Popular Science Monthly/Volume 29/May 1886/The Factors of Organic Evolution II
|THE FACTORS OF ORGANIC EVOLUTION.|
By HERBERT SPENCER.
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 continuance in an active state of such individual when others have been overthrown. Mr. Darwin was conscious of these misleading implications. In the introduction to his Animals and Plants under Domestication (p. 7) he says:
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; 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 interdependent 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 great 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.
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 organization 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.
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. 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 in 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 far 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 Philæ, 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 eight or ten 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 these 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 Algæ as the Conjugatæ; 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 says of the lowest Algæ 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 mothercell 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 Equisetaceæ "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 protoplasmic 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 some 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 amœba, 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 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 Monadinæ. 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"; and the genus Scytomonas he says "differs from Monas only in its persistent shape and accompanying greater rigidity of the peripheral or ectoplasmic layer." Describing generally such low and minute forms, some of which 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." 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-amœboid and repent state." 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, 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.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
- 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.
- It is true that while not deliberately admitted by Mr. Darwin, these effects are not denied by him. In his Animals and Plants under Domestication (vol. ii, 281), he refers to certain chapters in the Principles of Biology, in which I have discussed this general inter-action of the medium and the organism, and ascribed certain most general traits to it. But though, by his expressions, he implies a sympathetic attention to the argument, he does not in such way adopt the conclusion as to assign to this factor any share in the genesis of organic structures—much less that large share which I believe it has had. I did not myself at that time, nor indeed until quite recently, see how extensive and profound have been the influences on organization which, as we shall presently see, are traceable to the early results of this fundamental relation between organism and medium. I may add that it is in an essay on "Transcendental Physiology," first published in 1857, that the line of thought here followed out in it wider bearings, was first entered upon.
- Text-Book of Botany, &c. by Julius Sachs. Translated by A. W. Bennett and W. T. T. Dyer.
- A Manual of the Infusoria, by W. Saville Kent. Vol. i, p. 232.
- lb. Vol. i, p. 241.
- Kent. Vol. i, p. 56.
- lb., 1. c. Vol. i, p. 57.
- The Elements of Comparative Anatomy, by T. H. Huxley, pp. 7-9.