Popular Science Monthly/Volume 42/April 1893/The Inadequacy of Natural Selection I

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THE INADEQUACY OF "NATURAL SELECTION."
By HERBERT SPENCER.

Students of psychology are familiar with the experiments of Weber on the sense of touch. He found that different parts of the surface differ widely in their ability to give information concerning the things touched. Some parts, which yielded vivid sensations, yielded little or no knowledge of the size or form of the thing exciting it; whereas other parts, from which there came sensations much less acute, furnished clear impressions respecting tangible characters, even of relatively small objects. These unlikenesses of tactual discriminativeness he ingeniously expressed by actual measurements. Taking a pair of compasses, he found that if they were closed so nearly that the points were less than one twelfth of an inch apart, the end of the forefinger could not perceive that there were two points: the two points seemed one. But when the compasses were opened so that the points were one twelfth of an inch apart, then the end of the forefinger distinguished the two points. On the other hand, he found that the compasses must be opened to the extent of two and a half inches before the middle of the back could distinguish between two points and one. That is to say, as thus measured, the end of the forefinger has thirty times the tactual discriminativeness which the middle of the back has.

Between these extremes he found gradations. The inner surfaces of the second joints of the fingers can distinguish separateness of positions only half as well as the tip of the forefinger. The innermost joints are still less discriminating, but have a power of discrimination equal to that of the tip of the nose. The end of the great toe, the palm of the hand, and the cheek, have alike one fifth of the perceptiveness which the tip of the forefinger has; and the lower part of the forehead has but one half that possessed by the cheek. The back of the hand and the crown of the head are nearly alike in having but a fourteenth or a fifteenth of the ability to perceive positions as distinct, which is possessed by the finger-end. The thigh, near the knee, has rather less, and the breast less still; so that the compasses must be more than an inch and a half apart before the breast distinguishes the two points from one another.

What is the meaning of these differences? How, in the course of evolution, have they been established? If "natural selection" or survival of the fittest is the assigned cause, then it is required to show in what way each of these degrees of endowment has advantaged the possessor to such extent that not infrequently life has been directly or indirectly preserved by it. We might reasonably assume that in the absence of some differentiating process, all parts of the surface would have like powers of perceiving relative positions. They can not have become widely unlike in perceptiveness without some cause. And if the cause alleged is natural selection, then it is necessary to show that the greater degree of the power possessed by this part than by that, has not only conduced to the maintenance of life, but has conduced so much that an individual in whom a variation had produced better adjustment to needs, thereby maintained life when some others lost it; and that among the descendants inheriting this variation, there was a derived advantage such as enabled them to multiply more than the descendants of individuals not possessing it. Can this, or anything like this, be shown?

That the superior perceptiveness of the forefinger-tip has thus arisen, might be contended with some apparent reason. Such perceptiveness is an important aid to manipulation, and may have sometimes given a life-saving advantage. In making arrows or fish-hooks, a savage possessing some extra amount of it may have been thereby enabled to get food where another failed. In civilized life, too, a seamstress with well-endowed finger-ends might be expected to gain a better livelihood than one with finger-ends which were obtuse; though this advantage would not be so great as appears. I have found that two ladies whose finger-ends were covered with glove-tips, reducing their sensitiveness from one twelfth of an inch between compass points to one seventh, lost nothing appreciable of their quickness and goodness in sewing. An experience of my own here comes in evidence. Toward the close of my salmon-fishing days, I used to observe what a bungler I had become in putting on and taking off artificial flies. As the tactual discriminativeness of my finger-ends, recently tested, comes up to the standard specified by Weber, it is clear that this decrease of manipulative power, accompanying increase of age, was due to decrease in the delicacy of muscular co-ordination and sense of pressure—not to decrease of tactual discriminativeness. But not making much of these criticisms, let us admit the conclusion that this high perceptive power possessed by the forefinger-end may have arisen by survival of the fittest; and let us limit the argument to the other differences.

How about the back of the trunk and its face? Is any advantage derived from possession of greater tactual discriminativeness by the last than by the first? The tip of the nose has more than three times the power of distinguishing relative positions which the lower part of the forehead has. Can this greater power be shown to have any advantage? The back of the hand has scarcely more discriminative ability than the crown of the head, and has only one fourteenth of that which the finger-tip has. Why is this? Advantage might occasionally be derived if the back of the hand could tell us more than it does about the shapes of the surfaces touched. Why should the thigh near the knee be twice as perceptive as the middle of the thigh? And, last of all, why should the middle of the forearm, middle of the thigh, middle of the back of the neck, and middle of the back, all stand on the lowest level, as having but one thirtieth of the perceptive power which the tip of the forefinger has? To prove that these differences have arisen by natural selection, it has to be shown that such small variation in one of the parts as might occur in a generation—say one tenth extra amount—has yielded an appreciably greater power of self-preservation, and that those inheriting it have continued to be so far advantaged as to multiply more than those who, in other respects equal, were less endowed with this trait. Does any one think he can show this?

But if this distribution of tactual perceptiveness can not be explained by survival of the fittest, how can it be explained? The reply is that, if there has been in operation a cause which it is now the fashion among biologists to ignore or deny, these various differences are at once accounted for. This cause is the inheritance of acquired characters. As a preliminary to setting forth the argument showing this, I have made some experiments.

It is a current belief that the fingers of the blind, more practiced in tactual exploration than the fingers of those who can see, acquire greater discriminativeness: especially the fingers of those blind who have been taught to read from raised letters. Not wishing to trust to this current belief, I recently tested two youths, one of fifteen and the other younger, at the School for the Blind in Upper Avenue Road, and found the belief to be correct. Instead of being unable to distinguish between points of the compasses until they were opened to one twelfth of an inch apart, I found that both of them could distinguish between points when only one fourteenth of an inch apart. They had thick and coarse skins; and doubtless, had this intervening obstacle so produced been less, the discriminative power would have been greater. It afterward occurred to me that a better test would be furnished by those whose finger-ends are exercised in tactual perceptions, not occasionally, as by the blind in reading, but all day long in pursuit of their occupations. The facts answered expectation. Two skilled compositors, on whom I experimented, were both able to distinguish between points when they were only one seventeenth of an inch apart. Thus we have clear proof that constant exercise of the tactual nervous structures leads to further development.[1]

Now if acquired structural traits are inheritable, the various contrasts above set down are obvious consequences; for the gradations in tactual perceptiveness correspond with the gradations in the tactual exercise of the parts. Save by contact with clothes, which present only broad surfaces having but slight and indefinite contrasts, the trunk has but little converse with external bodies, and it has but small discriminative power; but what discriminative power it has is greater on its face than on its back, corresponding to the fact that the chest and abdomen are much more frequently explored by the hands: this difference being probably in part inherited from inferior creatures, for, as we may see in dogs and cats, the belly is far more accessible to feet and tongue than the back. No less obtuse than the back are the middle of the back of the neck, the middle of the forearm, and the middle of the thigh; and these parts have but rare experiences of irregular foreign bodies. The crown of the head is occasionally felt by the fingers, as also the back of one hand by the fingers of the other; but neither of these surfaces, which are only twice as perceptive as the back, is used with any frequency for touching objects, much less for examining them. The lower part of the forehead, though more perceptive than the crown of the head, in correspondence with a somewhat greater converse with the hands, is less than one third as perceptive as the tip of the nose; and manifestly, both in virtue of its relative prominence, in virtue of its contacts with things smelt at, and in virtue of its frequent acquaintance with the handkerchief, the tip of the nose has far greater tactual experience. Passing to the inner surfaces of the hands, which, taken as wholes, are more constantly occupied in touching than are the back, breast, thigh, forearm, forehead, or back of the hand, Weber's scale shows that they are much more perceptive, and that the degrees of perceptiveness of different parts correspond with their tactual activities. The palms have but one fifth the perceptiveness possessed by the forefinger-ends; the inner surfaces of the finger-joints next the palms have but one third, while the inner surfaces of the second joints have but one half. These abilities correspond with the facts that whereas the inner parts of the hand are used only in grasping things, the tips of the fingers come into play not only when things are grasped, but when such things, as well as smaller things, are felt at or manipulated. It needs but to observe the relative actions of these parts in writing, in sewing, in judging textures, etc., to see that above all other parts the finger-ends, and especially the forefinger-ends, have the most multiplied experiences. If, then, it be that the extra perceptiveness acquired from extra tactual activities, as in a compositor, is inheritable, these gradations of tactual perceptiveness are explained.

Doubtless some of those who remember Weber's results, have had on the tip of the tongue the argument derived from the tip of the tongue. This part exceeds all other parts in power of tactual discrimination: doubling, in that respect, the power of the forefinger-tip. It can distinguish points that are only one twenty-fourth of an inch apart. Why this unparalleled perceptiveness? If survival of the fittest be the ascribed cause, then it has to be shown what the advantages achieved have been; and, further, that those advantages have been sufficiently great to have had effects on the maintenance of life.

Besides tasting, there are two functions conducive to life, which the tongue performs. It enables us to move about food during mastication, and it enables us to make many of the articulations constituting speech. But how does the extreme discriminativeness of the tongue-tip aid these functions? The food is moved about, not by the tongue-tip, but by the body of the tongue; and even were the tip largely employed in this process, it would still have to be shown that its ability to distinguish between points one twenty-fourth of an inch apart, is of service to that end, which can not be shown. It may, indeed, be said that the tactual perceptiveness of the tongue-tip serves for detection of foreign bodies in the food, as plum-stones or as fish-bones. But such extreme perceptiveness is needless for the purpose—a perceptiveness equal to that of the finger-ends would suffice; and further, even were such extreme perceptiveness useful, it could not have caused survival of individuals who possessed it in slightly higher degrees than others. It needs but to observe a dog crunching small bones, and swallowing with impunity the sharp-angled pieces, to see that but a very small amount of mortality would be prevented.

But what about speech? Well, neither here can there be shown any advantage derived from this extreme perceptiveness. For making the s and z, the tongue has to be partially applied to a portion of the palate next the teeth. Not only, however, must the contact be incomplete, but its place is indefinite—may be half an inch further back. To make the sh and zh, the contact has to be made, not with the tip, but with the upper surface of the tongue; and must be an incomplete contact. Though, for making the liquids, the tip of the tongue and the sides of the tongue are used, yet the requisite is not any exact adjustment of the tip, but an imperfect contact with the palate. For the th, the tip is used along with the edges of the tongue; but no perfect adjustment is required, either to the edges of the teeth, or to the junction of the teeth with the palate, where the sound may equally well be made. Though for the t and d complete contact of the tip and edges of the tongue with the palate is required, yet the place of contact is not definite, and the tip takes no more important share in the action than the sides. Any one who observes the movements of his tongue in speaking, will find that there occur no cases in which the adjustments must have an exactness corresponding to the extreme power of discrimination which the tip possesses: for speech, this endowment is useless. Even were it useful, it could not be shown that it has been developed by survival of the fittest; for though perfect articulation is useful, yet imperfect articulation has rarely such an effect as to impede a man in the maintenance of his life. If he is a good workman, a German's interchanges of b's and p's do not disadvantage him. A Frenchman who, in place of the sound of th, always makes the sound of z, succeeds as a teacher of music or dancing, no less than if he achieved the English pronunciation. Nay, even such an imperfection of speech as that which arises from cleft palate, does not prevent a man from getting on if he is capable. True, it may go against him as a candidate for Parliament, or as an "orator" of the unemployed (mostly not worth employing). But in the struggle for life he is not hindered by the effect to the extent of being less able than others to maintain himself and his offspring. Clearly, then, even if this unparalleled perceptiveness of the tongue-tip is required for perfect speech, this use is not sufficiently important to have been developed by natural selection.

How, then, is this remarkable trait of the tongue-tip to be accounted for? Without difficulty, if there is inheritance of acquired characters. For the tongue-tip has, above all other parts of the body, unceasing experiences of small irregularities of surface. It is in contact with the teeth, and either consciously or unconsciously is continually exploring them. There is hardly a moment in which impressions of adjacent but different positions are not being yielded to it by either the surfaces of the teeth or their edges; and it is continually being moved about from some of them to others. No advantage is gained. It is simply that the tongue's position renders perpetual exploration almost inevitable; and by perpetual exploration is developed this unique power of discrimination. Thus the law holds throughout, from this highest degree of perceptiveness of the tongue-tip to its lowest degree on the back of the trunk; and no other explanation of the facts seems possible.

"Yes, there is another explanation," I hear some one say: "they may be explained by panmixia." Well, in the first place, as the explanation by panmixia implies that these gradations of perceptiveness have been arrived at by the dwindling of nervous structures, there lies at the basis of the explanation an unproved and improbable assumption; and, even were there no such difficulty, it may with certainty be denied that panmixia can furnish an explanation. Let us look at its pretensions.

It was not without good reason that Bentham protested against metaphors. Figures of speech in general, valuable as they are in poetry and rhetoric, can not be used without danger in science and philosophy. The title of Mr. Darwin's great work furnishes us with an instance of the misleading effects produced by them. It runs:—The Origin of Species by means of Natural Selection, or the preservation of Favored Races in the Struggle for Life. Here are two figures of speech which conspire to produce an impression more or less erroneous. The expression "natural selection" was chosen as serving to indicate some parallelism with artificial selection—the selection exercised by breeders. Now selection connotes volition, and thus gives to the thoughts of readers a wrong bias. Some increase of this bias is produced by the words in the second title, "favored races;" for anything which is favored implies the existence of some agent conferring a favor. I do not mean that Mr. Darwin himself failed to recognize the misleading connotations of his words, or that he did not avoid being misled by them. In chapter iv of the Origin of Species he says that, considered literally, "natural selection is a false term," and that the personification of Nature is objectionable; but he thinks that readers, and those who adopt his views, will soon learn to guard themselves against the wrong implications. Here I venture to think that he was mistaken. For thinking this there is the reason that even his disciple, Mr. Wallace—no, not his disciple, but his co-discoverer, ever to be honored—has apparently been influenced by them. When for example, in combating a view of mine, he says that "the very thing said to be impossible by variation and natural selection has been again and again effected by variation and artificial selection"; he seems clearly to imply that the processes are analogous and operate in the same way. Now this is untrue. They are analogous only within certain narrow limits; and, in the great majority of cases, natural selection is utterly incapable of doing that which artificial selection does.

To see this it needs only to de-personalize Nature, and to remember that, as Mr. Darwin says, Nature is "only the aggregate action and product of many natural laws [forces]." Observe its relative shortcomings. Artificial selection can pick out a particular trait, and, regardless of other traits of the individuals displaying it, can increase it by selective breeding in successive generations. For, to the breeder or fancier, it matters little whether such individuals are otherwise well constituted. They may be in this or that way so unfit for carrying on the struggle for life, that, were they without human care, they would disappear forthwith. On the other hand, if we regard Nature as that which it is, an assemblage of various forces, inorganic and organic, some favorable to the maintenance of life and many at variance with its maintenance—forces which operate blindly—we see that there is no such selection of this or that trait, but that there is a selection only of individuals which are, by the aggregate of their traits, best fitted for living. And here I may note an advantage possessed by the expression "survival of the fittest"; since this does not tend to raise the thought of any one character which, more than others, is to be maintained or increased; but tends rather to raise the thought of a general adaptation for all purposes. It implies the process which Nature can alone carry on—the leaving alive of those which are best able to utilize surrounding aids to life, and best able to combat or avoid surrounding dangers. And while this phrase covers the great mass of cases in which there are preserved well-constituted individuals, it also covers those special cases which are suggested by the phrase "natural selection," in which individuals succeed beyond others in the struggle for life by the help of particular characters which conduce in important ways to prosperity and multiplication. For now observe the fact which here chiefly concerns us, that survival of the fittest can increase any serviceable trait only if that trait conduces to prosperity of the individual, or of posterity, or of both, in an important degree. There can be no increase of any structure by natural selection unless, amid all the slightly varying structures constituting the organism, increase of this particular one is so advantageous as to cause greater multiplication of the family in which it arises than of other families. Variations which, though advantageous, fail to do this, must disappear again. Let us take a case.

Keenness of scent in a deer, by giving early notice of approaching enemies, subserves life so greatly that, other things equal, an individual having it in an unusual degree is more likely than others to survive, and, among descendants, to leave some similarly endowed or more endowed, who again transmit the variation with, in some cases, increase. Clearly this highly useful power may be developed by natural selection. So also, for like reasons, may quickness of vision and delicacy of hearing. Though it may be remarked in passing that since this extra sense-endowment, serving to give early alarm, profits the herd as a whole, which takes the alarm from one individual, selection of it is not so easy, unless it occurs in a conquering stag. But now suppose that one member of the herd—perhaps because of more efficient teeth, perhaps by greater muscularity of stomach, perhaps by secretion of more appropriate gastric juices—is enabled to eat and digest a not uncommon plant which the others refuse. This peculiarity may, if food is scarce, conduce to better self-maintenance, and better fostering of young, if the individual is a hind. But unless this plant is abundant, and the advantage consequently great, the advantages which other members of the herd gain from other slight variations may be equivalent. This one has unusual agility and leaps a chasm which others balk at. That one develops longer hair in winter, and resists the cold better. Another has a skin less irritated by flies, and can graze without so much interruption. Here is one which has an unusual power of detecting food under the snow; and there is one which shows extra sagacity in the choice of a shelter from wind and rain. That the variation giving the ability to eat a plant before unutilized, may become a trait of the herd, and eventually of a variety, it is needful that the individual in which it occurs shall have more descendants, or better descendants, or both, than have the various other individuals severally having their small superiorities. If these other individuals severally profit by their small superiorities, and transmit them to equally large numbers of offspring, no increase of the variation in question can take place: it must soon be canceled. Whether in the Origin of Species Mr. Darwin has recognized this fact, I do not remember, but he has certainly done it by implication in his Animals and Plants under Domestication. Speaking of variations in domestic animals, he there says that, "Any particular variation would generally be lost by crossing, reversion, and the accidental destruction of the varying individuals, unless carefully preserved by man" (Vol. II, 292). That which survival of the fittest does in cases like the one I have instanced is to keep all faculties up to the mark, by destroying such as have faculties in some respect below the mark; and it can produce development of some one faculty only if that faculty is predominantly important. It seems to me that many naturalists have practically lost sight of this, and assume that natural selection will increase any advantageous trait. Certainly a view now widely accepted assumes as much.

The consideration of this view, to which the foregoing paragraph is introductory, may now be entered upon. This view concerns, not direct selection, but what has been called, in questionable logic, "reversed selection"—the selection which effects, not increase of an organ, but decrease of it. For as, under some conditions, it is of advantage to an individual and its descendants to have some structure of larger size, it may be, under other conditions—namely, when the organ becomes useless—of advantage to have it of smaller size; since, even if it is not in the way, its weight and the cost of its nutrition are injurious taxes on the organism. But now comes the truth to be emphasized. Just as direct selection can increase an organ only in certain cases, so can reversed selection decrease it only in certain eases. Like the increase produced by a variation, the decrease produced by one must be such as will sensibly conduce to preservation and multiplication. It is, for instance, conceivable that were the long and massive tail of the kangaroo to become useless (say by the forcing of the species into a mountainous and rocky habitat filled with brushwood), a variation which considerably reduced the tail might sensibly profit the individual in which it occurred; and, in seasons when food was scarce, might cause survival when individuals with large tails died. But the economy of nutrition must be considerable before any such result could occur. Suppose that in this new habitat the kangaroo had no enemies; and suppose that, consequently, quickness of hearing not being called for, large ears gave no greater advantage than small ones. Would an individual with smaller ears than usual survive and propagate better than other individuals in consequence of the economy of nutrition achieved? To suppose this is to suppose that the saving of a grain or two of protein per day would determine the kangaroo's fate.

Long ago I discussed this matter in the Principles of Biology (§ 166), taking as an instance the decrease of the jaw implied by the crowding of the teeth, and now proved by measurement to have taken place. Here is the passage:—

"No functional superiority possessed by a small jaw over a large jaw, in civilized life, can be named as having caused the more frequent survival of small-jawed individuals. The only advantage which smallness of jaw might be supposed to give, is the advantage of economized nutrition; and this could not be great enough to further the preservation of men possessing it. The decrease of weight in the jaw and co-operative parts that has arisen in the course of many thousands of years, does not amount to more than a few ounces. This decrease has to be divided among the many generations that have lived and died in the interval. Let us admit that the weight of these parts diminished to the extent of an ounce in a single generation (which is a large admission); it still can not be contended that the having to carry an ounce less in weight, or the having to keep in repair an ounce less of tissue, could sensibly affect any man's fate. And if it never did this—nay, if it did not cause a frequent survival of small-jawed individuals where large-jawed individuals died, natural selection could neither cause nor aid diminution of the jaw and its appendages."

When writing this passage in 1864, I never dreamt that a quarter of a century later, the supposable cause of degeneration here examined and excluded as impossible, would be enunciated as not only a cause, but the cause, and the sole cause. This, however, has happened. Weismann's theory of degeneration by panmixia, is that when an organ previously maintained of the needful size by natural selection, is no longer maintained at that size, because it has become useless (or because a smaller size is equally useful), it results that among the variations in the size, which take place from generation to generation, the smaller will be preserved continually, and that so the part will decrease. And this is concluded without asking whether the economy in nutrition achieved by the smaller variation, will sensibly affect the survival of the individual, and the multiplication of its stirp. To make clear his hypothesis, and to prepare the way for criticism, let me quote the example he himself gives when contrasting the alleged efficiency of dwindling by panmixia with the alleged inefficiency of dwindling from disuse. This example is furnished him by the Proteus.

Concerning the "blind fish and amphibia" found in dark places, which have but rudimentary eyes "hidden under the skin," he argues that "it is difficult to reconcile the facts of the case with the ordinary theory that the eyes of these animals have simply degenerated through disuse." After giving instances of rapid degeneration of disused organs, he argues that if "the effects of disuse are so striking in a single life, we should certainly expect, if such effects can be transmitted, that all traces of an eye would soon disappear from a species which lives in the dark." Doubtless this is a reasonable conclusion. To explain the facts on the hypothesis that acquired characters are inheritable seems very difficult. One possible explanation may indeed be named. It appears to be a general law of organization that structures are stable in proportion to their antiquity; that while organs of relatively modern origin have but a comparatively superficial root in the constitution, and readily disappear if the conditions do not favor their maintenance, organs of ancient origin have deep-seated roots in the constitution, and do not readily disappear. Having been early elements in the type, and having continued to be reproduced as parts of it during a period extending throughout many geological epochs, they are comparatively persistent. Now the eye answers to this description as being a very early organ.[2] But waiving possible interpretations, let us admit that here is a difficulty—a difficulty like countless others which the phenomena of evolution present, as, for instance, the acquirement of such a habit as that of the Vanessa larva, hanging itself up by the tail and then changing into a chrysalis which usurps its place—a difficulty which, along with multitudes, has to await future solution, if any can be found. Let it be granted, I say, that here is a serious obstacle in the way of the hypothesis; and now let us turn to the alternative hypothesis, and observe whether it is not met by difficulties which are much more serious. Weismann writes:

"The caverns in Carniola and Carinthia, in which the blind Proteus and so many other blind animals live, belong geologically to the Jurassic formation; and although we do not exactly know when, for example, the Proteus first entered them, the low organization of this amphibian certainly indicates that it has been sheltered there for a very long period of time, and that thousands of generations of this species have succeeded one another in the caves.
 

"Hence there is no reason to wonder at the extent to which the degeneration of the eye has been already carried in the Proteus, even if we assume that it is merely due to the cessation of the conserving influence of natural selection.

 
"But it is unnecessary to depend upon this assumption alone, for when a useless organ degenerates, there are also other factors which demand consideration—namely, the higher development of other organs which compensate for the loss of the degenerating structure, or the increase in size of adjacent parts. If these newer developments are of advantage to the species, they finally come to take the place of the organ which natural selection has failed to preserve at its point of highest perfection."[3]

On these paragraphs let me first remark that one cause is multiplied into two. The cause is stated in the abstract, and it is then re-stated in the concrete, as though it were another cause. Manifestly, if by decrease of the eye an economy of nutriment is achieved, it is implied that the economized nutriment is turned to some advantageous purpose or other; and to specify that the nutriment is used for the further development of compensating organs, simply changes the indefinite statement of advantage into a definite statement of advantage. There are not two causes in operation, though the matter is presented as though there were.

But passing over this, let us now represent to ourselves in detail this process which Prof. Weismann thinks will, in thousands of generations, effect the observed reduction of the eyes: the process being that at each successive stage in the decrease, there must take place variations in the size of the eye, some larger, some smaller, than the size previously reached, and that in virtue of the economy, those having the smaller will continually survive and propagate, instead of those having the larger. Properly to appreciate this supposition, we must use figures. To give it every advantage we will assume that there have been only two thousand generations, and we will assume that, instead of being reduced to a rudiment, the eye has disappeared altogether. What amounts of variation shall we suppose? If the idea is that the process has operated uniformly on each generation, the implication is that some advantage has been gained by the individuals having the eyes 12000 less in weight; and this will hardly be contended. Not to put the hypothesis at this disadvantage, let us then imagine that there take place, at long intervals, decreasing variations considerable in amount—say 120, once in a hundred generations. This is an interval almost too long to be assumed; but yet if we assume the successive decrements to occur more frequently, and therefore to be smaller, the amount of each becomes too insignificant. If, seeing the small head, we assume that the eyes of the Proteus originally weighed some ten grains each, this would give us, as the amount of the decrement of 120, occurring once in a hundred generations, one grain. Suppose that this eel-shaped amphibian, about a foot long and more than half an inch in diameter, weighs three ounces—a very moderate estimate. In such case the decrement would amount to 11440 of the creature's weight; or, for convenience, let us say that it amounted to 11000 which would allow of the eyes being taken at some fourteen grains each.[4] To this extent, then, each occasional decrement would profit the organism. The economy in weight to a creature having nearly the same specific gravity as its medium, would be infinitesimal. The economy in nutrition of a rudimentary organ, consisting of passive tissues, would also be but nominal. The only appreciable economy would be in the original building up of the creature's structures; and the hypothesis of Weismann implies that the economy of this thousandth part of its weight, by decrease of the eyes, would so benefit the rest of the creature's organization as to give it an appreciably greater chance of survival, and an appreciably greater multiplication of descendants. Does any one accept this inference?

Of course the qualifications of data above set down can be only approximate; but I think no reasonable changes of them can alter the general result. If, instead of supposing the eyes to have disappeared wholly, we recognize them as being in fact rudimentary, the case is made worse. If, instead of two thousand generations, we assume ten thousand generations, which, considering the probably great age of the caverns, would be a far more reasonable assumption than the other, the case is made still worse. And if we assume larger variations—say decreases of one fourth—to occur only at intervals of many hundreds or thousands of generations, which is not a very reasonable assumption, the implied conclusion would still remain indefensible. For an economy of 1200 part of the creature's weight could not appreciably affect its survival and the increase of its posterity.

Is it not then, as said above, that the use of the expression, "natural selection," has had seriously perverting effects? Must we not infer that there has been produced in the minds of naturalists, the tacit assumption that it can do what artificial selection does—can pick out and select any small advantageous trait; while it can, in fact, pick out no traits, but can only further the development of traits which, in marked ways, increase the general fitness for the conditions of existence? And is it not inferable that, failing to bear in mind the limiting condition, that to become established an advantageous variation must be such as will, other things remaining equal, add to the prosperity of the stirp, many naturalists have been unawares led to espouse an untenable hypothesis?—Contemporary Review.

[To be concluded.]
  1. Let me here note in passing a highly significant implication. The development of nervous structures which in such cases takes place, can not be limited to the finger-ends. If we figure to ourselves the separate sensitive areas which severally yield independent feelings, as constituting a network (not, indeed, a network sharply marked out, but probably one such that the ultimate fibrils in each area intrude more or less into adjacent areas, so that the separations are indefinite), it is manifest that when, with exercise, the structure has become further elaborated, and the meshes of the network smaller, there must be a multiplication of fibers communicating with the central nervous system. If two adjacent areas were supplied by branches of one fiber, the touching of either would yield to consciousness the same sensation: there could be no discrimination between points touching the two. That there may be discrimination, there must be a distinct connection between each area and the tract of gray matter which receives the impressions. Nay more, there must be, in this central recipient tract, an added number of the separate elements which, by their excitement, yield separate feelings. So that this increased power of tactual discrimination implies a peripheral development, a multiplication of fibers in the trunk-nerve, and a complication of the nerve-center. It can scarcely be doubted that analogous changes occur under analogous conditions throughout all parts of the nervous system—not in its sensory appliances only, but in all its higher co-ordinating appliances up to the highest.
  2. While the proof of this article is in hand, I learn that the Proteus is not quite blind, and that its eyes have a use. It seems that when the underground streams it inhabits are unusually swollen, some individuals of the species are carried out of the caverns into the open (being then sometimes captured). It is also said that the creature shuns the light; this trait being, I presume, observed when it is in captivity. Now obviously, among individuals carried out into the open, those which remain visible are apt to be carried off by enemies; whereas, those which, appreciating the difference between light and darkness, shelter themselves in dark places, survive. Hence the tendency of natural selection is to prevent the decrease of the eyes beyond that point at which they can distinguish between light and darkness. Thus the apparent anomaly is explained.
  3. Essays upon Heredity, p. 87.
  4. I find that the eye of a small smelt (the only appropriate small fish obtainable here, St. Leonards) is about 1180 of its weight; and since in young fish the eyes are disproportionately large, in the full-grown smelt the eye would be probably not more than 1200 of the creature's weight. On turning to highly-finished plates, published by the Bibliographisches Institut of Leipzig, of this perenni-branchiate Proteus, and other amphibians, I find that in the nearest ally there represented, the caducibranchiate axolotl, the diameter of the eye, less than half that of the smelt, bears a much smaller ratio to the length of the body; the proportion in the smelt being 126 of the length, and in the axolotl about 156 (the body being also more bulky than that of the smelt). If, then, we take the linear ratio of the eye to body in this amphibian as one half the ratio which the fish presents, it results that the ratio of the mass of the eye to the mass of the body will be but one eighth. So that the weight of the eye of the amphibian will be but 11600 of that of the body. It is a liberal estimate, therefore, to suppose that its original weight in the Proteus was 11000 of that of the body. I may add that any one who glances at the representation of the axolotl, will see that, were the eye to disappear entirely by a single variation, the economy achieved could not have any appreciable physiological effect on the organism.