Weismann August
Essays Upon Heredity and Kindred Biological Problems
This example very clearly indicates that the degeneration of an organ does not depend upon its disuse; for although our domestic poultry very rarely make use of their wings, the muscles of flight have not disappeared, and, at any rate in the goose, do not seem to have undergone any marked degeneration.
The numerous and exact observations conducted by Darwin upon the weight and measurement of the bones in domestic fowls, seem to me to possess a significance beyond that which he attributed to them.
If the weight of the wing-bones of the domestic duck bears a smaller proportion to the weight of the leg-bones than in the wild duck, and if, as Darwin rightly assumes, this depends not only upon the diminution of the wings, but also upon the increase of the legs, it by no means follows that this latter increase in organs which are now more frequently used, is dependent upon hereditary influences alone.
It is quite possible that it depends, on the one hand, upon the suspension of natural selection, or panmixia (and these effects would be transmitted), and on the other hand upon the direct influence of increased use during the course of a single life. We do not yet know with any accuracy, the amount of change which may be produced by increased use in the course of a single life. If it is desired to prove that use and disuse produce hereditary effects without the assistance of natural selection, it will be necessary to domesticate wild animals (for example the wild duck) and preserve all their descendants, thus excluding the operation of natural selection. If then all individuals of the second, third, fourth and later generations of these tame ducks possess identical variations, which increase from generation to generation, and if the nature of these changes proves that they must have been due to the effect of use or disuse, then perhaps the transmission of such effects may be admitted; but it must always be remembered that domestication itself influences the organism,—not only directly, but also indirectly, by the increase of variability as a result of the suspension of natural selection. Such experiments have not yet been carried out in sufficient detail49.
It is usually considered that the origin and variation of instincts are also dependent upon the exercise of certain groups of muscles and nerves during a single life-time; and that the gradual improvement which is thus caused by practice, is accumulated by hereditary transmission. I believe that this is an entirely erroneous view, and I hold that all instinct is entirely due to the operation of natural selection, and has its foundation, not upon inherited experiences, but upon the variations of the germ.
Why, for instance, should not the instinct to fly from enemies have arisen by the survival of those individuals which are naturally timid and easily startled, together with the extermination of those which are unwary? It may be urged in opposition to this explanation that the birds of uninhabited islands which are not at first shy of man, acquire in a few generations an instinctive dread of him, an instinct which cannot have arisen in so short a time by means of natural selection. But, in this case are we really dealing with the origin of a new instinct, or only with the addition of one new perception (‘Wahrnehmung,’ Schneider)50, of the same kind as those which incite to the instinct of flight—an instinct which had been previously developed in past ages but had never been called forth by man? Again, has any one ascertained whether the young birds of the second or third generation are frightened by man? May it not be that the experience of a single life-time plays a great part in the origin of the habit? For my part, I am inclined to believe that the habit of flying from man is developed in the first generation which encounters him as a foe51. We see how wary and cautious a flock of birds become as soon as a few shots have been fired at them, and yet shortly before this occurrence they were perhaps playing carelessly close to the sportsmen. Intelligence plays a considerable part in the life of birds, and it by no means follows that the transmission of individual habits explains the above-mentioned phenomena. The long-continued operation of natural selection may very well have been necessary before the perception of man could awake the instinct to flee in young, inexperienced birds. Unfortunately the observations upon these points are far too indefinite to enable us to draw conclusions.
There is again the frequently-quoted instance of the young pointer, ‘which, untrained, and without any example which might have been imitated, pointed at a lizard in a subtropical jungle, just as many of its forefathers had pointed at partridges on the plain of St. Denis,’ and which, without knowing the effect of a shot, sprang forward barking, at the first discharge, to bring in the game. This conduct must not be attributed to the inheritance of any mental picture, such as the effect of a shot, but to the inheritance of a certain reflex mechanism. The young pointer does not spring forward at the shot because he has inherited from his forefathers a certain association of ideas,—shot and game,—but because he has inherited a reflex mechanism, which impels him to start forward on hearing a report. We cannot yet determine without more experiments how such an impulse due to perception (‘Wahrnehmungstrieb,’ Schneider) has arisen; but, in my opinion, it is almost inconceivable that artificial breeding has had nothing to do with it; and that we are here concerned—not with the inheritance of the effects of training—but with some predisposition on the part of the germ, which has been increased by artificial selection.
The necessity for extreme caution in appealing to the supposed hereditary effects of use, is well shown in the case of those numerous instincts, which only come into play once in a lifetime, and which do not therefore admit of improvement by practice. The queen-bee takes her nuptial flight only once, and yet how many and complex are the instincts and the reflex mechanisms which come into play on that occasion. Again, in many insects the deposition of eggs occurs but once in a life-time, and yet such insects always fulfil the necessary conditions with unfailing accuracy, either simply dropping the eggs into water, or carefully fixing them on the surface of the earth beneath some stone, or laying them on a particular part of a certain species of plant; and in all these cases the most complicated actions are performed. It is indeed astonishing to watch one of the Cynipidae (Rhodites rosae) depositing her eggs in the tissue of a young bud. She first carefully examines the bud on all sides, and feels it with her legs and antennae. Then she slowly inserts her long ovipositor between the closely-rolled leaves of the bud, but if it does not reach exactly the right spot, she will withdraw and re-insert it many times, until at length, when the proper place has been found, she will slowly bore deep into the very centre of the bud, so that the egg will reach the exact spot, where the necessary conditions for its development alone exist.
But each Cynips lays eggs many times, and it may be argued that practice may have led to improvement in this case; we cannot however, as a matter of fact, expect much improvement in a process which is repeated, perhaps a dozen times, at short intervals of time, and which is of such an excessively complex nature.
It is the same with the deposition of eggs in most insects. How can practice have had any influence upon the origin of the instinct which leads one of our butterflies—(Vanessa levana)—to lay its green eggs in single file, as columns, which project freely from the stem or leaf, so that protection is gained by their close resemblance to the flower-buds of the stinging-nettle, which forms the food-plant of their caterpillars?
Of course the butterfly is not aware of the advantage which follows from such a proceeding; intelligence has no part in the process. The entire operation depends upon certain inherent anatomical and physiological arrangements:—on the structure of the ovary and oviducts, on the simultaneous ripening of a certain number of eggs, and on certain very complex reflex mechanisms which compel the butterfly to lay its eggs on certain parts of certain plants. Schneider is certainly right when he maintains that this mechanism is released by a sensation, arising from the perception (whether by sight or smell, or both together) of the particular plant or part of the plant upon which the eggs are to be laid52. At any rate, we cannot, in such cases, appeal to the effects of constant use and the transmission of acquired characters, as an explanation; and the origin of the impulse can only be understood as a result of the process of natural selection.
The protective cocoons by which the pupae of many insects are surrounded also belong to the same category, and improvement by practice is entirely out of the question, for they are only constructed once in the course of a life-time. And yet these cocoons are often remarkably complex: think, for instance, of the cocoon spun by the caterpillar of the emperor moth (Saturnia carpini), which is so tough that it can hardly be torn, and which the moth would be unable to leave, if an opening were not provided for the purpose; while, on the other hand, the pupa would not be defended against enemies if the opening were not furnished with a circle of pointed bristles, converging outwards, on the principle of the lobster pot, so that the moth can easily emerge, although no enemy can enter. The impulse which leads to the production of such a structure can only have arisen by the operation of natural selection—not, of course, during the history of a single species, but during the development of numerous, consecutive species—by gradual and unceasing improvements in the initial stages of cocoon-building. A number of species exists at the present day, of which the cocoons can be arranged in a complete series, becoming gradually less and less complex, from that described above, down to a loosely-constructed, spherical case in which the pupa is contained.
The cocoon spun by the larva of Saturnia carpini differs but little in complexity from the web of the spider, and if the former is constructed without assistance from the experience of the single individual—and this must certainly be admitted—it follows that the latter may be also built without the aid of experience, while there is neither reason nor necessity for appealing to the entirely unproved transmission of acquired skill in order to explain this and a thousand other operations.
It may be objected that, in man, in addition to the instincts inherent in every individual, special individual predispositions are also found, of such a nature that it is impossible that they can have arisen by individual variations of the germ. On the other hand, these predispositions—which we call talents—cannot have arisen through natural selection, because life is in no way dependent upon their presence, and there seems to be no way of explaining their origin except by an assumption of the summation of the skill attained by exercise in the course of each single life. In this case, therefore, we seem at first sight to be compelled to accept the transmission of acquired characters.
Now it cannot be denied that all predispositions may be improved by practice during the course of a life-time,—and, in truth, very remarkably improved. If we could explain the existence of great talent, such as, for example, a gift for music, painting, sculpture, or mathematics, as due to the presence or absence of a special organ in the brain, it follows that we could only understand its origin and increase (natural selection being excluded) by accumulation, due to the transmission of the results of practice through a series of generations. But talents are not dependent upon the possession of special organs in the brain. They are not simple mental dispositions, but combinations of many dispositions, and often of a most complex nature: they depend upon a certain degree of irritability, and a power of readily transmitting impulses along the nerve-tracts of the brain, as well as upon the especial development of single parts of the brain. In my opinion, there is absolutely no trustworthy proof that talents have been improved by their exercise through the course of a long series of generations. The Bach family shows that musical talent, and the Bernoulli family that mathematical power, can be transmitted from generation to generation, but this teaches us nothing as to the origin of such talents. In both families the high-water mark of talent lies, not at the end of the series of generations, as it should do if the results of practice are transmitted, but in the middle. Again, talents frequently appear in some single member of a family which has not been previously distinguished.
Gauss was not the son of a mathematician; Handel’s father was a surgeon, of whose musical powers nothing is known; Titian was the son and also the nephew of a lawyer, while he and his brother, Francesco Vecellio, were the first painters in a family which produced a succession of seven other artists with diminishing talents. These facts do not, however, prove that the condition of the nerve-tracts and centres of the brain, which determine the specific talent, appeared for the first time in these men: the appropriate condition surely existed previously in their parents, although it did not achieve expression. They prove, as it seems to me, that a high degree of endowment in a special direction, which we call talent, cannot have arisen from the experience of previous generations, that is, by the exercise of the brain in the same specific direction.
It appears to me that talent consists in a happy combination of exceptionally high gifts, developed in one special direction. At present, it is of course impossible to understand the physiological conditions which render the origin of such combinations possible, but it is very probable that the crossing of the mental dispositions of the parents plays a great part in it. This has been admirably and concisely expressed by Goethe in describing his own characteristics—
Vom Vater hab’ ich die Statur
Des Lebens ernstes Führen,
Vom Mütterchen die Frohnatur
Die Lust zum Fabuliren, etc.
The combination of talents frequently found in one individual, and the appearance of different remarkable talents in the various branches of one and the same family, indicate that talents are only special combinations of certain highly-developed mental dispositions which are found in every brain. Many painters have been admirable musicians, and we very frequently find both these talents developed to a slighter extent in a single individual. In the Feuerbach family we find a distinguished jurist, a remarkable philosopher, and a highly-talented artist; and among the Mendelssohns a philosopher as well as a musician.
The sudden and yet widespread appearance of a particular talent in correspondence with the general intellectual excitement of a certain epoch points in the same direction. How many poets arose in Germany during the period of sentiment which marked the close of the last century, and how completely all poetic gifts seem to have disappeared during the Thirty Years’ War. How numerous were the philosophers that appeared in the epoch which succeeded Kant; while all philosophic talent seemed to have deserted the German nation during the sway of the antagonistic ‘exact science,’ with its contempt for speculation.
Wherever academies are founded, there the Schwanthalers, Defreggers, and Lenbachs emerge from the masses which had shown no sign of artistic endowment through long periods of time53. At the present day there are many men of science who, had they lived at the time of Bürger, Uhland, or Schelling, would probably have been poets or philosophers. And the man of science also cannot dispense with that mental disposition directed in a certain course, which we call talent, although the specific part of it may not be so obvious: we may, indeed, go further, and maintain that the Physicist and the Chemist are characterized by a combination of mental dispositions which differ from those of the Botanist and the Zoologist. Nevertheless, a man is not born a physicist or a botanist, and in most cases chance alone determines whether his endowments are developed in either direction.
Lessing has asked whether Raphael would have been a less distinguished artist had he been born without hands: we might also enquire whether he might not have been as great a musician as he was painter if, instead of living during the historical high-water mark of painting, he had lived, under favourable personal influences, at the time of highly-developed and widespread musical genius. A great artist is always a great man, and if he finds the outlet for his talent closed on one side, he forces his way through on the other.
From all these examples I wish to show that, in my opinion, talents do not appear to depend upon the improvement of any special mental quality by continued practice, but they are the expression, and to a certain extent the bye-product, of the human mind, which is so highly developed in all directions.
But if any one asks whether this high mental development, acquired in the course of innumerable generations of men, is not dependent upon the hereditary effects of use, I would remind him that human intelligence in general is the chief means and the chief weapon which has served and still serves the human species in the struggle for existence54. Even in the present state of civilization—distorted as it is by numerous artificial encroachments and unnatural conditions—the degree of intelligence possessed by the individual chiefly decides between destruction and life; and in a natural state, or still better in a state of low civilization, this result is even more striking.
Here again, therefore, we encounter the effects of natural selection, and to this power we must attribute, at any rate, a great part of the phenomena we have been discussing, and it cannot be shown that—in addition to its operation—the transmission of characters acquired by practice plays any part in nature.
I only know of one class of changes in the organism which is with difficulty explained by the supposition of changes in the germ; these are the modifications which appear as the direct consequence of some alteration in the surroundings. But our knowledge on this subject is still very defective, and we do not know the facts with sufficient precision to enable us to pronounce a final verdict as to the cause of such changes: and for this reason, I do not propose to consider the subject in detail.
These changes—such, for example, as are produced by a strange climate—have been always looked at under the supposition that they are transmitted and intensified from generation to generation, and for this reason the observations are not always sufficiently precise. It is difficult to say whether the changed climate may not have first changed the germ, and if this were the case the accumulation of effects through the action of heredity would present no difficulty. For instance, it is well known that increased nourishment not only causes a plant to grow more luxuriantly, but it alters the plant in some distinct way, and it would be wonderful indeed if the seeds were not also larger and better furnished with nutritive material. If the increased nourishment be repeated in the next generation, a still further increase in the size of the seed, in the luxuriance of the plant, and in all other changes which ensue, is at any rate conceivable if it is not a necessity. But this would not be an instance of the transmission of acquired characters, but only the consequence of a direct influence upon the germ-cells, and of better nourishment during growth.
A similar interpretation explains the converse change. When horses of normal size are introduced into the Falkland Islands, the next generation is smaller in consequence of poor nourishment and the damp climate, and after a few generations they have deteriorated to a marked extent. In such a case we have only to assume that the climate which is unfavourable and the nutriment which is insufficient for horses, affect not only the animal as a whole, but also its germ-cells. This would result in the diminution in size of the germ-cells, the effects upon the offspring being still further intensified by the insufficient nourishment supplied during growth. But such results would not depend upon the transmission by the germ-cells of certain peculiarities due to the unfavourable climate, which only appear in the full-grown horse.
It must be admitted that there are cases, such as the climatic varieties of certain butterflies, which raise some difficulties against this explanation. I myself, some years ago, experimentally investigated one such case55, and even now I cannot explain the facts otherwise than by supposing the passive acquisition of characters produced by the direct influence of climate.
It must be remembered, however, that my experiments, which have been repeated upon several American species by H. W. Edwards, with results confirmatory of my own in all essential respects, were not undertaken with the object of investigating the question from this point of view alone. New experiments, under varying conditions, will be necessary to afford the true explanation of this aspect of the question; and I have already begun to undertake them.
Leaving on one side, for the moment, these doubtful, and insufficiently investigated cases, we may still maintain that the assumption that changes induced by external conditions in the organism as a whole, are communicated to the germ-cells after the manner indicated in Darwin’s hypothesis of pangenesis,—is wholly unnecessary for the explanation of these phenomena. Still we cannot exclude the possibility of such a transmission occasionally occurring, for, even if the greater part of the effects must be attributed to natural selection, there might be a smaller part in certain cases which depends on this exceptional factor.
A complete and satisfactory refutation of such an opinion cannot be brought forward at present: we can only point out that such an assumption introduces new and entirely obscure forces, and that innumerable cases exist in which we can certainly exclude all assistance from the transmission of acquired characters. In most cases of variation in colour we have no explanation but the survival of the fittest56, and the same holds good for all changes of form which cannot be influenced by the will of the animal. Very numerous adaptations, such, for instance, as occur in the eggs of animals,—the markings, and appendages which conceal them from enemies, the complex coverings which prevent them from drying up or protect them from the injurious influence of cold,—must have all arisen entirely independently of any expression of will, or of any conscious or unconscious action on the part of the animals. I will not mention here the case of plants, which as every one knows are unconscious, for they are beyond my province. In this matter, there can be no suggestion of adaptation depending upon a struggle between the various parts of the organism (Roux)57. Natural selection cannot operate upon the different epithelial cells which secrete the egg-shell of Apus, since it is of no consequence to the animal which secretes the egg-shell whether a good or a bad shell is produced. Natural selection first operates among the offspring, and the egg with a shell incapable of resisting cold or drought is destroyed. The different cells of the same individual are not selected, but the different individuals themselves.
In all such cases we have no explanation except the operation of natural selection, and if we cannot accept this, we may as well abandon any attempt at a natural explanation. But, in my opinion, there is no reason why natural selection should be considered inadequate to the task. It is true that the objection has been lately urged, that it is inconceivable that all the wonderful adaptations of the organism to its surroundings can have arisen through the selection of individuals; and that for this purpose an infinite number of individuals and infinite time would be required; and stress is laid upon the fact that the wished-for useful changes can only arise singly and very rarely among a great number of individuals.
This last objection to the modern conception of natural selection has apparently some weight, for, as a matter of fact, useful variations of a conspicuous kind seldom appear, and are often entirely absent for many generations. If we expect to find that qualitative changes take place by sudden leaps, we can never escape this difficulty. But, I think, we must not look for conspicuous variations—such as occur among domesticated animals and plants—in the process of the evolution of species as it goes on in nature. Natural selection does not deal with qualitative but quantitative changes in the individual, and the latter are always present.
A simple example will make this clearer. Let us suppose that it was advantageous to some species—for instance the ancestors of the giraffe—to lengthen some part of the body, such as the neck: this result could be obtained in a relatively short time, for the members of the species already possessed necks of varying length, and the variations which form the material for natural selection were already in existence. Now all the organs of every species vary in size, and any one of them will undergo constant and progressive increase, as soon as it acquires exceptional usefulness. But not only will the organ fluctuate as a whole, but also the parts composing it will become larger or smaller under given conditions, will increase or diminish by the operation of natural selection. I believe that qualitative variations always depend upon differences in the size and number of the component parts of the whole. A skin appears to be naked, when it is really covered with a number of small fine hairs: if these grow larger and increase in number, a thick covering is formed, and we say that the skin is woolly or furry. In the same way the skin of many worms and Crustacea is apparently colourless, but the microscope reveals the presence of a number of beautiful pigment spots; and not until these have increased enormously does the skin appear coloured to the naked eye. The presence or absence of colour and its quality when present are here dependent upon the quantity of the most minute particles, and on the distance at which the object in question is observed. Again, the first appearance of colour, or the change from a green to a yellow or red colour depends upon slight variations in the position or in the number of the oxygen atoms which enter into the chemical combination in question. Fluctuations in the chemical composition of the molecules of a unicellular organism (for example) must continually arise, just as fluctuations are always occurring in the number of pigment granules in a certain cell, or in the number of pigment cells in a certain region of the body, or even in the size of the various parts of the body.
All these quantitative relations are exposed to individual fluctuations in every species; and natural selection can strengthen the fluctuations of any part, and thus cause it to develope further in any given direction.
From this point of view, it becomes less astonishing and less inconceivable that organisms adapt themselves—as we see that they obviously do—in all their parts to any condition of existence, and that they behave like a plastic mass which can be moulded into almost any imaginable form in the course of time.
If we ask in what lies the cause of this variability, the answer must undoubtedly be that it lies in the germ-cells. From the moment when the phenomena which precede segmentation commence in the egg, the exact kind of organism which will be developed is already determined—whether it will be larger or smaller, more like its father or its mother, which of its parts will resemble the one and which the other, even to the minutest detail. In spite of this, there still remains a certain scope for the influence of external conditions upon the organism. But this scope is limited, and forms but a small area round the fixed central point which is determined by heredity. Abundant nourishment can make the body large and strong, but can never make a giant out of the germ-cell destined to become a dwarf. Unhealthy sedentary habits or insufficient nourishment makes the factory-hand pale and stunted; life on board ship, with plenty of exercise and sea air, gives the sailor bodily strength and a tanned skin; but when once the resemblance to father or mother, or to both, is established in the germ-cell it can never be effaced, let the habit of life be what it will.
But if the essential nature of the germ-cell dominates over the organism which will grow from it, so also the quantitative individual differences, to which I referred just now, are, by the same principle, established in the germ, and—whatever be the cause which determines their presence—they must be looked upon as inherent in it. It therefore follows that, although natural selection appears to operate upon the qualities of the developed organism alone, it in truth works upon peculiarities which lie hidden in the germ-cells. Just as the final development of any predisposition in the germ, and just as any character in the mature organism vibrates with a certain amplitude around a fixed central point, so the predisposition of the germ itself fluctuates, and it is on this that the possibility of an increase of the predisposition in question, and its average result, depends.
