Essays: Scientific, Political, and Speculative, Volume I

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 amœba-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 à priori reasoning is dangerous; but there can be no danger in considering whether its results coincide with those reached by reasoning à 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 Encyclopædia 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 palæontology, 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.

A COUNTER-CRITICISM

[First published in The Nineteenth Century, for February, 1888.]

While I do not concur in sundry of the statements and conclusions contained in the article entitled "A Great Confession," contributed by the Duke of Argyll to the last number of this Review, yet I am obliged to him for having raised afresh the question discussed in it. Though the injunction "Rest and be thankful," is one for which in many spheres much may be said – especially in the political, where undue restlessness is proving very mischievous; yet rest and be thankful is an injunction out of place in science. Unhappily, while politicians have not duly regarded it, it appears to have been taken to heart too much by naturalists; in so far, at least, as concerns the question of the origin of species.

The new biological orthodoxy behaves just as the old biological orthodoxy did. In the days before Darwin, those who occupied themselves with the phenomena of life, passed by with unobservant eyes the multitudinous facts which point to an evolutionary origin for plants and animals; and they turned deaf ears to those who insisted on the significance of these facts. Now that they have come to believe in this evolutionary origin, and have at the same time accepted the hypothesis that natural selection has been the sole cause of the evolution, they are similarly unobservant of the multitudinous facts which cannot rationally be ascribed to that cause; and turn deaf ears to those who would draw their attention to them. The attitude is the same; it is only the creed which has changed.

But, as above implied, though the protest of the Duke of Argyll against this attitude is quite justifiable, it seems to me that many of his statements cannot be sustained. Some of these concern me personally, and others are of impersonal concern. I propose to deal with them in the order in which they occur.

On page 144 the Duke of Argyll quotes me as omitting "for the present any consideration of a factor which may be distinguished as primordial;" and he represents me as implying by this "that Darwin's ultimate conception of some primordial 'breathing of the breath of life' is a conception which can be omitted only 'for the present.'" Even had there been no other obvious interpretation, it would have been a somewhat rash assumption that this was my meaning when referring to an omitted factor; and it is surprising that this assumption should have been made after reading the second of the two articles criticised, in which this factor omitted from the first is dealt with: this omitted third factor being the direct physico-chemical action of the medium on the organism. Such a thought as that which the Duke of Argyll ascribes to me, is so incongruous with the beliefs I have in many places expressed that the ascription of it never occurred to me as possible.

Lower down on the same page are some other sentences having personal implications, which I must dispose of before going into the general question. The Duke says "it is more than doubtful whether any value attaches to the new factor with which he [I] desires to supplement it [natural selection]"; and he thinks it "unaccountable" that I "should make so great a fuss about so small a matter as the effect of use and disuse of particular organs as a separate and a newly-recognised factor in the development of varieties." I do not suppose that the Duke of Argyll intended to cast upon me the disagreeable imputation, that I claim as new that which all who are even slightly acquainted with the facts know to be anything rather than new. But his words certainly do this. How he should have thus written in spite of the extensive knowledge of the matter which he evidently has, and how he should have thus written in presence of the evidence contained in the articles he criticizes, I cannot understand. Naturalists, and multitudes besides naturalists, know that the hypothesis which I am represented as putting forward as new, is much older than the hypothesis of natural selection – goes back at least as far as Dr. Erasmus Darwin. My purpose was to bring into the foreground again a factor which has, I think, been of late years improperly ignored; to show that Mr. Darwin recognized this factor in an increasing degree as he grew older (by showing which I should have thought I sufficiently excluded the supposition that I brought it forward as new); to give further evidence that this factor is in operation; to show there are numerous phenomena which cannot be interpreted without it; and to argue that if proved operative in any case, it may be inferred that it is operative on all structures having active functions.