Part 12

According to my view this would be a case of the reversion of the Triton to the immediately anterior phyletic stage, _i.e._ to the perennibranchiate stage, and in the present instance the majority of zoologists who take their stand by the theory of descent, would certainly concur in this view. I should at least consider it to be a useless play upon words did we here speak of larval reproduction, and thereby believe that we had explained something. The animal certainly becomes sexually mature in the same condition as that in which it first appears as a larva, but we first get an insight into the nature of this process by considering that this so-called “sexually mature larva” has the precise structure which must have been possessed by the preceding phyletic stage of the species, and that an individual reversion to the older phyletic stage of the species is consequently before us. I maintain that Duméril is in error in regarding this case of the Triton as parallel with the true larval reproduction of Wagner’s _Cecidomyia_ larva. In this last case it is certainly not reversion to an older phyletic stage that confers the power of reproduction upon the larvæ, since the latter do not represent an older phyletic stage of the species, but must have arisen contemporaneously with this last stage. The enormous structural difference between the larvæ and the imagines is not explained by the latter having arisen from the former supplementarily as a finished production, but by both having been contemporaneously adapted to continually diverging conditions of life.[245] Considered phyletically, these larvæ are by no means necessarily transitional to the origination of the flies. They could have been quite different without the form of the imagines having been thereby modified, since the stages of insect metamorphosis vary independently of each other in accordance with the conditions of life to which they are subjected, and exert scarcely any, or only a very small form-determining influence upon each other, as has been amply proved in the preceding essay. In any case the power of these larvæ (the _Cecidomyiæ_) to propagate themselves asexually was first acquired as a secondary character, as appears from the fact that there exist numerous species of the same genus which do not “nurse.” In the form which they now possess they could never have played the part of the final stage of the ontogeny, nor could they formerly have possessed the power of sexual reproduction.[246] In brief, we are here concerned with true larval reproduction, whilst in Triton we have reversion to an older phyletic stage.[247]

I cannot agree with my friend Professor Haeckel when he occasionally designates the reversion of the Tritons as an “adaptation” to a purely aqueous existence.[248] We could here only speak of “adaptation” if we took the word in a quite different sense to that in which it was first introduced into science by Darwin and Wallace. These naturalists thereby designate a gradual bodily transformation appearing in the course of generations in correspondence with the new requirements of altered conditions of life or, in other words, the action of natural selection, and not the result of a suddenly and direct acting transforming cause exerted but once on a generation.

Just because the word “adaptation” can be used in ordinary language in many senses, it is desirable that it should have only _one_ precise signification, and above all that we should not speak of adaptation where scarcely any morphological change occurs, but only a kind of functional change in the sense used by Dohrn.[249] This is the case for example, when Forel[250] shows that fresh water Pulmonifera, the organization of which is attributed to the direct respiration of air, can nevertheless become settled in the greatest depths of mountain lakes through their lungs being again employed as gills. That not the least change in the lungs hereby takes place is shown by the observations of Von Siebold,[251] who saw the shallow water Pulmonifera using their lungs alternately for direct aërial and aquatic respiration, according to the amount of air contained in the water. If with Von Siebold we merely apply the word “adaptation” to such cases, this expression would lose the special sense which it originally conveyed, and the word would have to be abandoned as a _terminus technicus_; still, such cases may perhaps be spoken of as _physiological_ adaptation.

In any case the reproductive “larvæ” of the Tritons as little present a case of true adaptation as the Axolotl, which occasionally becomes transformed into an Amblystoma. In both cases the transformation referred to is by no means indispensable to the life of the individual. Mature Tritons (devoid of gills) can exist, as I have myself seen, for many months, and probably also for a year in deep water, although adapted for purely pulmonary respiration; whilst Axolotls, as I have already mentioned, can live well for a year in shallow water poor in air. If their gills by this means become shrivelled up or completely disappear, even this is not adaptation in the Darwinian sense, but the effect of directly acting external influences, and chiefly of diminished use.

A case entirely analagous to that of Filippi’s was observed by Jullien in 1869. Four female larvæ of _Lissotriton Punctatus_ (Bell)--(synonymous with _Triton Tæniatus_, Schnd.), taken from a pool, proved to be sexually mature. They contained mature eggs in their ovaria ready for laying, and two of them actually deposited eggs. Four male larvæ found in the same pool, appeared to be equally developed with respect to size, but their testicles contained no free spermatozoa, but only sperm-cells.[252]

I have met with a third case of a similar kind mentioned by Leydig in his memoir, rich in interesting details, “on the tailed Amphibians of the Wurtemburg fauna.”[253] Schreibers, the former director of the Vienna Museum, also found “larvæ” of Tritons with well-developed gills, but of the size of the “adult male individuals,” and, as shown by anatomical investigation, with well “developed sexual organs,” the ovaria especially being distended with eggs.

It is thus established that species which long ago reached the salamander stage in phyletic development, may occasionally degenerate to the perennibranchiate stage. This fact obviously makes my conception of the Axolotl as a reversion form appear much less paradoxical--indeed, the cases of reversion in Triton are precisely analagous to the process which I suppose to have taken place in the Axolotl. We have only to substitute Amblystomas for Tritons, to imagine the pool in which De Filippi found his “sexually mature Triton larvæ” enlarged to the size of the Lake of Mexico, and to conceive the unknown, and perhaps here transitory, causes of the reversion to be permanent, and we have all that is necessary, so far as we at present know, for the restoration of the Axolotl; we obtain a perennibranchiate population of the lake.

It has not yet been determined whether the perennibranchiate form of the Triton actually prevailed permanently in De Filippi’s pool, since, so far as I know, this has not since been examined.

Let us, however, assume for an instant that this is really the case, and that there exists at that spot a colony of sexually reproductive perennibranchiate Tritons: should we wonder if a true Triton occasionally appeared among their progeny, or if we were able to induce the majority of the individuals of this brood to become metamorphosed into Tritons by keeping them in shallow water? According to my view this is precisely the case of the Mexican Axolotl.

I need not, however, restrict myself to this in order to support my hypothesis, but must also directly combat the view hitherto received, since the latter is in contradiction with facts.

Did there really exist in the Axolotl a tendency to sudden phyletic advancement, then one fact would remain quite incomprehensible, viz. the sterility of the Amblystomas.

Out of about thirty Amblystomas obtained by Duméril down to the year 1870, there was not one in a state of sexual maturity; neither copulation nor deposition of eggs took place, and the anatomical investigation of single specimens showed that the eggs were immature, and that the spermatozoa, although present, were without the undulating membrane characteristic of the salamanders, but were not devoid of all power of movement, only, as established by Quatrefages, were “incompletely motile.”[254]

So also the five Amblystomas about which I have been writing, show up to the present time no appearance of reproduction.

The objection raised by Sacc,[255] that the sterility of the Amblystomas bred from Axolotls is attributable to “bad nourishment,” is obviously of but little avail. How is it that the Axolotls, which are fed in a precisely similar manner, propagate so readily? Moreover, I am able to expressly assert that my Amblystomas were very well fed. It is true that they have as yet scarcely reached the age of two years, but the Axolotl propagates freely in the second year, and some of Duméril’s Amblystomas were five years old in 1870.

This fact of the sterility is strongly opposed to the idea that these Amblystomas are the regular precursors of the phyletically advancing genus _Siredon_.[256] I will by no means assert that my theory of reversion actually explains the sterility, but it is at least not directly opposed to it. Mere reversion forms may die off without propagating themselves; but a _new form_ called forth by the action of a phyletic vital force should not be sterile, because this is the precise “aim” which the vital force had in view. The conception of a vital force comprises that of teleology.

The sterility of Amblystoma moreover, although not completely explicable from our standpoint, can be shown to be a phenomenon not entirely isolated. In the above mentioned case of _Lissotriton Punctatus_, the female “larvæ” were certainly sexually mature and laid eggs, but the males of the same period contained in their testicles no fully developed spermatozoa.

Other cases of this kind are unknown to me; at the time when I made the experiments with butterflies already recorded (see the first essay), this point of view was remote, and I therefore neglected to examine the artificially bred reversion forms with respect to their organs of reproduction. But general considerations lead to the supposition that atavistic forms may easily remain sterile.

Darwin[257] finds the proximate causes of sterility in the first place in the action of widely diverging conditions of life, and in the next place in the crossing of individuals widely different in constitution. Now it is certainly deviating conditions of life which lead to the metamorphosis of the Axolotl, and from this point of view it cannot be surprising if we find those individuals sterile which show themselves so especially affected by these changed conditions as to revert to the salamander form.

By this it is not in any way meant to be asserted that reversion is invariably accompanied by sterility, and one cannot raise as an objection to my interpretation of the metamorphosis of the Axolotl, that a reproductive colony of Axolotls could never have arisen by reversion. On the contrary, Jullien’s egg-depositing female Triton larvæ show that also with reversion the power of reproduction may be completely preserved.[258] From the above-mentioned general causes of sterility, it may even be inferred that fertility can be lost in different degrees, and it can be further understood to a certain extent why this fertility is more completely lost by reversion to the Amblystoma, than by the reversion of the Triton to the perennibranchiate form.

If in these cases the reversion is brought about by a change in the conditions of life, we may perhaps suppose that the magnitude of this change would determine the degree of fertility, and the preservation of the reversion form. Still more, however, would the fertility be influenced by the extent of the morphological difference resulting from the reversion. We know that the blending of very different constitutions (_e.g._ the crossing of different species) produces sterility. Something similar results from the sudden reversion to a stage of development widely different in its whole structure. Here also we have in a certain sense the union of two very different constitutions in one individual--a kind of crossing.

From this point of view it can in some measure be comprehended why sterility may be a result of reversion; on the other hand, we thereby obtain no explanation why, with the same amount of morphological difference, in one case complete sterility, and in another relative fertility occurs. The morphological difference between Axolotl and Amblystoma is exactly the same as between Triton and its “sexually mature larva;” the difference between the two cases of reversion depends entirely upon the direction of the leap, that taken in the former case being precisely opposite in direction to that taken in the latter.

Herein might be sought the explanation of the different strength with which the reproductive power is affected; not indeed in the direction of the leap itself, but in the differences in the ontogeny which are determined by the differences in the direction of the leap. The reversion of the Triton to an older phyletic stage coincides with the arrest at a younger ontogenetic stage; or, in other words, the older stage of the phylogeny to which reversion takes place is still entirely comprised in the ontogeny of each individual. Each Triton is perennibranchiate throughout a long period of its life; the reverting individual simply reverts to the older phyletic stage by remaining at the larval stage of its individual development.

But it is quite different with the reversion of the Axolotl to the formerly acquired, but long since abandoned Amblystoma form. This is not retained in the ontogeny of Axolotl, but has been completely lost; for a long series of generations--so must we suppose--the ontogeny has always only attained to the perennibranchiate form. Now if at the present time certain individuals were compelled to revert to the Amblystoma form, certainly no greater leap would have been made from a morphological point of view, than in the reversion of Triton to the perennibranchiate form, but at the same time the leap would be in another direction, viz. over a long series of generations back to a form which the species had not produced for a long period, and which had to a certain extent become foreign to it. We should thus have here also the grafting of a widely different constitution upon that of the Axolotl, or, if one prefers it, the commingling of two widely different constitutions.

Of course I am far from wishing to pretend that this “explanation” is exact; it is nothing more than an attempt to point out the direction in which the causes affecting the reproductive powers in different degrees are to be looked for. A deeper penetration into and special demonstration of the manner in which these causes bring about such results, must be reserved for a future period. For the present it must suffice to have indicated that there is an essential distinction between the two kinds of reversion, and to have made it to some extent comprehensible that this distinction may be the determining impulse with respect to the question of sterility. Perhaps the law here concealed from us may one day be thus formulated:--Atavistic individuals lose the power of reproduction the more completely, the greater the number of generations of their ancestors whose ontogeny no longer comprises the phyletically older stage to which the reversion takes place.

The hypothesis which interprets the transformation of the Axolotl as a case of reversion, thus holds out the possibility of our being able to comprehend the sterility of the Amblystomas arising in this manner, whilst, on the other hand, for the adherents of a phyletic vital force, not only is this observed sterility as Duméril expresses it “un véritable énigme scientifique,” but an absolute paradox. We should expect such a directive and inciting principle to call into existence new forms having vitality and not destined to perish, the more so when it is concerned with a combination of structural characters which, when originating in another manner (viz. from other species of _Siredon_), have long since shown themselves to have vitality and reproductive power. We are indeed acquainted with species of _Amblystoma_ which propagate as such, and each of which arises from an Axolotl-like larva. Thus we cannot regard the sterile Amblystomas produced by the Paris Axolotls as abortive attempts of a vital force--an interpretation which is certainly in itself already sufficiently rash.

Now if it be asked what change in the conditions of life could have led to the reversion in the Lake of Mexico[259] of the Amblystoma to the Siredon form, I must admit that I can only offer a conjectural reply, having but a conditional value so long as it is not supported by a precise knowledge of the conditions there obtaining, and of the habits both of the Axolotl and of the Amblystoma.

It may be supposed generally that reversion is brought about by the same external conditions as those which formerly produced the perennibranchiate stage. This supposition is in the first place supported by the experiments here recorded, since it is evidently the inducement to aërial respiration which causes the young Axolotl to revert to the Amblystoma form, _i.e._ the inciting cause under whose domineering influence the Amblystoma form must have arisen.

Here again the case is quite similar to that of seasonally dimorphic butterflies. Reversion of the summer brood to the winter form is there most easily caused by the action of cold, _i.e._ by the same influence as that under whose sway the winter form was developed.

We know indeed that reversion may also arise by the crossing of races and species, and I have attempted to show that reversion in butterflies may also be brought about by other influences than cold; but still the most probable supposition obviously is, that reversion would be caused by the persistent action of the same influences as those which in a certain sense created the perennibranchiate form. That the latter was produced under the influence of an aquatic life there can be no doubt, and thus, in accordance with my supposition, the hypothetical _Amblystoma Mexicanum_, the supposed ancestral form of the Axolotl of the Mexican Lake, might have been caused to revert to the perennibranchiate form by a reduction in the possibilities of its living upon land, and by its being compelled to frequent the water.

I will not here return to the consideration of every other opinion _ab initio_. It is very advisable to distinguish between the mere impulses which are able to produce sudden reversion, and between actual transforming causes which result directly or indirectly in the remodelling of a species. Thus, it is conceivable _à priori_ that reversion may occur by the action of an inciting cause having nothing to do with the origin of the phyletically older form. Temperature can certainly have played no part, or only a very small part, in the formation of the perennibranchiate form; nevertheless cold may well have been one of the inciting causes which induced the Amblystoma at one time to revert to the _Siredon_ form, and we cannot at present consider De Saussure to be incorrect when he maintains that the low temperature of the Mexican winter might prevent that transformation (of the Axolotl into the Amblystoma) which would occur “in the warm reptile-house” of the Jardin des Plantes. He supports this view by stating that “Tschudi has found the Amblystoma” (of course another species) “in the hottest parts of the United States.” “On the Mexican plateau, however, it snows every winter, and if the lake does not actually freeze, its temperature must fall very considerably in the shallowest parts.”

But although this view is not opposed by any theoretical considerations, I still hold it to be incorrect. I doubt whether it is temperature that has brought about the reverse transformation of the Amblystoma into the Axolotl, or which, according to De Saussure’s conception, at the present time prevents the transformation of the Axolotl in the Lake of Mexico. I doubt this because Amblystomas are now known from all parts of the United States as far north as New York, a proof that a winter cold considerably greater than that of the Mexican plateau is no hindrance to the metamorphosis of the Axolotl, and that the genus does not show itself to be in this respect more sensitive than our native genera of _Salamandridæ_.

The following observations of De Saussure, in which he calls attention to the nature of the Mexican Lake, appear to me to be more worthy of consideration:--“The bottom of this lake is shallow, and one passes imperceptibly from the lake into extensive marshy regions before reaching solid ground; perhaps this circumstance makes the Axolotl incapable of reaching dry land, and prevents the transformation.”

In any case the Lake of Mexico offers very peculiar conditions for Amphibian life. My esteemed friend Dr. v. Frantzius has called my attention to the fact that this lake--as well as many other Mexican lakes--is slightly saline. At the time of the conquest of Mexico by Ferdinand Cortez, this circumstance led to the final surrender of the city, as the Spaniards cut off the supply of water to the besieged, and the water of the lake is undrinkable. The ancient Mexicans had laid down water-conduits from the distant mountains, and the city is still supplied with water brought through conduits.

Now this saltness cannot in itself be the cause of the degeneration to the perennibranchiate form, but it may well be so in combination with other pecularities of the lake. The narrowest part of the lake is the eastern, and it is only in this part that the Axolotl lives. Now in winter, violent easterly gales rush down from the mountains and blow continuously, driving the water before them to such an extent that it becomes heaped up in the western portion of the lake, where it frequently causes floods, whilst 2000 feet of the shallow eastern shore are often laid completely dry.[260]

Now if we consider these two peculiarities, viz. salineness and periodical drying up of a part of the bottom of the lake through continuous gales, we certainly have for the Axolotl, conditions of life which are only to be found in few species. One might certainly attempt to apply these facts in a quite opposite sense, and to regard them as unfavourable to my theory, since the retreat of the water from a great portion of the bottom of the lake would--so one might think--rather facilitate transition to a life upon land, and indeed compel the adoption of such a mode of existence. But we should thus forget that the exposed bottom of the lake is a sterile surface without food or place of concealment, and, above all, without vegetation; and further, that owing to the considerable salineness of the water (specific gravity = 1.0215),[261] the whole of the exposed surface must be incrusted with salt, a circumstance which would render it quite impossible for the creatures to feed upon land. Sodic chloride and carbonate are dissolved in the water in such considerable quantities, that they are regularly deposited upon the shores of the lake as a crust, which is collected during the dry season of the year and sent into the market under the name of “tequisquite” (Mühlenpfordt).[262]