But Mr. Darwin's admission, or rather his assertion, that the inheritance of functionally-produced modifications has been a factor in organic evolution, is made clear not by these passages alone and by kindred ones. It is made clearer still by a passage in the preface to the second edition of his Descent of Man. He there protests against that current version of his views in which this factor makes no appearance. The passage is as follows.

"I may take this opportunity of remarking that my critics frequently assume that I attribute all changes of corporeal structure and mental power exclusively to the natural selection of such variations as are often called spontaneous; whereas, even in the first edition of the 'Origin of Species,' I distinctly stated that great weight must be attributed to the inherited effects of use and disuse, with respect both to the body and mind."

Nor is this all. There is evidence that Mr. Darwin's belief in the efficiency of this factor, became stronger as he grew older and accumulated more evidence. The first of the extracts above given, taken from the sixth edition of the Origin of Species, runs thus:—

"I think there can be no doubt that use in our domestic animals has strengthened and enlarged certain parts, and disuse diminished them; and that such modifications are inherited."

Now on turning to the first edition, p. 134, it will be found that instead of the words—"I think there can be no doubt," the words originally used were—"I think there can be little doubt." That this deliberate erasure of a qualifying word and substitution of a word implying unqualified belief, was due to a more decided recognition of a factor originally under-estimated, is clearly implied by the wording of the above-quoted passage from the preface to the Descent of Man; where he says that "even in the first edition of the 'Origin of Species,'" &c.: the implication being that much more in subsequent editions, and subsequent works, had he insisted on this factor. The change thus indicated is especially significant as having occurred at a time of life when the natural tendency is towards fixity of opinion.

During that earlier period when he was discovering the multitudinous cases in which his own hypothesis afforded solutions, and simultaneously observing how utterly futile in these multitudinous cases was the hypothesis propounded by his grandfather and Lamarck, Mr. Darwin was, not unnaturally, almost betrayed into the belief that the one is all-sufficient and the other inoperative. But in the mind of one so candid and ever open to more evidence, there naturally came a reaction. The inheritance of functionally-produced modifications, which, judging by the passage quoted above concerning the views of these earlier enquirers, would seem to have been at one time denied, but which as we have seen was always to some extent recognized, came to be recognized more and more, and deliberately included as a factor of importance.

* * * * *

Of this reaction displayed in the later writings of Mr. Darwin, let us now ask—Has it not to be carried further? Was the share in organic evolution which Mr. Darwin latterly assigned to the transmission of modifications caused by use and disuse, its due share? Consideration of the groups of evidences given above, will, I think, lead us to believe that its share has been much larger than he supposed even in his later days.

There is first the implication yielded by extensive classes of phenomena which remain inexplicable in the absence of this factor. If, as we see, co-operative parts do not vary together, even when few and close together, and may not therefore be assumed to do so when many and remote, we cannot account for those innumerable changes in organization which are implied when, for advantageous use of some modified part, many other parts which join it in action have to be modified.

Further, as increasing complexity of structure, accompanying increasing complexity of life, implies increasing number of faculties, of which each one conduces to preservation of self or descendants; and as the various individuals of a species, severally requiring something like the normal amounts of all these, may individually profit, here by an unusual amount of one, and there by an unusual amount of another; it follows that as the number of faculties becomes greater, it becomes more difficult for any one to be further developed by natural selection. Only where increase of some one is predominantly advantageous does the means seem adequate to the end. Especially in the case of powers which do not subserve self-preservation in appreciable degrees, does development by natural selection appear impracticable.

It is a fact recognized by Mr. Darwin, that where, by selection through successive generations, a part has been increased or decreased, its reaction upon other parts entails changes in them. This reaction is effected through the changes of function involved. If the changes of structure produced by such changes of function, are inheritable, then the re-adjustment of parts throughout the organism, taking place generation after generation, maintains an approximate balance; but if not, then generation after generation the organism must get more and more out of gear, and tend to become unworkable.

Further, as it is proved that change in the balance of functions registers its effects on the reproductive elements, we have to choose between the alternatives that the registered effects are irrelevant to the particular modifications which the organism has undergone, or that they are such as tend to produce repetitions of these modifications. The last of these alternatives makes the facts comprehensible; but the first of them not only leaves us with several unsolved problems, but is incongruous with the general truth that by reproduction, ancestral traits, down to minute details, are transmitted.

Though, in the absence of pecuniary interests and the interests in hobbies, no such special experiments as those which have established the inheritance of fortuitous variations have been made to ascertain whether functionally-produced modifications are inherited; yet certain apparent instances of such inheritance have forced themselves on observation without being sought for. In addition to other indications of a less conspicuous kind, is the one I have given above—the fact that the apparatus for tearing and mastication has decreased with decrease of its function, alike in civilized man and in some varieties of dogs which lead protected and pampered lives. Of the numerous cases named by Mr. Darwin, it is observable that they are yielded not by one class of parts only, but by most if not all classes—by the dermal system, the muscular system, the osseous system, the nervous system, the viscera; and that among parts liable to be functionally modified, the most numerous observed cases of inheritance are furnished by those which admit of preservation and easy comparison—the bones: these cases, moreover, being specially significant as showing how, in sundry unallied species, parallel changes of structure have occurred along with parallel changes of habit.

What, then, shall we say of the general implication? Are we to stop short with the admission that inheritance of functionally-produced modifications takes place only in cases in which there is evidence of it? May we properly assume that these many instances of changes of structure caused by changes of function, occurring in various tissues and various organs, are merely special and exceptional instances having no general significance? Shall we suppose that though the evidence which already exists has come to light without aid from a body of inquirers, there would be no great increase were due attention devoted to the collection of evidence? This is, I think, not a reasonable supposition. To me the ensemble of the facts suggests the belief, scarcely to be resisted, that the inheritance of functionally-produced modifications takes place universally. Looking at physiological phenomena as conforming to physical principles, it is difficult to conceive that a changed play of organic forces which in many cases of different kinds produces an inherited change of structure, does not do this in all cases. The implication, very strong I think, is that the action of every organ produces on it a reaction which, usually not altering its rate of nutrition, sometimes leaves it with diminished nutrition consequent on diminished action, and at other times increases its nutrition in proportion to its increased action; that while generating a modified consensus of functions and of structures, the activities are at the same time impressing this modified consensus on the sperm-cells and germ-cells whence future individuals are to be produced; and that in ways mostly too small to be identified, but occasionally in more conspicuous ways and in the course of generations, the resulting modifications of one or other kind show themselves. Further, it seems to me that as there are certain extensive classes of phenomena which are inexplicable if we assume the inheritance of fortuitous variations to be the sole factor, but which become at once explicable if we admit the inheritance of functionally-produced changes, we are justified in concluding that this inheritance of functionally-produced changes has been not simply a co-operating factor in organic evolution, but has been a co-operating factor without which organic evolution, in its higher forms at any rate, could never have taken place.

Be this or be it not a warrantable conclusion, there is, I think, good reason for a provisional acceptance of the hypothesis that the effects of use and disuse are inheritable; and for a methodic pursuit of inquiries with the view of either establishing it or disproving it. It seems scarcely reasonable to accept without clear demonstration, the belief that while a trivial difference of structure arising spontaneously is transmissible, a massive difference of structure, maintained generation after generation by change of function, leaves no trace in posterity. Considering that unquestionably the modification of structure by function is a vera causa, in so far as concerns the individual; and considering the number of facts which so competent an observer as Mr. Darwin regarded as evidence that transmission of such modifications takes place in particular cases; the hypothesis that such transmission takes place in conformity with a general law, holding of all active structures, should, I think, be regarded as at least a good working hypothesis.

* * * * *

But now supposing the broad conclusion above drawn to be granted—supposing all to agree that from the beginning, along with inheritance of useful variations fortuitously arising, there has been inheritance of effects produced by use and disuse; do there remain no classes of organic phenomena unaccounted for? To this question I think it must be replied that there do remain classes of organic phenomena unaccounted for. It may, I believe, be shown that certain cardinal traits of animals and plants at large are still unexplained; and that a further factor must be recognized. To show this, however, will require another paper.

II.

Ask a plumber who is repairing your pump, how the water is raised in it, and he replies—"By suction." Recalling the ability which he has to suck up water into his mouth through a tube, he is certain that he understands the pump's action. To inquire what he means by suction, seems to him absurd. He says you know as well as he does, what he means; and he cannot see that there is any need for asking how it happens that the water rises in the tube when he strains his mouth in a particular way. To the question why the pump, acting by suction, will not make the water rise above 32 feet, and practically not so much, he can give no answer; but this does not shake his confidence in his explanation.

On the other hand an inquirer who insists on knowing what suction is, may obtain from the physicist answers which give him clear ideas, not only about it but about many other things. He learns that on ourselves and all things around, there is an atmospheric pressure amounting to about 15 pounds on the square inch: 15 pounds being the average weight of a column of air having a square inch for its base and extending upwards from the sea-level to the limit of the Earth's atmosphere. He is made to observe that when he puts one end of a tube into water and the other end into his mouth, and then draws back his tongue, so leaving a vacant space, two things happen. One is that the pressure of air outside his cheeks, no longer balanced by an equal pressure of air inside, thrusts his cheeks inwards; and the other is that the pressure of air on the surface of the water, no longer balanced by an equal pressure of air within the tube and his mouth (into which part of the air from the tube has gone) the water is forced up the tube in consequence of the unequal pressure. Once understanding thus the nature of the so-called suction, he sees how it happens that when the plunger of the pump is raised and relieves from atmospheric pressure the water below it, the atmospheric pressure on the water in the well, not being balanced by that on the water in the tube, forces the water higher up the tube, so that it follows the plunger. And now he sees why the water cannot be raised beyond the theoretic limit of 32 feet: a limit made much lower in practice by imperfections in the apparatus. For if, simplifying the conception, he supposes the tube of the pump to be a square inch in section, then the atmospheric pressure of 15 pounds per square inch on the water in the well, can raise the water in the tube to such height only that the entire column of it weighs 15 pounds. Having been thus enlightened about the pump's action, the action of a barometer becomes intelligible. He perceives how, under the conditions established, the weight of the column of mercury balances that of an atmospheric column of equal diameter; and how, as the weight of the atmospheric column varies, there is a corresponding variation in the weight of the mercurial column,—shown by change of height. Moreover, having previously supposed that he understood the ascent of a balloon when he ascribed it to relative lightness, he now sees that he did not truly understand it. For he did not recognize it as a result of that upward pressure caused by the difference between the weight of the mass formed by the gas in the balloon plus the cylindrical column of air extending above it to the limit of the atmosphere, and the weight of a similar cylindrical column of air extending down to the under surface of the balloon: this difference of weight causing an equivalent upward pressure on the under surface.

Why do I introduce these familiar truths so entirely irrelevant to my subject? I do it to show, in the first place, the contrast between a vague conception of a cause and a distinct conception of it; or rather, the contrast between that conception of a cause which results when it is simply classed with some other or others which familiarity makes us think we understand, and that conception of a cause which results when it is represented in terms of definite physical forces admitting of measurement. And I do it to show, in the second place, that when we insist on resolving a verbally-intelligible cause into its actual factors, we get not only a clear solution of the problem before us, but we find that the way is opened to solutions of sundry other problems. While we rest satisfied with unanalyzed causes, we may be sure both that we do not rightly comprehend the production of the particular effects ascribed to them, and that we overlook other effects which would be revealed to us by contemplation of the causes as analyzed. Especially must this be so where the causation is complex. Hence we may infer that the phenomena presented by the development of species, are not likely to be truly conceived unless we keep in view the concrete agencies at work. Let us look closely at the facts to be dealt with.

* * * * *

The growth of a thing is effected by the joint operation of certain forces on certain materials; and when it dwindles, there is either a lack of some materials, or the forces co-operate in a way different from that which produces growth. If a structure has varied, the implication is that the processes which built it up were made unlike the parallel processes in other cases, by the greater or less amount of some one or more of the matters or actions concerned. Where there is unusual fertility, the play of vital activities is thereby shown to have deviated from the ordinary play of vital activities; and conversely, if there is infertility. If the germs, or ova, or seed, or offspring partially developed, survive more or survive less, it is either because their molar or molecular structures are unlike the average ones, or because they are affected in unlike ways by surrounding agencies. When life is prolonged, the fact implies that the combination of actions, visible and invisible, constituting life, retains its equilibrium longer than usual in presence of environing forces which tend to destroy its equilibrium. That is to say, growth, variation, survival, death, if they are to be reduced to the forms in which physical science can recognize them, must be expressed as effects of agencies definitely conceived—mechanical forces, light, heat, chemical affinity, &c.

This general conclusion brings with it the thought that the phrases employed in discussing organic evolution, though convenient and indeed needful, are liable to mislead us by veiling the actual agencies. That which really goes on in every organism is the working together of component parts in ways conducing to the continuance of their combined actions, in presence of things and actions outside; some of which tend to subserve, and others to destroy, the combination. The matters and forces in these two groups, are the sole causes properly so called. The words "natural selection," do not express a cause in the physical sense. They express a mode of co-operation among causes—or rather, to speak strictly, they express an effect of this mode of co-operation. The idea they convey seems perfectly intelligible. Natural selection having been compared with artificial selection, and the analogy pointed out, there apparently remains no indefiniteness: the inconvenience being, however, that the definiteness is of a wrong kind. The tacitly implied Nature which selects, is not an embodied agency analogous to the man who selects artificially; and the selection is not the picking out of an individual fixed on, but the overthrowing of many individuals by agencies which one successfully resists, and hence continues to live and multiply. Mr. Darwin was conscious of these misleading implications. In the introduction to his Animals and Plants under Domestication (p. 6) he says:—

"For brevity sake I sometimes speak of natural selection as an intelligent power; ... I have, also, often personified the word Nature; for I have found it difficult to avoid this ambiguity; but I mean by nature only the aggregate action and product of many natural laws,—and by laws only the ascertained sequence of events."

But while he thus clearly saw, and distinctly asserted, that the factors of organic evolution are the concrete actions, inner and outer, to which every organism is subject, Mr. Darwin, by habitually using the convenient figure of speech, was, I think, prevented from recognizing so fully as he would otherwise have done, certain fundamental consequences of these actions.

Though it does not personalize the cause, and does not assimilate its mode of working to a human mode of working, kindred objections may be urged against the expression to which I was led when seeking to present the phenomena in literal terms rather than metaphorical terms—the survival of the fittest;[42] for in a vague way the first word, and in a clear way the second word, calls up an anthropocentric idea. The thought of survival inevitably suggests the human view of certain sets of phenomena, rather than that character which they have simply as groups of changes. If, asking what we really know of a plant, we exclude all the ideas associated with the words life and death, we find that the sole facts known to us are that there go on in the plant certain inter-dependent processes, in presence of certain aiding and hindering influences outside of it; and that in some cases a difference of structure or a favourable set of circumstances, allows these inter-dependent processes to go on for longer periods than in other cases. Again, in the working together of those many actions, internal and external, which determine the lives or deaths of organisms, we see nothing to which the words fitness and unfitness are applicable in the physical sense. If a key fits a lock, or a glove a hand, the relation of the things to one another is presentable to the perceptions. No approach to fitness of this kind is made by an organism which continues to live under certain conditions. Neither the organic structures themselves, nor their individual movements, nor those combined movements of certain among them which constitute conduct, are related in any analogous way to the things and actions in the environment. Evidently the word fittest, as thus used, is a figure of speech; suggesting the fact that amid surrounding actions, an organism characterized by the word has either a greater ability than others of its kind to maintain the equilibrium of its vital activities, or else has so much greater a power of multiplication that though not longer lived than they, it continues to live in posterity more persistently. And indeed, as we here see, the word fittest has to cover cases in which there may be less ability than usual to survive individually, but in which the defect is more than made good by higher degrees of fertility.

I have elaborated this criticism with the intention of emphasizing the need for studying the changes which have gone on, and are ever going on, in organic bodies, from an exclusively physical point of view. On contemplating the facts from this point of view, we become aware that, besides those special effects of the co-operating forces which eventuate in the longer survival of one individual than of others, and in the consequent increase through generations, of some trait which furthered its survival, many other effects are being wrought on each and all of the individuals. Bodies of every class and quality, inorganic as well as organic, are from instant to instant subject to the influences in their environments; are from instant to instant being changed by these in ways that are mostly inconspicuous; and are in course of time changed by them in conspicuous ways. Living things in common with dead things, are, I say, being thus perpetually acted upon and modified; and the changes hence resulting, constitute an all-important part of those undergone in the course of organic evolution. I do not mean to imply that changes of this class pass entirely unrecognized; for, as we shall see, Mr. Darwin takes cognizance of certain secondary and special ones. But the effects which are not taken into account, are those primary and universal effects which give certain fundamental characters to all organisms. Contemplation of an analogy will best prepare the way for appreciation of them, and of the relation they bear to those which at present monopolize attention.

An observant rambler along shores, will, here and there, note places where the sea has deposited things more or less similar, and separated them from dissimilar things—will see shingle parted from sand; larger stones sorted from smaller stones; and will occasionally discover deposits of shells more or less worn by being rolled about. Sometimes the pebbles or boulders composing the shingle at one end of a bay, he will find much larger than those at the other: intermediate sizes, having small average differences, occupying the space between the extremes. An example occurs, if I remember rightly, some mile or two to the west of Tenby; but the most remarkable and well-known example is that afforded by the Chesil bank. Here, along a shore some sixteen miles long, there is a gradual increase in the sizes of the stones; which, being at one end but mere pebbles, are at the other end immense boulders. In this case, then, the breakers and the undertow have effected a selection—have at each place left behind those stones which were too large to be readily moved, while taking away others small enough to be moved easily. But now, if we contemplate exclusively this selective action of the sea, we overlook certain important effects which the sea simultaneously works. While the stones have been differently acted upon in so far that some have been left here and some carried there; they have been similarly acted upon in two allied, but distinguishable, ways. By perpetually rolling them about and knocking them one against another, the waves have so broken off their most prominent parts as to produce in all of them more or less rounded forms; and then, further, the mutual friction of the stones simultaneously caused, has smoothed their surfaces. That is to say in general terms, the actions of environing agencies, so far as they have operated indiscriminately, have produced in the stones a certain unity of character; at the same time that they have, by their differential effects, separated them: the larger ones having withstood certain violent actions which the smaller ones could not withstand.

Similarly with other assemblages of objects which are alike in their primary traits but unlike in their secondary traits. When simultaneously exposed to the same set of actions, some of these actions, rising to a certain intensity, may be expected to work on particular members of the assemblage changes which they cannot work in those which are markedly unlike; while others of the actions will work in all of them similar changes, because of the uniform relations between these actions and certain attributes common to all members of the assemblage. Hence it is inferable that on living organisms, which form an assemblage of this kind, and are unceasingly exposed in common to the agencies composing their inorganic environments, there must be wrought two such sets of effects. There will result a universal likeness among them consequent on the likeness of their respective relations to the matters and forces around; and there will result, in some cases, the differences due to the differential effects of these matters and forces, and in other cases, the changes which, being life-sustaining or life-destroying, eventuate in certain natural selections.

I have, above, made a passing reference to the fact that Mr. Darwin did not fail to take account of some among these effects directly produced on organisms by surrounding inorganic agencies. Here are extracts from the sixth edition of the Origin of Species showing this.

"It is very difficult to decide how far changed conditions, such as of climate, food, &c., have acted in a definite manner. There is reason to believe that in the course of time the effects have been greater than can be proved by clear evidence.... Mr. Gould believes that birds of the same species are more brightly coloured under a clear atmosphere, than when living near the coast or on islands; and Wollaston is convinced that residence near the sea affects the colours of insects. Moquin-Tandon gives a list of plants which, when growing near the sea-shore, have their leaves in some degree fleshy, though not elsewhere fleshy" (pp. 106-7). "Some observers are convinced that a damp climate affects the growth of the hair, and that with the hair the horns are correlated" (p. 159).

In his subsequent work, Animals and Plants under Domestication, Mr. Darwin still more clearly recognizes these causes of change in organization. A chapter is devoted to the subject. After premising that "the direct action of the conditions of life, whether leading to definite or indefinite results, is a totally distinct consideration from the effects of natural selection;" he goes on to say that changed conditions of life "have acted so definitely and powerfully on the organisation of our domesticated productions, that they have sufficed to form new sub-varieties or races, without the aid of selection by man or of natural selection." Of his examples here are two.

"I have given in detail in the ninth chapter the most remarkable case known to me, namely, that in Germany several varieties of maize brought from the hotter parts of America were transformed in the course of only two or three generations." (Vol. ii, p. 277.) [And in this ninth chapter concerning these and other such instances he says "some of the foregoing differences would certainly be considered of specific value with plants in a state of nature." (Vol. i, p. 321.)] "Mr. Meehan, in a remarkable paper, compares twenty-nine kinds of American trees, belonging to various orders, with their nearest European allies, all grown in close proximity in the same garden and under as nearly as possible the same conditions." And then enumerating six traits in which the American forms all of them differ in like ways from their allied European forms, Mr. Darwin thinks there is no choice but to conclude that these "have been definitely caused by the long-continued action of the different climate of the two continents on the trees." (Vol. ii, pp. 281-2.)

But the fact we have to note is that while Mr. Darwin thus took account of special effects due to special amounts and combinations of agencies in the environment, he did not take account of the far more important effects due to the general and constant operation of these agencies.[43] If a difference between the quantities of a force which acts on two organisms, otherwise alike and otherwise similarly conditioned, produces some difference between them; then, by implication, this force produces in both of them effects which they show in common. The inequality between two things cannot have a value unless the things themselves have values. Similarly if, in two cases, some unlikeness of proportion among the surrounding inorganic agencies to which two plants or two animals are exposed, is followed by some unlikeness in the changes wrought on them; then it follows that these several agencies taken separately, work changes in both of them. Hence we must infer that organisms have certain structural characters in common, which are consequent on the action of the medium in which they exist: using the word medium in a comprehensive sense, as including all physical forces falling upon them as well as matters bathing them. And we may conclude that from the primary characters thus produced there must result secondary characters.

Before going on to observe those general traits of organisms due to the general action of the inorganic environment upon them, I feel tempted to enlarge on the effects produced by each of the several matters and forces constituting the environment. I should like to do this not only to give a clear preliminary conception of the ways in which all organisms are affected by these universally-present agents, but also to show that, in the first place, these agents modify inorganic bodies as well as organic bodies, and that, in the second place, the organic are far more modifiable by them than the inorganic. But to avoid undue suspension of the argument, I content myself with saying that when the respective effects of gravitation, heat, light, &c., are studied, as well as the respective effects, physical and chemical, of the matters forming the media, water and air, it will be found that while more or less operative on all bodies, each modifies organic bodies to an extent immensely greater than the extent to which it modifies inorganic bodies.

* * * * *

Here, not discriminating among the special effects which these various forces and matters in the environment produce on both classes of bodies, let us consider their combined effects, and ask—What is the most general trait of such effects?

Obviously the most general trait is the greater amount of change wrought on the outer surface than on the inner mass. In so far as the matters of which the medium is composed come into play, the unavoidable implication is that they act more on the parts directly exposed to them than on the parts sheltered from them. And in so far as the forces pervading the medium come into play, it is manifest that, excluding gravity, which affects outer and inner parts indiscriminately, the outer parts have to bear larger shares of their actions. If it is a question of heat, then the exterior must lose it or gain it faster than the interior; and in a medium which is now warmer and now colder, the two must habitually differ in temperature to some extent—at least where the size is considerable. If it is a question of light, then in all but absolutely transparent masses, the outer parts must undergo more of any change producible by it than the inner parts—supposing other things equal; by which I mean, supposing the case is not complicated by any such convexities of the outer surface as produce internal concentrations of rays. Hence then, speaking generally, the necessity is that the primary and almost universal effect of the converse between the body and its medium, is to differentiate its outside from its inside. I say almost universal, because where the body is both mechanically and chemically stable, like, for instance, a quartz crystal, the medium may fail to work either inner or outer change.

Of illustrations among inorganic bodies, a convenient one is supplied by an old cannon-ball that has been long lying exposed. A coating of rust, formed of flakes within flakes, incloses it; and this thickens year by year, until, perhaps, it reaches a stage at which its exterior loses as much by rain and wind as its interior gains by further oxidation of the iron. Most mineral masses—pebbles, boulders, rocks—if they show any effect of the environment at all, show it only by that disintegration of surface which follows the freezing of absorbed water: an effect which, though mechanical rather than chemical, equally illustrates the general truth. Occasionally a "rocking-stone" is thus produced. There are formed successive layers relatively friable in texture, each of which, thickest at the most exposed parts, and being presently lost by weathering, leaves the contained mass in a shape more rounded than before; until, resting on its convex under-surface, it is easily moved. But of all instances perhaps the most remarkable is one to be seen on the west bank of the Nile at Philae, where a ridge of granite 100 feet high, has had its outer parts reduced in course of time to a collection of boulder-shaped masses, varying from say a yard in diameter to six or eight feet, each one of which shows in progress an exfoliation of successively-formed shells of decomposed granite: most of the masses having portions of such shells partially detached.

If, now, inorganic masses, relatively so stable in composition, thus have their outer parts differentiated from their inner parts, what must we say of organic masses, characterized by such extreme chemical instability?—instability so great that their essential material is named protein, to indicate the readiness with which it passes from one isomeric form to another. Clearly the necessary inference is that this effect of the medium must be wrought inevitably and promptly, wherever the relation of outer and inner has become settled: a qualification for which the need will be seen hereafter.

* * * * *

Beginning with the earliest and most minute kinds of living things, we necessarily encounter difficulties in getting direct evidence; since, of the countless species now existing, all have been subject during millions upon millions of years to the evolutionary process, and have had their primary traits complicated and obscured by those endless secondary traits which the natural selection of favourable variations has produced. Among protophytes it needs but to think of the multitudinous varieties of diatoms and desmids, with their elaborately-constructed coverings; or of the definite methods of growth and multiplication among such simple Algae as the Conjugatae; to see that most of their distinctive characters are due to inherited constitutions, which have been slowly moulded by survival of the fittest to this or that mode of life. To disentangle such parts of their developmental changes as are due to the action of the medium, is therefore hardly possible. We can hope only to get a general conception of it by contemplating the totality of the facts.

The first cardinal fact is that all protophytes are cellular—all show us this contrast between outside and inside. Supposing the multitudinous specialities of the envelope in different orders and genera of protophytes to be set against one another, and mutually cancelled, there remains as a trait common to them—an envelope unlike that which it envelopes. The second cardinal fact is that this simple trait is the earliest trait displayed in germs, or spores, or other parts from which new individuals are to arise; and that, consequently, this trait must be regarded as having been primordial. For it is an established truth of organic evolution that embryos show us, in general ways, the forms of remote ancestors; and that the first changes undergone, indicate, more or less clearly, the first changes which took place in the series of forms through which the existing form has been reached. Describing, in successive groups of plants, the early transformations of these primitive units, Sachs[44] says of the lowest Algae that "the conjugated protoplasmic body clothes itself with a cell-wall" (p. 10); that in "the spores of Mosses and Vascular Cryptogams" and in "the pollen of Phanerogams" ... "the protoplasmic body of the mother-cell breaks up into four lumps, which quickly round themselves off and contract, and become enveloped by a cell-membrane only after complete separation" (p. 13); that in the Equisetaceae "the young spores, when first separated, are still naked, but they soon become surrounded by a cell-membrane" (p. 14); and that in higher plants, as in the pollen of many Dicotyledons, "the contracting daughter-cells secrete cellulose even during their separation" (p. 14). Here, then, in whatever way we interpret it, the fact is that there quickly arises an outer layer different from the contained matter. But the most significant evidence is furnished by "the masses of protoplasm that escape into water from the injured sacs of Vaucheria, which often instantly become rounded into globular bodies," and of which the "hyaline protoplasm envelopes the whole as a skin" (p. 41) which "is denser than the inner and more watery substance" (p. 42). As in this case the protoplasm is but a fragment, and as it is removed from the influence of the parent-cell, this differentiating process can scarcely be regarded as anything more than the effect of physico-chemical actions: a conclusion which is supported by the statement of Sachs that "not only every vacuole in a solid protoplasmic body, but also every thread of protoplasm which penetrates the sap-cavity, and finally the inner side of the protoplasm-sac which encloses the sap-cavity, is also bounded by a skin" (p. 42). If then "every portion of a protoplasmic body immediately surrounds itself, when it becomes isolated, with such a skin," which is shown in all cases to arise at the surface of contact with sap or water, this primary differentiation of outer from inner must be ascribed to the direct action of the medium. Whether the coating thus initiated is secreted by the protoplasm, or whether, as seems more likely, it results from transformation of it, matters not to the argument. Either way the action of the medium causes its formation; and either way the many varied and complex differentiations which developed cell-walls display, must be considered as originating from those variations of this physically-generated covering which natural selection has taken advantage of.

The contained protoplasm of a vegetal cell, which has self-mobility and when liberated sometimes performs amoeba-like motions for a time, may be regarded as an imprisoned amoeba; and when we pass from it to a free amoeba, which is one of the simplest types of first animals, or Protozoa, we naturally meet with kindred phenomena. The general trait which here concerns us, is that while its plastic or semi-fluid sarcode goes on protruding, in irregular ways, now this and now that part of its periphery, and again withdrawing into its interior first one and then another of these temporary processes, perhaps with some small portion of food attached, there is but an indistinct differentiation of outer from inner (a fact shown by the frequent coalescence of the pseudopodia in Rhizopods); but that when it eventually becomes quiescent, the surface becomes differentiated from the contents: the passing into an encysted state, doubtless in large measure due to inherited proclivity, being furthered, and having probably been once initiated, by the action of the medium. The connexion between constancy of relative position among the parts of the sarcode, and the rise of a contrast between superficial and central parts, is perhaps best shown in the minutest and simplest Infusoria, the Monadinae. The genus Monas is described by Kent as "plastic and unstable in form, possessing no distinct cuticular investment; ... the food-substances incepted at all parts of the periphery";[45] and the genus Scytomonas he says "differs from Monas only in its persistent shape and accompanying greater rigidity of the peripheral or ectoplasmic layer."[46] Describing generally such low forms, some of which are said to have neither nucleus nor vacuole, he remarks that in types somewhat higher "the outer or peripheral border of the protoplasmic mass, while not assuming the character of a distinct cell-wall or so-called cuticle, presents, as compared with the inner substance of that mass, a slightly more solid type of composition."[47] And it is added that these forms having so slightly differentiated an exterior, "while usually exhibiting a more or less characteristic normal outline, can revert at will to a pseud-amoeboid and repent state."[48] Here, then, we have several indications of the truth that the permanent externality of a certain part of the substance, is followed by transformation of it into a coating unlike the substance it contains. Indefinite and structureless in the simplest of these forms, as instance again the Gregarina,[49] the limiting membrane becomes, in higher Infusoria, definite and often complex: showing that the selection of favourable variations has had largely to do with its formation. In such types as the Foraminifera, which, almost structureless internally though they are, secrete calcareous shells, it is clear that the nature of this outer layer is determined by inherited constitution. But recognition of this consists with the belief that the action of the medium initiated the outer layer, specialized though it now is; and that even still, contact with the medium excites secretion of it.

A remarkable analogy remains to be named. When we study the action of the medium in an inorganic mass, we are led to see that between the outer changed layer and the inner unchanged mass, comes a surface where active change is going on. Here we have to note that, alike in the plant-cell and in the animal-cell, there is a similar relation of parts. Immediately inside the envelope comes the primordial utricle in the one case, and in the other case the layer of active sarcode. In either case the living protoplasm, placed in the position of a lining to the cuticle of the cell, is shielded from the direct action of the medium, and yet is not beyond the reach of its influences.

* * * * *

Limited, as thus far drawn, to a certain common trait of those minute organisms which are mostly below the reach of unaided vision, the foregoing conclusion appears trivial enough. But it ceases to appear trivial on passing into a wider field, and observing the implications, direct and indirect, as they concern plants and animals of sensible sizes.

Popular expositions of science have so far familiarized many readers with a certain fundamental trait of living things around, that they have ceased to perceive how marvellous a trait it is, and, until interpreted by the Theory of Evolution, how utterly mysterious. In past times, the conception of an ordinary plant or animal which prevailed, not throughout the world at large only but among the most instructed, was that it is a single continuous entity. One of these livings things was unhesitatingly regarded as being in all respects a unit. Parts it might have, various in their sizes, forms, and compositions; but these were components of a whole which had been from the beginning in its original nature a whole. Even to naturalists fifty years ago, the assertion that a cabbage or a cow, though in one sense a whole, is in another sense a vast society of minute individuals, severally living in greater or less degrees, and some of them maintaining their independent lives unrestrained, would have seemed an absurdity. But this truth which, like so many of the truths established by science, is contrary to that common sense in which most people have so much confidence, has been gradually growing clear since the days when Leeuwenhoeck and his contemporaries began to examine through lenses the minute structures of common plants and animals. Each improvement in the microscope, while it has widened our knowledge of those minute forms of life described above, has revealed further evidence of the fact that all the larger forms of life consist of units severally allied in their fundamental traits to these minute forms of life. Though, as formulated by Schwann and Schleiden, the cell-doctrine has undergone qualifications of statement; yet the qualifications have not been such as to militate against the general proposition that organisms visible to the naked eye, are severally compounded of invisible organisms—using that word in its most comprehensive sense. And then, when the development of any animal is traced, it is found that having been primarily a nucleated cell, and having afterwards become by spontaneous fission a cluster of nucleated cells, it goes on through successive stages to form out of such cells, ever multiplying and modifying in various ways, the several tissues and organs composing the adult.

On the hypothesis of evolution this universal trait has to be accepted not as a fact that is strange but unmeaning. It has to be accepted as evidence that all the visible forms of life have arisen by union of the invisible forms; which, instead of flying apart when they divided, remained together. Various intermediate stages are known. Among plants, those of the Volvox type show us the component protophytes so feebly combined that they severally carry on their lives with no appreciable subordination to the life of the group. And among animals, a parallel relation between the lives of the units and the life of the group is shown us in Uroglena and Syncrypta. From these first stages upwards, may be traced through successively higher types, an increasing subordination of the units to the aggregate; though still a subordination leaving to them conspicuous amounts of individual activity. Joining which facts with the phenomena presented by the cell-multiplication and aggregation of every unfolding germ, naturalists are now accepting the conclusion that by this process of composition from Protozoa, were formed all classes of the Metazoa[50]—(as animals formed by this compounding are now called); and that in a similar way from Protophyta, were formed all classes of what I suppose will be called Metaphyta, though the word does not yet seem to have become current.

And now what is the general meaning of these truths, taken in connexion with the conclusion reached in the last section. It is that this universal trait of the Metazoa and Metaphyta, must be ascribed to the primitive action and re-action between the organism and its medium. The operation of those forces which produced the primary differentiation of outer from inner in early minute masses of protoplasm, pre-determined this universal cell-structure of all embryos, plant and animal, and the consequent cell-composition of adult forms arising from them. How unavoidable is this implication, will be seen on carrying further an illustration already used—that of the shingle-covered shore, the pebbles on which, while being in some cases selected, have been in all cases rounded and smoothed. Suppose a bed of such shingle to be, as we often see it, solidified, along with interfused material, into a conglomerate. What in such case must be considered as the chief trait of such conglomerate; or rather—what must we regard as the chief cause of its distinctive characters? Evidently the action of the sea. Without the breakers, no pebbles; without the pebbles, no conglomerate. Similarly then, in the absence of that action of the medium by which was effected the differentiation of outer from inner in those microscopic portions of protoplasm constituting the earliest and simplest animals and plants, there could not have existed this cardinal trait of composition which all the higher animals and plants show us.

So that, active as has been the part played by natural selection, alike in modifying and moulding the original units—largely as survival of the fittest has been instrumental in furthering and controlling the combination of these units into visible organisms, and eventually into large ones; yet we must ascribe to the direct effect of the medium on the first forms of life, that character of which this everywhere-operative factor has taken advantage.

* * * * *

Let us turn now to another and more obvious attribute of higher organisms, for which also there is this same general cause. Let us observe how, on a higher platform, there recurs this differentiation of outer from inner—how this primary trait in the living units with which life commences, re-appears as a primary trait in those aggregates of such units which constitute visible organisms.

In its simplest and most unmistakable form, we see this in the early changes of an unfolding ovum of primitive type. The original fertilized single cell, having by spontaneous fission multiplied into a cluster of such cells, there begins to show itself a contrast between periphery and centre; and presently there is formed a sphere consisting of a superficial layer unlike its contents. The first change, then, is the rise of a difference between that outer part which holds direct converse with the surrounding medium, and that inclosed part which does not. This primary differentiation in these compound embryos of higher animals, parallels the primary differentiation undergone by the simplest living things.

Leaving, for the present, succeeding changes of the compound embryo, the significance of which we shall have to consider by-and-by, let us pass now to the adult forms of visible plants and animals. In them we find cardinal traits which, after what we have seen above, will further impress us with the importance of the effects wrought on the organism by its medium.

From the thallus of a sea-weed up to the leaf of a highly developed phaenogam, we find, at all stages, a contrast between the inner and outer parts of these flattened masses of tissue. In the higher Algae "the outermost layers consist of smaller and firmer cells, while the inner cells are often very large, and sometimes extremely long;"[51] and in the leaves of trees the epidermal layer, besides differing in the sizes and shapes of its component cells from the parenchyma forming the inner substance of the leaf, is itself differentiated by having a continuous cuticle, and by having the outer walls of its cells unlike the inner walls.[52] Especially significant is the structure of such intermediate types as the Liverworts. Beyond the differentiation of the covering cells from the contained cells, and the contrast between upper surface and under surface, the frond of Marchantia polymorpha clearly shows us the direct effect of incident forces; and shows us, too, how it is involved with the effect of inherited proclivities. The frond grows from a flat disc-shaped gemma, the two sides of which are alike. Either side may fall uppermost; and then of the developing shoot, the side exposed to the light "is under all circumstances the upper side which forms stomata, the dark side becomes the under side which produces root-hairs and leafy processes."[53] So that while we have undeniable proof that the contrasted influences of the medium on the two sides, initiate the differentiation, we have also proof that the completion of it is determined by the transmitted structure of the type; since it is impossible to ascribe the development of stomata to the direct action of air and light. On turning from foliar expansions, to stems and roots, facts of like meaning meet us. Speaking generally of epidermal tissue and inner tissue, Sachs remarks that "the contrast of the two is the plainer the more the part of the plant concerned is exposed to air and light."[54] Elsewhere, in correspondence with this, it is said that in roots the cells of the epidermis, though distinguished by bearing hairs, "are otherwise similar to those of the fundamental tissue" which they clothe,[55] while the cuticular covering is relatively thin; whereas in stems the epidermis (often further differentiated) is composed of layers of cells which are smaller and thicker-walled: a stronger contrast of structure corresponding to a stronger contrast of conditions. By way of meeting the suggestion that these respective differences are wholly due to the natural selection of favourable variations, it will suffice if I draw attention to the unlikeness between imbedded roots and exposed roots. While in darkness, and surrounded by moist earth, the outermost protective coats, even of large roots, are comparatively thin; but when the accidents of growth entail permanent exposure to light and air, roots acquire coverings allied in character to the coverings of branches. That the action of the medium causes these and converse changes, cannot be doubted when we find, on the one hand, that "roots can become directly transformed into leaf-bearing shoots," and, on the other hand, that in some plants certain "apparent roots are only underground shoots," and that nevertheless "they are similar to true roots in function and in the formation of tissue, but have no root-cap, and, when they come to the light above ground, continue to grow in the manner of ordinary leaf-shoots."[56] If, then, in highly developed plants inheriting pronounced structures, this differentiating influence of the medium is so marked, it must have been all-important at the outset while types were undetermined.

As with plants so with animals, we find good reason for inferring that while the specialities of the tegumentary parts must be ascribed to the natural selection of favourable variations, their most general traits are due to the direct action of surrounding agencies. Here we come upon the border of those changes which are ascribable to use and disuse. But from this class of changes we may fitly exclude those in which the parts concerned are wholly or mainly passive. A corn and a blister will conveniently serve to illustrate the way in which certain outer actions initiate in the superficial tissues, effects of very marked kinds, which are related neither to the needs of the organism nor to its normal structure. They are neither adaptive changes nor changes towards completion of the type. After noting them we may pass to allied, but still more instructive, changes. Continuous pressure on any portion of the surface causes absorption, while intermittent pressure causes growth: the one impeding circulation and the passage of plasma from the capillaries into the tissues, and the other aiding both. There are yet further mechanically-produced effects. That the general character of the ribbed skin on the under surfaces of the feet and insides of the hands is directly due to friction and intermittent pressure, we have the proofs:—first, that the tracts most exposed to rough usage are the most ribbed; second, that the insides of hands subject to unusual amounts of rough usage, as those of sailors, are strongly ribbed all over; and third, that in hands which are very little used, the parts commonly ribbed become quite smooth. These several kinds of evidence, however, full of meaning as they are, I give simply to prepare the way for evidence of a much more conclusive kind.

Where a wide ulcer has eaten away the deep-seated layer out of which the epidermis grows, or where this layer has been destroyed by an extensive burn, the process of healing is very significant. From the subjacent tissues, which in the normal order have no concern with outward growth, there is produced a new skin, or rather a pro-skin; for this substituted outward-growing layer contains no hair-follicles or other specialities of the original one. Nevertheless, it is like the original one in so far that it is a continually renewed protective covering. Doubtless it may be contended that this make-shift skin results from the inherited proclivity of the type—the tendency to complete afresh the structure of the species when injured. We cannot, however, ignore the immediate influence of the medium, on recalling the facts above named, or on remembering the further fact that an inflamed surface of skin, when not sheltered from the air, will throw out a film of coagulable lymph. But that the direct action of the medium is a chief factor we are clearly shown by another case. Accident or disease occasionally causes permanent eversion, or protrusion, of mucous membrane. After a period of irritability, great at first but decreasing as the change advances, this membrane assumes the general character of ordinary skin. Nor is this all: its microscopic structure changes. Where it is a mucous membrane of the kind covered by cylinder-epithelium, the cylinders gradually shorten, becoming finally flat, and there results a squamous epithelium: there is a near approach in minute composition to epidermis. Here a tendency towards completion of the type cannot be alleged; for there is, contrariwise, divergence from the type. The effect of the medium is so great that, in a short time, it overcomes the inherited proclivity and produces a structure of opposite kind to the normal one.

With but little break we come here upon a significant analogy, parallel to an analogy already described. As was pointed out, an inorganic body that is modifiable by its medium, acquires, after a time, an outer coat which has already undergone such change as surrounding agencies can effect; has a contained mass which is as yet unchanged, because unreached; and has a surface between the two where change is going on—a region of activity. And we saw that alike in the vegetal cell and the animal cell there exist analogous distributions: of course with the difference that the innermost part is not inert. Now we have to note that in those aggregates of cells constituting the Metaphyta and Metazoa, analogous distributions also exist. In plants they are of course not to be looked for in leaves and other deciduous portions, but only in portions of long duration—stems and branches. Naturally, too, we need not expect them in plants having modes of growth which early produce an outer practically dead part, that effectually shields the inner actively living part of the stem from the influence of the medium—long-lived acrogens such as tree-ferns and long-lived endogens such as palms. But in the highest plants, exogens, which have the actively living part of their stems within reach of environing agencies, we find this part,—the cambium layer,—is one from which there is a growth inwards forming wood, and a growth outwards forming bark: there is an increasingly thick covering (where it does not scale off) of tissue changed by the medium, and inside this a film of highest vitality. In so far as concerns the present argument, it is the same with the Metazoa, or at least all of them which have developed organizations. The outer skin grows up from a limiting plane, or layer, a little distance below the surface—a place of predominant vital activity. Here perpetually arise new cells, which, as they develop, are thrust outwards and form the epidermis: flattening and drying up as they approach the surface, whence, having for a time served to shield the parts below, they finally scale off and leave younger ones to take their places. This still undifferentiated tissue forming the base of the epidermis, and existing also as a source of renewal in internal organs, is the essentially living substance; and facts above given imply that it was the action of the medium on this essentially living substance, which, during early stages in the organization of the Metazoa, initiated that protective envelope which presently became an inherited structure—a structure which, though now mainly inherited, still continues to be modifiable by its initiator.

Fully to perceive the way in which these evidences compel us to recognize the influence of the medium as a primordial factor, we need but conceive them as interpreted without it. Suppose, for instance, we say that the structure of the epidermis is wholly determined by the natural selection of favourable variations; what must be the position taken in presence of the fact above named, that when mucous membrane is exposed to the air its cell-structure changes into the cell-structure of skin? The position taken must be this:—Though mucous membrane in a highly-evolved individual organism, thus shows the powerful effect of the medium on its surface; yet we must not suppose that the medium had the effect of producing such a cell-structure on the surfaces of primitive forms, undifferentiated though they were; or, if we suppose that such an effect was produced on them, we must not suppose that it was inheritable. Contrariwise, we must suppose that such effect of the medium either was not wrought at all, or that it was evanescent: though repeated through millions upon millions of generations it left no traces. And we must conclude that this skin-structure arose only in consequence of spontaneous variations not physically initiated (though like those physically initiated) which natural selection laid hold of and increased. Does any one think this a tenable position?

* * * * *

And now we approach the last and chief series of morphological phenomena which must be ascribed to the direct action of environing matters and forces. These are presented to us when we study the early stages in the development of the embryos of the Metazoa in general.

We will set out with the fact already noted in passing, that after repeated spontaneous fissions have changed the original fertilized germ-cell into that cluster of cells which forms a gemmule or a primitive ovum, the first contrast which arises is between the peripheral parts and the central parts. Where, as with lower creatures which do not lay up large stores of nutriment with the germs of their offspring, the inner mass is inconsiderable, the outer layer of cells, which are presently made quite small by repeated subdivisions, forms a membrane extending over the whole surface—the blastoderm. The next stage of development, which ends in this covering layer becoming double, is reached in two ways—by invagination and by delamination; but which is the original way and which the abridged way, is not quite certain. Of invagination, multitudinously exemplified in the lowest types, Mr. Balfour says:—"On purely a priori grounds there is in my opinion more to be said for invagination than for any other view";[57] and, for present purposes, it will suffice if we limit ourselves to this: making its nature clear to the general reader by a simple illustration.

Take a small india-rubber ball—not of the inflated kind, nor of the solid kind, but of the kind about an inch or so in diameter with a small hole through which, under pressure, the air escapes. Suppose that instead of consisting of india-rubber its wall consists of small cells made polyhedral in form by mutual pressure, and united together. This will represent the blastoderm. Now with the finger, thrust in one side of the ball until it touches the other: so making a cup. This action will stand for the process of invagination. Imagine that by continuance of it, the hemispherical cup becomes very much deepened and the opening narrowed, until the cup becomes a sac, of which the introverted wall is everywhere in contact with the outer wall. This will represent the two-layered "gastrula"—the simplest ancestral form of the Metazoa: a form which is permanently represented in some of the lowest types; for it needs but tentacles round the mouth of the sac, to produce a common hydra. Here the fact which it chiefly concerns us to remark, is that of these two layers the outer, called in embryological language the epiblast, continues to carry on direct converse with the forces and matters in the environment; while the inner, called the hypoblast, comes in contact with such only of these matters as are put into the food-cavity which it lines. We have further to note that in the embryos of Metazoa at all advanced in organization, there arises between these two layers a third—the mesoblast. The origin of this is seen in types where the developmental process is not obscured by the presence of a large food-yolk. While the above-described introversion is taking place, and before the inner surfaces of the resulting epiblast and hypoblast have come into contact, cells, or amoeboid units equivalent to them, are budded off from one or both of these inner surfaces, or some part of one or other; and these form a layer which eventually lies between the other two—a layer which, as this mode of formation implies, never has any converse with the surrounding medium and its contents, or with the nutritive bodies taken in from it. The striking facts to which this description is a necessary introduction, may now be stated. From the outer layer, or epiblast, are developed the permanent epidermis and its out-growths, the nervous system, and the organs of sense. From the introverted layer, or hypoblast, are developed the alimentary canal and those parts of its appended organs, liver, pancreas, &c., which are concerned in delivering their secretions into the alimentary canal, as well as the linings of those ramifying tubes in the lungs which convey air to the places where gaseous exchange is effected. And from the mesoblast originate the bones, the muscles, the heart and blood-vessels, and the lymphatics, together with such parts of various internal organs as are most remotely concerned with the outer world. Minor qualifications being admitted, there remain the broad general facts, that out of that part of the external layer which remains permanently external, are developed all the structures which carry on intercourse with the medium and its contents, active and passive; out of the introverted part of this external layer, are developed the structures which carry on intercourse with the quasi-external substances that are taken into the interior—solid food, water, and air; while out of the mesoblast are developed structures which have never had, from first to last, any intercourse with the environment. Let us contemplate these general facts.

Who would have imagined that the nervous system is a modified portion of the primitive epidermis? In the absence of proofs furnished by the concurrent testimony of embryologists during the last thirty or forty years, who would have believed that the brain arises from an infolded tract of the outer skin, which, sinking down beneath the surface, becomes imbedded in other tissues and eventually surrounded by a bony case? Yet the human nervous system in common with the nervous systems of lower animals is thus originated. In the words of Mr. Balfour, early embryological changes imply that—

"the functions of the central nervous system, which were originally taken by the whole skin, became gradually concentrated in a special part of the skin which was step by step removed from the surface, and has finally become in the higher types a well-defined organ imbedded in the subdermal tissues.... The embryological evidence shows that the ganglion-cells of the central part of the nervous system are originally derived from the simple undifferentiated epithelial cells of the surface of the body."[58]

Less startling perhaps, though still startling enough, is the fact that the eye is evolved out of a portion of the skin; and that while the crystalline lens and its surroundings thus originate, the "percipient portions of the organs of special sense, especially of optic organs, are often formed from the same part of the primitive epidermis" which forms the central nervous system.[59] Similarly is it with the organs for smelling and hearing. These, too, begin as sacs formed by infoldings of the epidermis; and while their parts are developing they are joined from within by nervous structures which were themselves epidermic in origin. How are we to interpret these strange transformations? Observing, as we pass, how absurd from the point of view of the special-creationist, would appear such a filiation of structures, and such a round-about mode of embryonic development, we have here to remark that the process is not one to have been anticipated as a result of natural selection. After numbers of spontaneous variations had occurred, as the hypothesis implies, in useless ways, the variation which primarily initiated a nervous centre might reasonably have been expected to occur in some internal part where it would be fitly located. Its initiation in a dangerous place and subsequent migration to a safe place, would be incomprehensible. Not so if we bear in mind the cardinal truth above set forth, that the structures for holding converse with the medium and its contents, arise in that completely superficial part which is directly affected by the medium and its contents; and if we draw the inference that the external actions themselves initiate the structures. These once commenced, and furthered by natural selection where favourable to life, would form the first term of a series ending in developed sense organs and a developed nervous system.[60]

Though it would enforce the argument, I must, for brevity's sake, pass over the analogous evolution of that introverted layer, or hypoblast, out of which the alimentary canal and attached organs arise. It will suffice to emphasize the fact that having been originally external, this layer continues in its developed form to have a quasi-externality, alike in its digesting part and in its respiratory part; since it continues to deal with matters alien to the organism. I must also refrain from dwelling at length on the fact already adverted to, that the intermediate derived layer, or mesoblast, which was at the outset completely internal, originates those structures which ever remain completely internal, and have no communication with the environment save through the structures developed from the other two: an antithesis which has great significance.

Here, instead of dwelling on these details, it will be better to draw attention to the most general aspect of the facts. Whatever may be the course of subsequent changes, the first change is the formation of a superficial layer or blastoderm; and by whatever series of transformations the adult structure is reached, it is from the blastoderm that all the organs forming the adult originate. Why this marvellous fact?

Meaning is given to it if we go back to the first stage in which Protozoa, having by repeated fissions formed a cluster, then arranged themselves into a hollow sphere, as do the protophytes forming a Volvox. Originally alike all over its surface, the hollow sphere of ciliated units thus formed, would, if not quite spherical, assume a constant attitude when moving through the water; and hence one part of the spheroid would more frequently than the rest come in contact with nutritive matters to be taken in. A division of labour resulting from such a variation being advantageous, and tending therefore to increase in descendants, would end in a differentiation like that shown in the gemmules of various low types of Metazoa, which, ovate in shape, are ciliated over one part of the surface only. There would arise a form in which the cilium-bearing units effected locomotion and aeration; while on the others, assuming an amoeba-like character, devolved the function of absorbing food: a primordial specialization variously indicated by evidence.[61] Just noting that an ancestral origin of this kind is implied by the fact that in low types of Metazoa a hollow sphere of cells is the form first assumed by the unfolding embryo, I draw attention to the point here of chief interest; namely that the primary differentiation of this hollow sphere is in such case determined by a difference in the converse of its parts with the medium and its contents; and that the subsequent invagination arises by a continuance of this differential converse.

Even neglecting this first stage and commencing with the next, in which a "gastrula" has been produced by the permanent introversion of one portion of the surface of the hollow sphere, it will suffice if we consider what must thereafter have happened. That which continued to be the outer surface was the part which from time to time touched quiescent masses and occasionally received the collisions consequent on its own motions or the motions of other things. It was the part to receive the sound-vibrations occasionally propagated through the water; the part to be affected more strongly than any other by those variations in the amounts of light caused by the passing of small bodies close to it; and the part which met those diffused molecules constituting odours. That is to say, from the beginning the surface was the part on which there fell the various influences pervading the environment, the part by which there was received those impressions from the environment serving for the guidance of actions, and the part which had to bear the mechanical re-actions consequent upon such actions. Necessarily, therefore, the surface was the part in which were initiated the various instrumentalities for carrying on intercourse with the environment. To suppose otherwise is to suppose that such instrumentalities arose internally where they could neither be operated on by surrounding agencies nor operate on them,—where the differentiating forces did not come into play, and the differentiated structures had nothing to do; and it is to suppose that meanwhile the parts directly exposed to the differentiating forces remained unchanged. Clearly, then, organization could not but begin on the surface; and having thus begun, its subsequent course could not but be determined by its superficial origin. And hence these remarkable facts showing us that individual evolution is accomplished by successive in-foldings and in-growings. Doubtless natural selection soon came into action, as, for example, in the removal of the rudimentary nervous centres from the surface; since an individual in which they were a little more deeply seated would be less likely to be incapacitated by injury of them. And so in multitudinous other ways. But nevertheless, as we here see, natural selection could operate only under subjection. It could do no more than take advantage of those structural changes which the medium and its contents initiated.

See, then, how large has been the part played by this primordial factor. Had it done no more than give to Protozoa and Protophyta that cell-form which characterizes them—had it done no more than entail the cellular composition which is so remarkable a trait of Metazoa and Metaphyta—had it done no more than cause the repetition in all visible animals and plants of that primary differentiation of outer from inner which it first wrought in animals and plants invisible to the naked eye; it would have done much towards giving to organisms of all kinds certain leading traits. But it has done more than this. By causing the first differentiations of those clusters of units out of which visible animals in general arose, it fixed the starting place for organization, and therefore determined the course of organization; and, doing this, gave indelible traits to embryonic transformations and to adult structures.

* * * * *

Though mainly carried on after the inductive method, the argument at the close of the foregoing section has passed into the deductive. Here let us follow for a space the deductive method pure and simple. Doubtless in biology a priori reasoning is dangerous; but there can be no danger in considering whether its results coincide with those reached by reasoning a posteriori.

Biologists in general agree that in the present state of the world, no such thing happens as the rise of a living creature out of non-living matter. They do not deny, however, that at a remote period in the past, when the temperature of the Earth's surface was much higher than at present, and other physical conditions were unlike those we know, inorganic matter, through successive complications, gave origin to organic matter. So many substances once supposed to belong exclusively to living bodies, have now been formed artificially, that men of science scarcely question the conclusion that there are conditions under which, by yet another step of composition, quaternary compounds of lower types pass into those of highest types. That there once took place gradual divergence of the organic from the inorganic, is, indeed, a necessary implication of the hypothesis of Evolution, taken as a whole; and if we accept it as a whole, we must put to ourselves the question—What were the early stages of progress which followed, after the most complex form of matter had arisen out of forms of matter a degree less complex?

At first, protoplasm could have had no proclivities to one or other arrangement of parts; unless, indeed, a purely mechanical proclivity towards a spherical form when suspended in a liquid. At the outset it must have been passive. In respect of its passivity, primitive organic matter must have been like inorganic matter. No such thing as spontaneous variation could have occurred in it; for variation implies some habitual course of change from which it is a divergence, and is therefore excluded where there is no habitual course of change. In the absence of that cyclical series of metamorphoses which even the simplest living thing now shows us, as a result of its inherited constitution, there could be no point d'appui for natural selection. How, then, did organic evolution begin?

If a primitive mass of organic matter was like a mass of inorganic matter in respect of its passivity, and differed only in respect of its greater changeableness; then we must infer that its first changes conformed to the same general law as do the changes of an inorganic mass. The instability of the homogeneous is a universal principle. In all cases the homogeneous tends to pass into the heterogeneous, and the less heterogeneous into the more heterogeneous. In the primordial units of protoplasm, then, the step with which evolution commenced must have been the passage from a state of complete likeness throughout the mass to a state in which there existed some unlikeness. Further, the cause of this step in one of these portions of organic matter, as in any portion of inorganic matter, must have been the different exposure of its parts to incident forces. What incident forces? Those of its medium or environment. Which were the parts thus differently exposed? Necessarily the outside and the inside. Inevitably, then, alike in the organic aggregate and the inorganic aggregate (supposing it to have coherence enough to maintain constant relative positions among its parts), the first fall from homogeneity to heterogeneity must always have been the differentiation of the external surface from the internal contents. No matter whether the modification was physical or chemical, one of composition or of decomposition, it comes within the same generalization. The direct action of the medium was the primordial factor of organic evolution.

* * * * *

And now, finally, let us look at the factors in their ensemble, and consider the respective parts they play: observing, especially, the ways in which, at successive stages, they severally give place one to another in degree of importance.

Acting alone, the primordial factor must have initiated the primary differentiation in all units of protoplasm alike. I say alike, but I must forthwith qualify the word. For since surrounding influences, physical and chemical, could not be absolutely the same in all places, especially when the first rudiments of living things had spread over a considerable area, there necessarily arose small contrasts between the degrees and kinds of superficial differentiation effected. As soon as these became decided, natural selection came into play; for inevitably the unlikenesses produced among the units had effects on their lives: there was survival of some among the modified forms rather than others. Utterly in the dark though we are respecting the causes which set up that process of fission everywhere occurring among the minutest forms of life, we must infer that, when established, it furthered the spread of those which were most favourably differentiated by the medium. Though natural selection must have become increasingly active when once it had got a start; yet the differentiating action of the medium never ceased to be a co-operator in the development of these first animals and plants. Again taking the lead as there arose the composite forms of animals and plants, and again losing the lead with that advancing differentiation of these higher types which gave more scope to natural selection, it nevertheless continued, and must ever continue, to be a cause, both direct and indirect, of modifications in structure.

Along with that remarkable process which, beginning in minute forms with what is called conjugation, developed into sexual generation, there came into play causes of frequent and marked fortuitous variations. The mixtures of constitutional proclivities made more or less unlike by unlikenesses of physical conditions, inevitably led to occasional concurrences of forces producing deviations of structure. These were of course mostly suppressed, but sometimes increased, by survival of the fittest. When, along with the growing multiplication in forms of life, conflict and competition became continually more active, fortuitous variations of structure of no account in the converse with the medium, became of much account in the struggle with enemies and competitors; and natural selection of such variations became the predominant factor. Especially throughout the plant-world its action appears to have been immensely the most important; and throughout that large part of the animal world characterized by relative inactivity, the survival of individuals that had varied in favourable ways, must all along have been the chief cause of the divergence of species and the occasional production of higher ones.

But gradually with that increase of activity which we see on ascending to successively higher grades of animals, and especially with that increased complexity of life which we also see, there came more and more into play as a factor, the inheritance of those modifications of structure caused by modifications of function. Eventually, among creatures of high organization, this factor became an important one; and I think there is reason to conclude that, in the case of the highest of creatures, civilized men, among whom the kinds of variation which affect survival are too multitudinous to permit easy selection of any one, and among whom survival of the fittest is greatly interfered with, it has become the chief factor: such aid as survival of the fittest gives, being usually limited to the preservation of those in whom the totality of the faculties has been most favourably moulded by functional changes.

Of course this sketch of the relations among the factors must be taken as in large measure a speculation. We are now too far removed from the beginnings of life to obtain data for anything more than tentative conclusions respecting its earliest stages; especially in the absence of any clue to the mode in which multiplication, first agamogenetic and then gamogenetic, was initiated. But it has seemed to me not amiss to present this general conception, by way of showing how the deductive interpretation harmonizes with the several inferences reached by induction.

* * * * *

In his article on Evolution in the Encyclopaedia Britannica, Professor Huxley writes as follows:—

"How far 'natural selection' suffices for the production of species remains to be seen. Few can doubt that, if not the whole cause, it is a very important factor in that operation.... On the evidence of palaeontology, the evolution of many existing forms of animal life from their predecessors is no longer an hypothesis, but an historical fact; it is only the nature of the physiological factors to which that evolution is due which is still open to discussion."

With these passages I may fitly join a remark made in the admirable address Prof. Huxley delivered before unveiling the statue of Mr. Darwin in the Museum at South Kensington. Deprecating the supposition that an authoritative sanction was given by the ceremony to the current ideas concerning organic evolution, he said that "science commits suicide when it adopts a creed."

Along with larger motives, one motive which has joined in prompting the foregoing articles, has been the desire to point out that already among biologists, the beliefs concerning the origin of species have assumed too much the character of a creed; and that while becoming settled they have been narrowed. So far from further broadening that broader view which Mr. Darwin reached as he grew older, his followers appear to have retrograded towards a more restricted view than he ever expressed. Thus there seems occasion for recognizing the warning uttered by Prof. Huxley, as not uncalled for.

Whatever may be thought of the arguments and conclusions set forth in this article and the preceding one, they will perhaps serve to show that it is as yet far too soon to close the inquiry concerning the causes of organic evolution.

NOTE.

[The following passages formed part of a preface to the small volume in which the foregoing essay re-appeared. I append them here as they cannot now be conveniently prefixed.]

Though the direct bearings of the arguments contained in this Essay are biological, the argument contained in its first half has indirect bearings upon Psychology, Ethics, and Sociology. My belief in the profound importance of these indirect bearings, was originally a chief prompter to set forth the argument; and it now prompts me to re-issue it in permanent form.

Though mental phenomena of many kinds, and especially of the simpler kinds, are explicable only as resulting from the natural selection of favourable variations; yet there are, I believe, still more numerous mental phenomena, including all those of any considerable complexity, which cannot be explained otherwise than as results of the inheritance of functionally-produced modifications. What theory of psychological evolution is espoused, thus depends on acceptance or rejection of the doctrine that not only in the individual, but in the successions of individuals, use and disuse of parts produce respectively increase and decrease of them.

Of course there are involved the conceptions we form of the genesis and nature of our higher emotions; and, by implication, the conceptions we form of our moral intuitions. If functionally-produced modifications are inheritable, then the mental associations habitually produced in individuals by experiences of the relations between actions and their consequences, pleasurable or painful, may, in the successions of individuals, generate innate tendencies to like or dislike such actions. But if not, the genesis of such tendencies is, as we shall see, not satisfactorily explicable.

That our sociological beliefs must also be profoundly affected by the conclusions we draw on this point, is obvious. If a nation is modified en masse by transmission of the effects produced on the natures of its members by those modes of daily activity which its institutions and circumstances involve; then we must infer that such institutions and circumstances mould its members far more rapidly and comprehensively than they can do if the solo cause of adaptation to them is the more frequent survival of individuals who happen to have varied in favourable ways.

I will add only that, considering the width and depth of the effects which acceptance of one or other of these hypotheses must have on our views of Life, Mind, Morals, and Politics, the question—Which of them is true? demands, beyond all other questions whatever, the attention of scientific men.

* * * * *

After the above articles were published, I received from Dr. Downes a copy of a paper "On the Influence of Light on Protoplasm," written by himself and Mr. T.P. Blunt, M.A., which was communicated to the Royal Society in 1878. It was a continuation of a preceding paper which, referring chiefly to Bacteria, contended that—

"Light is inimical to, and under favourable conditions may wholly prevent, the development of these organisms."

This supplementary paper goes on to show that the injurious effect of light upon protoplasm results only in presence of oxygen. Taking first a comparatively simple type of molecule which enters into the composition of organic matter, the authors say, after detailing experiments:—

"It was evident, therefore, that oxygen was the agent of destruction under the influence of sunlight."

And accounts of experiments upon minute organisms are followed by the sentence—

"It seemed, therefore, that in absence of an atmosphere, light failed entirely to produce any effect on such organisms as were able to appear."

They sum up the results of their experiments in the paragraph—

"We conclude, therefore, both from analogy and from direct experiment, that the observed action on these organisms is not dependent on light per se, but that the presence of free oxygen is necessary; light and oxygen together accomplishing what neither can do alone: and the inference seems irresistible that the effect produced is a gradual oxidation of the constituent protoplasm of these organisms, and that, in this respect, protoplasm, although living, is not exempt from laws which appear to govern the relations of light and oxygen to forms of matter less highly endowed. A force which is indirectly absolutely essential to life as we know it, and matter in the absence of which life has not yet been proved to exist, here unite for its destruction."

What is the obvious implication? If oxygen in presence of light destroys one of these minutest portions of protoplasm, what will be its effect on a larger portion of protoplasm? It will work an effect on the surface instead of on the whole mass. Not like the minutest mass made inert all through, the larger mass will be made inert only on its outside; and, indeed, the like will happen with the minutest mass if the light or the oxygen is very small in quantity. Hence there will result an envelope of changed matter, inclosing and protecting the unchanged protoplasm—there will result a rudimentary cell-wall.

FOOTNOTES:

[Footnote 41: It is probable that this shortening has resulted not directly but indirectly, from the selection of individuals which were noted for tenacity of hold; for the bull-dog's peculiarity in this respect seems due to relative shortness of the upper jaw, giving the underhung structure which, involving retreat of the nostrils, enables the dog to continue breathing while holding.]

[Footnote 42: Though Mr. Darwin approved of this expression and occasionally employed it, he did not adopt it for general use; contending, very truly, that the expression Natural Selection is in some cases more convenient. See Animals and Plants under Domestication (first edition) Vol. i, p. 6; and Origin of Species (sixth edition) p. 49.]