The Whence and the Whither of Man Part 18
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In Nageli's theory initial tendency is overwhelmingly dominant; in Weismann's, natural selection is almighty.
Weismann's followers have received the name of Neo-Darwinians. The so-called Neo-Lamarckian school believes in the transmissibility of acquired characteristics, and of at least particular effects of use and disuse. The one theory is neither more nor less Darwinian than the other. For while Darwin emphasized natural selection, he accepted to a certain extent the transmission of special effects of use and disuse.
A special theory of heredity, pangenesis, has been accepted by many of the Neo-Lamarckian school. The theory of pangenesis, as propounded by Mr. Darwin, may be very briefly stated as follows: The cells in all parts of the body are continually throwing off germinal particles, or "gemmules." These become scattered through the body, grow, and multiply by division. On account of mutual attraction they unite in the reproductive glands to form eggs or spermatozoa. The germ-cells are thus the bearers of heredity because they contain samples, so to speak, of all the organs of the body.
In heredity, according to Weismann's theory, the egg is the centre of control, the continuous germ-plasm the source of all transmitted changes; according to Darwin's theory, the body is the source, and the egg is derived in great part at least from it. If you put to the two the time-honored question, Which is first, the owl or the egg?
Weismann would announce, with emphasis, The egg; Darwin would say, The owl. One proposition is the converse of the other, and most facts accord almost equally well with both theories.
In any family, devoted for generations to literary or artistic pursuits, the children show, as a rule, an apt.i.tude for such pursuits not manifested by those of other families. According to the Neo-Lamarckian view, this inherited apt.i.tude is to a certain extent the result of the constant exercise of these faculties through a series of generations. The active efforts and voluntary disposition of the parents have given an increased predisposition to the child.
"Quite the reverse," says Weismann, "the increase of an organ in the course of generations does not depend upon the summation of exercise taken during single lives, but upon the summation of more favorable predispositions in the germ." "An organism cannot acquire anything unless it already possesses the predisposition to acquire it."[A]
[Footnote A: Weismann, Essays, pp. 85 and 171.]
We may accept or deny this last statement, but it is evident that facts like these, and indeed the origin of most or all characteristics involving use or disuse, may be explained almost equally well by either theory.
But as far as the transmission of effects of somatic changes is concerned, if protozoa undergo special modifications under the influence of external conditions, will not the germ-cells undergo special modification under the influence of changes in the somatoplasm which forms their immediate environment? We must never forget the close relations.h.i.+p between all the cells of the body, and how slight a change in the body or its surroundings may conduce to sterility or fertility. Such isolation and independence in the body, on the part of the germ-cells, is opposed to all that we know of the organic unity of the body, whose cells have arisen by the differentiation of, and division of labor between, cells primitively alike. The facts of bud-variation, of changes in the parent stock due to grafting, and others, of which Mr. Darwin has given a summary in the eleventh chapter of the first volume of his "Plants and Animals under Domestication," have never been adequately explained by Weismann in accordance with his theory. He has perhaps succeeded in parrying their force by showing that some such explanation is conceivable; they still point strongly against him.
Wilson has good reason for his "steadily growing conviction that the cell is not a self-regulating mechanism in itself, that no cell is isolated, and that Weismann's fundamental proposition is false."
But, granting the force of these criticisms, the question still remains, Is the special effect of use or disuse transmissible? Would the blacksmith's son have a stronger right arm?
1. The isolation and independence of the germ-cells, which Weismann postulates as opposing this, can hardly be as great as he thinks. 2.
It is in his view impossible to conceive how these acquired characteristics can in any way reach and affect the germ-cells in such a manner as to reappear in the next generation. 3. All variations can be explained by his own theory without such transmission. Why then believe that acquired characteristics can in some inconceivable way affect the germ-cells so as to reappear in the next generation, as long as all the facts can be explained in a more simple and easily conceivable manner?
As to his second argument, I would readily acknowledge that it is at present difficult or impossible for me to conceive how any cell can act upon another, except through the nutrient or other fluids which it can produce. But though I cannot conceive how one cell can affect another, I may be compelled to believe that it does so. And this Weismann readily acknowledges.
Driesch changed by pressure the relative position of the cells of a very young embryo, so that those which in a normal embryo would have produced one organ were now compelled, if used at all, to form quite a different one. And yet these displaced cells formed the organ required of cells normally occupying this new position, not the one for which they were normally intended. And the organ which they would have builded in a normal embryo was now formed by other cells transferred to their rightful place.
What made them thus change? Not change of substance or structure, for the slight pressure could hardly have modified this. Not change of nutriment. The only visible or easily conceivable change was in position relative to other cells of the embryo.
Let us in imagination simplify Driesch's experiment, for the sake of gaining a clearer view of its meaning. In a certain embryo at an early stage are certain cells whose descendants should form the lining of the intestine and be used in the adult for digestion. A second set of cells should form muscle endowed mainly with contractility. When these two sets of cells, or some of them, exchange positions in the embryo, they exchange lines of development. The first set now form muscle, the second digestive tissue. The only change has been in their relative positions.
Driesch maintains, therefore, that the goal of development in any embryonic cell is determined not by structure or nutriment but by position. And this would seem to be true of the cells of the earliest embryonic stages.
Certain other experiments point in the same direction. Cut a hydra into equal halves and each half will form a complete animal. The lower half forms a new top, with mouth and tentacles; the upper half, a new base. Cut the other hydra a hair's-breadth farther up.
The same layer of cells which in the first animal formed the lower exposed surface of the upper half now forms the upper exposed surface of the lower half. And with this change of position it has changed its line of development; it will now give rise to a new upper half, not a base as before. The same experiment can be tried on certain worms with similar results, only head and tail differ far more than top and base of hydra. Difference in the position of cells has made vast difference in their line of development. Now in both embryo and adult there must be some directing influence guiding these cells. What is it?
An army is more than a mob of individuals; it is individuals plus organization, discipline, authority. A republic is not square miles of territory and thousands or millions of inhabitants. It is these plus organization, central government. Webster claimed that the central government was, and had to be, before the states. The organism cannot exist without its parts; it has a very real existence in and through them. It can coerce them. The state may be an abstraction, but it is one against which it is usually fatal to rebel, and which can say to a citizen, Go and be hanged, and he straightway mounts the scaffold. Now these are a.n.a.logies and prove nothing. But in so far as they throw light on the essential idea of an organism, they may aid us in gaining a right view of our "cell republic."
Says Whitman in a very interesting article on the "Inadequacy of the Cell-Theory": "That organization precedes cell-formation and regulates it, rather than the reverse, is a conclusion that forces itself upon us from many sides." "The structure which we see in a cell-mosaic is something superadded to organization, not itself the foundation of organization. Comparative embryology reminds us at every turn that the organism dominates cell-formation, using for the same purpose one, several, or many cells, ma.s.sing its material and directing its movements, and shaping its organs as if cells did not exist, or as if they existed only in complete subordination to its will, if I may so speak. The organization of the egg is carried forward to the adult as an unbroken physiological unity, or individuality, through all modifications and transformations." And Wilson, Whitman, Hertwig, and others urge "that the organism as a whole controls the formative processes going on in each part" of the embryo. And many years ago Huxley wrote, "They (the cells) are no more the producers of the vital phenomena than the sh.e.l.ls scattered along the sea-beach are the instruments by which the gravitative force of the moon acts upon the ocean. Like these, the cells mark only where the vital tides have been, and how they have acted."[A]
[Footnote A: See articles by Whitman and Wilson, Journal of Morphology, vol. viii., pp. 649, 607, etc.]
"Interaction of cells" can help us but little. For how can neighboring cells direct others placed in a new position? The expression, if not positively misleading and untrue, is at the best only a restatement of fact. It certainly offers no explanation.
Flood-tide is not due to the interaction of particles of water, though this may influence the form of the waves.
The centre of control is therefore not to be sought in individual cells, whether germ-cells or somatic, but in the organism. And it is the whole organism, one and indivisible, which controls in germ, embryo, and adult, in egg and owl. This individuality, or whatever you will call it, impresses itself upon developing somatic cells, moulding them into appropriate organs, and upon germ-cells in process of formation, moulding them so that they may continue its sway. The muscle, modified by use or disuse, is a better expression of the individuality of its possessor, and the same individuality moulds similarly and simultaneously the germ-cells. Both are different expressions or manifestations of the same individuality.
Only slowly does the individuality mould the muscles and nerves of the adult body to its use. Still more slow may be the moulding of the still more refractory germ-plasm, if such there be. But the moulding process goes on parallel in the two cases.
But Weismann's argument rests not merely upon any difficulty or impossibility of the transmissibility of acquired characteristics.
His argument is rather that all facts can be better explained by his theory without postulating or accepting such transmission, cases of which have never been absolutely proven. But the question is not whether his theory offers a possible explanation of the facts, but whether it is the most probable explanation of all the facts. No one would deny, I think, that the continuity of the germ-plasm offers the best and most natural explanation of heredity; and that variations could be produced by the influence on the germ-plasm of external conditions seems entirely probable.
But when we consider the aggregation of these variations in a process of evolution, his theory seems unsatisfactory. We have already seen that what we commonly call a variation involves not one change, but a series of changes, each term of which is necessary.
Muscle, nerve, and ganglion must all vary simultaneously and correspondingly. Correlation and combination are just as essential as variation. And evolution often demands the disappearance of less fit structures just as much as the advance of the fittest. Says...o...b..rne, "It is misleading to base our theory of evolution and heredity solely upon entire organs; in the hand and foot we have numerous cases of muscles in close contiguity, one steadily developing, the other degenerating." Weismann offers the explanation that "if the average amount of food which an animal can a.s.similate every day remains constant for a considerable time, it follows that a strong influx toward one organ must be accompanied by a drain upon others, and this tendency will increase, from generation to generation, in proportion to the development of the growing organ, which is favored by natural selection in its increased blood-supply, etc.; while the operation of natural selection has also determined the organ which can bear a corresponding loss without detriment to the organism as a whole."[A]
[Footnote A: Weismann, Essays, p. 88.]
Here again natural selection of individuals, not the diminished supply of nutriment, has to determine which of many muscles shall be poorly fed and which favored. But natural selection can favor special organs only indirectly through the individuals which possess such organs. Variation is fortuitous, and there is nothing, except natural selection, to combine or direct them. And, I think, we have already seen that any theory which neglects or excludes such directing and combining agencies must be unsatisfactory and inadequate. Weismann has promised us an explanation of correlation of variation in accordance with his theory; and if such an explanation can be made, it would remove one of the strongest objections. But for the present the objection has very great weight.
Furthermore, as...o...b..rne has insisted, linear variations, or variations proceeding along certain single and well-marked lines, would seem inexplicable by, if not fatal to, Weismann's theory. And yet Osborne, Cope, and others have shown that the teeth of mammals have developed steadily along well-marked lines. They have apparently not resulted at all by selection from a host of fortuitous variations.
Says...o...b..rne in his "Cartwright Lectures"[A]: "It is evident that use and disuse characterize all the centres of evolution; that changes of structure are slowly following on changes of function or habit. In eight independent regions of evolution in the human body there are upward of twenty developing organs, upward of thirty degenerating organs." Now this parallelism, through a long series of generations, between the evolution of organs, their advance or degeneration, and the use or disuse of these same organs, that is, of the habits of the individual, is certainly of great significance.
It must have an explanation; and the most natural one would seem to be the transmission of the effects of use and disuse.
[Footnote A: American Naturalist, vols. xxv. and xxvi.]
On the whole Osborne's verdict would seem just: The Neo-Lamarckian theory fails to explain heredity, Weismann's theory does not explain evolution. But, if the effects of use and disuse are transmitted, correlation of variation is to be expected. Muscle, nerve, and ganglion all vary in correlation because they are all used together and in like degree. Evolution and degeneration of muscles in hand and foot go on side by side, because some are used and some are disused. Centres of use and disuse must be centres of evolution. And there would be as many distinct centres of evolution in different parts of the body as there were centres of use and disuse. And between these centres there might be no correlation except that of use and disuse. Brain, muscles, and jaws would develop simultaneously in the ancestors of insects. And the effects of use and disuse, transmitted through a series of generations, would be c.u.mulative. The species advances rapidly because all its members have in general the same habits; the same parts are advancing or degenerating, although at different rates, in all its individuals.
An animal having an organ highly developed is far less likely to pair with one having a lower development of the same organ. The Neo-Lamarckian theory supplies thus what is lacking in the Neo-Darwinian.
In lower forms, like hydra, of simple structure and comparatively few possibilities of variation, natural selection is dominant. In higher forms, like vertebrates, and especially in man, it is of decidedly subordinate value as a promoter of evolution. For man, as we have seen, is a marvellously complex being. The great difficulty in his case is not so much to quickly gain new and favorable variations as to keep all the organs and powers of the body steadily advancing side by side. Natural selection has in man the important but subordinate position of the judge in a criminal court, to p.r.o.nounce the death verdict on the hopeless and incorrigible.
Both Neo-Darwinians and Neo-Lamarckians have erred in being too exclusively mechanical in their theories. It is the main business of the scientific man to discover and study mechanisms. But he must remember that mechanism does not produce force, it only transmits it. If he maintains that he has nothing to do with anything outside of mechanism, that the invisible and imponderable force lies outside of his domain, he has handed over to metaphysics the fairest and richest portion of his realm. In our fear of being metaphysical we have swung to another extreme, and have lost sight of valuable truth which lay at the bottom of the old vitalistic theories. Cells, tissues, and organs are but channels along which the flood of life-force flows. Boveri has well said, "There is too much intelligence (Verstand) in nature for any purely mechanical theory to be possible."
Each theory contains important truth. Nageli's view of the importance of initial tendencies, inherent in the original living substance, is too often undervalued. My own conviction, at least, is steadily strengthening that, without some such original tendency or aim, evolution would never have reached its present culmination in man. His error lies in emphasizing this factor too exclusively. The fundamental proposition of Weismann's theory, that heredity is due to continuity of germ-plasm, seems to contain important truth. But we need not therefore accept his theory of a germ-plasm so isolated and independent as to be beyond control or influence by the habits of the body. The importance of use and disuse, and the transmissibility of their effects, would seem to supply a factor essential to evolution. Weismann has done good service in emphasizing the stability of the germ-plasm. Evolution is always slow, and, for that very reason, sure.
If these conclusions are correct, they have an important practical bearing. Struggle and effort are essential to progress. Not inborn talent alone, but the use which one makes of it, counts in evolution. The effects of use and disuse are c.u.mulative. The hard-fought battle of past generations becomes an easy victory in the present, just because of the strength acquired and handed down from the past struggle. Persistent variation toward evil is in time weeded out by natural selection. And, while evil remains in the world, we are to lay up stores of strength for ourselves and our descendants by st.u.r.dily fighting it. But the effects of right living through a hundred generations are not overcome by the criminal life of one or two. Evil surroundings weigh more in producing criminals than heredity, and their children are not irreclaimable.
The struggles and victories of each one of us encourage the rest.
There is, to borrow Mr. Huxley's language, not only a survival of the fittest, but a fitting of as many as possible to survive. And in the midst of the hardest struggle there is the peace which comes from the a.s.surance of a glorious triumph.
The Morse Lectures for 1895
THE WHENCE AND WHITHER OF MAN
A BRIEF HISTORY OF MAN'S ORIGIN AND DEVELOPMENT, AND OF THE EVOLUTION OF HIS MORAL AND RELIGIOUS CAPACITIES THROUGH CONFORMITY TO ENVIRONMENT
By JOHN M. TYLER Professor of Biology, Amherst College
CHARLES SCRIBNER'S SONS, PUBLISHERS
This work is a solidification of some new matter with the substance of the ten Morse Lectures delivered at Union Theological Seminary in the spring of 1895. Professor Tyler aims to trace the development of man from the simple living substance to his position at present, paying attention to incidental facts merely as incidental and contributory. He keeps always in view the successive accomplishments of life as they appear in the person of accepted general truth, rather than in the guise of the facts of progress.
The Whence and the Whither of Man Part 18
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