughly averaged from a variety of measurements in different species) will demonstrate this relationship. Composition of the Salts in Per Cent Sea Water Blood Serum Na 30.59 3.79 1.20 1.11 55.27 7.66 0.21 0.19 39 Mg Ca 0.4 1 K 2.7 CI 45 S04 C03 12 Br P,Os 0.4 1 See the interesting paper by Macallum, Transactions of the Royal Society of Canada, 1908, II, p. 145. 188 THE FITNESS OF THE ENVIRONMENT The gaps in the table do not indicate that substances are lacking, but merely that the amounts are small. In short, the same sub- stances are present in both cases, and in both cases sodium chloride largely predominates. The importance of carbonic acid in metabo- lism accounts for the large amount of sodium bicarbonate in the blood, and this raises the amounts of both sodium and carbonic acid. It is also to be noted that the regulatory processes in the ocean and in the organism are in one or two aspects similar, e.g. tempera- ture regulation by evaporation, and regula- tion of the alkalinity. Of course no impor- tance attaches to such resemblances, beyond the fact that both regulations are highly favorable, because of the special fitness of water in one case and of carbonic acid in the other. But it is at least worthy of mention that the regulation of the ocean in general bears a striking resemblance to a physiolog- ical regulatory process, although such physi- ological processes are supposed to be the result of organic evolution alone. Very much this same idea occurred to Palitzsch in the course of his investigation of the alkalinity of the ocean.1 The resemblance is more ob- vious still when the stability of all the more 1 See note above, p. 153. THE OCEAN 189 important physical conditions of the ocean is taken into account. Indeed, however difficult it may be to make out those subtle traits of physiological processes which account for their efficiency, their adaptability, and their exactness, I feel sure that no one who is thor- oughly conversant with the general char- acteristics of the life process can fail to see a rough counterpart in the means by which conditions in the ocean are regulated. It is certainly a salient, and hardly a mean- ingless fact that the processes of inorganic and organic evolution have a similar out- come in complex, exact, and almost ideally efficient activities. Is it not possible that in the case of the organic processes some have now and then been regarded as adaptations which in reality arose automatically and quite inevitably ? The existence of efficient regulation of the ocean, establishing its most important physico- chemical characteristics as constants, is of far greater importance in the sciences of nature, especially for living organisms, than could formerly have been guessed. Such natural processes were perhaps even necessary to make life possible in the birthplace of life. I cannot undertake to explain the very great importance which to-day the physical chemist 190 THE FITNESS OF THE ENVIRONMENT attaches to the regulation of the conditions of a chemical process. The only way to gain an idea of this is to examine a work on physi- cal chemistry. Certainly, however, nothing has lately arisen more essential to biology than the understanding of the influence of temperature, pressure, reaction, concentration, ionization, etc., upon all physico-chemical structures and changes, whether inorganic or vital. Thus the fitness of the ocean appears as an embodiment of the physical fitnesses of water and carbonic acid, resulting directly and in- evitably from these and other natural phenom- ena, and providing a lodgment for life and a medium for its earlier development upon the earth. No philosopher's or poet's fancy, no myth of a primitive people has ever exag- gerated the importance, the usefulness, and above all the marvelous beneficence of the ocean for the community of living things. CHAPTER VI THE CHEMISTRY OF THE THREE ELEMENTS ORGANIC CHEMISTRY A HUNDRED years ago the firm belief was held by all chemists that whatever substance is synthesized within the body of the living organism possesses special and pe- culiar characteristics of its own, which mark it off from all inorganic bodies, and divide chemistry into the two great and perfectly distinct departments of Organic Chemistry and Inorganic Chemistry. To be sure, even then many organic substances had been sep- arated from the organism, purified, and sub- jected to the usual experiments of the labora- tory, without at any stage manifesting unique properties. But, as Berzelius believed, a special vital force had presided over their formation, and this, therefore, he supposed to be impossible under any other circum- stances. 191 192 THE FITNESS OF THE ENVIRONMENT In the course of time, however, a long series of successful syntheses of undoubted constitu- ents of animals and plants, among which Woh- ler's preparation of urea in 1828 is the most famous, completely destroyed the old erro- neous assumption. The compounds of organic chemistry gradually came to be recognized as different from inorganic substances only in the special characteristics of the elements carbon, hydrogen, and oxygen when in chem- ical union with one another, just as the com- pounds of any other elements have their own specific characteristics. No other difference remains; every principle of chemical science applies to organic and inorganic substances alike; and accordingly life has been for- ever subjected to the general laws of chem- istry. As syntheses multiplied, the organic chem- ist found many fields for investigation where life was not concerned. The application of his new substances in the arts, as well as many fascinating theoretical problems, led him on, until, about the middle of the century or a little later, it became clear that organic sub- stances in the original sense are but a small part of his scope. His occupation had be- come the study of all the compounds of carbon, wherever and however they might CHEMISTRY 193 occur, and as a rule he had little to do with physiological or biological chemistry. Not that he was now ever disposed to distinguish between substances which happened to occur in living organisms and others ; for at length he had completely accepted the view that, apart possibly from a few complicated sub- stances like the proteins, such distinctions are thoroughly irrational. But the nature of the subject and the historical accidents of its development directed his attention in the main elsewThere. Nevertheless, the distinction between or- ganic chemistry as the science of all the com- pounds of carbon, and inorganic chemistry as the science of all other chemical compounds whatever has persisted, and not without sound reasons. In the course of the wonder- ful development of organic chemistry, which must ever be counted as one of the greatest achievements of the nineteenth century, enor- mous numbers of new chemical substances were discovered. In 1883 the number of carbon compounds had reached 20,000, in 1899, 74,000, and in 1902 it exceeded lOO^OO.1 1 See M. M. Richter, "Lexikon der Kohlenstoffverbindun- gen," Hamburg and Leipzig, 1900, continued in supplemen- tary volumes. This work catalogues all the compounds of carbon as they come to light. o 194 THE FITNESS OF THE ENVIRONMENT This is the sufficient practical ground for pre- serving organic chemistry as a separate sci- ence. The subject is so vast that in fact it is impossible to incorporate it with other departments of chemistry. Even the com- pounds of carbon and hydrogen alone are counted by hundreds, those of carbon, hydro- gen, and oxygen, by thousands, and the number of possible compounds of the three elements is almost unlimited. The mere number of organic compounds is, however, far from constituting the only dis- tinction between the two departments of descriptive chemistry. The unique variety of compounds containing carbon, hydrogen, and oxygen, and, in a small proportion of cases a few other elements besides, must obviously rest upon the nature of the elements them- selves, especially of course upon the nature of carbon, upon the properties which are pe- culiar to them and which mark them off from other elements, just as the properties of argon, of the metals of the alkalies, or of the halogens determine their own chemical be- havior. Moreover, such characteristics must and do contribute properties to the com- pounds of carbon which are theirs as a class, which distinguish them from the compounds of other elements in somewhat the same way CHEMISTRY 195 that the anatomical characteristics of one class of animals distinguish such a class from other classes of animals. In short, the carbon compounds are not unique merely because they are numerous; they are uniquely nu- merous because they are compounds of carbon with hydrogen, oxygen, and in some cases certain other elements. They possess, more- over, other less obvious class properties as well, though of these, it must be admitted, chemistry is even yet far from a clear under- standing. But unquestionably that is due to the incompleteness of the science, for the peculiar methods of organic chemistry are a sufficient guarantee of the existence of such class peculiarities.1 In our present investigation a study of the possibilities of chemical union between the elements carbon, hydrogen, and oxygen is of great importance, and accordingly we must now examine some of the results of synthetic organic chemistry. 1 See the introductory chapter to Meyer and Jacobson's "Lehrbuch der Organischen Chemic," Leipzig, 1907. 196 THE FITNESS OF THE ENVIRONMENT VALENCE The principal theoretical foundation of organic chemistry is the idea of valence. Let us consider the chemical formulas of a number of the simple compounds of hydrogen, e.g. HC1, H20, NH3, CH4, HI, HBr. It is evident that in such formulas a single atom of hydrogen is never represented as in union with more than one atom of another element. There are, however, cases where one atom of hydrogen is in union with a single other atom, e.g. HC1, HBr, HI ; or two atoms of hydro- gen may unite with a single other atom, e.g. H20 ; or three atoms of hydrogen with one other, e.g. NH3 ; or four hydrogens with one other, e.g. CH4. If the assumption be made that discrete bonds or forces take part in the union of atoms, hydrogen must possess but a single such bond or valence. Otherwise com- pounds of the type X — H — X, X — HOH CH2OH CHOH CHOH CH2OH CHOH II C II CHOH CH2OH i CH2OH CHoOH 1 COH II CH2 i C III COH CHOH 1 CH2OH COH li CHOH CH3 1 CH2 1 CHO CH3 1 CO 1 CH3 CH3 1 CH II CO CHO 1 CH II CH2 CO II C II CH2 CHO 1 c III CH CH3 CHO CHO CO II c II CO CHO 1 CO 1 CHO CH2 1 CHO CH II CO CO 1 CHO CH3 COOH COOH i COOH CH2 1 COOH CH II CH2 i c III CH CH2 1 COOH CH3 CHoOH CH2OH I CH3 1 CH2OH 1 CHO 1 CHOH 1 CHO CH2 1 CHO CO 1 CH3 COH II CO CH II CO COH II CH2 CHEMISTRY 205 CHO I CH II CHOH CHOH II C II CO CHO I C III COH CH2OH CH2OH CHoOH CHO CH2OH CHO I I I I I I CHOH CO COH COH CO CHOH I I II II I I CHO CH2OH CO CHOH CHO CHO CHO COH II CO CH3 CH2OH COOH COOH COOH CHoOH I I I I I I CHOH CH2 COH CH C CHOH I I II II III I COOH COOH CH2 COOH COOH CHOH COH COOH I COH II CHOH CHOH COOH CH3 CHO 1 COOH 1 COOH CO CH2 CH II CO CO COOH COOH COOH CH,OH CHO COOH I I I CO CHOH COH I I II COOH COOH C = O 206 THE FITNESS OF THE ENVIRONMENT This is relatively a simple case. As the number of carbon atoms in the molecule increases, the number of possible oxygen derivatives multiplies in a far more rapid progression than in the case of the simplest hydrocarbons, which has been stated above. Accordingly there can be no doubt that in addition to the many thousands now known, the existence of countless millions of com- pounds consisting of carbon, hydrogen, and oxygen alone is possible. In a large propor- tion of cases the only difficulties involved in their preparation are to obtain suitable start- ing materials, and the enormous labor of the process. There are, for instance, hundreds of thousands of possible hydroxyl derivatives alone of the paraffine hydrocarbons of the formula Ci4H30, but only one of these is now known.1 Yet all, or at least a vast majority, would unquestionably be stable bodies if once formed. Not less important than the number and variety of such substances is their diversity of physical and chemical characteristics. The following are, for example, individual chemi- cal compounds of at least moderate purity, made up of the three elements alone : al- cohol, formaldehyde, acetic acid, carbolic 1 Me thai, a constituent of spermaceti. CHEMISTRY 207 acid, oxalic acid, acetone, ether, lactic acid, sugar, cotton, glycerine, olive oil, camphor, tannin, ophiotoxin (the chief poisonous con- stituent of cobra venom), starch, vanilline (the flavoring constituent of the vanilla bean), oil of wintergreen, salol, benzoic acid, digita- line. Here is a variety that baffles description ; but description is hardly necessary, for the facts explain themselves. In short, the com- pounds of the three elements which compose water and carbon dioxide exist in enormous numbers and in unparalleled diversity of chem- ical and physical characteristics. They in- clude substances of the greatest stability, and others of exceeding instability ; liquids, solids, and gases ; chemically active and chemically inert bodies ; acids and neutral substances ; substances which are readily oxidized and others which are oxidized only with great difficulty. In a very large proportion of cases these compounds are capable of enter- ing into reactions with one another. They are, moreover, capable of forming still more complex substances, in still greater variety by entering into union with other elements, notably with nitrogen and sulphur. 208 THE FITNESS OF THE ENVIRONMENT D OTHER ORGANIC COMPOUNDS The organic substances which contain nitrogen are very numerous and exceedingly diverse in their properties. A few of the principal classes of such compounds are the following : — Amines R-NH2, R2NH, R3N Nitrocompounds R — N02 Nitriles R-C = N Isonitriles R — NC Amino-acids R-CHNH2COOH R-N. Azoxy compounds I /O R-Nx Azo compounds R — N = N — R Hydrazo compounds R — NH — NH — R Derivatives of purine, pyridine, and other ring systems, etc. The nitrogenous organic substances include classes of compounds which differ in their properties from any of the non-nitrogenous substances. Of such special properties the most conspicuous is perhaps alkalinity. Like ammonia, of which it is a derivative, the amino group (— NH2), and various other groups containing nitrogen possess this char- CHEMISTRY 209 acteristic. Such compounds, accordingly, supplement those which contain the acid carboxyl group (- COOII) and make possible the fundamental relations of acid, base, and salt among organic compounds, corresponding to those of inorganic chemistry. There exist also many compounds of sul- phur, of chlorine, bromine, and iodine, as well as of various less common elements among organic substances; but in all such cases the complexity and variety of the compounds depend primarily upon the ca- pacity of carbon, hydrogen, and oxygen, or carbon and hydrogen together, to form nu- merous, varied, and complex compounds on which, as it were, the further complexity is superposed. E THE CHARACTERISTICS OF ORGANIC SUBSTANCES Thus the great diversity of organic sub- stances depends in the first instance upon the quadrivalence of carbon, which makes of the carbon atom in the organic molecule a focus, from which chains of atoms may ex- tend in four different directions; or, in the case of double or treble ties, in three directions ; 1 or in but two: «-C->, ^C=>, «=C=»>, 210 THE FITNESS OF THE ENVIRONMENT <-C=>. Next comes the fact that carbon atoms, when otherwise exclusively in com- bination with hydrogen, and under other circumstances in lesser degree, possess an almost unlimited capacity to join together and form chains and rings in great variety. The longest carbon chain yet synthesized oc- curs in the compound hexakontane,1 C60H122, a substance whose constitution is probably as follows : — CH3CH2CH2: CH2 -CHs-CHa. (CH2)54 The stability of this substance justifies a belief that even far longer chains of carbon atoms can exist, and, in fact, there is no known limit to the possibility of stringing carbon atoms together. No other element is believed to share both of these characteristics, and there are various reasons to suppose that the resulting pecul- iarity of the system of organic compounds is really unique. Turning to the periodic classification of the elements, it will be seen that carbon is a member of the first series. Several of the elements of this series, unlike all the other elements except hydrogen, pos- sess very definite individual properties, which 1 " Hell und Hagele," Berichte, 22, 502 (1899). CHEMISTRY 211 mark them off sharply from other substances. Thus carbon bears relatively little resem- blance to its neighbors silicon or titanium, nitrogen to phosphorus or vanadium, oxy- gen to sulphur or chromium; while hydro- gen, of course, has a place quite apart in the classification, and as an element appears to be correspondingly unique. It is, therefore, in the highest degree prob- able that compounds made from elements of such positive chemical characteristics and very unusual properties will be unlike com- pounds formed from other elementary sub- stances. In this manner the periodic classi- fication confirms our confidence in the results of many decades of experience, which lead us to believe that other elements are exceedingly unlikely readily to form compounds com- parable in number, variety, and complexity with those of organic chemistry as we know it. For more evidence we may turn to certain further data of organic chemistry. I refer principally to the character of the organic radicals composed exclusively of carbon and hydrogen. In making a rational classifica- tion of the carbon compounds it has been found convenient to commence with that series of hydrocarbons, called paraffines, with which the present discussion was begun. 212 THE FITNESS OF THE ENVIRNOMENT From them other series may be derived by making a substitution in the molecule. Thus the substitution of a hydroxyl radical for a single hydrogen atom leads from the paraf- fine hydrocarbons, CnH2n+2, to the alcohols CnII2ri+iOH; the substitution of a carboxyl radical — COOH, for the methyl group — CIT3, leads from the paraffine hydrocarbons CnH2n+1CH3 to the acids CnH2n+1COOH. Moreover, the classes of compounds thus defined chemically fulfill the logical require- ments of a class. They are collections of well-characterized and very similar individual things which differ greatly, and in well-marked manner, from all other things. In other words, growing complexity of the molecule, when it consists only in increase in complexity of the simple radical comprised of carbon and hydrogen, of the formula CnH2n+u CH3-, CH3CH2-, CH3CH2CH2-, CH3 CH2 CH2 CH2- CH3x CH3. >CH-, >CH-CH2- ch/ ch/ CHs-CH. >CH- CH/ CH3v CHr)C - CH3/ has very little effect upon the properties of the molecule. Thus the compound methane, CHEMISTRY 213 CH4, very closely resembles normal butane, CH3 * CH2 * CH2 ' CHa ; and again propionic acid, CII3 • CII2 • COOII, and heptylic acid, CH3CH2CII2- CH2- CIL- Clio- COOII, are very much alike. Quite different is the case when any other radical accumulates in the molecule. For instance, propyl alcohol, CII3*CII2*CII2()II, which closely resembles ordinary alcohol, CH3*CH2OH, is very different in its behavior from glycerine, CH2OH CHOH CH2OII, and similarly, acetic acid, CII3*COOH, differs materially in properties from oxalic acid COOH* COOII. Even more marked are the differences when a radical accumulates upon a single carbon atom. In successive stages of oxidation ethane, CH3*CH3, yields alcohol, CH3CH2OH, aldehyde, CH3CH(OH)2, which by a secondary transformation goes over into the more stable form CH3*CHO, and acetic acid, CH3*C(OH)3, which similarly becomes CH3*COOH. These changes correspond to the conversion of ethane, that is monomethvl methane, into dimethyl methane, trimethyl and tetramethyl methane : — CH3-CII3->CH3CH2CH3-+ CH,yH _ rw _> cn3\ r/CH3 ch3Ah "Ltl3 ch/l\chi. 214 THE FITNESS OF THE ENVIRONMENT In the latter case the introduction of methyl groups in place of hydrogen produces no appre- ciable change in the general characteristics of the substances ; in the former, the succes- sive introduction of hydroxyl groups forms substances belonging to three different classes of compounds, — alcohols, aldehydes, and acids, which have nothing in common. In short, variation in the number and arrangement of such groups as occur in the paraffine hydro- carbon, I I I — C — CH3, — C — CH2— -C I I I C-, is without manifest effect upon the more important properties of the molecule, but variation in the number and arrangement of any other groups produces complete change in its characteristic properties. CHEMISTRY 215 It may perhaps he urged that this argu- ment is fallacious, in that the increase of the relative amount of hydroxy! in the above cases is larger than the relative change in radicals of the types — CH3, = CH2, =CH, and == C. But, in the first place, it is evi- dent that the latter four radicals are actually different, and a priori there is no reason to suppose that they should not greatly differ in their effect upon the properties of a mole- cule, for instance, to render dissimilar the compounds normal pentane CH3CH2CH2* CH2*CH3 and tetramethyl methane, CH3\ /CH3 CH3/^\CH3, which is not the case. In the second place, the change from methane, CH4, to ethane, CH3CH3, is a larger proportional change in the molecule than the change from alcohol, CH3CH2OH, to glycol, CH2OHCH2OH, or aldehyde, CH3CHO, both of which produce far greater changes in the properties. In fact, the union of carbon with hydrogen in organic compounds is a unique and peculiar chemical relationship, upon which the proper- ties of the carbon compounds, their number, variety, and complexity largely depend. It seems to make no important difference whether 216 THE FITNESS OF THE ENVIRONMENT a carbon atom is attached to four hydrogen atoms, or to one carbon and three hydrogens, or to two carbons and two hydrogens, or to three carbons and one hydrogen, or to four carbon atoms ; in all such cases the effect of the radical upon the general characteristics of the molecule varies very little. There are a great number of phenomena which might be employed further to illustrate the nature of the case, but two will suffice. The acidity of acetic acid, CH3*COOH, is only slightly and slowly changed by the accumulation of hydrocarbon radicals ; thus the compounds propionic acid, CH3*CH2' COOH, and butyric acid, CHg-CHa'CHv COOH, are only a little less acid than acetic acid itself, because nearlv all the effect of such larger radicals as they contain is already exerted by the methyl group. Ionization Constants of Acids Acetic acid, CH3 COOH 0.000018 Propionic acid, CH3 • CH2 COOH 0.000014 Butyric acid, CH3 CH2 CH> COOH 0.000016 Glycolic acid, CH2OH COOH 0.00015 Chloracetic acid, CH2C1 COOH 0.0015 Dickloracetic acid, CHC12 COOH 0.05 Trichloracetic acid, CC13 COOH 1.2 Glycocoll, CH2NH2 COOH 0.00000000018 Oxalic acid, COOH COOH 0.1 CHEMISTRY 217 But if a hydroxy! group be substituted, as in glycolic acid, CH2OHCOOII, or a chlorine atom, as in monochloracetic arid, CHaCl' COOH, tlie acidity is greatly increased, while the compound trichloracetic acid, CC13- COOH, is a strong acid. On the other hand, aminoacetic acid, CH2*NH2*COOH, is scarcely acid at all. The effect of introducing a carboxyl group in place of a methyl group into any paraffine hydrocarbon, regardless of its constitution, e.g. CH3CH3->CH3COOH, is to diminish the heat of combustion of the molecule almost exactly 157 calories ; but the conversion of acetic acid into oxalic acid, CH3*COOH-> COOH-COOH, structurally an identical change, diminishes the molecular heat of combustion only 147 calories.1 In both these instances it is certain that the nature of the influence of the radicals consisting of carbon and hydrogen exclusively is nearly independ- ent of their size and configuration. Any other group, however, by its presence at once modifies the nature of the case, though un- concerned in the process or property. Since it can be shown that such effects, like the difference between monochloracetic acid and 1 Stohmann, Zeitschrift fur Physikalischc Chemic. II, 29, 1888. 218 THE FITNESS OF THE ENVIRONMENT trichloracetic acid, depend upon the number of such foreign groups and their arrangement,1 it is evident that the hydrocarbon radicals have a constancy of influence upon the prop- erties of the molecule which is not shared by other radicals. The indifference of effect of hydrogen and carbon, when linked to carbon, upon the properties of the molecule is undoubtedly a principal cause of the stability of complex organic substances. Through this peculiar- ity of the two elements the integrity of the valence energy of carbon is preserved, and the long carbon chains are stable. Whenever the molecule becomes overloaded with radi- cals of other kinds the strength of the tie between carbon atoms diminishes and the compound becomes unstable. The proper- ties of the carbohydrates, which will be later discussed, admirably illustrate such instabil- ity. In short, organic compounds are in some respects properly to be regarded as compounds of carbon and hydrogen jointly, for it is not the properties of carbon alone, but those of carbon and hydrogen together which chiefly make them possible. 1 Henderson, Journal of Physical Chemistry, IX, 40, 1905 ; Proceedings of the American Academy of Arts and Sciences, XLII, 639, 1907; Zeitschrijt filr Physikalische Chemie, LX, 413, 1907. CHEMISTRY 219 In the more complex substances, such as the various ring systems of organic chemistry, it is not possible to discuss such problems of molecular mechanics. There too, however, hydrogen predominates over all other elements except carbon, and that may well be taken as a sufficient indication of its continued im- portance. All of these considerations taken together suffice, I believe, to prove, or at least to make it exceedingly probable, that organic chemistry is in truth a unique field, and that no other elements can form compounds in such variety, complexity, and number as carbon, hydrogen, and oxygen. At any rate there can be no possible doubt that the compounds of organic chemistry are in these respects very remark- able indeed, and that similar cases must be extremely rare among all the possible systems of compounds made up of all the known elements. It follows from the peculiarities just ex- plained that the first great factor in the com- plexity of living organisms as we know them, the complexity and variety of their chemical constituents, depends principally upon the nature of the elements which compose such substances, and is most probably a unique, certainly a very rare characteristic of matter. 220 THE FITNESS OF THE ENVIRONMENT That the very elements which make up water and carbonic acid, and apparently they alone, should possess this wonderful property is, when taken together with the physical prop- erties of water and carbonic acid and their place in cosmic evolution as constituents of the atmosphere, a fact which cannot lightly be set aside. Not less valuable for the organism than the multiplicity of organic substances, and the diversity of their properties, are the great variety of chemical changes which they can undergo, and that characteristic instability which renders such great complexity of chem- ical behavior easily attainable. In short, organic substances are uniquely fitted not only to provide complexity of structure to the organism, but also, through their instability and manifold transformations, to endow it with diverse chemical activities, with com- plexity of physiological function. One factor in determining the complexity of chemical changes which organic chemical substances manifest is the enormous number of simple structural relationships which every substance bears to others. This may be readily illustrated by the formulas of some of the derivatives of propane which possess biological importance : — CHExMISTRY 221 CH, CH3 CH3 i CH3 i CH2 CHo CO CO CH2OH COOH CH3 CHO Propyl alcohol, Propionic acid, Acetone, Methyl glyoxal. CH3 CHoOH CHoOH CHoOH CHOH CHOH CHOH CO COOH CH.OH CHO CHoOH Lactic acid, Glycerine Glycerine aldehyde, Dioxvacetone CH3 CHoOH CHo SH CH3 CH NH2 CH NH2 1 CH NHo CH SH COOH COOH COOH COOH Alanine, Serine, Cysteine, a-Thiolactic acid Of the above substances, acetone, lactic acid, glycerine, glycerine aldehyde, dioxyacetone, alanine, serine, and cysteine are of the great- est moment in physiological processes. Such complexity of chemical relationships results automatically, so to speak, in a variety of chemical transformations. But the trans- formations are greatly facilitated by thai characteristic instability of organic substances, which is perhaps the chief distinguishing fea- ture of their behavior. For example, many inorganic substances may be subjected to very 222 THE FITNESS OF THE ENVIRONMENT high temperatures without undergoing chem- ical change, but there are hardly any organic substances which can survive such treatment. Organic substances are also peculiarly liable to modification from the action of light and air. These are, however, but rough indica- tions of instability, and a special case will help more clearly to define the real characteristic. THE SUGARS In accordance with the researches of Emil Fischer, the following constitution is ordina- rily assigned to glucose.1 H I C = 0 I H-C-O-H I H-O-C-H I H-C-O-H I H-C-O-H CH2OH 1 A discussion of this formula may be found in any text- book of organic chemistry. CHEMISTRY 223 It must be noted that carbon atoms to which four different groups are attached are asymmetric, that is to say, they can exist in two forms which resemble each other as the right hand resembles the left (Pasteur, LeBel, van't Hoff). This characteristic results in further increase in the number and variety of organic compounds. It is therefore neces- sary, in writing the formula, to represent the form of the molecule as it exists in space (in three dimensions), and this is actually accom- plished by imagining the three-dimensional formula to be projected upon the paper so that when the hydrogen atom is written to the right of the carbon atom one asymmetric form of the latter is designated, when the hydrogen atom appears to the left, the other. It has long been known that when glucose is dissolved in water its optical activity, as the power of a substance to rotate the plane of polarization of light is loosely termed, changes slowly for some time before reaching a constant value. Recently it has been shown that this phenomenon probably depends upon the existence in solution of three different forms of glucose, which pass freely into one another and ultimately attain a state of equilibrium.1 1 This subject has been fully discussed by Hudson, Journal of the American Chemical Society, XXXII, 889, 1910. 224 THE FITNESS OF THE ENVIRONMENT OH H-C— I H-C-OH I HO-C-H I H-C O H-C-OH I CH2OH CHO I H-C-OH I HO-C-H I H-C-OH I H-C-OH I CH2OH H I HO-C H-C-OH I HO-C-H H-C O H-C-OH I CHoOH When to such a solution of glucose a small quantity of alkali is added, certain remarkable further changes occur, as was first demon- strated by Lobry de Bruyn.1 These pro- cesses result in the formation of mannose and levulose, probably according to the accom- panying diagram. Moreover, like glucose, levulose and man- nose both exist in solution in three different forms, so that the resulting solution contains at least ten chemical individuals. But it is almost certain that other changes simultane- ously occur and that the solution is actually still more complex, even from the outset. Upon a continued increase of alkalinity, or even slowly under the original conditions, a multitude of other changes set in. These 1 See the numerous papers by Lobry de Bruyn and Al- berda van Eckenstein in Recueil des Travaux Chimiques des Pays Bas, XIV-XIX. CHEMISTRY 21 CHO HCOH HOCH HCOH 1 HCOH CH,OH 1 CH2OH 1 CHOH II C-OH CHO CO HO- -C-H HO-C-H =^= HO-C-H ^ HO- -C-H H-C-OH H-C-OH 1 H- -C-OH 1 H-C-OH 1 H-C-OH H- -C-OH 1 CH2OH CH2OH CH2OH Levulose Mannose are known sometimes to lead to the formation of lactic acid, CH3CHOHCOOH, methyl- glyoxal, CH3*CO*CHO, and formaldehyde, H-CHO. Further, as Neff has shown,1 many other substances may also be formed. Such bodies chiefly belong to the class of oxyacids. It is also certain that a great variety of other simple sugars resembling glucose, levulose, iNeff, "Liebig's Annallen," 376, p. 1, 1910, and 357, p. 294, 1907. Q 226 THE FITNESS OF THE ENVIRONMENT and mannose are produced, and, all told, the constituents of such a solution probably number at least two hundred, all produced from glucose alone, under the influence of a slight excess of hydroxyl ions. Among these substances the greatest diversity of chemical behavior is to be distinguished. Alcohol, aldehvde and ketone, and acid radicals occur in great profusion and variety of combinations ; compounds possessing forked chains are pres- ent ; and double bonds between carbon atoms add to the complexity. Moreover, all these substances themselves possess great chemical activity. A single case may perhaps illustrate this point. It has been shown by Windaus and Knoop l that in such solutions, in the presence of ammonia, one molecule of methyl glyoxal, one of formaldehyde, and two of ammonia unite to form the cyclic compound, methyl imidazol, a substance related to histidine, the latter being an important constituent of the protein molecule : — CH3-C = 0 NH3 /H CH3-C-N/H I C^O -> II ^>C-H C=0 NH3 XH H— C-N^ H 1 Knoop and Windaus, Berichte, XXXVI, 1166, 1905. Hofmeister's Beitrage, VI, 392, 1905. CHEMISTRY 227 The instability of glucose and of all the simple sugars is indeed exceptional in char- acter, and the resulting processes arc perhaps far more intricate and numerous than in any other similar case. However, this very case is of exceptional physiological importance, be- cause carbohydrates are the direct result of that synthetic action of chlorophyll, 6 C02 + 6 H20 - C6H1206 + 6 02, which is the source of all organic substances and of all the energy of the organic cycle in plants and animals. Carbohydrates, more- over, are the chief constituents of plants and the chief food of animals. Turning to this synthesis of carbohydrate in the plant, we find much that is important in the present study. The details of the chemical transformation by which water and carbonic acid and solar energy are changed to sugars and oxygen still remain unknown, in spite of many careful investigations. But, at all events, it is possible to see that two things must somehow be done in order to accomplish the synthesis: — (1) Carbonic acid and water must be re- duced. That is to say, oxygen must be sepa- rated from both of these compounds so that free valences may exist to unite carbon and 228 THE FITNESS OF THE ENVIRONMENT hydrogen in one molecule, and so that, further, the relative proportions of the three elements may become what they are in the simple carbohydrates, C:H:0 = 1:2:1. (2) Somehow individual carbon atoms must be joined together until there are six in each molecule, where formerly there was but one. Theoretically, it might be possible by reduc- tion to form from carbon dioxide and water without union of carbon atoms the following substances: carbon monoxide, CO; formic acid, H-COOH; formaldehyde, HCHO; methyl alcohol, CH3OH ; and methane, CH4. Of such reductions the formation of formic acid, formaldehyde, and carbon monoxide has been directly realized by laboratory experiment.1 The most familiar theory of the formation of carbohydrates in the leaf is that of von Baeyer, which assumes a polymerization of one of the above substances, formaldehyde, leading directly to the formation of sugar, according to the reaction 6HCHO=C6H1206. This process also has been carried out ex- perimentally. Indeed, as a result of the investigations of Butlerow, O. Loew, and ^ee Meyer u. Jacobson's "Lehrbuch der Organischen Chemie," Leipzig, 1907, p. 688, 693-696. CHEMISTRY 229 E. Fischer, it is known that in distinctly alka- line solutions formaldehyde spontaneously goes over into a mixture of sugars which resemble glucose. Moreover, such a solution is un- questionably made up of much the same variety of substances as a glucose solution which has been subjected to the action of alkali in the same concentration. As stated above, it is evident that if any- thing is to be done chemically with a mix- ture of carbon dioxide and water, oxygen must be split off from both carbon and hydrogen so that they may enter into the same mole- cule. If this chemical change, which, to be sure, is no easy one in the laboratory, be ac- complished, formaldehyde among other things results ; and in alkaline solution formaldehvde produces carbohydrates and leads to that amazing tangle of substances and reactions, whose nature has been briefly indicated above. In short, the one chemical process ivhich is open, if any transformations whatever are to be accomplished with carbonic acid and water, leads directly and to all appearances necessarily to the greatest complexity that has been found in any one chemical process; to a system made up of possibly two hundred substances or more, most of which possess very great chemical activity. 230 THE FITNESS OF THE ENVIRONMENT It must not be forgotten that the share of the organism in such processes is also impor- tant. If possible, the smoothness with which chemical reactions are carried out in the leaf, perhaps quite without any stop at the formal- dehyde stage, and the certainty with which definite substances in large amounts instead of mixtures in very small amounts are pro- duced, seem more remarkable than the under- lying chemical facts. Needless to say, one great factor in such processes is the action of enzymes. Otherwise we are at a loss for a description of the ways and means by which the organism operates, though the brilliant studies of chlorophyll which have recently been carried out by Willstatter promise great achievements in the future.1 The underlying chemical facts, however, remain ; carbohydrates are among the natural products of carbon dioxide and water; they manifest in solution, especially in such con- ditions as obtain in protoplasm or the ocean,2 unparalleled instability and variety of reac- tions ; and they produce spontaneously an enormous number of very active chemical substances. It is easy to see that, given 1 See his many papers of recent years in "Liebig's Annal- len." 2 Henderson, Journal of Biological Chemistry ', X, 3, 1911. CHEMISTRY 231 an enzyme possessing the power to select and catalyze any one of the reactions, the for- mation of any special one of the many possible products in comparative purity is an auto- matic result. Such processes are in nature carried out with a perfection which to the chemist is almost inconceivable, by means of organic structures of the highest intricacy ; but in the last analysis they rest upon the native properties of the three elements. The important consideration, I repeat, is this : that reduction, the necessary first change of carbonic acid and water, can lead directly by a single continuous chemical transforma- tion, of which the exact control is no whit more remarkable than the accurate control of the other processes of chemical physiology,1 to the full intricacy of organic chemistry ; to the very most notable instance of number, variety, and activity of substances, all formed inevitably, in the nature of the case, which has yet come to light. It is of no consequence that most of the substances must be formed in mere traces by the spontaneous synthesis, for in highest degree the organism possesses the power, by enzymatic catalysis, to select 1 Consult the work of Bayliss on Enzymes. London, Longmans, Green & Company. 232 THE FITNESS OF THE ENVIRONMENT any one of a group of simultaneous reactions that may serve its needs, and make that one predominant. In short, the fitness is recip- rocal; the unique chemical adaptability of the process and the unique chemical powers of the living organism interlock. Such, in brief, are the superlative advan- tages which the properties of the compounds of the three elements contribute to the organic mechanism. They include number of sub- stances, variety of substances, variety of properties, variety of reactions, facility of reactions (instability), and the remarkable relationship between carbon dioxide and water and the carbohydrates. And they insure that extreme variety of chemical relationship which especially fits organic substances, once created, to be, throughout the various forms of life, the source of still other bodies, and the source of energy, by means of far-reaching chemical changes rapidly accomplished. G HYDROLYSIS In the course of digestion the principal foodstuffs, carbohydrates, fats, and proteins, undergo a series of changes which are substan- tially the same for all. Such processes are known as hydrolytic cleavage, or more loosely CHEMISTRY 233 as hydrolysis. Essentially they amount to successive splittings of the large molecules of the native substances, each cleavage being accompanied by the addition of a molecule of water, until finally from starches and like substances the simple sugars like glucose result; from the fats, fatty acids and glycerine; from the proteins, the so-called amino acids. The cleavage of fats closely resembles the hydrolysis of a simple ester; the cleavage of proteins and carbohydrates a little more remotely resembles the same process. Accord- ingly, the hydrolysis of the simplest ester, methyl formate, may serve as an illustration of the nature of the reaction: — H-C-O^CH3+HO lH = H-C-0-H+H-0-CH, II : I II o o This process is nothing less than the typical reaction between water and organic sub- stances. Accordingly, it is not surprising that such reactions are by no means confined to the digestion of food. Once formed, the prod- ucts of digestion are absorbed, the more readily because of their simplicity, and, also because of their simplicity, they carry into the body no trace of the organism in which they previously existed. But, if they are to 234 THE FITNESS OF THE ENVIRONMENT be built up into the tissues of the animal, they must now be turned back into such fats, carbohydrates, and proteins as are character- istic of his physical structure ; into glycogen, haemoglobin, fibrinogen, etc. Accordingly, they undergo a process which is the exact reverse of the digestive change, in the simple case: — H-C-0-H + H-0-CH3 = H-C-0-CH3 + H-0-H II II o o But this is by no means the end of the matter. For example, glycogen thus formed in the liver from the glucose of the portal blood is soon torn down to glucose again. More- over, there are a host of other special cases of the same hydrolytic cleavage, or the re- verse process, in mammalian physiology. For instance, the formation of hippuric acid from benzoic acid and glycocoll, and the formation of urea itself from ammonium carbonate belong to this same class. In fact, such reactions make up a large part of all the chemical changes which take place within the organism. It must not be imagined, however, that hydrolytic cleavages are infrequent outside the organism, or that the types of processes CHEMISTRY 235 which occur within the organism are the only ones of this class. There is no more common and universally important reaction in organic chemistry, and many compounds and classes of compounds which have nothing to do with the organism undergo hydrolysis. Moreover, generally speaking, all reactions of this class are very similar in their principal character- istics, resembling one another both dvnami- cally and statically. Spontaneously they oc- cur not at all, or very slowly. Under the influence of enzymes, of acids, and of alkalies acting catalytically, that is to say, facilitating the process without in the end taking part in it, much as oil facilitates the action of a machine, they progress rather slowly and very smoothly. By-products are not formed ; the reactions are simple, uncomplicated, and reliable. Hence they enable the organism to make all sorts of rearrangements and recon- structions of chemical substances efficiently and without loss of material. The chief cause of such traits in hydrolysis is the fact that the energy transformation which accompanies the process is almosl exactly nil. For it has been found in general that chemical reactions which liberate much energy are violent, hard to regulate, often complicated by intricate side reactions, and 236 THE FITNESS OF THE ENVIRONMENT complete. In contrast, those which are with- out energy change generally proceed smoothly, slowly, and without complication to a state of equilibrium in which the reaction is very in- complete.1 Under the latter circumstances slight changes of conditions make possible a reversal of the delicately balanced process ; the reaction can be made to run in either direction at will. The absence of transformation of energy accompanying hydrolysis may be illustrated by a few typical cases chosen from the data of simple substances. Calories Per Cent + 1.9 +4.3 -3.7 +2.0 0.2 0.3 0.3 0.3 Such measurements of heats of reaction fall well within the limits of error of the method of investigation, and there can be no doubt that in all such cases the heat of reaction is so small that it cannot be detected by the ordinary methods of measurement.2 1 van't Hoff, "Acht Vortrage liber Physikalische Chemie." Brunswick, 1902, Lecture 6. 2 Stohmann, Zeitschrift fur Physikalische Chemie, II, 29, 1888 (see also Ostwald's "Lehrbuch der Allgemeinen Che- CHEMISTRY -237 Thus it is evident that the process possesses another advantage. In the course of such rearrangements no energy is lost. This con- clusion is thoroughly confirmed by the studies of the energy transformations of metabolism. The body may carry on such processes as it will, in the greatest variety and complexity, rearranging and modifying its chemical struc- tures to any extent, and there will never be an appreciable wastage of precious material or of equally precious energy in the process. This process, as we have seen, is the char- acteristic reaction between water and the organic compounds. As such it is necessarily one of our chief concerns ; its maximal fitness as a means of regulation, and otherwise, there- fore assumes real importance in the present discussion. n INORGANIC CHEMISTRY With the survey of organic chemistry we have exhausted the compounds of carbon ; mie"). These data are the most accurate now in existence which permit an estimate of the heat of hydrolysis of non- nitrogenous compounds. Numerous studies of protein deriv- atives from Fischer's laboratory prove that the facts are the same for these substances, and direct measurements confirm the measurements of heats of combustion. 238 THE FITNESS OF THE ENVIRONMENT not so those of hydrogen and oxygen. In almost equal frequency the latter elements take part in the reactions of inorganic chem- istry, and help to form its molecular struc- tures. As an illustration of their importance in this department of the science I have counted the compounds and the classes of compounds mentioned in the table of con- tents of the second edition of Erdmann's * * Lehrbuch der Anorganischen Chemie. ' : In all 435 substances are referred to ; of these 259, approximately 60 per cent, contain oxygen ; 130, or 30 per cent, contain hydrogen. There seems to be little doubt that this is a fair test, for the work is compendious, and all im- portant substances and classes of substances are mentioned. Even if the acids, and the small number of bodies which are referred to in connection with their water of crystalli- zation, be eliminated from the above count the great importance of the two elements remains clearly evident. Only about one fourth of all the compounds mentioned contain neither hydrogen nor oxy- gen. A very large proportion of these con- sist of the chlorides, bromides, iodides, sul- phides, fluorides, and other similar binary compounds, whose importance certainly does not depend upon the variety of chemical CHEMISTRY 039 reactions into which they may enter, while their formation unquestionably docs depend upon the intervention of both hydrogen and oxygen. All told, the chemical substances which contain neither carbon, nor hydrogen, nor oxygen make up only a few per cent of known bodies. It is also clear that an especially large pro- portion of the most active inorganic com- pounds contain either hydrogen or oxygen. All acids contain hydrogen; most of them oxygen as well. All bases contain oxygen. Moreover, the most important classes of re- actions of inorganic chemistry are probably oxidations and reductions, and the formation of salts from acids and bases. In such pro- cesses both oxygen and hydrogen are con- cerned. In addition to the oxides and resulting bases and acids, a few other important sub- stances wThich contain hydrogen or oxygen may be cited : ozone 03, hydrogen peroxide H202, ammonia NH3, hydrazine N2H4, hydrox- ylamine NH2OH, sulphuretted hydrogen H2S, hydrochloric acid HC1, nitrosyl chloride NOC1, thionyl chloride SOCl2, phosgene COCl2, phos- phine PH3, phosphorus oxychloride POCl3, arsine AsH3. Such compounds, and many other similar ones, are of great importance on 240 THE FITNESS OF THE ENVIRONMENT account of the variety of chemical processes into which they can enter. They make up the active agents of inorganic chemistry, and it is safe to assume that their activity depends in great part upon the properties of oxygen and hydrogen. The importance of oxygen and hydrogen in inorganic chemistry possesses a double significance in the present inquiry. In the first place it provides further confirmation of the view that the elements which make up water and carbon dioxide are unique. For the data of inorganic chemistry prove that hydrogen and oxygen are likely to confer great chemical activity wherever they are, and that they are quite unrivaled in this respect. Secondly, the occurrence of hydrogen and oxygen as primary factors of the metabolic process and as the chief constituents of the environment and of the living organism enables the latter to make use of other elements at need. Without hydrogen and oxygen, op- portunities for the introduction of such other elements into the physiological processes would be necessarily much restricted, and in many cases the physiological utility of compounds containing the elements of inorganic chemistry is very great. Chlorophyll, for example, contains mag- CHEMISTRY 241 nesium, and it is thought that the process of reduction in the leaf may depend upon the characteristic properties of this element ; at all events, in organic chemistry, magnesium, when employed in Grignard's reaction, is one of the most effective agents to accomplish reductions. In like manner, haemoglobin contains iron, and the capacity of haemoglobin to unite with oxygen, and as oxy haemoglobin to carry it from the lungs to the tissues is unquestion- ably due to the chemical behavior of that metal. Other similar metallic elements, no- tably copper in the class of compounds known as haemocyanines, fulfill a similar function in lower animals. Phosphorus in organic union is an essential constituent of a great variety of the chemical structures of living organisms, — the nucleic acids, which appear to be not less important than fats, carbohydrates, and proteids them- selves in both animal and plant cells, contain phosphorus as an essential constituent. Thus phosphorus follows close upon nitrogen, after carbon, oxygen, and hydrogen, as structural material in biological chemistry. This same element also occurs in many other compounds, the simplest derivative of such bodies being glycerophosphoric acid, 242 THE FITNESS OF THE ENVIRONMENT CH2OH I CHOH \ />-H 0-PM3 X0-H, a compound of phosphorus, hydrogen, and oxygen with glycerine. Sulphur is a constituent of the proteins, and occurs in many other important com- pounds. In the metabolic process of the animal sulphur is converted from a derivative of hydrogen sulphide, H2S, by oxidation, into sulphuric acid, H2S04, and in the plant the process is reversed. Iodine occurs in the thyroid and in many marine organisms. The availability of this element depends upon its existence in nature as iodides, that is to say, upon its capacity to unite with hydrogen to form hydriodic acid, HI. Finally, it is the analogous com- pound of chlorine with hydrogen, hydro- chloric acid, which contributes acidity to the gastric juice. This list might be much further extended, but I think that the nature of the case is now established. The conclusion seems inevitable CHEMISTRY £43 that active, diverse, and important inorganic substances usually contain oxygen or hydro- gen, and that it is the union of other elements with these two which renders them available and useful to the organism. Ill THERMOCHEMISTRY Every chemical change consists in simul- taneous rearrangements of matter and energy. The true nature of the chemical process is to be sought neither in the one nor in the other of these two phenomena, but in both together ; and properly energy is as much the chemist's concern as matter itself. Thus far in the present investigation, con- siderations regarding energy have been avoided except in the case of hydrolytic cleavages, and these constitute a unique class of reactions* No other large and important class is char- acterized by inappreciable heat of reaction, for it is as heat that chemical energy commonly manifests itself when liberated. It is evident, however, in accordance with the fundamental postulates, that the organism must have energy to actuate as well as matter to form its mechanism. Therefore the nature of the energy transformations, which make up one 244 THE FITNESS OF THE ENVIRONMENT aspect of the chemical reactions into which carbon, hydrogen, and oxygen enter, must be now noticed. It has been shown above that the one pos- sible chemical process by means of which any- thing can be made out of the primary con- stituents of the environment is reduction, — the more or less complete tearing off of oxygen from carbon and hydrogen atoms in the mole- cules of carbon dioxide and water. As a function of the extent of the reduction the energy change involved in the process will vary. In all cases, however, the process is accompanied by large absorption of heat, as the following table of the energy absorbed per gram of the resulting substance, when reduction begins with water and carbonic acid, may indicate: — H2 34.5 Cal. C 8.1 CH4 13.3 CH3OH 5.3 HCHO 4.2 C6H1206 3.74 CO 2.4 HCOOH 1.4 Such new compounds hold their energy only so long as they persist unchanged, and CHEMISTRY 245 upon oxidation they yield it all back again, just as water vapor on condensing yields back the latent heat which it has taken up during evaporation. In this manner, every gram of glucose or other monosaccharide is necessarily a temporary depository of solar energy amounting to about 3.74 calories, taking up just that amount of energy when synthesized by chlorophyll, yielding it back when burned in the muscle. Compounds of carbon and hydrogen are especially well qualified to be reservoirs of energy which may be liberated by oxidation, as the following table shows: — HEATS OF COMBUSTION OF ELEMENTS PER GRAM Hydrogen 34.5 Cal. Carbon 8.1 Sulphur (to S02) 2.3 Sulphur (to S03) 3.2 Nitrogen (to N02) 0.2 Phosphorus 5.9 Boron 12.3 Silicon 3.3 Potassium 1.3 Calcium 3.3 Aluminium 7.0 Hydrogen, it will be seen, far exceeds any other element in the amount of heat that it 246 THE FITNESS OF THE ENVIRONMENT yields upon oxidation; carbon is surpassed by but one other element, boron. Although necessarily a good deal of this heat cannot be stored in the compounds of the two elements which still contain some oxygen, yet enough remains to make the common constituents of the organism greater reservoirs of energy than most of the other elements themselves, far greater than compounds of any other elements. Thus the heat of combustion of carbohydrates ranges from about 3.7 calories to about 4.2 calories per gram, that of the proteins from about 5 calories to about 6 calories, that of the fats from about 9.2 calo- ries to about 9.5 calories. On account of the small quantity of oxygen and the large quan- tity of hydrogen which they contain, the fats are a richer source of energy than carbon itself, or than any other element except hydro- gen and boron. There remains one other equally important consideration to be dealt with: the very great energy change which is involved in processes of oxidation and reduction compared with other chemical processes. The following table, showing the amount of heat liberated in the process of formation of certain binary compounds from their several elements, illus- trates the case: — CHEMISTRY 1A HEATS OF FORMATION H20 3.83 Cal. CSa -0.25 Cal C02 2.22 Nad 1.67 HC1 0.60 LiCl 2.20 HF 1.97 NaBr 0.87 NPI3 1.23 NaF 2.64 02V>12 0.08 1 Na2S 1.14 ecu 0.49 SiH4 -0.21 PI3 0.26 SiF4 2.31 BC13 0.79 NS -0.69 Oxygen, as will be seen, far surpasses the other chemical elements (except fluorine) in the amount of energy liberated in the process of its chemical union with other substances. Accordingly, it may be concluded that, on the whole, oxidations are the best chemical source of energy ; reductions the best means of storing energy by chemical processes; and that among oxidations and reductions those of hydrogen especially, and then those of carbon, are associated with the largest energy transformations. This is the last argument which I have to present, but it is one of the most potent. The very chemical changes, which for so many other reasons seem to be best fitted to be- come the processes of physiology, turn out to be the very ones which can divert the greatest 248 THE FITNESS OF THE ENVIRONMENT flood of energy into the stream of life; and these are the reactions automatically provided for by the cosmic process. From the materialistic and the energetic standpoint alike, carbon, hydrogen, and oxy- gen, each by itself, and all taken together, possess unique and preeminent chemical fit- ness for the organic mechanism. They alone are best fitted to form it and to set it in motion ; and their stable compounds, water and car- bonic acid, which make up the changeless environment, protect and renew it, forever drawing fresh energy from the sunshine. CHAPTER VII THE ARGUMENT THE statement of evidence for the bio- logical fitness of the environment is at length completed. Whatever favorable prop- erties of water, carbonic acid, and the com- pounds of the three elements, whatever results favorable to life, I have succeeded in finding, have been set forth. Now, therefore, we may return to the exam- ination of this evidence in the manner sug- gested in Chapter II. We may inquire into the exhaustiveness of the preceding treatment of important physical properties, seeking to discover what things have been overlooked. Thus it may be possible to decide the weighty question whether another group of elements can possess another group of equally impor- tant properties. Next, we may consider if there be other elements or compounds which rival carbon, hydrogen and oxygen, water and carbonic acid, in the qualities which make these fit for the organic mechanism, taking such properties as a whole. Unfortunately, 249 250 THE FITNESS OF THE ENVIRONMENT in adopting this somewhat rigid logical method, tedious and perhaps unnecessary repetition is involved, but the advantages of care at this stage of the inquiry seem to be very great, for it is not easy to survey so large a field, and at best certainty that impor- tant oversights have been avoided is obviously impossible. For example, peculiarities like the anomalous expansion of water, or the relation of carbonic acid and water to the carbohy- drates are not to be foreseen. On the other hand, the more general characteristics of matter are well known and, for the most part, must reveal themselves to diligent search. ANALYSIS OF THE EVIDENCE First the natural phenomena which seem to be concerned in fitness may be brought to- gether analytically, and their effect briefly summarized. NATURAL PHENOMENA WHICH PROMOTE FITNESS IN THE ENVIRONMENT I. The occurrence of great quantities of water and carbon dioxide outside the solid crust of an astronomical body. THE ARGUMENT J51 II. Properties of Water. a. Specific Heat. b. Freezing Point. c. Latent Heat of Fusion. d. Latent Heat of Vaporization. e. Vapor Tension. /. (Thermal Conductivity.) g. Expansion before Freezing. h. (Expansion in Freezing.) i. Solvent Power. j. Dielectric Constant. k. Ionizing Power. /. Surface Tension. III. Properties of Carbon Dioxide. a. Solubility in Water. b. Ionization Constant. IV. Properties of the Ocean. a. Number and Variety of Constituents. b. Quantity of Dissolved Material. c. Mobility. d. Constancy of Temperature. e. Constancy of Osmotic Pressure. /. Constancy of Alkalinity. g. Constancy of Composition. V. Chemical Properties of Carbon, Hydro- gen, and Oxygen. a. Number of Compounds. b. Variety of Compounds. c. Complexity of Compounds. 252 THE FITNESS OF THE ENVIRONMENT d. Number of Reactions. e. Variety of Reactions. /. Complexity of Reactions. g. The Evenness and Lack of Energy Change of the Process of Hydro- lytic Cleavage. h. The Chemical Relationship of Car- bonic Acid and Water to the Sugars. i. Instability of the Sugars. j. Variety and Reactions of the Sugars. k. Heats of Reaction in Organic Chemis- try. L The Number and Variety of Com- pounds and Reactions of Oxygen with Other Elements. m. The Number and Variety of Com- pounds and Reactions of Hydro- gen with Other Elements. All the properties or other phenomena noted in the above table (except II /, and II h) are in character or in magnitude either unique or nearly so, and are in their effect favorable to the organism as defined in the fundamental postulates. Indeed, they constitute or bring about an extraordinary set of conditions favorable to life, — ubiquity, abundance, va- riety, stability, mobility, constancy of composi- tion, and in variance of physico-chemical con- THE ARGUMENT ££9 ditions in the environment; number, variety, complexity, adaptability, availability, activity, and richness in energy of the substances which take part in the metabolic processes and in the chemical and physical format ion of the organism ; constancy of physico-chemical con- ditions, such as temperature, alkalinity, col- loidal disperseness, etc., within the organism; the efficiency of many physiological processes ; the availabilitv of electrical forces, etc. In short, by many independent and united actions the above catalogued natural char- acteristics of the environment promote and favor complexity, regulation, and metabo- lism, the three fundamental characteristics of life upon which all our discussion has been based. II THE EXHAUSTIVENESS OF THE TREATMENT One manner of judging the completeness with which different types of phenomena and properties, different elements and compounds, have been considered in the descriptive chap- ters preceding is to glance at the several departments of physical science, chemistry, mechanics, heat, sound, light, magnetism, elec- tricity, and physical chemistry. 254 THE FITNESS OF THE ENVIRONMENT In setting out to consider physical and chemical properties we may perhaps begin with chemical phenomena in the narrowest sense. Such phenomena depend, according to the atomic theory, upon rearrangements of atoms within molecules. They result in the conversion of individual substances into one another, and they are accompanied by rearrangements of energy. In the first place, it is to be noted that enormous quantities of carbon, hydrogen, and oxygen, as water and carbonic acid, are, during a very long period of time, apparently inevitable constituents of the atmosphere of an astronomical body of sufficient size, after cooling has led to the formation of a crust. Further, it has been shown that in number, variety, complexity of forms and changes, and in the magnitude of the accompanying trans- formations of energy the known substances made up of carbon and hydrogen, and those made up of carbon, hydrogen, and oxygen far surpass the compounds of any other elements. Likewise the known compounds of oxygen and hydrogen with other elements are the most numerous and important among inorganic substances. Two peculiarities of the carbon compounds, the formation and properties of the carbohydrates, and the nature of the pro- THE ARGUMENT 155 cess known as hydrolytic cleavage, add to this list of chemical characteristics which make for fitness. These facts appear to indicate that in gen- eral chemical behavior, in certain special characteristics as well, and in the magnitude of the quantity of energy rendered available by their chemical changes, the elements car- bon, hydrogen, and oxygen are uniquely and most highly fitted to be the stuff of which life is formed and of the environment in which it exists. Mechanics has taken a place subordinate to chemistry in the present work. Neverthe- less, it has been noted that the unique proper- ties of water are the cause of the admirable mobility of that substance and of the whole environment, and therefore of the dynamical processes of geology, meteorology, etc., in- cluding soil formation ; that it is surface ten- sion which holds water in the soil ; that the efficacv of water as a means of dissolving the greatest variety of substances in the greatest amounts, makes possible high osmotic pres- sures, as wTell as mobility of all the elements; and there are a host of other considerations which have been discussed above. In all such cases the properties of water have been found to be favorable influences for the wel- 256 THE FITNESS OF THE ENVIRONMENT fare of the organism. Considering the com- parative unimportance of mechanics in rela- tion to the fundamental postulates, it seems clear that this department has not been over- looked. Thermal processes and thermal effects are perhaps more conspicuous in the table. The thermochemical characteristics of organic com- pounds and the thermal properties of water are all very favorable to life. Stores of heat for the organism, constancy of temperature of both organism and environment, the per- manence of bodies of water, and a multitude of other most important results flow from these properties and bear witness to their unique fitness. Sound, light, and magnetism have not been considered, for they appear to bear only a secondary relation to the fundamental postu- lates. In electricity no phenomena are more im- portant than those of ionization in solution. To bring about ionization and thus make possible electrochemical processes, water is the very best medium, and the possibility of such processes is probably necessary to the organic mechanism. In addition to the topics of physical chem- istry already referred to under chemistry THE ARGUMENT 257 and the several departments of physics, the colloids and the ions of hydrogen and hydroxy] remain to be mentioned. It has been shown that the properties of water arc exceptionally favorable to the existence and stability of colloidal systems; also that the properties of carbonic acid result in automatic regulation of the concentration of hydrogen and hydroxy] ions in the ocean and in the organism. So far, then, as it is possible to judge by telling over the departments of physical science, our examination of physical and chem- ical properties has not been incomplete. This conclusion may be further tested with the help of the ideas which underlie YVillard Gibbs's " Phase Rule." l According to this rule, the condition of equilibrium in any material system depends upon the number of its com- ponents, the number of its phases, temperature, pressure, and, in general, the concentrations of all the components. Without entering upon an explanation of the exact mathemat- ical notions which determine the meaning of the terms "component' and "phase' it will here suffice to say that in general the number of components increases as the num- ber of separate chemical individuals increases, 1 See, for instance, Findlay, "The Phase Rule and its Applications." London, 1911, 3d ed. 8 258 THE FITNESS OF THE ENVIRONMENT and that a phase is any solid, liquid, or gas- eous part of the whole system which possesses homogeneity of composition. For instance, if a system is made up of sand, salt solution, ice, and aqueous vapor, each of these separate parts, in that it is homogeneous, is a phase. Now the properties of water have the result that more readily than other substances it exists simultaneously and in large quanti- ties in the three phases of solid, liquid, and gas as ice, water, and aqueous vapor. This depends upon the high latent heats of fusion and vaporization, the high freezing point of water, and its vapor tension. Water en- hances the complexity of the environment, and is one principal factor in the mobility of the environment as a whole. Further, it makes for stability ; other things being equal, the greater the number of phases, the less the tendency to change. Among phases the dis- perse colloidal type is unique and of very great importance — almost the sole basis, indeed, of great physical complexity — and, as above shown, the peculiar properties of water highly favor the colloidal condition. The solvent power of water much increases the number of components which may enter into a system of which it is a part ; hence the large number of components of sea water, THE ARGUMENT 250 blood plasma, etc. The variety of compounds, both organic and inorganic, which contain carbon, hydrogen, or oxygen also causes enor- mous increase in the number of components of biological systems like protoplasm. The effects of the properties of water above enumerated to regulate temperature are almost too numerous to mention. The specific heat of water, its latent heats of fusion and vapori- zation, and the high freezing point all con- tribute to the restriction of temperature range within the organism, in the waters, and over the whole surface of the earth. The vapor pressure of water has been shown to possess great and exceptional variability wTith change of temperature. This is the most impor- tant property of water meteorologically, and is the necessary condition for its ample cir- culation. The ratio between the gas pressure of carbonic acid and its concentration in water (absorption coefficient) has been shown to be the great factor in establishing the mo- bility of that substance. The total atmos- pheric pressure has not entered into our dis- cussion, for it seems to have no important special relation to the properties of the three elements. In short, the properties of water and of the carbon compounds provide for number, va- 260 THE FITNESS OF THE ENVIRONMENT riety, and complexity of phases and compo- nents, and for constancy of temperature, while equally important and unique relationships between the properties of water and carbonic acid and their vapor or gas pressures exist, and exert much influence upon the meteoro- logical cycle. Thus, judged by the phase rule, the actual characteristics of the environment may be shown to contribute the factors which make for complexity and regulation of material sys- tems. Now there can be no doubt that, when feasible, the ideal method — from the physico- chemical point of view — to describe a ma- terial system is in the terms of the phase rule.1 Hence the characteristics which that 1 "Ten years after the law of mass action was propounded by Guldberg and Waage, Willard Gibbs, Professor of Physics in Yale University, showed how, in a perfectly general manner, free from all hypothetical assumptions as to the molecular condition of the participating substances, all cases of equi- librium could be surveyed and grouped into classes, and how similarities in the behavior of apparently different kinds of systems, and differences in apparently similar systems, could be explained. " As the basis of his theory of equilibria, Gibbs adopted the laws of thermodynamics, a method of treatment which had first been employed by Horstmann. In deducing the law of equilibrium, Gibbs regarded a system as possessing only three independently variable factors — temperature, pressure, and the concentration of the components of the system — and he enunciated the general theorem now usually known as the THE ARGUMENT 261 rule contemplates are in certain respects the most important of characteristics. Accord- ingly the above test is a valuable indication of the adequacy of the preceding analysis of physical and chemical properties. In order if possible to discover the nature of such properties of matter as may have been omitted in our study of fitness, I have ex- amined the index of Landolt and Bdrnstein's "Physikalisch-chemischen Tabellen," a very extensive and comprehensive work. In addi- tion to information regarding the arbitrary units of physical science, I find mention of the following properties which have not been considered in the present discussion: — The Mechanical Equivalent of Heat. The Dimensions of the Angles of Crystals. The Refraction of Light. Compressibility. The Dimensions of the Molecules of Gases. Elasticity. The Electromagnetic Rotation of the Plane of Polarization of Light. Color. Phase Rule, by which he defined the conditions of equilibrium as a relationship between the number of what arc called the phases and the components of the system." — Findlay, "The Phase Rule and its Applications." London, 1911, 3d ed., p. 8. 262 THE FITNESS OF THE ENVIRONMENT Viscosity. Torsion. The Velocity of the Molecules of Gases. Hardness. Magnetism. The Velocity of Light. Optical Activity. Friction. The Velocity of Sound. The Wave Length of Light. The Length of the Path of a Gaseous Particle. To these may be added the phenomena of radioactivity, etc. It is clear that in the present state of knowledge the consideration of most of these properties is uncalled for. However, it may perhaps be noted in passing that the com- pressibility of water is remarkably small, that of protoplasm even less.1 Hence even great changes in pressure do not readily damage the organism, and, indeed, a frog's muscle appears to function normally after undergoing a pressure of 500 atmospheres.2 Further, it is of decided consequence for many reasons that the optical properties of water are such 1 Henderson and Brink, American Journal of Physiology, XXI, 248, 1908. 2 Henderson, Leland, and Means, American Journal of Physiology, XXII, 48, 1908. THE ARGUMENT 268 that light readily penetrates it to considerable depths. As for color, landscape and modern chemical industry alike testify to the availa- bility of carbon compounds as its source. A final test of thoroughness may be based upon a consideration of other compounds and elements. Accepting the decision thai no other properties can be so important to an active, complex, and regulated mechan- ism as those possessed nearly or quite as maxima by water, carbonic acid, and the compounds of the three elements, what are the possibilities of obtaining the same char- acteristics from other substances ? So far as chemical substances are now known, the only compound which can be even considered on this score as a substitute for water in the environment is ammonia, and in many respects, no doubt, ammonia might serve as well.1 However, chemical proces- s 1 A full discussion of the properties of ammonia which qual- ify it as a substitute for water in the role of solvent and other- wise will be found in the article by E. C. Franklin, 'The Ammonia System of Acids, Bases, and Salts," American Chem- ical Journal, Vol. 47, p. 28.5, 191-2. In this paper the re- sults of a long series of investigations an- brought together. Especially important for the present purpose are the intro- ductory remarks. "The many striking analogies between liquid ammonia and water as electrolytic solvents have been emphasized by the writer and Ins co-workers in papers which have appeared from time to time during the past decade. In 264 THE FITNESS OF THE ENVIRONMENT being what they are, it is impossible to im- agine the presence of vast amounts of am- monia in an atmosphere, while the loss of the greater part of the energy which can be stored by tearing apart hydrogen and oxygen would be a very serious difficulty ; but the loss of substantially all the incomparable chemical activity of oxygen is to all appear- ances an insurmountable obstacle to the sub- stitution of ammonia for water in biological processes. From time to time, loose discussion has arisen among chemists as to the possibility of substituting another element for carbon in the organic cycle. Such speculations have never been serious, but they have at least all those properties which give to water its unique position among solvents, such as its abnormally high boiling point, its high specific heat, its high heat of volatilization, its high critical temperature and pressure, its high association con- stant,-its high dielectric constant, and its low boiling-point elevation constant, its power as an electrolytic solvent, and the facility with which it forms compounds with salts, liquid ammonia shows a remarkable similarity to water." " While the boiling point of liquid ammonia is 33.46° below zero, it still appears abnormally high when compared with the boiling temperatures of phosphine, arsine, stibine, me- thane, ethylene, hydrogen sulphide, hydrochloric acid, etc. The specific heat of liquid ammonia and the heat of fusion of the solid are greater than the corresponding constants for water or any other known substance, while its heat of volatilization, with the one exception of water, is the highest THE ARGUMENT 265 demonstrated that very few elements, prob- ably only silicon, and perhaps boron, can even be imagined in such a role. It h;i>. moreover, just been shown that there are many facts leading to the conclusion that only carbon among elements, and carbon itself only in conjunction with hydrogen, has the power to form the skeletons of compounds numerous, complex, and varied like those of organic chemistry. But, apart from this conclusion, it is certain that silicon and boron could not be mobilized like carbon. Quartz, of any known liquid. The critical temperature of ammonia is abnormally high, and its critical pressure — the more char- acteristic constant — is higher than that of any other liquid excepting water. Ammonia is an associated liquid, and its dielectric constant, though much below that of water, is still high when compared with that of non-electrolytic solvents. Its boiling-point elevation constant is the lowest of any known liquid, namely 3.4, as compared with .V2 for water. In its tendency to unite with salts and other compounds, it probably exceeds water, since salts with ammonia of crystal- lization are perhaps even more numerously recorded in the literature than are salts with water of crystallization. As a solvent for salts it is generally much inferior to water, though some salts, for example the iodides and bromides of mercury, lead, and silver, dissolve very much more abundantly in ammonia than they do in water, and it far surpasses the latter solvent in its ability to dissolve the compounds of carbon. Finally it exhibits conspicuous power as an ionizing solvent, the more dilute ammonia solutions at .'>:>. 5° being very much better conductors of electricity than aqueous solutions of the same concentration at 18°." £66 THE FITNESS OF THE ENVIRONMENT the oxide of silicon, is the most inert and immobile of rocks : the oxide of boron is only less available s a movable constituent of the environment : and there is no other stable compound of either element which can be compared with carbonic acid for its mobility. It must be remembered that this property is the result of two independent characteris- tics of the latter substance, its gaseous na- ture, and the precise degree of its solubility in water. Finallv. the regulation of the reaction of aqueous solutions by means of carbonic acid has to be taken into account. Hence it mav be concluded that hvdrosen, oxvsen. and carbon, water and carbonic acid, are not to be rivaled in their own qual- ities, even as these cannot be balanced bv others which they do not posse — On the whole, then, we mav believe that the m physico-chemical characteristics of material systems and material processes have been com- prehensively examined in the course of the present study. Accordingly, we may finally conclude that the fitness of water, carbonic acid, and the three elements make up a unique ensemble of fitness for the organic mechanism. The search, however incomplete, has certainly not overlooked properties so important and jmerous. or compounds and elements so TEZ ;i arises, of water, carbonic acid, m F : t 11 r : r 15 11: ri: -__.;_ lii rrl It 1 11 f . . - 1111 r H. Iifr a iMcfcww Inm point of vie-a- of p A - 1_r 268 THE FITNESS OF THE ENVIRONMENT a. Complex (physically, chemically, physiologically) . b. Durable, hence well regulated physico-chemically. This con- clusion applies to — 1. The Organism. 2. The Environment. c. Endowed with a metabolism. Hence there must be ex- change with the environment of — 1. Matter. 2. Energy. III. The primary constituents of the natural environment are — a. Water. b. Carbonic acid. IV. In places where life is possible the primary constituents of the environment are necessarily and automatically formed in vast amounts by the cosmic process. V. Water, carbonic acid, and their con- stituent elements manifest great fitness for their biologi- cal role. a. Water possesses a great number of unique or very unusual prop- THE AR(,!Mi;\T ?f>{) erties, e.g. thermal proper- ties, solvent power, dielectric constant, surface tension, which together result in max- imal fitness in certain respects, e.g. mobility, ubiquity, con- stancy of temperature and richness of the environment, richness of the organism in chemical constituents, variety of chemical processes, electri- cal phenomena, colloidal phe- nomena. b. Carbon dioxide possesses very un- usual properties, e.g. mag- nitude of absorption coeffi- cient, strength as acid, which together result in maximal fit- ness in certain respects, e.g. mobility, ubiquity, richness of the environment and organ- ism in other elements and compounds, constancy of re- action, etc. c. Chemical compounds containing carbon, hydrogen, and oxygen possess unique properties, e.g. number, variety, complexity, activity, variety of chemical 270 THE FITNESS OF THE ENVIRONMENT relations and reactions, heats of reaction, instability, etc., which together result in maxi- mal fitness in certain respects, e.g. as sources of matter and energy for the processes of metabolism, as sources of com- plex structures, as the means of establishing complex func- tions, etc. VI. Oceans are formed automatically in the cosmic process. VII. The ocean possesses unique properties, e.g. mobility, richness in dis- solved substances, durability, and stability of physico-chem- ical conditions, depending chiefly upon the properties of water and carbonic acid, which together result in maximal fit- ness in certain respects, e.g. as milieu, and as source of matter for the processes of metabolism, to moderate and equalize temperature, etc. VIII. The physical and chemical properties which have been taken into consideration include nearly all those which are known to THE ARGUMENT 271 be of biological importance or which appear to be related to complexity, regulation, and metabolism. IX. There are no other compounds which share more than a small part of the qualities of fitness of water and carbonic acid ; no other elements which share those of carbon, hydrogen, and oxygen. X. None of the characteristics of these substances is known to be unfit, or seriously inferior to the same characteristic in any other substance. XI. Therefore the fitness of the environ- ment is both real and unique. In drawing this final conclusion I mean to assert the following propositions: — I. The fitness of the environment is one part of a reciprocal relationship of which the fitness of the organism is the other. This relationship is completely and perfectly recip- rocal; J the one fitness is not less important 1 This is not to be understood as an assertion that the rela- tionship is symmetrical. The fad is that each uranism fits its particular environment, while tli«- environment in its moat general and universal characteristics fits the meal general and universal characteristics of the organic mechanism. 272 THE FITNESS OF THE ENVIRONMENT than the other, nor less invariably a constitu- ent of a particular case of biological fitness; it is not less frequently evident in the char- acteristics of water, carbonic acid, and the compounds of carbon, hydrogen, and oxygen than is fitness from adaptation in the char- acteristics of the organism. II. The fitness of the environment results from characteristics which constitute a series of maxima — unique or nearly unique prop- erties of water, carbonic acid, the compounds of carbon, hydrogen, and oxygen and the ocean — so numerous, so varied, so nearly complete among all things which are concerned in the problem that together they form cer- tainly the greatest possible fitness. No other environment consisting of primary constitu- ents made up of other known elements, or lacking water and carbonic acid, could possess a like number of fit characteristics or such highly fit characteristics, or in any manner such great fitness to promote complexity, durability, and active metabolism in the organic mechanism which we call life. It must not be forgotten that the possibility of such conclusions depends upon the universal character of physics and chemistry. Out of the properties of universal matter and the characteristics of universal energy has arisen THE ARGUMENT 073 mechanism, as the expression of physico- chemical activity and the instrument of physico- chemical performance. Given matter, energy, and the resulting necessity thai life shall be a mechanism, the conclusion follows that the atmosphere of solid bodies docs actually pro- vide the best of all possible environments for life. CHAPTER VIII LIFE AND THE COSMOS THE SIGNIFICANCE OF FITNESS A HALF century has passed since Darwin wrote "The Origin of Species," and once again, but with a new aspect, the relation between life and the environment presents itself as an unexplained phenomenon. The problem is now far different from what it was before, for adaptation has won a secure posi- tion among the greatest of natural processes, a position from which we may suppose it is certainly never to be dislodged ; and natural selection is its instrument, even if, as many think, not the only one.1 Yet natural selec- 1 Natural selection remains still a vera causa in the origin of species; but the function ascribed to it is practically reversed. It exchanges its former supremacy as the supposed sole determinant among practically indefinite possibilities of structure and function, for the more modest position of simply accelerating, retarding, or terminating the process of otherwise determined change. It furnishes the brake rather than the steam or the rails for the journey of life ; or in better 274 LIFE AND THE COSMOS tion does but mold the organism ; the environ- ment it changes only secondarily, without truly altering the primary quality of environ- mental fitness. This latter component of fit- ness, antecedent to adaptations, a natural result of the properties of matter and tli<" characteristics of energy in the course of cosmic evolution, is as yet nowise accounted for. It exists, however, and must not be dismissed as gross contingency. The mind balks at such a view. Coincidences so nu- merous and so remarkable as those which we have met in examining the properties of matter as they are related to life, must be the metaphor, instead of guiding the ramifications of the tree of life, it would, in Mivart's excellent phrase, do little more than apply the pruning knife to them. In other words, its functions are mainly those of the third Fate, not the first, of Siva, not of Brahma. — Patrick Geddes and J. Arthttb Thomson, 'Evolution." New York, Home University I.i brary, 1911, p. 248. "But as my conclusions have lately been much misrep- resented, and it has been stated that 1 attribute the modifica- tion of species exclusively to natural selection, I may be permitted to remark that in the first edition <»f this work, and subsequently, I placed in a most conspicuous position — namely, at the close of the Introduction — the following words: 'I am convinced that natural selection has been the main but not the exclusive means of modification." Charles Darwin, "The Origin of Species by Means of Natural Selection." New York, reprinted from the Sixth London Edition, The Home Library, pp. 495-496. 276 THE FITNESS OF THE ENVIRONMENT orderly results of law, or else we shall have to turn them over to final causes ! and the phi- losopher. There is, in truth, not one chance in count- less millions of millions that the many unique properties of carbon, hydrogen, and oxygen, and especially of their stable compounds water and carbonic acid, which chiefly make up the atmosphere of a new planet, should simultaneously occur in the three elements otherwise than through the operation of a natural law which somehow connects them together. There is no greater probability that these unique properties should be with- out due cause uniquely favorable to the organic mechanism. These are no mere acci- dents ; an explanation is to seek. It must be admitted, however, that no explanation is at hand. For the coincidence of properties itself a rational explanation based upon known laws of nature is perhaps conceivable. Attention has already been called to the interconnection of such properties as latent heat of vaporiza- tion, thermal conductivity, molecular vol- ume, the value of the van der Waals constant a, 1 Bacon compared final causes to vestal virgins. "Like them," he says, "they are dedicated to God, and are barren." — "The Advancement of Learning," Book II, p. 142. LIFE AND THE COSMOS 077 the dielectric constant, and ionizing power. Further, it is of course most probable that numerous other properties are necessarily associated with these; and finally it is nol surprising that elements of low atomic weight, which become concentrated in the atmos- phere on account of the small specific gravity of their gases, should possess unusual proper- ties, like high specific heat, or if one property leads to another, many unusual properties. Be that as it may, chemical science is still a very long way from accounting for the simul- taneous occurrence of the various character- istics of water, especially if we include such things as heat of formation, solvent power, the process of hydrolytic cleavage, the degree of solubility of carbon dioxide, the anomalous expansion on cooling near the freezing point, etc. There is, in fact, exceedingly little ground for hope that any single explanation of these coincidences can arise from current hypotheses and laws. But if to the coincidence of the unique properties of water we add that of the chemical properties of the three elements, a problem results under which the science of to-day must surely break down. If these taken as a whole are ever to be understood, it will be in the future, when research has pene- 278 THE FITNESS OF THE ENVIRONMENT trated far deeper into the riddle of the prop- erties of matter. Nevertheless an explana- tion cognate with known laws is conceivable, and in the light of experience it would be folly to think it impossible or even improbable. Such an explanation once attained might, however, avail the biologist little; for a further problem, apparently more difficult, re- mains. How does it come about that each and all of these many unique properties should be favorable to the organic mechanism, should fit the universe for life ? And for the answer to this question existing knowledge provides, I believe, no clew.1 Thus regarded, our new form of the old riddle appears twofold, and, on that account, for the present the more unanswerable. There is but one immediate compensation for this complexity ; a proof that somehow, beneath adaptations, peculiar and unsuspected relation- ships exist between the properties of matter and the phenomena of life; that the process of cosmic evolution is indissolubly linked with the fundamental characteristics of the organism; that logically, in some obscure 1 The great difficulty appears to be that there is here no possibility of interaction. In our solar system, at least, the fitness of the environment far precedes the existence of the living organisms. LIFE AND THE COSMOS 27Q manner, cosmic and biological evolution are one. In short, we appear to be led to t he assumption that the genetic or evolutionary processes, both cosmic and biological, when considered in certain aspects, constitute a single orderly development thai yields results not merely contingent, but resembling those which in human action we recognize as pur- poseful. For, undeniably, two things which are related together in a complex manner by reciprocal fitness make up in a very real sense a unit, — something quite different from the two alone, or the sum of the two, or the re- lationship between the two.1 In human af- fairs such a unit arises only from the effective operation of purpose. Now it is most clearly evident from the experience of centuries that ordinary teleoloirv is dangerous doctrine in science, and in the past, accidents apart, it has been invariably sterile.2 A statement that the legs have been formed for the purpose of locomotion, no doubt possesses scientific validity, if it be properly interpreted. But the real scientific concern is for the bones and muscles, the 1 This appears logically to correspond with the "schOp- ferische Synthese" of Wuixlt. 2 Interesting discussions bearing upon flu's lubjed will be found in Pearson's well-known "Grammar of Science." 280 THE FITNESS OF THE ENVIRONMENT tendons and ligaments which are employed in walking, and for the evolutionary process by which they have been adapted to their use. Nevertheless, biological science has not been able to escape the recognition of a natural formative tendency, which Darwin identified as the result of natural selection. And now it appears to be necessary to postulate a like tendency in the evolution of inorganic nature. We have found that the properties of the environment, biologically considered, present the same fitness as the properties of life. In each case the fitness results, at least in part, from an evolutionary process. Through the main lines of later development these are both known, though in both cases we stop short, perhaps far short, of the origins — the origin of life and the origin of the universe — if indeed they have ever originated.1 Can we then deny that in the one as in the other process there is a tendency, a bent, a direc- tion of flow or development?2 I think not, 1 It is hardly necessary to point out that the properties of the elements are themselves quite free from variation of any sort. 2 "Alike in the external and the internal worlds, the man of science sees himself in the midst of perpetual changes of which he can discover neither the beginning nor the end. If, trac- ing back the evolution of things, he allows himself to enter- tain the hypothesis that the universe once existed in a dif- LIFE AND THE COSMOS Si and it seems clear thai the facts of physical science call for an explanation of the tend- ency to fitness of the environment in the same way that formerly the facts of biologi- cal science called for an explanation of the tendency to fitness of the organism. To postulate such a tendency is, however, in itself rather a philosophical than a scientific act, and so, too, must he conjecture regarding the origin of fitness. It is open to any one who may be so minded speculatively to enrich this tendency with characteristics of any sort. He may follow the lead of M. Bergson and call it impetus, with all which that term now implies, or he may turn to natural theology and regard it as proof of supernatural purpose and design, or he may find a model for teleolog- ical views in many other quarters. But one thing is certain, no such discussion, be it ever so important to the philosopher or the theo- logian, can directly contribute to scientific knowledge and comprehension of the under- lying phenomena, which arc the sole positive and certain knowledge of the subject that fused form, he finds it utterly impossible to conceive how this came to !><■ bo; and equally, if he speculates on tin- rutu he can assign no limit to the L,rrand succession of phenomena ever unfolding themselves before him.*' Herbert Spbn< nt, "First Principles." Ww York, reprinted from the Fifth Loudon Edition, 1880. The Home Library, j>. 57. 282 THE FITNESS OF THE ENVIRONMENT we possess. For these facts an explanation of a different sort would be necessary, some- thing logically resembling natural selection, a natural process acting automatically through the properties of matter and energy, and never overstepping the limits of matter and energy, space and time; neither supernatural nor metaphysical, but purely mechanistic. Lacking any indication of what such an expla- nation may be, or how it is to be sought, we shall do well to turn to other considerations. n VITALISM All the skill of trained biologists, multi- plying and refining our knowledge of the forms of life, has even yet not availed to make clear the fundamental ideas of the science. Complexity exists here in the very nature of the case, and here, if at all, the complete subjugation of natural phenomena to physi- cal science may be expected to fail. In an earlier chapter the painful advance of physics and chemistry into the domain of biology has been sketched, and it was then shown how progress is beset with well-nigh insuperable obstacles. Thus it is that bio- logical thought has never attained to that LIFE AND THE COSMOS &9 finality which appears, at least by contrast, to characterize the greater body of opinions in physical science. In particular two extreme views, though often commingled, have continually striven for the mastery. The one of these, purely scientific and wholly positive, declares the phenomena of life to be, while partly unknown, ultimately knowable as manifestations of matter and energy. According to this view life is a mechanism and nothing more, in its positive scientific aspects at least. Without necessarily denying such assertions, the other view sees the unique properties of life to be dependent upon an equally unique force or tendency, operating in or through its physico-chemical organization. Either there is a peculiar vital force; or there is manifest in the organism a peculiar tendency ; or at any rate life patently follows the path into which it was propelled by an original impetus, peculiar to life, unknown in other phenomena. All such views inherently par- take of metaphysics, and have, therefore, ever aroused most determined opposition among the more orthodox devotees of science. Descartes appears to have been the first person to adopt the modern scientific attitude toward life, and from him a very large pro- 284 THE FITNESS OF THE ENVIRONMENT portion of French biologists, as well as those of other nations like Huxley and Du Bois- Reymond, derive their philosophical views concerning their science. Descartes per- ceived, apparently the first among the mod- erns, that the scientific explanation of vital phenomena must be a physical one, in terms of matter and motion. Far in advance of his time he applied such ideas to the nervous system, thereby establishing the nature of reflex action and invading the very citadel of animism. Outside natural science, how- ever, Descartes was far from being a mechan- ist. Since the early seventeenth century the conflict between vitalism and mechanism has ranged over the whole field of biology, and its history is most complicated. After Des- cartes, Lavoisier, by his studies of combustion within and without the body, made the next very important step. He was then followed by Liebig, Wohler, and a host of later chem- ists. In the main the growth of exact science has steadily delivered over one vitalistic strong- hold after another to the mechanists. And though in the first flush of triumph mechan- ism has sometimes seemed to gain more in a particular engagement than later proved to be the case, vitalism has perhaps not had a LIFE AND HIE COSMOS 285 positive success in three centuries. Such a history no doubt depends upon the very na- ture of the situation; upon the inherent and inevitable weakness, within the domain of science, of vital istic views. Experience seems to show that the only kind of hypothesis which can find conclusive scien- tific support, or sound basis in the phenomena of matter and energy, is a mechanistic hy- pothesis. Exact and positive knowledge can demonstrate scientifically the truth of no other hypothesis with the finality which char- acterizes its proof of a mechanistic theory. Hence, so far as it ventures into the field of science at all, a vitalistic theory, when attacked by science, cannot effectually avail itself of the weapons of the assailant, and can never make a powerful counter attack. Its only method consists in a determined resistance, yielding little by little before the advance of positive knowledge and never gaining new territory, nor, except by accident, regaining what it has lost. Where this process is to end; in what respect and how far life is des- tined ever to remain a scientific riddle, can only be surmised. The chief definitive triumphs of the mech- anistic view are two: the elimination of vital force and of a belief in peculiarity of chemical 286 THE FITNESS OF THE ENVIRONMENT composition from organic chemistry, through the actual successes of the laboratory in new syntheses; and the final recognition, based upon understanding of the principle of the conservation of energy, that, whatever else "vital force" may be, it is certainly not force, — a form of energy. Thus limited, vitalism has been obliged to take refuge in a more restricted belief; namely, that the organ- ism is somehow governed by a directive tendency which, like an architect, presides over its development ; but that meanwhile the manifold processes of life and evolution go on within the world of physical science just as the work of the builder conforms to the laws of mechanics, though following the plan of the architect. This view has been well stated by another great Frenchman, Claude Bernard: "Life is the directive idea or evolutive force of the being ; . . . but it would be an error to believe that this metaphysical force operates after the manner of a physical force. . . . The metaphysical evolutive force by which we may characterize life is useless to science, because, existing apart from physical forces, it can exercise no influence upon them. Hence we must here separate the world of meta- physics from the world of positive phenomena LIFE AND THE COSMOS 287 which serves it as foundation, but which has nothing to contribute to it. . . . Summaris- ing, if we can define life with the help of a special metaphysical conception, it is none the less true that mechanical, physical, and chemical forces are the sole effective agents of the living organism, and that the physiologist has to take account of their action alone. We shall say with Descartes, 'One thinks meta- physically, but one lives and acts physically.' ] Thus restricted, vitalism can apply only to formative processes and the like, though the vitalist still sees in the state of the organism 1 "Claude Bernard, 'La Science Experimentale,' 3me <■< art on pense metaphysiquement, mais on vit et on agit physique- ment.'" — Merz, "A History of European Thought in the Nineteenth Century." Edinburgh and London. L906, Vol. II, pp. 379-380. 288 THE FITNESS OF THE ENVIRONMENT effects of vitalistic control of its evolution, just as we perceive in a house not only the material structure, but the idea of the archi- tect. Further, the origin of life itself remains shrouded in mystery. Meanwhile, for most men physiology has become merely biophys- ics and biochemistry, and mechanism is un- doubtedly firmly established throughout every department of the science. Such limitations of the vitalistic hypothesis, damaging though they may be, do not de- stroy its claim to consideration as a controlling factor of the processes of evolution, embry- ology, repair, etc., in spite of the fact that even here it has suffered serious though less complete reverses. In 1859 Darwin's natural selection offered itself as a possible substi- tute for vitalism in a part or the whole of this field, and soon gained very general accept- ance. The survival of the fittest has now become in the judgment of all biologists an unquestioned force in the molding of life. Therefore, at best, but a restricted scope within its restricted field remains to vital- ism. From the earliest days of the new hypothesis it has been widely recognized that to accept the survival of the fittest as one factor in the adaptation of life to its environment is quite LIFE AND THE COSMOS ^S9 a different matter from proving il to be the only force which directs evolution. An early eulogy by Du Bois-Reymond upon the work of Darwin clearly discloses the nature of the situation: 'Here is the knot, here the great difficulty that tortures the intellect which would understand the world. Whoever does not place all activity wholesale under the sway of Epicurean chance, whoever gives only his little finger to teleology, will inevitably arrive at Paley's discarded 'Natural Theology,' and so much the more necessarily, the more clearly he thinks and the more independent his judgment . . . the physiologist may define his science as a doctrine of the changes which take place in organisms from internal causes. . . . No sooner has he, so to speak, turned his back on himself than he discovers himself talking again of functions, performances, ac- tions, and purposes of the organs. The possi- bility, ever so distant, of banishing from na- ture its seeming purpose, and putting a blind necessity everywhere in the place of final causes, appears, therefore, as one of the greatest advances in the world of thought, from which a new era will be dated in the treatment of these problems. To have some- what eased the torture of the intellect which ponders over the world-problem will, as long u 290 THE FITNESS OF THE ENVIRONMENT as philosophical naturalists exist, be Charles Darwin's greatest title to glory." 1 Recently the work of de Vries, "The Muta- tion Theory," has at length set forth a num- ber of trustworthy observations of the origin of species in plants with which natural selec- tion, in the restricted original sense at least, can have nothing to do. The origin of species by mutation consists in a sudden discontinu- ous variation, and selection, therefore, has no opportunity to operate upon a series of numerous minute variations which them- selves display no tendency of any sort what- ever, in the manner demanded by the Darwin- ian hypothesis.2 Hence it appears certain that natural selection cannot be regarded as completely master of the situation; apart from the origin of life there remains a lacuna in biology which for the present no existing mechanistic hypothesis can fill. Moreover, among other things, the ordinary processes of regeneration and repair have frequently been brought forward with some success as purposeful activities inexplicable 1 Du Bois-Reymond, "Darwin versus Galiani," "Reden," Vol. I, p. 211. Quoted from Merz, "History of European Thought in the Nineteenth Century," Vol. II, p. 435. To the same source I am indebted for several other quotations. 2 Hugo de Vries, "The Mutation Theory." Chicago, 2 vols., 1909, 1910 (trans. Farmer and Darbishire). LIFE AND THE COSMOS 29] by natural selection.1 Tims Du Bois-Rey- mond: "One of the greatest difficulties pre- sents itself in physiology in the so-called re- generative power, and — what is allied to it — the natural power of healing; {his may now be seen in the healing of wounds, in the delim- itation and compensation of morbid processes or, at the farthest end of the series, in the re-formation of an entire fresh-water polyp out of one of the two halves into which it had been divided. This artifice could surely not have been learned by natural selection, and here it appears impossible to avoid the assump- 1 "Still less explicable in any way thus far proposed are certain remedial actions seen in animals. An example of them was furnished in §67, where 'false joints' were de- scribed — joints formed at places where the ends of a brok. n bone, failing to unite, remain movable one upon the other. According to the character of the habitual motions there re- sults a rudely formed hinge-joint or a ball-and-socket joint, either having the various constituent parts — periosteum, fibrous tissue, capsule, ligaments. Now Darwin's hypothesis, contemplating only normal structures, fails to account for this formation of an abnormal structure. Neither can we ascribe this local development to determinants: there were no appropriate ones in the germ-plasm, since no such struc- ture was provided for. Xor does the hypothesis of phil- ological units, as presented in preceding chapters, yield an interpretation. These could have qo other tendency than to restore the normal form of the limb, and mighl be expected to oppose the genesis of these new parts." — HsBBBBT Spencer, "The Principles of Biology," Vol. I. New York and London, 1909, revised and enlarged edition, p. :>f Bergson'a impetus can be clearly perceived, it must remain scientifically an unsound hypothesis. 296 THE FITNESS OF THE ENVIRONMENT In such speculations the properties of matter and the process of cosmic evolution have no place.1 Bergson, indeed, very defi- nitely, and it would seem gratuitously, puts aside cosmic evolution and also, with certain slight reservations, the properties of matter as of no essential consequence in organic evolu- tion; e.g. "This twofold result has been ob- tained in a particular way on our planet. But it might have been obtained by entirely dif- ferent means. It was not necessary that life should fix its choice mainly upon the carbon of carbonic acid. What was essential for it was to store solar energy ; but, instead of ask- ing the sun to separate, for instance, atoms of oxygen and carbon, it might (theoretically at least, and, apart from practical difficulties 1 Driesch, to be sure, has considered the problem of uni- versal teleology, but unsuccessfully and with obvious vitalistic preconceptions such as individuality. His nearest approach to the thesis of the present work is to be found in the follow- ing lines: "I do not hesitate to confess that, apart from historical teleology relating to the sequence of one state of poli- tics or economy upon another, and apart from phylogeny, there seems to me to be a certain sound foundation in the concept of the general harmony between organic and inor- ganic nature, a something which seems to show that nature is nature for a certain purpose. But I confess at the same time that I am absolutely unable to consider this purpose in any other than a purely anthropomorphic manner." — L.c, Vol. II, pp. 348-349. LIFE AND Till: COSMOS possibly insurmountable) have put forth otln r chemical elements, which would then have had to be associated or dissociated 1>\* entirely dif- ferent physical means. And if the element characteristic of the substances that supply energy to the organism had been oilier than carbon, the element characteristic of the plastic substances would probably have been other than nitrogen, and the chemistry of living bodies would then have been radically different from what it is. The result would have been living forms without any analogy to those we know, whose anatomy would have been different, whose physiology also would have been different. Alone, the sensori-motor function would have been preserved, if not its mechanism, at least in its effects. It is there- fore probable that life goes on in other planets, in other solar systems also, under forms of which we have no idea, in physical conditions to which it seems to us, from the point of view of our physiology, to be absolutely op- posed. If its essential aim is to eat eh up usable energy in order to expend it in explo- sive actions, it probably chooses, in each solar system and on each planet, as it does on the earth, the fittest means to get this result in the circumstances with which it i^ confronted. That is at least what reasoning 298 THE FITNESS OF THE ENVIRONMENT by analogy leads to, and we use analogy the wrong way when we declare life to be impos- sible wherever the circumstances with which it is confronted are other than those on the earth. The truth is that life is possible wher- ever energy descends the incline indicated by Carnot's law and where a cause of inverse direction can retard the descent — that is to say, probably, in all the worlds suspended from all the stars. We go further : it is not even necessary that life should be concen- trated and determined in organisms properly so called, that is, in definite bodies present- ing to the flow of energy ready-made though elastic canals. It can be conceived (although it can hardly be imagined) that energy might be saved up, and then expended on varying lines running across a matter not yet solidified. Every essential of life would still be there, since there would still be slow accumulation of energy and sudden release." * B VITALISM AND TELEOLOGY These conclusions appear to be based upon decisions regarding the essential physico-chem- ical conditions and characteristics of life arbi- 1 Bergson, l.c.y pp. %55, 256. LIFE AND THE COSMOS trarily reached in accordance with precon- ceived views, and quite without scientific justification. There is certainly no reason to ascribe greater importance to energy than to matter in the vital processes, and in the light of the facts with which the preceding chapters are concerned, such views seem absurd. Indeed, whoever is disposed to spec- ulate about biological fitness — and not even the incomparable finesse of M. Bergson's dia- lectic can make fitness other than the most general result of the process of organic evolu- tion— must now weigh well the cosmic pro- cesses. For, if allowance be made for the results of natural selection in the organic world, fitness of the environment has the greater claim to be considered. The two fitnesses are complementary; are they then single or dual in origin ? The simple view would be to imagine one common impe- tus operating upon all matter, inorganic and organic, through all stages of its evolution, in all its states and forms, and leading to worlds like our own through paths apparently purposeful and really not yet explained. Such, it seems to me, is the natural hypothesis for the vitalist to adopt. But then vitalism vanishes, only teleology remains; for the unique characteristic of life is gone. Vet, 300 THE FITNESS OF THE ENVIRONMENT putting aside mechanistic differences, is it not now lost in any case ? Has not modern vital- ism in accepting the limitation to entelechies or impetus destroyed itself ? The situation, briefly, seems to be as fol- lows : two evolutionary processes independ- ently result in two complementary fitnesses; hence they are related. In the one process the origin of fitness is in part explained by a mechanistic hypothesis. Nevertheless, many philosophers, as is their right, declare that in this process a further extraphysical influence is to be assumed. But any one who makes such an assumption for the one process must certainly now make it for the other; thus he will be led to see impetus or entelechies every- where. Under these circumstances it may be doubted if his acquaintance with the na- ture of his impetus or entelechies is so inti- mate that he will be able to distinguish the inorganic from the organic, for he has surren- dered to science all the positive physico-chem- ical differences between organic and inorganic bodies and processes. Hence, unless he is to make an arbitrary and unintelligible distinc- tion, or to indulge in the spinning of cob- webs, his vitalism has ceased to be exclusively organic, in short has ceased to be vitalism at all, and has become mere universal teleology. LIFE AND THE COSMOS 301 III COSMIC EVOLUTION But, for the scientist, these are matters of little moment, lie, at least, is not obliged to take any stand concerning them. This could hardly be better illustrated than by our new facts themselves. For it seems to be clear that where science is most self-sufficient, at the very basis of physical science itself, if anyiohere, teleology is at work. Yet it is certain that physical science needs no tele- ology to explain its phenomena and pro- cesses. These are mechanisms, and since the publication of Newton's "Principia" no one has seriously doubted the fact.1 To-day there is as little room for doubt thai a complete description of cosmic evolution in terms of matter and energy is possible; for it is sound scientific doctrine that what exists in the finished solar system depends upon what already existed in the nebula. The forms and states and quantities of matter and energy in the nebula determine the re- sulting solar system. Further, since both nebulae and solar systems are common oc- 1 Laplace's reply to a question of Napoleon's, "Why the name of God did not occur in his ' Rlecanique Odette,' will be recalled : "Sire, je n'ai pas besom de cet hypothei 302 THE FITNESS OF THE ENVIRONMENT currences, it is evident that nebulae them- selves are in a general way determined by other antecedent conditions and phenomena, which turn out to be collisions between stars. Thus arises the suspicion that cosmic evolu- tion may be in truth a cyclic process which had no beginning and can have no end.1 An alternative hypothesis regards the pres- 1 Such a view, until quite recently, was universally rejected because it appeared to conflict with the second law of thermo- dynamics, — that of the degradation of energy. But lately it has been put forth by no less an authority than Arrhenius, who has advanced a theory to explain away the difficulty of the second law. "The recognition of the indestructibility of energy seemed to accentuate the difficulties of the cosmogonic prob- lems. The theses of Mayer and of Helmholtz, on the man- ner in which the Sun replenished its losses of heat, have had to be abandoned. My explanation is based upon chemical reactions in the interior of the Sun in accordance with the second law of thermodynamics. The theory of the 'degra- dation' of energy appeared to introduce a still greater diffi- culty. That theory seems to lead to the inevitable conclu- sion that the Universe is tending towards the state which Clausius has designated as 'Warme Tod' (heat death), when all the energy of the Universe will be uniformly distributed through space in the shape of movements of the smallest particles. That would imply an absolutely inconceivable end of the development of the Universe. The way out of this difficulty which I propose comes to this : the energy is * de- graded' in bodies which are in the solar state, and the energy is 'elevated,' raised to a higher level, in bodies which are in the nebular state." — Arrhenius, "Worlds in the Making," translated by Boras. New York and London, 1908, p. xiii. LIFE AND THE COSMOS 303 ent form of our universe as the result of a gradual evolution from an earlier unknown form, the development of successive solar systems being mere incidents of the larger process, the evolution as a whole directively governed by the law of the degradation of energy. A THE PERIODIC SYSTEM In either hypothesis the remarkable sys- tematic relationship between the elements which is manifest in the periodic classifica- tion has a peculiar place. If the second hy- pothesis be accepted, there seems to be little room for doubt that at an early period the chief cosmic process wras the evolution of the elements themselves; and in the first theory the nebula, wdiose properties depend almost wholly upon chemical constitution and chem- ical and molecular energy, occupies a unique position, like the leaf in the organic cycle, or spring among the seasons. Thus, whether or not the periodic system is to be regarded as the one remaining plain result of a process bv which the elements were evolved, at leas! it takes precedence over the other properties of matter, and lies at the very foundation of the known processes of evolution. Clearly, 304 THE FITNESS OF THE ENVIRONMENT no one can doubt that upon the properties of matter as determined by the periodic system, and upon the relative amounts of the different elements, the actual process of cos- mic evolution from nebula to solar system is dependent.1 Hence, in accordance with the general method of science, we must assume that the origin of environmental fitness lies at least as far back as the phenomena of the periodic system, at least as far back as the evolution of the elements, if they were ever evolved. We simply cannot doubt that the origin of a body like the earth depends exclusively upon chance plus the properties of the elements, their relative amounts, the indestructible forces of nature, and the other known factors of mechanism. The perfect induction of phys- ical science, based upon each and all of its countless successes in every department of physics and chemistry, conclusively proves that the whole process of cosmic evolution from its earliest conceivable state to the pres- ent is pure mechanism.2 1 The same considerations apply to any other scientific hypothesis of the genesis of the solar system. 2 Not only is this proved by all experience of physical science, it has also ever been the necessary working hypoth- esis of physicists and chemists. LIFE and THE COSMOS 805 B TKU.nl ni;v If, then, cosmic evolution be pure mechan- ism and yet issue in fitness, why imi organic evolution as well? Mechanism is enough in physical science, which no less than bioloj ical science appears to manifesl teleology; it must therefore suffice in biology. Thus once more we arrive a1 the negation of vitalism. For this conclusion we possi two arguments: the argumenl that in such aspects as concern physical science, and apart from differences scientifically explicable, or- ganic and inorganic phenomena arc alike and therefore a specifically vital teleology is un- necessary; and the argument thai inorganic science unquestionably has no need of non- mechanistic teleology. Hence we are obliged to conclude that all metaphysical teleology is to be banished from the whole domain of natural science.1 What then becomes of fitness? Clearly there are two logical possibilities. Either there exists an unknown mechanistic explanation of that common issue of the organic and cosmic 1 Such at least is the simplest provisional hypothesis, :m gratuitous assumptions. It ii therefore the one which must DOW !><• adopted. 306 THE FITNESS OF THE ENVIRONMENT evolutionary processes, or there does not. If such an explanation be possible, at least it must be admitted that it is very hard to con- ceive. Yet, recalling the difficulty before the idea of natural selection arose of imagining any mechanistic explanation whatever of fitness, we shall do well not to decide against such a possibility. On the other hand, it is conceivable that a tendency could work parallel with mechanism without interfering with it, according to a view which has been held by such thorough- going mechanists as Descartes, Claude Ber- nard, Virchow, DuBois-Reymond, and many another. Although I have no intention of here seeking a choice between these two hy- potheses, being in fact convinced that now, at all events, no choice is scientifically possi- ble, and doubting if properly speaking they are alternatives at all,1 I do feel concerned to 1 "Either the multitudinous kinds of organisms which now exist, and the far more multitudinous kinds which have existed during past geologic eras have been from time to time sep- arately made, or they have arisen by insensible steps, through actions such as we see habitually going on. Both hypotheses imply a Cause. The last, certainly as much as the first, recognizes this Cause as inscrutable. The point at issue is, how this inscrutable Cause has worked in the production of living forms. This point, if it is to be decided at all, is to be decided only by examination of evidence. Let us inquire which of these antagonistic hypotheses is most congruous LIFE AND THE COSMOS 807 remove from the latter view, if I may, some <>f the objections which are commonly raised against it in scientific circles, conscious thai in this attempt I am overstepping the bound- aries of natural science. It is evident that a perfect mechanistic description of the building <>f a house may be conceived. Within the world of physical science the whole process is logically complete without consideration of the architect's de- sign and purpose. Yet such desij and purpose, whether or not in themselves of mechanistic origin, are at one and the same time determining factors in the result, ;md nowise components of the physical process. Now it seems clear that a similar effect of a tendency working steadily through the whole process of evolution is also at least conceive- able, however small its bearing upon science, provided, like time itself, it be a perfectly independent variable, making up. therefore, with time the constant environment, so to speak, of the evolutionary process. The tend- ency must not be demonstrable either by weighing or by measuring, else it would amount to an interference within the mech- with establish^! facts." — HSBBMBT SpENCBB, 'The Prin- ciples of Biology." Men York And London, L909, V6L I. revised and enlarged edition, p. 116. 308 THE FITNESS OF THE ENVIRONMENT anistic process, and it must not be itself liable to any kind of variation whose detec- tion would directly reveal it. Where then can the origin of such a tendency be located? Why clearly, if we accept the induction in favor of mechanism, only where Bergson has shrewdly placed his vital impetus, at the very origin of things, just before mechanism begins to act. In short, our new teleology cannot have originated in or through mechanism, but it is a necessary and preestablished asso- ciate of mechanism. Matter and energy have an original property, assuredly not by chance, which organizes the universe in space and time. This is in very truth a metaphysical doc- trine; but it has strong claims to sympa- thetic regard from men of science. In the first place, it leaves mechanism with the perfectly free hand which that process has undoubtedly earned in the world of phe- nomena. Secondly, it does but add one further riddle, and that an old and familiar one, to those two already tacitly recognized by most scientists : the existence of the universe and the existence of life. Given the universe, life, and the tendency, mechanism is inductively proved sufficient to account for all phenomena. The existence of the universe, on the other LIFE and thi: COSMOS hand, is no concern of the scientist. What- ever else it may achieve, mechanism can never explain, cannot even face the problem of the existence of matter and energy, Within the world of science these are conserved; only outside thai world can they have originated or not originated. As for the existence of life, in spite of our utter ignorance, if must be admitted that a half century has greatly diminished the number of substantial biolo- gists who really look forward to its scientific explanation, and the greatesl chemists have ever shared such a view. Liebig is reported by Lord Kelvin to have replied to the ques- tion whether he believed that a leaf or a Bower could be formed or could grow by chemical forces, "I would more readily believe thai a book on chemistry or on botany could grow out of dead matter."1 Darwin, too, once said, "It is mere rubbish tihinlrfng at present of the origin of life; one might as well think of the origin of matter."1 Since Liebig's day the chemical organization of the cell lias become in scientific knowledge vastly more complex than it was before, and I know of DO biological chemist to whom the spontaneous, 1 Lord Kelvin, "On the I)i>sipat ion of Energy," IV. ;.ii!ar Lectures, Vol. III. p, 164. 2Merz, Vol. II. p. I 06. 310 THE FITNESS OF THE ENVIRONMENT that is to say, the mechanistic, origin of a cell is scientifically imaginable,1 though all believe that once formed, cells exist as mechanisms in a mechanistic universe.2 Thus the chem- ist puts his mind at rest regarding the exist- ence of life, just as the physicist calms his regarding the existence of matter, simply by turning his back on the problem. Thereby he suffers nothing in his practical task as a man of science. Returning now to fitness, we may be sure that, whatever successes science shall in future celebrate within the domain of teleology, the philosopher will never cease to perceive the wonder of a universe which moves onward from chaos to very perfect harmonies, and, quite apart from any possible mechanistic 1 This is not to express an opinion concerning the problem of abiogenesis ; all admit that we cannot disprove such a theory. But while biophysicists like Professor Schafer follow Spencer in assuming a gradual evolution of the organic from the inorganic, biochemists are more than ever unable to per- ceive how such a process is possible, and without taking any final stand prefer to let the riddle rest. But if life has originated by an evolutionary process from dead matter, that is surely the crowning and most wonderful instance of teleology in the whole universe. 2 See, for instance, F. Hofmeister, "Die Chemische Organ- isation der Zelle," Vieweg, Brunswick, 1901, and Alsberg, "Mechanisms of Cell Activity," Science, pp. 97-105, July 28, 1911. LIFE AM) THE ( OSMOfi ! 1 explanation of origin and fulfillment, to feel it a worthy subjecl of reflect ion. From t bis point of view, however, science need i cf no inter- ference, but without any lasl v< of former shackles may pursue the search after mechan- istic explanations of all natural phenomena At length we have reached the conclusion which I was concerned to establish. Science has finally put the old teleology to death. Its disembodied spirit, freed from vitalism and all material ties, immortal, alone li\- - on, and from such a ghost science has noth- ing to fear. The man of science is not even obliged to have an opinion concerning its reality, for it dwells in another world where he as scientist can never enter. 1 "An evolution is a series of events that in itself M M ri* ■ is purely physical, — a set of necessary occurrences in the world of space and time. An egg develops into s chick; a poet grows up from infancy; a nation emerges from bar- barism; a planet condenses from the Quid state, and develops the life that for millions of years makes it so prondrous s pin Look upon all these things descriptively, and you shall - nothing but matter moving instant after instant, each con- taining in its full description the necessity of ps - into the next. Nowhere will there be. for descriptive seien any genuine novelty or any discontinuity admissible. Hut look at the whole appreciatively, historically, synthetically, as a musician listens to ■ symphony, as s a drama. Now you shall seem to hs n. in phenoi form, a story." — ROYCU, "The Spirit of Modern I'hilosoph; Boston and New York, 1890, 8th ed., p. 1 312 THE FITNESS OF THE ENVIRONMENT I cannot hope to have provided more than a very imperfect illumination of certain as- pects of teleology in this venture upon the foreign field of metaphysics, and I should wish to be understood as very doubtful of my success in stating what seem to me some of the philosophical conclusions to be drawn from the fitness of the environment. There is, however, one scientific conclusion which I wish to put forward as a positive and, I trust, fruitful outcome of the present investigation. The properties of matter and the course of cosmic evolution are now seen to be intimately related to the structure of the living being and to its activities; they become, therefore, far more important in biology than has been previously suspected. For the whole evolutionary process, both cos- mic and organic, is one, and the biologist may now rightly regard the universe in its very essence as biocentric. jam/an xaauww PKOPERTT LMAST K. C Sf '*«' TNDEX Absorption coefficient, 13i tables, 137. Acetylene, 200. Acidity, 140, 142, 143. A. ids, 144, 157, 212, 213, 216, 217. Adaptation, 5, 36, 66, loll. 274. Adsorption, 128, 129, 130. Air, 135. tables, 135. Alcohols, 212. Algol, 46, 47. Alkalinity, 142, 143, 155, 167-170, 188. blood, 155-159. sea water, 167-170. table, 169. Ammonia, 66, 110, 263, 264. Analysis of evidence, 250-253. Animism, 284. Astronomy, 38-49. Asymmetric carbon atom, 223. Atlantic, 167, 168. Atmosphere, 55-60, 134, 135. tables, 135. Atomic volume, 10, 11. curve, 11. Atomic weights, 14. Avogadro's Hypothesis, 177. Balanced solutions, 175. Baltic, 168. Bicarbonates in blood, 157. Biocentric point of view, 110, 312. Biological chemistry, 19 ; Black Sea, 168, 169. Blood, 115, 116, 153, 155-15S, 161. alkalinity, 155-158, 187. serum, 116. tables, 116, 187. Boron, 265. Bosphorue, 168. Bottom water, l( Boyle'i Law, 177. Bromine, 209. Calcium oarbonate, 172, 173. Caloric, 81. Cane sugar, l Capillary action, 78, 126, I Carbohydrates, 218, complexity of reaction instability, 223-221 mutarotation, 21 photosynthesis, 227 Carbon, 55, 56, 64, 211. 246. constituent of environment, I in stars, 55, ">(">. source of energy, 2 16. unique chemical properties, 21 1 . Carbon chains, 210. Carbon oompoun 194, 246. Carbon dioxide, Chapter IV (133-163), 56, 61, 64, 65, 61 absorption coefficient, 136-140. acidity, 140 168. atmospheric, 184, 186. distribution, 138, I ration, 189, 140, in sea witter. 1 70. metabolism, 182, 188. necessary cmnpom itmos- phen r. gul it* - neutrality, 1 17, solubility, 186 I M) Carl ><>nie acid iioridr. < '- lestial i 163. ( Sbaracteristiol of lif Inoompleteness <>f. ( 'hernieal propert 20-.' -:. 313 314 INDEX Chemistry, Chapter VI (191-248). inorganic, 237-243. organic, 191-237. Chlorine, 209, 242. Chlorophyll, 230, 240, 241. Circulation of water, 91, 180-182. Classification of organic com- pounds, 212. Climate, 87. table, 87. Coagulation, 90. Coal, 57. Colloids, 77, 123, 128, 129, 130. Color, 263. Combustion, 25. Complexity, 31. Compounds of carbon, hydrogen, and oxygen, 202-207. Compressibility, 262. Concentration, 171. Conditions, 31, 164-180. Conservation of energy, 15, 18, 25, 286. Constitutional formulas, 197-207. Contraction, see Expansion. Copper, 241. Cosmic evolution, 131, 301-312. Cosmography, 39, 60, 61. Cycle of matter, 26-28. Cyclic compounds, 200, 201, 219. Degradation of energy, 18. Design, 85, 307. Dew, 105. Diabetic coma, 157. Dielectric constant, 121, 122, 125. table, 125. Digestion, 232. Distribution, 128, 138. Double bonds, 200. Dulong and Petit's Law, 82, 83, 84. Durability, 31. Dynamics, 2, 16. Earth, 61-63. Electricity, 256. Electro-physiology, 124. Elements, 72, 73. chemical, 9-15. figure, 11. table, 14. Energy, 15-19, 68, 69, 237. Entelechy, 293, 300. Environment, Chapter II (38- 71), 8-21, 32, 33, 183-190. possible, 49-51. primary constituents of, 61-63. Enzymes, 90, 230, 231. Ethane, 198. Ethylene, 200. Evaporation, 92, 97, 98, 101, 102, 103, 127, 174, 180. latent heat of, 97-103. table, 99, 100. Evolution, 278, 279. Excretion, 139. Exhaustiveness of treatment, 253- 267. Expansion, 106-110. table, 107. Fauna, 186. Figure, the atomic volume curve, 11. Final causes, 4, 276. Fitness, Chapter I (1-37), 4-8, 65, 66, 69, 131, 132, 266, 267, 274-282, 305-307. explained by Darwin, 5. Flora, 186. Form, 24, 26, 32. Formaldehyde, 225, 228, 229. Freezing point, 93, 94, 178, 179. depression, 178, 179. tables, 178, 179. table, 94. Frog's muscle, 262. Fusion, 92-94. latent heat, 92-97. table, 95, 96. Gastric juice, 242. Gay-Lussac's Law, 177. Geology, 73, 112, 173. Geophysics, 52-55. Germanium, 12, 13. table, 13. Glucose, 160, 222-227. formation, 227. mutarotation, 223. reactions, 224-226. Glycerophosphoric acid, 241, 242. INDIA 9 1 5 Glycogen, 234. Gulf Stream, 1*83. Hsamocyanine, 241. Haemoglobin, 'J 1 1 . Heat of combustion, 80, 217, 246. table, 246. Hoat of formation, 244 247. tables, 244, 247. Hoat of reaction, 235, 236, 243 247. table, 236. Helium, 41, 42, 58. Henry's Law, 137. Herring, 175. Heterogeneous equilibrium, 151, 158. Hexakontanr. 210. Hoxanos, 198, 199. Hippuric acid, 234. Histidine, 226. Homogeneous equilibrium, 149. Hydrocarbons, 197-202. Hydrogen, 55, 56, 58, 64, 211, 237-243, 245. ions, 141-143, 169. in sea water, table, 169. Hydrogen sulphide, 152. Hydrolysis, 159, 232-237. Hydroxyl ions, 141-143. Impetus, 281, 293-295. Inactivity, 147. Inorganic chemistry, 287—243. Instability, 221. Interstellar Bpace, 60. Iodine, 209, 242. Ionization, 118 126, 155. constant, 140-163, 216. tables, 144, 216. water, 141-143. Iron, 241. Lactic acid, 2l'"> Latent neat, 92 106, 12 tables, 95. 96, 99, 100. Lesina, 166. table, 16 Levulose, 22 I -26. Life, origin of, 288, 309, 310. bic1 Liu . 1*6. I rj dc Bruyn'a phenomenon. 224. ium, 24 1 . Mam, - i .,f, li.v M <.-- Law, 16 Mlltter, 8 15. Me. ha, Mechanism, 2£ 01. Mediterranean, i • . }, 174. Melting m. Melting point, m I ■ ■ wimQ point. Metabolism, 24-2J 168. urology, 57, 5S : Mi th.-.r,,-, L97. Methyl formate, Methyl glyoxal, . Methyl imidasol, 226. Mobility, 139, 184. Mobilisation, 1 15, 141. Molecular constitution, 28, 196- 210. Molecular volume, 125. Morphology, - Mutarotataon, Mutation, - iral selection, l, 166 2 si iral the.,1, v bubs, 4 Niptuiii-m. 73. 'it;.. 1 12, 1 I! 1 If ilataon, l '< ■ Nitrogen, 211. impounds of, . 19. North Sea, 168, 169. Nucleic acids, 241 . m, Chapter V • » 190). 18 li.;. 114, l.v alkalinity. 167 171. table, I- as environment, lv'%-190. 316 INDEX Ocean, concentration, 171-176. table, 171. currents, 88, 180-182. mobility, 180-182. osmotic pressure, 176-179. tables, 178, 179. regulation of conditions, 164- 180. temperature, 165-167. tables, 165, 166, 167. Oceanography, 165. (Edema, 161. Optical activity, 223. Order, 1. Organic chemistry, 28-30, 191- 237. Organism, 21-36, 63, 76. Osmotic pressure, 176-180. Oxidation, 25, 246, 247. Oxygen, 64, 211, 237-243. Paleocrystic ice, 108. Panspermia, 50. Paraffine hydrocarbons, 212, 214. Partial pressure, 137. Periodic system, 9, 11, 14, 210, 211, 303, 304. figure, 11. table, 14. Pharmacological action, 175. Phase Rule, 257-261. Phosphoric acid, 146, 147, 156. Phosphorus, 241, 242. Photosynthesis, 27, 227-231. Physical chemistry, 256, 257. Physiology, 22, 123, 124. Planets, 46, 60. Possible environments, 49-51. Primary constituents of the en- vironment, 61-63. Propane, derivatives of, 203-205. Properties of matter, 70. interconnection, 276, 277. omitted, 261, 262. Proteins, 156, 242. Protoplasm, 26, 129, 130, 153, 155, 156. Purpose, 1, 307. Radicals, 211-218. Rain, 105. Red Sea, 174. Reduction, 227, 228, 231, 244, 247. Reflex action, 284. Regeneration, 290-292. Regulation, 31, 164-180, 186, 189. Repair, 290-292. Respiratory center, 157. Rivers, 113, 172, 180. composition, 113. table, 113. Salinity, 113, 114, 171-175, 187. table, 171. Salt, 113, 114, 172, 173. Sea urchin, 175. Sea water, 114, 153, 154, 155, 185. a balanced solution, 175. Silicon, 66, 265. Skager-Rack, 168. Soil, 78, 127. Solar system, 60, 61. Solidification, 108. Solubility, 136-138, 140. of carbonic acid, 136-140. Solvent, 68, 79, 111-118, 121. Sound, 256. Space, 19-21. Specific heat, 67, 68, 80-91. Stability, 78, 150, 164, 186, 218. Stars, 41-44, 50. Sugars, 218, 222-232. Sulphur, 209, 242. Sulphureted hydrogen, 152. Sulphurous acid, 169. Summary, 267-273. Sun, 44-46, 50, 60. Surface temperature, 166. Surface tension, 126-130. Survival of fittest, 288. Synthesis, 192, 286. Tables: Absorption coefficients, 137. Alkalinity and acidity, 148, 150. Alkalinity of sea water, 169. Comparison of blood and sea water, 187. Comparison of properties, 125. Composition of air, 135. INDEX 317 Composition of blood strum, 116. Composition of river water, 113. Composition of sea water, 171. Derivatives of propane, 203- 205. Elements, 14. Expansion of water, 107. Fitness, 250-252. Freezing point of blood serum, 178, 179. Germanium, 13. Heat conductivity, 100. Heat loss of dog, 103. Heats of combustion, 245. Heats of formation, 247. Heats of reaction, 236. Heats of reduction, 244. Ionization constants, 144, 216. Latent heat of fusion, 95, 96. Latent heat of vaporization, 99, 100. Melting points, 94. Normal temperatures, 87. Properties omitted, 261, 282. Range of temperature, 165, 166, 167. Specific heats, 81, 83. Surface tension, 126. Vapor tension, 105. Teleology, 279-282, 289, 298-300, 301, 305-312. Temperature, 67, 137, 153, 165, 166, 167. 170, 188. range, 165, 166, 167. tables, 165, 166, 167. tables, 87, 165, 166, 167. Temperature regulation, 70, 86, 87, 89, 91, 95,98, 102, 108, 109, 167. Theory of solution, 177, 179. Thermal conductivity, 106, 125. table, 106. Thermal properties, 80-110. Th.-rmorhpniistrv Therniodynamics, 16, 18, vj, ; Tim.-, 10 21. :<■ winds, 181. Treble bonk, 200. Orea, 102, 284. Urine, 117. Valence, 100, 197. Y:ui (l.-r Waals's <•<»! Vaporization, Vapor t.-ii.-ioii, 104, ! table, 106. Variable stars, 40, 47. Variety, 08, 207, 220. Velocity of reaction, 90, 94, 159, 170, 171. Vital force, 101, 2E Vitalism, 282-800, 806. of Bi-rgaon. - «. Volcanoes, 56, 134. Water, Chapter III (72-182), 57, 61-67, 131 ; transpan d 100; as element, 72, : universal important 78; Quantity and distri- bution, ~ ohettiosl inertness, 7'.' ; thermal pi erties, SO 1 10 ; beat, sl> 01 t upon tnnperaturr, s»i \>\ : Lit