The causes and course of organic evolution, a study in bioenergics.

Preface

In sending out the present volume as an addition to the literature of evolutionary science, the writer feels that a few words of personal explanation may be appropriate.

As a university student he was deeply impressed, during the epochal period from 1875 to 1882, by the varied—often conflicting—views on evolution presented by his teachers. So he early formed a resolve to investigate quietly, but as extensively as possible, the problems presented by organisms from the simplest types up to man himself. Later as a teacher and investigator of biological and specially of botanical problems, during a period of well nigh forty years, he has had considerable, and in some respects exceptional, opportunities for becoming acquainted with organisms over extensive fields of the Old and New World; for studying the structure and behavior of these in laboratory, in experimental grounds, and in cultural houses; as well as for quiet study of biological literature personally and in conjunction with his graduate students.

Though a student and teacher of zoological as well as botanical science for about ten years of his earlier academic career, he would in many ways have preferred to confine attention to the latter science, that has been his chosen field for specialized investigation. But questions constantly presented themselves from the side of plant life, that seemed to demand continuous study and solution from the animal side as well. If one such case may be cited here, reference might be made to the past and present geographical distribution of plants as eminently favoring the view that these originated over fresh-water areas, and only secondarily spread into the sea. As the truth of this became increasingly evident, the writer was confronted by the universally accepted though opposing view of zoologists that animal life originated in the sea, and thence spread landward. Here then was a seeming biological anomaly that called for investigation. The results of such are shortly pre-

Vi vi PREFACE sented in succeeding chapters of this work, and though the " writer’s conclusions are at variance with past current opinion, he trusts that the facts advanced will be weighed in that spirit of fairness and impartiality that should characterize all scientific method.

But, as has often been emphasized during the past quarter century, it is impossible to bring together a book like the present, with pretension at the same time on the part of the author to the knowledge of a specialist in every department of biological inquiry. The facts of Botany and Zoology are now so detailed and elaborated that few can hope to cover more than a very limited field with the knowledge and discerning keenness of an extreme specialist. This need not militate however against the making of an effort to link together isolated facts or groups of facts, if thereby the bounds of biological knowledge can be extended. But in saying this the writer is deeply conscious that not a few shortcomings or mistakes may be revealed in the succeeding pages, when these are perused by the specialist. For, as a specialist himself in _— some departments of botanical science, he has ever found the above to be true of the efforts even of the most distinguished investigator. He would therefore ask a degree of consideration on the ground that his whole aim has been to advance truth and knowledge, even though at the cost of cherished opinions or convictions of the past.

During the progress of his investigations he has been increasingly impressed by three outstanding characteristics of organisms. First and of extreme importance is the principle of continuity as traced from the simplest organized body to the highest expressions of moral and religious organization in man. Second, as the subtitle of this work and practically every chapter of it emphasize, the outstanding phenomenon of organic existence is not passive inert matter but energy, that continuous but ever transformable and subtle agency that starts and effects all cosmic changes. So energy, continuity, evolution may be said to constitute the triune basis of ex- . istence, and together form the keynote to this volume.

PREFACE vil

The work as now presented was practically completed in June, 1914, and in that month a sealed copy was deposited in the vaults of one of the learned societies, where it now remains. But some statistics and results bearing on plant and animal distribution, as well as some questions dealt with in Chapter X, had to be supplied later. These were added to and deposited with the above in July, 1916. Since that time only a few footnotes have been appended in small type and while the work was passing through the press. So the conclusions presented in Chapters XXI to XXX have been in no way determined by the iniquitous international struggle of the past three years. The writer cannot but hope, however, that some of his views may find the more ready acceptance, in virtue of the melancholy spectacle presented by the nations in this Twentieth Century of, Christ’s Era.

The writer would now make hearty and grateful acknowledgment of the kind help and encouragement extended by his friends Dr. Henry W. Cattell and Mr. Roberts Le Boutillier, for without their sympathetic aid this volume could not now have seen the light. He also expresses sincerest thanks to his friend Dr. Joseph Hepburn, who willingly volunteered to revise proofs, and by his expert chemical knowledge has furnished not a few valuable corrections and suggestions from the chemical standpoint.

In conclusion the writer desires to state that he would gladly welcome suggestions from every source, whether these may differ from orconform to the views now expressed. For, if strength, length of days, and needed leisure are granted him, he trusts in future volumes to ney, and further extend present conclusions.

Philadelphia, ecm 1917.

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XXII.

XXIV.

XXV.

XXVI.

XXVII.

XXVIUI.

XXIX.

XXX.

Contents

Ether and Foner in the Evolution of Matter............... The Relations and Transformations of Energy aati the

BVolutioncou Watters chet yuk eer vanes mite Yee te etant The Relation of Inorganic to Organic Bodies................ The Energies of the Organic World—the Biotic, the Cognitic,

the Cogitic—and their Relation to Organisms............. PURE MBioticjotatel ol WRCLLV cco hie tes eilenis ci eter reer her Copnitic State OhiMerd vores cuntvek ee eave sticks wale Cs A Review of the Formed Constituents of Organisms.......... Pentamorphogeny, or the Five Cooperating Causes i in Organic

Chapter I Ether And Energy In The Evolution Of Matter

In many recent treatises on physical and physiological phenomena, all cosmic changes have been considered to be due to the existence and interrelation of two great factors, matter and energy. From the discoveries mainly of the past century the former was viewed as being composed of discrete particles or atoms that belonged to some one of about 70 elements, that were truly ultimate and elemental in their nature. Hydrogen, oxygen, mercury, potassium, carbon, and iron might be cited as examples. Such were regarded as passive or inert in themselves, but, owing to the all-pervading presence and activity of some form of energy, the atoms were supposed to be in constant motion. Their relation to each other and degree of motion, however, might vary greatly according as the atoms were in a gaseous, a liquid, or a solid state.

Further, such atoms might‘unite with each other in definite combining proportions, and with definite interchanges of energy, to form more complex bodies or molecules. Water (H,O), carbonate of lime (CaCO,), and sodium phosphate (Na,HPO,) might be named as types.

The energy that gave rise to baie activities and changes was considered to exist under several conditions, such as heat, light, chemical affinity, or electricity. These energies, however,

Q CAUSES AND CoursE OF OrGANIC EVOLUTION as proved by Mayer, were convertible into each other under appropriate relations, and so “‘the transformability of energy” was a fundamental law in physics.

But the epoch-making discoveries of the past decade, and especially the recognition of radio-active bodies, promise to revolutionize many of the views that physicists, chemists, and — biologists have alike founded on in the past. So it can truly be said that we live in a period of transition (/: 182), and that no one can predict what the ultimate views—not to say established facts—as to the constitution and relation of matter and energy may be.

The ancient opinion that a diffused and inert ether filled the world and equally sidereal space was revived during the past century, and it has been widely accepted that this ether is a necessary constituent of the world’s upbuild. It has been viewed as composed of separate entities or units of extreme elasticity. These units have been supposed to act as centers round which charges of energy—of varying velocity according to the substance studied—revolve at varying rates. So by some the ether units were viewed as interpenetrating the atoms of matter, and acting as vehicles for the transmission of energy between the atoms.

Le Bon (2: 13) says: ‘The ether is doubtless a mysterious agent which we have not yet learned to isolate, but its reality is manifest, since no phenomenon can be explained without it. Its existence now seems to several physicists more certain than even that of matter. It cannot be isolated, but it is impossible to say it cannot be seen or touched. It is, on the contrary, the substance we most often see and touch. When a body radiates the heat which warms or burns us, what constitutes this heat, if it be not the vibrations of the ether? When we see a green landscape on the ground glass of a camera obscura, What constitutes this image, if not the ether?”

But the past five years have brought to our notice some remarkable and far-reaching views. From the researches of the Curies (3) and later of Rutherford and of Ramsay (4) into the properties of radium and its direct relation to other bodies,

Eruer AND ENERGY IN EvouutTion or MATTER B3 it has been proved that at least some of the so-called material elements can be further split up, and so transformed into simpler elements. Thus the metal uranium gradually changes into ionium, and later into radium, while it again can be changed into helium. But experiment and atomic weight alike suggest strongly that the scarce element polonium becomes transformed into helium and likewise into lead. Furthermore, since it has been found that the characteristics of all of the so-called elements can be expressed in terms of their atomic weight, this would suggest that possibly a continuous and definite relation exists between all of the elements. Accordingly, some physicists from Prout’s time (1815) onward have regarded hydrogen —with atomic weight 1—as the sole element, from which, by increasingly complex unions and transfers of energy, the other now known 82 elements have resulted.

In reaching this conclusion, however, physicists have been almost insensibly carried another step backward, so as to suggest that the world—and possibly all sidereal space—is the arena for only two primitive and primeval entities, ether and energy. The former consists of highly elastic particles, absolutely inert in themselves, but when charged with definite amounts of energy they have properties conferred on them that have caused us in the past to name them ultimate atoms, of which hydrogen might be regarded as the simplest.

But the development of electro-chemistry has introduced a still more fundamental conception as to the ultimate material unit. From the researches of J. J. Thomson and of Larmor (5, 6) it has been shown that even the unit-element hydrogen, as well as others of higher atomic weight, when rendered incandescent, gives off extremely minute negatively electrified “‘corpuscles” or electrons. The radius of each of these is about the jm part of that of a hydrogen atom, and its mass about the gm part. So it may be said that the ultimate physical unit is the electron or. corpuscle, that is immensely smaller than the hydrogen atom. This probably has an inert but elastic ether core or center, and carries around it a charge of negative electricity. When brought in contact with a body charged with positive electricity, union occurs with formation of an atom. The atoms again, when electrified to varying degrees, or when acted on by other forms of energy, e. g., heat or light, unite into more complex particles or molecules, of which most inorganic and all organic bodies consist.

Such views as the above have weighty experimental facts to support them, but it seems difficult if not impossible at present to say whether they will be fully sustained, or whether some new discovery may not modify the viewpoint, for the future. Meanwhile, it may be accepted as a working hypothesis that facts suggest the probable primitive and universal existence of only two constituents, energy as a moving, impelling, molding agency, and ether particles, elastic but inert. The two together constitute an electron, and when groups of them vibrate or rotate at definite rates and in definite relations they give rise to those bodies that we have hitherto spoken of as the physico-chemical elements.

Striking features of these elements that suggest an ultimate association and possible origin in common are: that they exhibit stages of transition from gaseous through liquid to solid bodies; that when two or more unite they form a compound of invariable and predicable proportion (Dalton’s first law); that when two or more bodies unite in several proportions the combining weights of these always bear an exact relation (Dalton’s second law); also that the law of proportional numbers of Berzelius, the law of specific heat of Dulong and Petit, all confirm a definite progressive relationship between the so-called elements.

Of these elements the gaseous body hydrogen is lightest and has been accepted as the unit element.

In order to obtain, if possible, a correct picture of molecular potentialities amongst living bodies, it is necessary that we look into the fundamental qualities of those elements that enter most largely and importantly into the constitution of living matter. Of the 83 known elements, some like hydrogen, oxygen, and nitrogen exist free and abundantly in nature, but. only in the gaseous state. Others like argon, krypton, neon,

ErHEerR aND ENERGY IN EvoLtuTion or Matrrer 5 and helium agree thus, but they are scarce. Only two elements, mercury and bromine—both of infrequent occurrence—are liquid at ordinary temperatures. The majority are solids, and are either abundant and widely diffused, like carbon, sulphur, silicon, and many of the metals, or are relatively scarce, like the greater number of the metals, and in some cases are strikingly related to living bodies, like sulphur, iodine, and phosphorus.

By far the most abundant of the elements in organic bodies are carbon, hydrogen, oxygen, and nitrogen, all characteristic non-metallic bodies. As pointed out by H. Spencer (7: 4-10), the first of these shows a remarkable relation to the other three. Carbon is not only allomorphic as occurring naturally in the forms of charcoal, graphite, and diamond; it is also solid at ordinary and even high:temperatures. It likewise shows a striking degree of molecular stability and cohesion, since it unites with oxygen at a rather high temperature, it unites with difficulty with hydrogen, and still less readily with nitrogen. It might thus be termed the passive element of fundamental strength and tenacity, which acts as the basis for most organic molecules, and which caused organic chemistry, at an earlier stage of its history, to be spoken of as “the chemistry of the carbon compounds.”

Hydrogen, oxygen, and nitrogen, in marked contrast to carbon, are gaseous, mobile, and amongst the lightest of elements. But even these are diverse in behavior. Nitrogen is extremely inert chemically, and only when united with oxygen, or to a less degree with hydrogen, does it show a marked capacity for chemical unions with other elements. But, given the formation of these, the resulting bodies are often of most unstable equilibrium, or even readily and violently explosive. Hydrogen, though less inert than the last, only shows high chemical activity when in presence of oxygen, but then takes part with it, and by aid of it and carbon forms many substances, particularly those of organic origin. Oxygen, while gaseous, light, and mobile in physical relation, exhibits on the other hand extreme chemical activity, and might well be re-

6 CAUSES AND CoursE or OrcGAnic EvouuTiIon garded as the element that starts energizing action, and that passes on and distributes energy to the other three, so as to cause these to link up in ever-increasing complexity.

Of the metallic elements, potassium, calcium, magnesium, and iron are most abundant in living tissues. They exist there in various degrees of combining complexity.

We may now briefly glance at the views generally held by physicists and geologists of the present day, as to the evolution of the earth’s crust; the disposition of materials in it as gradual cooling took place; the relation of land, water, and air to each other as these became differentiated; also the environmental relations which probably prevailed when conditions had become favorable for the appearance of primitive organisms on it.

In the gradual condensation and cooling of the earth from an originally nebulous or gaseous and then incandescent planet, it seems possible and even probable from the studies made by Arrhenius (8) that the most abundant or the densest elements such as platinum, gold, bismuth, and iron tended to gravitate toward the center, to form there a gaseous core composed . very largely of iron, mixed in much smaller proportions with other and rarer metals such as platinum and gold, but all in a highly compressed though gaseous state. According to some, as this metalliferous core continued to condense, and to radiate off heat, it became a solid mass round which the lighter metals, and some of the nonmetals—notably aluminium, carbon, and silicon—collected either in pure state or in varying chemical combinations between themselves and the heavier metals within, or with the more gaseous bodies, such as oxygen and hydrogen.

But the metal iron, from its ppeponderating abundance in | the earth’s core, and from the strains taking place between the core and exterior, must have continually welled up, been ejected, or exposed in large amount. Thus the rocks of archzean or proterozoic age contain large deposits of iron, probably derived in part from decomposition of igneous rocks, in part owing to activity of primitive plant organisms (p. 16). Iron, therefore, in all subsequent ages up to the present, has been the leading

Eruer AND Enrercy In Evouution or MarrEer uf metal of the earth’s surface layers. So there originated an investing zone or crust round the earth’s metallic core, which, in virtue of its own heat energy, in virtue of the heat radiated out to it from the core, in virtue also of the active chemical affinity existing between many of the condensing bodies themselves, gave rise to extensive masses of carbonates, silicates, and other compounds of aluminium, iron, or manganese, which, when rearranged or erupted later, formed the igneous or foundational rocks of the earth’s crust.

But on the still hot, often it might be incandescent, surface crust, and aided by abundant chemical reactions that were then proceeding, union of the light hydrogen and oxygen of the gaseous envelope gave rise to.water vapor, that probably long remained as a dense shroud, and that was constantly added to by new formations.

Owing to continued radiation of heat into space, owing also doubtless to condensation and conversion of heat round the ether particles into intra-atomic energy, probably of an electric nature as Le Bon has pictured, the gaseous and vaporous envelope became condensed and cooled. But, during transition from a gaseous and highly-heated state to that of commencing formation of solid surface materials, the earth doubtless passed, as Arldt has well indicated (9: 553), through those successive phases of appearing among the other planets as a white star, later as a yellow star, still later as a red star, _ this last phase inaugurating formation of the beginning solid crust. As this crust condensed it must have been subject to considerable strains and displacements. So it would tend to be ridged up into evolving higher lands, and to collapse into depressions which, when the water vapor had sufficiently cooled or condensed as water, would start denudation action. This also, considering the active agencies then at work, would proceed comparatively rapidly, and would tend still further to mix up the already-formed constituents, and to evolve new chemical compounds in addition.

Contrary, however, to the older views of geologists, and in line with the suggestive studies of Arldt (9: 564-604), we now

8 CausEs AND CoursE or OrGaAnic EvoLutTion recognize that these elevations and depressions were fundamentally different in distribution, situation, and connection from those of the present day, or even of the mesozoic period.

Gradually, therefore, through intimate union of the cooling materials of the crust, there formed upon the igneous rock masses that extremely complex layer of fundamental solid rock that we know as granite. From all geological evidence, this is the primitive crystalline zone on which all the more recent strata have been deposited, and even from which in great part they have been derived, by subsequent decompositions and denudations. It may therefore be of value here to give a tabular view of the composition of granite in nine of its most abundant components, that together make up about 99.75 per cent. of its composition.

From the above table it will be seen that the most striking peculiarities of granite are: (1) the presence in it of all the elements needed for complete nutrition of plants and animals except those three fundamentally diverse elements already referred to, carbon, hydrogen, and nitrogen. But undoubtedly the three existed in abundance in the surrounding gaseous envelope that constituted the atmosphere of the evolving earth. - For CO, has all along played an important part in uniting with bases to form carbonates, while the masses of graphite that occur in the older crystalline rocks prove that it must have been abundant. Nitrogen must also have been a widely diffused gas in the atmosphere, but its presence otherwise is demonstrated by its frequent occurrence in cavities of granite, gneissose and basaltic rocks. The same is true for hydrogen.

Eruer AND ENERGY In Evouution or Matter 9

The element phosphorus, in many respects as typical for organisms as carbon, though present at times in minute quantity, is also present in all typical granites but only to the extent of 4 to » of one per cent. and so has been omitted from the above table. :

A second feature of granitic composition is*the important part which oxygen plays, in that it is a constituent of all of the granite components, and is the energizing element that would further cause these to enter into new combinations.

As bearing on the formation alike of crystalloid and of colloid bodies, and their subsequent presence in organic tissues, it may be convenient here to present, in tabular form, data regarding the earth’s crust. From comparative study of the substances entering into its composition, it is found that oxygen and silicon make up by far the largest percentage proportion. Thus oxygen, with atomic weight 16, constitutes about 50 per cent.; silicon, with atomic weight 28, constitutes about 27 per cent. of the entire mass; or these two together represent about 77 per cent. of its bulk. The subjoined table of the six most abundant nonmetals and of the six most abundant metals, with the atomic weight of each, graphically sets forth the preponderating importance of the twelve in the construction of the earth as a whole, since together they make up fully 99 per cent. of it.

Proportion in Proportion in Atomic outer crust Atoutic outer crust Nonmetals : including Metals 5 including weight weight crust, sea, crust, sea, atmosphere : atmosphere Oxygen 16 50 ‘ {| Aluminium | 27.30 7 AS Silicon 28 26 Tron 55.90 4.20 Hydrogen it 0.90 Calcium 39.90 3.25 Carbon 11.97 0.20 Magnesium! 23.94 2.35 Phosphorus | 30.96 0.08 Potassium | 39.04 Bt

These again, as already set forth for granite, enter primarily into union with oxygen to form oxides. The next table sets forth F. W. Clarke’s average results of the analysis of 830

1* typical samples of the primitive part of the earth’s crust; and from this it will be seen that the group of oxides given makes up about 99 per cent. of its total mass.

Silica (SiOs) red hoes ney ne a 59.71 ’ lamina (CAB Os) oo 4 tor ee core eee 15.41 Ferric oxide (He:O3). : vss foc oh Fee 2.63 Ferrous oxide (FeO)=; 3% 22 eace sees 3.52 Calcic oxide'(CaO) oe 2 Saas eee ee 4.90 Magnesia (MgO) >. 45 3 eer ets 4.36 Potash! (Re Oyo es ee eee 2.80 Soda: (Nac®) 0076 Fess. eee ee pia cee ee Nery Water: (H2O) 4%. 4:-.caus Seee cee 1.52 Phosphoric acid C2sO;))2 tera eet eee 0. 22 98.62

It is generally conceded that, during the early formation of this shell or crust, abundant and widespread volcanic activity on the one hand combined with increasing cooling of the crust on the other were proceeding, so that, with evolution of a definite enveloping atmosphere, subaerial denudation would, on the one hand, cause steady attrition, disintegration, and removal of the previously formed masses, while new accumulations of molten material, thrown out from the earth’s interior, would cause, by gradual cooling of these, steady upbuilding or integration of the crust.

At the close then of what may be termed the igneous or volcanic epoch of the earth’s history, the internal masses of gaseous iron and rarer metals, along with smaller quantities of other elements that now formed, on Arrhenius’s view, the earth’s core, were bounded by a transition layer of viscous and unstable material, that again was enveloped by a granite shell. Or to picture it from without inward in Geikie’s words: “The interior of the earth, therefore, with the exception of a solid crust about 40 kilos thick, consists of a molten magma 100 to 200 kilos in depth, which shades continually inward into a gaseous center. The liquids and gases in the interior possess a viscosity and incompressibility such as permit them to be regarded as solid bodies.”

Eruer anD ENERGY IN Evo.Lution or Marrer 11

The granite shell or crust contained either in its own constitution, or enclosed as graphite masses (in the case of carbon), or included as gaseous cavities, all of the elements that make up organic bodies. Furthermore, abundant and varied combinations had already taken place among the elements, so as to produce granitic compounds, from which, by subsequent alterations, disintegration, and reintegration, all of the more recent rock constituents could be derived.

If we now compare the views of physicists and geologists, as to the probable age of the earth’s crust from the time when the granitic rocks had been formed up to the present day, a considerable diversity of opinion is encountered. Kelvin, in his earlier calculations, based his estimates on three physical factors, and concluded that a period of from 20,000,000 to 100,000,000 of years has elapsed since subaerial denudation and deposition began to take place. But the more recent studies of G. Darwin, Woodward, and Perry, on the physical side, have caused them to concede at least 100,000,000 of years as a likely time-minimum. In this they have agreed with the earlier claims of geologists, who alike from the geodynamical and the paleontological standpoints claimed a greatly extended period.

From the geological standpoint J. Joly (10: 23) has estimated the amount of sodium annually carried to the ocean, since its earliest formation, in relation to the total quantity dissolved there, and concludes that the process of transfer has proceeded for 90 to 100 millions of years. Sollas (11: 24) again is inclined, also on geological grounds, to accept a time limit vaguely intermediate betw: een the minimum and maximum of Kelvin. -

Arldt in treating of the phases of the earth (9: 553) apportions the time limits as follows, beginning with the period when the earth had just passed from a nebulous state into that of a white star, and onward till recent date.

Period of white star evolution........ 135 millions of years

“. “* yellow star evolution........ 125 x ss } oeeeered star evolution... ..:..... QT SS apes:

12 Causes AND CoursE oF OrnGANIC EVOLUTION

Period of crust formation............ 105 millions of years ““ dry or non-oceanic crust..... 95 . 5 een “« _* Inorganic evolution; ........ 122345 — = ees “_ “ archeean rock formation..... 95 ie “eee “© naleeozoic rock formation.... 75 = bake Pe “« ~ ** mesozoic rock formation.... 71% “ fines “>. eenozoie rock formations... . oo sagen eae

This calculation—confessedly vague—would give about 180 millions of years for formation of the sedimentary crust.

The oldest stratified or semi-stratified rocks of the earth’s crust have variously been called the metamorphic, archzan, proterozoic, or pre-cambrian, and it is to the early or midarchezean epoch that all evidence tends, as the starting point for the dawn of organisms on the earth. The rocks then formed are usually quite unconformable with the masses of primitive granite on which they rest, thus indicating that a considerable —even extended—period of time had elapsed between deposit of the two systems. Only rarely do they seem to be continuous, but even then the composition of the older igneous or granitic and newer or archean rocks differs markedly (12: 863). Often — also the older archezan beds are broken up, distorted, and copiously interbedded with intrusive granitic masses, that indicate active and extensive alterations in the crust.

But, while the archzean strata or masses consist very largely of schistose and micaceous gneissic materials that have been highly metamorphosed by contact with granitic or other intrusive igneous flows, in the middle or later beds of the series there occur zones of graphite that may have had a vegetable origin; cherts and iron ores probably also of vegetable origin, as Geikie has suggested (12: 877) and as the writer has further elucidated (13: 257); as well as limestones that may have had an organic, possibly even a vegetable, origin; and all frequently interbedded with schists, serpentines, and other rocks.