[14] Aeussere Einflüsse als Entwickelungsreize, Jena, 1894.
Fig. 105. Ovary of a fertile Queen-Ant and
ovaries of a Worker. Od, oviduct. A, one ovary
of Myrmica lævinodis with many ovarian-tubes,
in each of which there is an almost ripe egg
(Ei) and a younger egg (Ei´). B, the ovaries of a
Worker of Lasius fuliginosus; each ovary has only
one ovarian-tube, and no ripening egg-cells. After
Elizabeth Bickford.
Against this proposition an observation by O. vom Rath has been cited. According to it, three drone larvæ which had been accidentally fed by the workers with royal food exhibited striking retrogressive peculiarities in their sexual organs. The testes contained only immature sperms (just before emergence from the pupa), and the copulatory organ was entirely wanting. That a certain degree of fatty degeneration of the testes should be caused by the 'unusual fattening' is not surprising, but it seems to me very questionable whether the absence of the copulatory organ can be referred to the abnormal diet; it ought to be definitely decided, by the investigation of numerous cases, whether some abnormal peculiarity in the constitution of the germ-plasm in these eggs was not the true cause. Hitherto, unfortunately, I have not been able to procure the fresh material necessary to decide this point[15]. From all this it must be evident that we are not justified in regarding either the absence of wings or the degeneration of the ovaries as a direct result of the inferior nourishment supplied to the workers in the larval state: but should any one still have doubt on this point I may mention that, among our indigenous ants, there are two species in which the workers are just as large as the fertile females, and that in tropical America a species (Myrmecocystus megalocola) occurs in which the workers are larger than the true females; this must mean that they have received more food than the females, though perhaps not the same mixture of food.
[15] Since completing my manuscript I note that the point was settled three years ago, when Koshewnikow had the opportunity of investigating drone-pupæ which were abnormally reared in royal cells, and therefore fed with royal food. He found their sexual organs perfectly normal, and agrees with me that the abnormalities in Vom Rath's case must have been due to some other cause. (See the report by Von Adelung on the Russian paper in Zool. Centralblatt, Sept. 10, 1901.)
From all the facts we have discussed we can confidently conclude that the differences in structure, which distinguish the workers from the true females, do not depend upon the influence, in the individual lifetime, of a poorer diet, but upon variation in the primary constituents of the germ; we must conceive of the germ-plasm of ants as containing, in addition to male and female ids, special ids of workers, in which the determinants of wing and ovary are degenerate in some degree, while the determinants of other parts, such as the brain, are more highly developed. The manner of feeding, however, and perhaps the mingling with the food of a special secretion of the salivary glands, acts as a stimulus which determines whether one kind of id or another is to be liberated, that is, to become active and to enter on the path of development.
A proof of this view is to be found, it seems to me, in the existence of transition forms between workers and true females, which was first brought to general knowledge by Forel. Perhaps it would be better to call these 'mongrel forms' for their various parts do not maintain a medium between the two types, but many parts follow the type of the worker, and others that of the true female. Thus Forel twice found a nest of the red wood-ant which contained a large number of these mixed forms, all of which possessed the small head and large curved thorax of the queen, but otherwise resembled the workers in size and appearance, and also in the degeneration of the ovaries. Many of them were very small, only 5 mm. in length, and had probably received very little food, and, according to the theory of direct influence, these should have been pure workers. That they possessed the head and thorax of a queen is a proof that the characters of both forms of individual were present in the germ-plasm as primary constituents, or indeed entire ids. In normal circumstances only one kind of these ids would have become active, either the worker-id or the queen-id, but in abnormal circumstances they might both be liberated to activity simultaneously, and then they would stamp one part of the body with the character of a queen, another with that of a worker. Forel observed one of these nests in two successive years, and both times found the mixed forms in large numbers[16]. In the second year he found a great number of newly-emerged individuals of this type. I have already inferred from this observation that the mixed forms were probably in both years the offspring of the same mother, and this may well have been the case. My further conclusion, that the mixed forms must be due to some abnormality in the constitution of the germ-plasm of the maternal eggs, no longer appears to me so convincing as it did formerly, because, in the interval, we have learnt, through that indefatigable investigator of ants, Pater Wasmann, that there is another possible explanation of these mixed forms; it, too, is based upon a hypothesis, but it is so interesting that I must briefly outline it to you.
[16] There are different kinds of 'mixed forms' among ants, which may owe their origin to a variety of conditions, as Forel, Wasmann, and Emery have shown in detail.
Like Forel and myself, Pater Wasmann had supposed that the reason of this kind of mixed form (the so-called pseudogynous worker) lay in an abnormality of the constitution of the germ-plasm, but he now regards it as the result of a change in the mode of rearing instituted by the workers with respect to the constitutionally female or queen larvæ, because there was a scarcity of workers. The hypothesis sounds very daring, but it is well founded, at least in so far that there really is a reason why a scarcity of females must occur at certain times in some colonies of ants, and this might certainly determine the workers in charge of the larvæ to feed females with worker food, so as to rear them to render the necessary assistance.
This reason lies in the occasional presence of a parasitic beetle, Lomechusa strumosa, whose larvæ, curiously enough, are cared for and fed by the ants as though they were their own, and in return they eat up the larvæ of the ants, often destroying them in large numbers. Wasmann informs us that the parasitic larvæ grow up just at the time at which the ants are rearing their workers, and it is these, therefore, which fall victims to the Lomechusa-larvæ, and the result is that a scarcity of young workers must soon make itself felt. The workers seek to make this good by rearing as workers all the larvæ previously destined for queens. But this only succeeds partially, because the development towards true females has already begun; thus mixed forms arise.