Essays Upon Heredity and Kindred Biological Problems

I believe that this is at any rate the direction in which we shall have to seek for the first meaning of conjugation and for its phyletic origin. This first result and meaning of conjugation may be provisionally expressed in the following formula:—conjugation originally signified a strengthening of the organism in relation to reproduction, which happened when from some external cause, such as want of oxygen, warmth, or food, the growth of the individual to the extent necessary for reproduction could not take place.

This explanation must not be regarded as equivalent to that afforded by the theory of rejuvenescence; for the latter process is said to be necessary for the continuance of reproduction, and ought therefore to occur periodically quite independently of external circumstances; while according to my theory, conjugation at first only occurred under unfavourable conditions, and assisted the species to overcome such difficulties.

But whatever the original meaning of conjugation may have been, it seems to have become already subordinated in the higher Protozoa, as is indicated by the changes in the course taken by this process. The higher Protozoa when conjugating do not as a rule coalesce completely and permanently190 in the manner followed by the lower Protozoa, and it seems to me possible, or even probable, that in the former the process has already gained the full significance of sexual reproduction, and is to be looked upon as a source of variability.

Whether this be so or not, I believe it is certain that sexual reproduction could not have been entirely abandoned at any period since the time when the Metazoa and Metaphyta first arose; for they derived this form of reproduction from their unicellular ancestors.

We know that organs and characters which have persisted through a long series of generations are transmitted with extreme tenacity, even when they have ceased to be of any direct use to their immediate possessors. The rudimentary organs in various animals, and not least in man, afford very strong proofs of the soundness of this conclusion. Another example has only recently been discovered in the sixth finger, which has been shown to exist in the human embryo191, a part which has only been present in a rudimentary form ever since the origin of the Amphibia192. Superfluous organs become rudimentary very slowly, and enormous periods must elapse before they completely disappear, while the older a character is, the more firmly it becomes rooted in the organism. What I have above called the physical constitution of a species is based upon these facts, and upon them depend the tout ensemble of inherited characters, which are adapted to one another and woven together into a harmonious whole. It is this specific nature of an organism which causes it to respond to external influences in a manner different from that followed by any other organism, which prevents it from changing in any way except along certain definite lines of variation, although these may be very numerous. Furthermore these facts ensure that characters cannot be taken at random from the constitution of a species and others substituted for them. Such a variation as a mammal wanting the firm axis of the backbone is an impossibility, not only because the backbone is necessary as a support to the body, but chiefly because this structure has been inherited from times immemorial, and has become so impressed upon the mammalian organization that any variation so great as to threaten its very existence cannot now take place. The view here set forth of the origin of hereditary variability by amphigonic reproduction, makes it clear that an organism is in a state of continual oscillation only upon the surface, so to speak, while the fundamental parts of its constitution, which have been inherited from extremely remote periods, remain unaffected.

Thus sexual reproduction itself did not cease after it had existed in the form of conjugation through innumerable generations of the vast numbers of species which have been included under the Protozoa; it did not cease even when its original physiological significance had lost its importance, either completely or in part. This process, however, had come to possess a new significance which ensured its continuance, in the enormous advantage conferred on a species by the power of adapting itself to new conditions of life, a power which could only be preserved by means of this method of reproduction. The formation of new species which among the lower Protozoa could be achieved without amphigony, could only be attained by means of this process in the Metazoa and Metaphyta. It was only in this way that hereditary individual differences could arise and persist. It was impossible for amphigony to disappear, for each species in which it was preserved was necessarily superior to those which had lost it, and must have replaced them in the course of time; for the former alone could adapt itself to the ever-changing conditions of life, and the longer sexual reproduction endured, the more firmly was it necessarily impressed upon the constitution of the species, and the more difficult its disappearance became.

Sexual reproduction has nevertheless been lost in some cases, although only at first in certain generations. Thus in the Aphidae and in many lower Crustacea, generations with parthenogenetic reproduction alternate with others which reproduce themselves by the sexual method. But in most cases it is clear that this partial loss of amphigony conferred considerable advantages upon the species by giving increased capabilities for the maintenance of existence. By means of partial parthenogenesis a much more rapid increase in the number of individuals could be attained in a given time, and this fact is of the highest importance for the peculiar circumstances under which these species exist. A species of Crustacean which inhabits rapidly drying pools, and developes from winter-eggs which have remained dried up in the mud, has, as a rule, only a very short time in which to secure the existence of succeeding generations. The few sexual eggs which have escaped the attacks of numerous enemies develope immediately after the first shower of rain; the animals attain their full size in a few days and reproduce themselves as virgin females. Their descendants are propagated in the same manner, and thus in a short time almost incredible numbers of individuals are formed, until finally the sexual eggs are again produced. If now the pool dries up again, the existence of the colony is secured, for the number of animals which produce sexual eggs is very large, and the eggs themselves are of course far more numerous, so that in spite of the destructive agencies to which they are subjected, there will be every chance of the survival of a sufficient number to produce a new generation at a later period. Here, therefore, sexual reproduction has not been abandoned accidentally or from any internal cause, but as an adaptation to certain definite necessities imposed upon the organism by its surroundings.

It is, however, well known that there are certain instances in which sexual reproduction has been altogether lost, and in which parthenogenesis is the only form of propagation. In the animal kingdom, such a condition chiefly occurs in species of which the closely-allied forms exhibit the above-mentioned alternation between parthenogenesis and amphigony, viz. in many Cynipidae and Aphidae, and also in certain freshwater and marine Crustacea. We may imagine that these parthenogenetic species have arisen from forms with alternating methods of reproduction, by the disappearance of the sexual phase.

In any particular case, it may be difficult to point out the motive by which this change has been determined; but it is most probable that the same conditions which originally caused the intercalation of a parthenogenetic stage have been efficient in causing the gradual disappearance of the sexual stage. If a species of Crustacean, with the above-described alternating method of reproduction (heterogeny), were killed off by its enemies on a larger scale than before, it is obvious that the threatened extinction of the species could be checked by the attainment of a correspondingly greater degree of fertility. Such increased fertility might well be produced by pure parthenogenesis (see Appendix V, p. 323), by means of which the number of egg-producing individuals in all the previous sexual generations would be doubled.

In a certain sense, this would be the last and most extreme method by means of which a species might secure continued existence, for it is a method for which it would have to pay very dearly at a later period. If my theory as to the causes of hereditary individual variability be correct, it follows that all species with purely parthenogenetic reproduction are sure to die out; not, indeed, because of any failure in meeting the existing conditions of life, but because they are incapable of transforming themselves into new species, or, in fact, of adapting themselves to any new conditions. Such species can no longer be subject to the process of natural selection, because, with the disappearance of sexual reproduction, they have also lost the power of combining and increasing those hereditary individual characters which they possess.

All the facts with which we are acquainted confirm this conclusion, for whole groups of purely parthenogenetic species or genera are never met with, as would certainly be the case if parthenogenesis had been the only method of reproduction through a successional series of species. We always find it in isolated instances, and under conditions which compel the conclusion that it has become predominant in the species in question, and has not been transmitted from any preceding species.

There still remains a very different class of facts which, so far as we can judge, are in accordance with my theory as to the significance of sexual reproduction, and which may be quoted in its support. I refer to the condition of functionless organs in species with parthenogenetic reproduction.

Under the supposition that acquired characters cannot be transmitted—and this forms the foundation of the views here set forth—organs which are of no further use cannot become rudimentary in the direct and simple manner in which it has been hitherto imagined that degeneration takes place. It is true that an organ which does not perform any function exhibits a marked decrease of strength and perfection in the individual which possesses it, but such acquired degradation is not transmitted to its descendants, and we must therefore look for some other explanation of the firmly established fact that organs do become rudimentary through a series of generations. In seeking this explanation, we shall have to start from the supposition that new forms are not only created by natural selection, but are also preserved by its means. In order that any part of the body of an individual of any species may be kept at the maximum degree of development, it is necessary that all individuals possessing it in a less perfect form must be prevented from propagation—they must succumb in the struggle for existence. I will illustrate this by a special instance. In species which, like the birds of prey193, depend for food upon the acuteness of their vision, all individuals with relatively weak eyesight must be exterminated, because they will fail in the competition for food. Such birds will perish before they have reproduced themselves, and their imperfect vision is not further transmitted. In this way the keen eyesight of birds of prey is kept up to its maximum.

But as soon as an organ becomes useless, the continued selection of individuals in which it is best developed must cease, and a process which I have termed panmixia takes place. When this process is in operation, not only those individuals with the best-developed organs have the chance of reproducing themselves, but also those individuals in which the organs are less well-developed. Hence follows a mixture of all possible degrees of perfection, which must in the course of time result in the deterioration of the average development of the organ. Thus a species which has retired into dark caverns must necessarily come to gradually possess less developed powers of vision; for defects in the structure of the eyes, which occur in consequence of individual variability, are not eliminated by natural selection, but may be transmitted and fixed in the descendants194. This result is all the more likely to happen, inasmuch as other organs which are of importance for the life of the species will gain what the functionless organ loses in size and nutrition. As at each stage of retrogressive transformation individual fluctuations always occur, a continued decline from the original degree of development will inevitably, although very slowly, take place, until the last remnant finally disappears. How inconceivably slowly this process goes on is shown by the numerous cases of rudimentary organs: by the above-mentioned embryonic sixth finger of man, or by the hind limbs of whales buried beneath the surface of the body, or by their embryonic tooth-germs. I believe that the very slowness with which functionless organs gradually disappear, agrees much better with my theory than with the one which has been hitherto held. The result of the disuse of an organ is considerable, even in the course of a single individual life, and if only a small fraction of such a result were transmitted to the descendants, the organ would be necessarily reduced to a minimum, in a hundred or at any rate in a thousand generations. But how many millions of generations may have elapsed since e. g. the teeth of the whalebone whales became useless, and were replaced by whalebone! We do not know the actual number of years, but we know that the whole material of the tertiary rocks has been derived from the older strata, deposited in the sea, elevated, and has been itself largely removed by denudation, since that time.

Now if this theory as to the causes of deterioration in disused organs be correct, it follows that rudimentary organs can only occur in species with sexual reproduction, and that they cannot be formed in species which are exclusively reproduced by the parthenogenetic method: for, according to my theory, variability depends upon sexual reproduction, while the deterioration of an organ when disused, no less than its improvement when in use, depends upon variability. There are therefore two reasons which lead us to expect that organs which are no longer used will remain unreduced in species with asexual reproduction: first, because only a very slight degree of hereditary variability can be present, viz. such a degree as was transmitted from the time when sexual reproduction was first abandoned by the ancestors; and, secondly, because even these slight degrees of variability are not combined, or, in other words, because panmixia cannot occur.

And the facts seem to point in the direction required by the theory, for superfluous organs do not become rudimentary in parthenogenetic species. For example, as far as my experience goes, the receptaculum seminis does not deteriorate, although it is, of course, altogether functionless when parthenogenesis has become established. I do not attach much importance to the fact that the Psychids and Solenobias—(genera of Lepidoptera which Siebold and Leuckart have shown to include species with parthenogenetic reproduction)—still retain the complete female sexual apparatus, because colonies containing males still occasionally occur in these species. Although the majority of colonies are now purely female, the occasional appearance of males points to the fact that the unisexuality of the majority cannot have been of very long duration. The process of transformation of the species from a bisexual into a unisexual form, only composed of females, is obviously incomplete, and is still in process of development. The case is similar with several species of Cynipidae, which reproduce by the parthenogenetic method. In these cases the occurrence of a very small proportion of males is the general rule, and is not confined to single colonies. Thus Adler195 counted 7 males and 664 females in the common Cynips of the rose.

In some Ostracodes, on the other hand, the males appear to be entirely wanting: at least, I have tried in vain for years to discover them in any locality or at any time of the year196.

Cypris vidua and Cypris reptans are such species. Now, although the transformation of these formerly bisexual species into purely unisexual female species appears to be complete197, yet the females still possess a large, pear-shaped receptaculum seminis, with its long spirally twisted duct, which is surrounded by a thick glandular layer. This is the more remarkable as the apparatus is very complicated in the Ostracodes, and retrogressive changes could be therefore easily detected. Furthermore among insects, in the genus Chermes the receptaculum seminis of the females has also remained unreduced, although the males appear to be entirely wanting, or at least have never been found, in spite of the united efforts of several acute observers198. The case is quite different in species which retain both sexual and parthenogenetic reproduction. Thus, the summer females of the Aphidae have lost the receptaculum seminis; and in these insects sexual reproduction has not ceased, but alternates regularly with parthenogenetic reproduction.

Certainly this proof of the truth of my theory as to the significance of sexual reproduction is far from settling the question: it only renders the theory highly probable. At present it is impossible to do more than this, because we do not yet possess a sufficient number of facts, for many of them could not have been sought for until after the theory had been suggested. We are here concerned with complicated phenomena, into which we cannot acquire an immediate insight, but can only attain it gradually.

But, nevertheless, I hope to have shown that the theory of natural selection is by no means incompatible with the theory of ‘the continuity of the germ-plasm;’ and, further, that if we accept this latter theory, sexual reproduction appears in an entirely new light: it has received a meaning, and has to a certain extent become intelligible.

The time in which men believed that science could be advanced by the mere collection of facts has long passed away: we know that it is not necessary to accumulate a vast number of miscellaneous facts, or to make as it were a catalogue of them; but we know that it is necessary to establish facts which, when grouped together in the light of a theory, will enable us to acquire a certain degree of insight into some natural phenomenon. In order to direct our attention to those new facts which are of immediate importance, it is absolutely necessary to seek the aid of some general theory for the arrangement and grouping of those which we already possess. This has been my object in the present paper.

But it may be perhaps objected that these phenomena are far too complicated to be attacked at the present time, and that we ought to wait quietly until the simpler phenomena have been resolved into their components. It may be asked whether the trouble and labour involved in the attempt to solve such questions as heredity or the transformation of species are not likely to be wasted and useless.

It is true that we sometimes meet with such opinions, but I believe that they are based upon a misunderstanding of the method which mankind has always followed in the investigation of nature, and which must therefore be founded upon the necessary relations existing between mankind and nature.

Science has often been compared to an edifice which has been solidly built by laying stone upon stone, until it has gradually risen to greater height and perfection. This comparison holds good up to a certain point, but it leads us to easily overlook the fact that this metaphorical building does not at any point rest upon the ground, and that, at least up to the present time, it has remained floating in the air. Not a single branch of science, not even Physics itself, has commenced building from below; all branches have begun to build at greater or less heights in the air, and have then built downwards: and even Physics has not yet reached the ground, for it is still very uncertain as to the nature of matter and force. In no single group of phenomena can we begin with the investigation of ultimate causes, because at this very point our means of reasoning stop short. We cannot begin with ultimate phenomena and gradually lead up to those which are more complicated: we cannot proceed synthetically and deductively, building up the phenomena from below; but we must as a rule proceed analytically and inductively, proceeding from above downwards.

No one will dispute these statements, but they are often forgotten, as is proved by the above-mentioned objection. If we were only permitted to attack the more complicated phenomena after gaining a complete insight into the simpler ones, then all scientists would be physicists and chemists, and not until Physics and Chemistry were done with should we be permitted to proceed to the investigation of organic nature. Under these circumstances we ought not to possess now any scientific theory of medicine; for the study of pathological physiology could not be commenced until normal physiology was completely known and understood. Yet how great a debt is owing by normal to pathological physiology! This is an example which enforces the conclusion that it is not only permissible, but in the highest degree advantageous, for the different spheres of phenomena to be attacked simultaneously.

Furthermore, if we had been compelled to proceed from the simple to the complex, what would have become of the Theory of Descent, the influence of which has advanced our knowledge of Biology to an altogether immeasurable extent?

But in this often repeated criticism that we are not yet ready to attack such complicated phenomena as heredity, is hidden still another fallacy, for it is implied that facts become less certain in proportion to the complexity of their causes. But is it less certain that the egg of an eagle developes into an eagle, or that the peculiarities of the father and mother are transmitted to the child, than that a stone falls to the ground when its support is taken away? Again, is it not possible to draw a perfectly distinct and certain conclusion as to the relative quantity of the material basis of heredity, present in the germ-cells of either parent, from the fact that the father and mother possess an equal or nearly equal share in heredity? But it is really unnecessary to argue in this way: why should we do more than re-affirm that such a method of procedure in scientific investigation is the only way by which we can gradually penetrate the hidden depths of natural phenomena?

No! Biology is not obliged to wait until Physics and Chemistry are completely finished; nor have we to wait for the investigation of the phenomena of heredity until the physiology of the cell is complete. Instead of comparing the progress of science to a building, I should prefer to compare it to a mining operation, undertaken in order to open up a freely branching lode. Such a lode must not be attacked from one point alone, but from many points simultaneously. From some of these we should quickly reach the deep-seated parts of the lode, from others we should only reach its superficial parts; but from every point some knowledge of the complex tout ensemble of the lode would be gained. And the more numerous the points of attack, the more complete would be the knowledge acquired, for valuable insight will be obtained in every place where the work is carried on with discretion and perseverance.

But discretion is indispensable for a fruitful result; or, leaving our metaphor, facts must be connected together by theories, if science is to advance. Just as theories are valueless without a firm basis of facts, so the mere collection of facts, without relation and without coherence, is utterly valueless. Science is impossible without hypotheses and theories: they are the plummets with which we test the depth of the ocean of unknown phenomena, and thus determine the future course to be pursued on our voyage of discovery. They do not give us absolute knowledge, but they afford us as much insight as it is possible for us to gain at the present time. To go on investigating without the guidance of theories, is like attempting to walk in a thick mist without a track and without a compass. We should get somewhere under these circumstances, but chance alone would determine whether we should reach a stony desert of unintelligible facts or a system of roads leading in some useful direction; and in most cases chance would decide against us.