Weismann August
Studies in the Theory of Descent, Volume II
Part III. ON THE FINAL CAUSES OF TRANSFORMATION
III. THE TRANSFORMATION OF THE MEXICAN AXOLOTL INTO AMBLYSTOMA
INTRODUCTION
Since the time when Duméril made known the transformation of a number of Axolotls into the so-called Amblystoma form, this Mexican Amphibian has been bred in many European aquaria, chiefly with the view to establish the conditions under which this transformation occurred, so as to be enabled to draw further conclusions as to the true causes of this exceptional and enigmatical metamorphosis.
Although the Amphibians propagated freely, the cases in which transformation occurred remained extremely rare, and it was not once possible to reply to the main question, viz. whether this metamorphosis was determined by external conditions or by purely internal causes; to say nothing of the possibility of there perhaps being discoverable certain definite external influences by means of which the metamorphosis could have been induced with certainty. But while these points are undecided all attempted theoretical interpretations of the phenomenon must be devoid of a solid basis.
It appeared to me from the first that the history of this transformation of the Axolotl was of special theoretical value; indeed I believed that it might possibly furnish a special case for deciding the truth of those ground-principles, according to which the origin of this species is represented by the two conflicting schools as a case of transformation or as one of heterogenesis. I therefore determined to make some experiments with the Axolotl myself, in the hopes of being fortunate enough to be able to throw some light upon the subject.
In the year 1872 Prof. v. Kölliker was so good as to leave with me five specimens of his Axolotls, bred in Würzburg, and these furnished a numerous progeny in the following year. With these I carried out the idea, the theoretical bearing of which will be shown subsequently, whether it would not be possible to force all the larvæ, or at any rate, the greater majority, to undergo transformation by exposing them to conditions of life which made the use of gills difficult, and that of lungs more easy; in other words, by compelling them to live partly on land at a certain stage of life.
During that year indeed I obtained no results, most of the larvæ perishing before the time for such an experiment had arrived, and the few survivors did not undergo transformation, but lived on to the following spring and then also died one after the other. Through long absence from Freiburg, necessitated by other labours, I had evidently left them without sufficient care and attention. I was thus led to the conviction, which was more fully confirmed subsequently, that no results can be obtained without the greatest care and attention in rearing, towards which single object all one’s interest should be concentrated, and it must not be considered irksome to have to devote daily for many months a large amount of time to this experiment. As it was evident that I could not afford this time without calling in other aid, I hailed with pleasure an opportunity of witnessing the experiment performed by other hands.
A lady living here (Freiburg), Fräulein v. Chauvin, undertook to rear a number of my larvæ of the following year which had just hatched, and in accordance with my idea to make the experiment of forcibly compelling them to adopt the Amblystoma form. How completely this was accomplished will be seen from the following notes by the lady herself, and it will no less appear that these results were only obtained by that care in treatment and delicacy of observation which she devoted to the experiments.
EXPERIMENTS
“I began the experiments on June 12th, 1874, with five larvæ about eight days old, these being the only survivors out of twelve. Owing to the extraordinary delicacy of these creatures, the quality and temperature of the water, and the nature and quantity of their food exerts the greatest influence, especially in early life, and one cannot be too cautious in their treatment.
“The specimens were kept in a glass globe of about thirty centimeters in diameter, the temperature of the water being regulated; as food at first Daphnids, and afterwards larger aquatic animals were introduced in large numbers. By this means all the five larvæ throve excellently. At the end of June the rudiments of the front legs appeared in the most vigorous specimens, and on the 9th of July the hind legs also became visible. At the end of November I noticed that one Axolotl remained constantly at the surface of the water, and this led me to suppose that the right period had now arrived for effecting the transformation into Amblystoma. For brevity I shall designate this as No. I., and the succeeding specimens by corresponding Roman numerals.
“In order to bring about this metamorphosis, on December 1st, 1874, No. I. was placed in a large-sized glass vessel containing earth arranged in such a manner that, when the vessel was filled with water, only one portion of the surface of the earth was entirely covered by the liquid, and the creature in the course of its frequent perigrinations was thus more or less exposed to the air. The water was gradually diminished on the following days, during which period the first changes made their appearance in the Amphibian —the gills commenced to shrivel up, and at the same time the creature showed a tendency to seek the shallowest spots. On December 4th, it took entirely to the land, and concealed itself among some damp moss which I had placed on a heap of sand on the highest portion of the earth in the glass vessel. At this period the first ecdysis occurred. Within the four days from the 1st to the 4th of December, a striking change took place in the external appearance of No. I., the gill-tufts shrivelled up almost entirely, the dorsal crest completely disappeared, and the tail, which had hitherto been broad, became rounded and similarly formed to that of a land salamander. The grey-brown colour of the body changed gradually into a blackish hue; isolated spots, at first of a dull white, made their appearance and these in time increased in intensity.
“When the Axolotl left the water on December 4th the gill-clefts were still open, but these closed gradually, and after about eight days were overgrown with skin and no longer to be seen.
“Of the other larvæ three appeared at the end of November (i. e. at the same time when No. I. came to the surface of the water) to have kept pace in development with No. I., an indication that for these also the right period had arrived for accelerating the developmental processes. They were therefore submitted to the same treatment as No. I. No. II. became transformed at the same time and exactly in the same manner as the latter; its gill-tufts were complete when it was first placed in the shallow water, but after four days these had almost entirely disappeared; in the course of about ten days after it took to the land, the overgrowth of skin on the gill-clefts and the complete assumption of the salamander form occurred. During this last period the creature took food, but only when urged to do so.
“In Nos. III. and IV. the development proceeded more slowly. Neither of these so frequently sought the shallow spots, nor did they as a rule remain so long exposed to the air, so that the greater part of January had expired before they took entirely to the land. Nevertheless the dessication of the gill-tufts did not take a longer time than in Nos. I. and II. as the first ecdysis occurred as soon as they took to the land.
“No. V. showed still more striking deviations in its transformation than Nos. III. and IV., but as this specimen appeared much weaker than the others from the beginning and was retarded in growth to a most notable extent, this is by no means surprising. It took fourteen instead of four days before the transformation had advanced far enough to enable it to leave the water. It was especially interesting to observe the behaviour of this specimen during this period. Its weak and delicate constitution evidently made it much more susceptible to all external influences than the others. If exposed to the air for too long a time it acquired a light colour, and when annoyed or alarmed it emitted a peculiar odour, similar to that of a salamander. As soon as these phenomena were observed it was at once placed in deeper water, into which it immediately plunged and gradually recovered itself, the gills always becoming again expanded. The same experiment was repeated several times and always led to the same result, from which we may venture to conclude that by accelerating the transformation too energetically, the process may come to a standstill, and even by continued compulsion may end in death.
“It yet remains to be mentioned with respect to Axolotl No. V. that this specimen, unlike all the others, did not emerge from the water at the first ecdysis, but at the time of the fourth.
“All the Axolotls are now (July, 1875) living, and are healthy and vigorous, so that with respect to their state of nourishment there is nothing to prevent their propagating. Of the first four the largest is fifteen centim. long; Axolotl No. V. measures twelve centim.
“The preceding statements appear to demonstrate the correctness of the views advanced in the Introduction: – Axolotl larvæ generally but not always complete their metamorphosis if, in the first place, they emerge sound from the egg and are properly fed; and if, in the next place, they are submitted to the necessary treatment for changing aquatic into aërial respiration. It is obvious that this treatment must only be applied very gradually, and in such a manner as not to overtax the vital energy of the Amphibian.”
* * * * *
To the foregoing remarks of Fräulein v. Chauvin I may add that in all five cases the transformation was complete, and not to be confounded with that change which occurs more or less in all Axolotls in the course of time when confined in small glass vessels. In this last case there frequently appear changes in the direction of the Amblystoma form without the latter being actually reached. In the five adult Axolotls which I possessed for a short time, and of which two were at least four years old, the gills were much shrivelled, but the aquatic tail and dorsal crest were unchanged. The crest may, however, also disappear, and the tail become shortened without these changes being due to a transformation into Amblystoma, as will be shown further on.
With respect to the duration of the transformation, this amounted in Axolotls Nos. I. to IV. altogether to twelve or fourteen days. Of these, four days were taken by the first changes which occurred while the creature was still in the water; the remaining time, to the completion of the metamorphosis, was passed on land. Duméril gives the duration of the metamorphosis as sixteen days.
The following results of the experiments just described appear to me to be especially noteworthy: – The five Axolotl larvæ which can alone be taken into consideration, the others having soon perished, all experienced metamorphosis, and without an exception became Amblystomas. Only one of them, No. I., by persistently swimming at the surface, as was observed at the end of six months, showed a decided tendency to undergo metamorphosis and to adopt aërial respiration. With respect to this specimen it may therefore be confidently assumed that it would have taken to the land, and that metamorphosis would have occurred without artificial aid, just as was the case in the thirty specimens which Duméril altogether observed.
Respecting Nos. II., III., and IV., on the other hand, such a supposition is but little probable. These three larvæ endeavoured to keep in deep water and avoided as long as possible the shallow places which would have enforced them to take entirely to lung breathing. Metamorphosis thus occurred more than a month later in these individuals.
Finally, there can scarcely be any doubt that No. V. would not have become transformed without forcible adaptation to an aërial life.
From these results we may venture to conclude that most Axolotl larvæ change into the Amblystoma form when, at the age of six to nine months, they are placed in such shallow water that they are compelled to respire chiefly by their lungs. The experiments before us are certainly at present but very few in number, but such a conclusion cannot be termed premature if we consider that out of several hundred Axolotls (the exact number is not given) Duméril obtained only about thirty Amblystomas, while v. Kölliker bred only one Amblystoma out of a hundred Axolotls.
It now only remains questionable whether each larva could have been forced to undergo metamorphosis, but this could only be decided by new experiments. It was originally my intention to have delayed the publication of the experiments till Fräulein v. Chauvin had repeated them in larger numbers, but as my Axolotls have not bred this year (1875) I must abandon my scheme, and this can be done the more readily because, for the theoretical consideration of the facts, it is immaterial whether all or only nearly all the Axolotls could have been compelled to undergo transformation. I must not, however, omit to mention that Herr Gehrig, the curator of our Zoological Museum, bred a considerable number of larvæ from the same brood as that with which Fräulein v. Chauvin experimented, and that of these larvæ six lived over the winter without undergoing metamorphosis. They were always kept in deep water and thus furnished the converse experiment to those recorded above; they further prove that this whole brood did not have a previous tendency to undergo metamorphosis.
If these new facts are to be made use of to explain the nature of this extraordinary process of transformation in accordance with our present conception, the data already known must in the first place be called to our aid.
It has first to be established that Siredon Mexicanus never, as far as we know, undergoes metamorphosis in its native country. This Amphibian is there only known in the Siredon form, a statement which I have taken from De Saussure,55 who has himself observed the Axolotl in the Mexican lakes. This naturalist never found a single Amblystoma in the neighbourhood of the lakes, “nevertheless the larva (Axolotl) is so common there that it is brought into the market by thousands.” De Saussure believes that in Mexico the Axolotl does not undergo transformation.56 The same statement is distinctly made by Cope,57 whose specimens of Siredon Mexicanus bred in America, even in captivity showed “no tendency to become metamorphosed.” On the other hand Tegetmeier observed58 that one out of five specimens obtained from the Lake of Mexico underwent metamorphosis, and this accordingly establishes the second fact, viz. that the true Axolotl becomes transformed under certain conditions into an Amblystoma when in captivity.
This last remark would be superfluous if, as was for a long time believed, the Paris Axolotls, of which the metamorphosis was first observed and which at the time made such a sensation, were actually Siredon Mexicanus, i. e. the Siredon which alone in its native country bears the name of Axolotl. In his first communication Duméril was himself of this opinion; he then termed the animal “Siredon Mexicanus vel Humboldtii,”59 but subsequently, in his amplified work60 on the transformation of the Axolotl observed in the Jardin des Plantes, he retracted this view, and after a critical comparison of the five described species of Siredon, he came to the conclusion that the species in the possession of the Paris Museum was probably Siredon Lichenoides (Baird). All the transformations of Axolotls observed in Europe must consequently be referred to this species, since they were – at least as far as I know – all derived from the Paris colony. My own experimental specimens were also indirectly descended from these.
Now it must be admitted that this does not coincide with the fact that the Amblystoma form which Duméril first obtained from his Axolotls agreed with Cope’s species, A. Tigrinum, while on the other hand we learn from Marsh61 that Siredon Lichenoides (Baird), when it does undergo metamorphosis, becomes transformed into Amblystoma Mavortium (Baird).
Marsh found Siredon Lichenoides in mountain lakes (7000 feet above the sea) in the southwest of the United States (Wyoming Territory), and obtained from them, by breeding in aquaria, Amblystoma Mavortium (Baird). He considers it indeed doubtful whether the Amphibian undergoes this transformation in its native habitat, although he certainly states this opinion without rigorous proof on purely theoretical considerations, because, according to his view, “the low temperature is there less favourable.”62
If I throw doubt upon this last statement it is simply because Amblystoma Mavortium is found native in many parts of the United States, viz: – in California, New Mexico, Texas, Kansas, Nebraska, and Minnesota. It is indeed by no means inconceivable that in the mountain lakes where Marsh obtained this species, it may behave differently with respect to metamorphosis than in other habitats, and this appears probable from certain observations upon Triton which will be subsequently referred to.
Meanwhile, in the absence of further observations, we must admit that the Paris Axolotls were not Siredon Lichenoides, but some nearly allied and probably new species. But little information is furnished by observing the course of the transformation, although it is at least established that this Axolotl in its native habitat does not undergo metamorphosis or does so as exceptionally as in Europe. Unfortunately in his papers Duméril gives no precise statement respecting the locality of this species imported from “Mexico” – it is probable that he was himself unacquainted with it, so that I can only state on the authority of Cope that Amblystoma has never been brought from south of the provinces of Tamaulipas and Chilhuahua, i. e. south of the Tropic of Cancer.63
This last statement, however, gives no certainty to the matter. Of much more importance is the above-mentioned fact, that the true Axolotl of the lakes about the city of Mexico does not, as a rule, become transformed into an Amblystoma in that locality, although this species in certain cases undergoes metamorphosis when in confinement. From this circumstance and from the fact that the Paris Axolotl likewise experienced but a very small percentage of metamorphosis in captivity, we may venture to conclude that this species also, in its native habitat, either does not become transformed at all or does so only exceptionally.
But there is yet another body of facts which come prominently into view on considering the history of the transformations. I refer to the existence of numerous species of Amblystoma in a natural state. In the “Revision of the genera of Salamandridæ,” published some years ago by Strauch,64 this author, following Cope,65 gives twenty species of Amblystoma as inhabiting North America. Although some of these species are based on single examples, and consequently, as Strauch justly remarks, “may well have to be reduced in the course of time,” there must nevertheless always remain a large number of species which live and propagate as true Amblystomas, and of which the habitat extends from the latitude of New York to that of New Mexico. There are therefore true species of Siredon which regularly assume the Amblystoma form under their natural conditions of life, and which propagate in this form, while, on the other hand, there are at least two species which, under their existing natural conditions of life, always propagate as Siredon. It is but another mode of expression for the same facts if we say that the Mexican Axolotl and the Paris Siredon– whether this is Lichenoides or some other species – stand at a lower grade of phyletic development than those species of Amblystoma which propagate in the salamander form. No one can raise any objection to this statement, while the alternative view maintained by all authors contains a theory either expressed or implied which is, as I believe, incorrect, viz. that the Mexican Axolotl has remained at an inferior stage of phyletic development.
All zoologists66 who have expressed an opinion upon the transformation of the Axolotl, and who are not, like the first observer of this fact, embarrassed by Cuvier’s views as to the immutability of species, regard the phenomenon as though a species, which owing to some special conditions had hitherto remained at a low stage of development, had now through some other influences been compelled to advance to a higher stage.
I believed for a long time that the phenomenon could not otherwise be comprehended, so little was I then in a position to bring all the facts into harmony with this view. Thus in the year 1872 I expressed myself as follows67: – “Why should not a sudden change in all the conditions of life (transference from Mexico to Paris) have a direct action on the organization of the Axolotl, causing it suddenly to reach a higher stage of development, such as many of its allies have already attained, and which obviously lies in the nature of its organization – a stage which it would perhaps itself have reached, although later, in its native country? Or is it inconceivable that the sudden removal from 8000 feet above the sea (Mexican plateau) to the altitude of Paris, may have given the respiratory organs an impetus in the direction of the transformation imminent? In all probability we have here to do with the direct action of changed conditions of life.”
That the substance of this last statement must still hold good is obvious from the experiments previously described, which show that by the application of definite external influences, we have it to a certain extent in our power to produce the transformation. It is precisely in this last point that there lies the new feature furnished by these experiments.
But are we also compelled to interpret the phenomenon in the above manner? i. e. as a sudden advance in the phyletic development of the species occurring, so to speak, at one stroke? I believe not.
What first made this view appear to me erroneous, was the appearance of the living Amblystomas bred from my Axolotl larvæ. These creatures by no means differed from the Axolotls merely in single characters, but they were distinct from the latter in their entire aspect; they differed in some measure in all their parts, in some but slightly and in other parts strongly – in brief, they had become quite different animals. In accordance with this, their mode of life had become completely modified; they no longer lived in the water, but remained concealed by day among the damp moss of the vivarium, coming forth only by night in search of food in dry places.
I had been able to perceive the great difference between the two stages of development from the anatomical data with which I had long been familiar, and which Duméril had made known with respect to the structure of his Amblystomas. But the collecting of numerous details gives no very vivid picture of the metamorphosis; it was the appearance of the living animal that first made me conscious how deep-seated was the transformation which we have here before us, and that this change not merely affected those parts which would be directly influenced by the change in the conditions of life, such as the gills, but that most if not all the parts of the animal underwent a transformation, which could in part be well explained as morphological adaptation to new conditions of life, and partly as a consequence of this adaptation (correlative changes), but could not possibly be regarded as the sudden action of these changed conditions.
Such at least is my view of the case, according to which a per saltum development of the species of such a kind as must here have taken place, is quite inconceivable.
I may venture to assume that most observers of the metamorphosis of Axolotl have, like myself, not been hitherto aware of the extent of the transformation, and it may thus be explained why the theoretical bearings of the case have on all sides been incorrectly conceived. We have here obviously a quite extraordinary case of the first order of importance. I believe that it can easily be shown that the explanation of the history of the metamorphosis of the Paris Axolotl which has hitherto been pretty generally accepted, necessarily comprises a very far-reaching principle. If this interpretation is correct, then in my opinion must also hold good the ideas of those who, like Kölliker, Askenasy, Nägeli, and, among the philosophers, Hartmann and Hübner, would refer the transformation of species in the first instance to a power innate in the organism, to an active, i. e. a self-urging “law of development” – a phyletic vital force.
Thus, if the Axolotls transformed into Amblystomas are regarded as individuals which, impelled by external influences, have anticipated the phyletic development of the others, then this advance can only be ascribed to a phyletic vital force, since the transformation is sudden, and leaves no time for gradual adaptation in the course of generations. The indirect influence of the external conditions of life, i. e. natural selection, is thus excluded from the beginning. But the direct action of the changed conditions of life by no means furnishes an explanation of the complete transformation of the whole structure, such as I have already alluded to, and which I will now enter into more closely.
The differences between the Paris Axolotl and its Amblystoma according to Duméril, Kölliker, and my own observations are as follow: —
1. The gills disappear; the gill-clefts close up, and of the branchial arches only the foremost remain, the posterior ones disappearing. At the same time the os hyoideum becomes changed (Duméril).
2. The dorsal crest completely disappears (Duméril).
3. The aquatic tail becomes transformed into one like that of the salamanders (Duméril), which, however, is not as in the salamander cylindrical, but somewhat compressed laterally (Weismann).
4. The skin becomes yellowish white, irregularly spotted on the sides and back (Duméril), whilst at the same time its former grey-black ground-colour changes into a shining greenish black (Weismann); it loses, moreover, the slimy secretion of the skin, and the cutaneous glands become insignificant (Kölliker).
5. The eyes become prominent and the pupils narrow (Kölliker), and eyelids capable of completely covering the eyes are formed; in Axolotl only a narrow annular fold surrounds the eyes, so that these cannot be closed (Weismann).
6. The toes become narrowed and lose their skin-like appendages (Kölliker), or more precisely, the half webs which connect the proximal extremities of the toes on all the feet (Weismann).
7. The teeth are disposed in this species, as in all other Amblystomæ, in transverse series; whilst in Axolotl, as in Triton larvæ, they are arranged at the sides of the jaw in the form of a curved arch-like band beset with several rows of teeth.68 (Duméril. See his fig., loc. cit. p. 279).
8. In Axolotl the lower jaw, in addition to the teeth on the upper edge of the bone, also bears “de très petites dents disposées sur plusieurs rangs;” these last disappear after metamorphosis (Duméril). I will add that the persistent teeth belong to the os dentale of the lower jaw, and those that are shed to the os operculare.69
9. The surface of the posterior moveable part of the body is slightly concave both before and after transformation; the anterior part is, however, less concave in Amblystoma than in Siredon (Duméril).
I have not yet been able to verify Duméril’s 7th and 9th statements, as I did not want to kill any of my living Amblystomas,70 simply in order to confirm the observations of a naturalist in whom one may certainly place complete confidence. Neither have I as yet observed the transformation of the branchial arches, but all the other statements of Kölliker and Duméril I can entirely corroborate.
The structural differences between Axolotl and Amblystoma are considerably greater and of more importance than those between allied genera, or indeed than between the families of the Urodela. The genus Siredon undoubtedly belongs to a different sub-order to the genus Amblystoma into which it occasionally becomes transformed. Strauch, the most recent systematic worker at this group, distinguishes the sub-order Salamandrida from that of the Ichthyodea by the possession of eyelids, and by the situation of the palatine teeth in single rows on the posterior edge of the palatal bone: in Ichthyodea the eyelids are wanting and the palatine teeth are either “situated on the anterior edge of the palatal bone,” or “cover the whole surface of the palatal plates in brush-like tufts.”
How is it possible to regard such widely divergent anatomical characters as changes suddenly produced by the action (but once exerted) of deviating conditions of life? Hand in hand with the shedding of the old and the appearance of new palatine teeth, there occurs a change in the anatomical structure of the vertebral column, and also – as we may fairly conclude from Kölliker’s correct observation of the cessation of the slimy secretion – in the histological structure of the skin. Who would undertake to explain all these profound modifications as the direct and sudden action of certain external influences? And if any one were inclined to explain such changes as a consequence of the disappearance of the gills, i. e. as correlative changes, what else is such a correlation than the phyletic vital force under another name?
If from one change arising from the direct action of external agencies, the whole body can in two days become transformed in all its parts, in the precise manner which appears best adapted for the new conditions of life under which it is henceforward to exist, then the word “correlation” is only a phrase which explains nothing, but which prevents any attempt at a better explanation, and it would be preferable to profess simply the belief in a phyletic vital force.
