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Ants and Some Other Insects: An Inquiry Into the Psychic Powers of These Animals
Even in many insects with movable antennæ and of less ærial habits, e. g., the chafers and bombycid moths, the antennal olfactory sense is evidently much better adapted to function at a distance, i. e., to the perception of odors from distant objects, than to the perception of space and trails. Such insects find their way by means of their eyes, but fly in the direction whence their antennæ perceive an odor that is being sought.
A genuine topochemical antennal sense is, therefore, probably best developed in all arthropods, whose antennæ are not only movable in the atmosphere, but adapted to feeling of objects. In these cases the still imperfect topochemical odor-sense for distances can be momentarily controlled by the contact-odor-sense and definitively fixed topographically, i. e., topochemically, as we see so extensively practised in the ants.
It would be possible to meet this view with the objection that a contact-odor sense could not accomplish much more than the tactile sense. I have made this objection to myself. But in the first place it is necessary to reckon with the facts. Now it is a fact that insects in touching objects with their antennæ mainly perceive and distinguish the chemical constitution of the objects touched and heed these very much more than they do the mechanical impacts also perceived at the same time. Secondly, the tactile sense gives only resistance and through this, form. On the other hand, the multiplicity of odors is enormous, and it is possible to demonstrate, as I have done for the ants, and Von Buttel-Reepen for the bees, that these animals in distinguishing their different nest-mates and their enemies, betray nothing beyond the perception of extremely delicate and numerous gradations in the qualities of odors.
In combination with topochemical space-perception, these numerous odor-qualities must constitute a spatial sense which is vastly superior to the tactile sense. The whole biology of the social Hymenoptera furnishes the objective proof of this assertion.
It would certainly be well worth while to investigate this matter in other groups of arthropods which possess complex instincts.
In conclusion I will cite an example, which I have myself observed, for the purpose of illustrating the capacity of the topochemical olfactory sense.
The American genus Eciton comprises predatory ants that build temporary nests from which they undertake expeditions for the purpose of preying on all kinds of insects. The Ecitons follow one another in files, like geese, and are very quick to detect new hunting grounds. As “ants of visitation,” like the Africo-Indian species of Dorylus, they often take possession of human dwellings, ferret about in all the crevices of the walls and rooms for spiders, roaches, mice, and even rats, attack and tear to pieces all such vermin in the course of a few hours and then carry the booty home. They can convert a mouse into a clean skeleton. They also attack other ants and plunder their nests.
Now all the workers of the African species of Dorylus and of many of the species of Eciton are totally blind, so that they must orient themselves exclusively by means of their antennal sense.
In 1899 at Faisons, North Carolina, I was fortunate enough to find a temporary nest of the totally blind little Eciton carolinense in a rotten log. I placed the ants in a bag and made them the subject of some observations. The Eciton workers carry their elongate larvæ in their jaws and extending back between their legs in such a position that the antennæ have full play in front.
Their ability to follow one another and to find their way about rapidly and unanimously in new territory without a single ant going astray, is incredible. I threw a handful of Ecitons with their young into a strange garden in Washington, i. e., after a long railway journey and far away from their nest. Without losing a moment’s time, the little animals began to form in files which were fully organised in five minutes. Tapping the ground continually with their antennæ, they took up their larvæ and moved away in order, reconnoitering the territory in all directions. Not a pebble, not a crevice, not a plant was left unnoticed or overlooked. The place best suited for concealing their young was very soon found, whereas most of our European ants under such conditions, i. e., in a completely unknown locality, would probably have consumed at least an hour in accomplishing the same result. The order and dispatch with which such a procession is formed in the midst of a totally strange locality is almost fabulous. I repeated the experiment in two localities, both times with the same result. The antennæ of the Ecitons are highly developed, and it is obvious that their brain is instinctively adapted to such rapid orientation in strange places.
In Colombia, to be sure, I had had opportunities of observing, not the temporary nests, but the predatory expeditions of larger Ecitons (E. Burchelli and hamatum) possessing eyes. But these in no respect surpassed the completely blind E. carolinense in their power of orientation and of keeping together in files. As soon as an ant perceives that she is not being followed, she turns back and follows the others. But the marvellous fact is the certainty of this recognition, the quickness and readiness with which the animals recognise their topochemical trail without hesitation. There is none of the groping about and wandering to and fro exhibited by most of our ants. Our species of Tapinoma and Polyergus alone exhibit a similar but less perfect condition. It is especially interesting, however, to watch the perpetuum mobile of the antennæ of the Ecitons, the lively manner in which these are kept titillating the earth, all objects, and their companions.
All this could never be accomplished by a tactile sense alone. Nor could it be brought about by an olfactory sense which furnished no spatial associations. As soon as an Eciton is deprived of its two antennæ it is utterly lost, like any other ant under the same circumstances. It is absolutely unable to orient itself further or to recognise its companions.
In combination with the powerful development of the cerebrum (corpora pedunculata) the topochemical olfactory sense of the antennæ constitutes the key to ant psychology. Feeling obliged to treat of the latter in the preceding lecture, I found it necessary here to discuss in detail this particular matter which is so often misunderstood.
[In his latest Souvenirs entomologiques (Seventh Series) J. H. Fabre has recorded a number of ingenious experiments showing the ability of the males of Saturnia and Bombyx to find their females at great distances and in concealment. He tried in vain (which was to have been foreseen) to conceal the female by odors which are strong even to our olfactories. The males came notwithstanding. He established the following facts: (1) Even an adverse wind does not prevent the males from finding their way; (2) if the box containing the female is loosely closed, the males come nevertheless; (3) if it is hermetically closed (e. g., with wadding or soldered) they no longer come; (4) the female must have settled for some time on a particular spot before the males come; (5) if the female is then suddenly placed under a wire netting or a bell-jar, though still clearly visible, the males nevertheless do not fly to her, but pass on to the spot where she had previously rested and left her odor; (6) the experiment of cutting off the antennæ proves very little. The males without antennæ do not, of course, come again; but even the other males usually come only once: their lives are too short and too soon exhausted.
At first Fabre did not wish to believe in smell, but he was compelled finally, as a result of his own experiments, to eliminate sight and hearing. Now he makes a bold hypothesis: the olfactory sense of insects has two energies, one (ours), which reacts to dissolved chemical particles, and another which receives “physical odor-waves,” similar to the waves of light and sound. He already foresees how science will provide us with a “radiography of odors” (after the pattern of the Roentgen rays). But his own results, enumerated above under (4) and (5) contradict this view. The great distances from which the Bombyx males can discern their females is a proof to him that this cannot be due to dissolved chemical particles. And these same animals smell the female only after a certain time and smell the spot where she had rested, instead of the female when she is taken away! This, however, would be inconceivable on the theory of a physical wave-sense, while it agrees very well with that of an extremely delicate, chemical olfactory sense.
It is a fact that insects very frequently fail to notice odors which we perceive as intense, and even while these are present, detect odors which are imperceptible to our olfactories. We must explain this as due to the fact that the olfactory papillæ of different species of animals are especially adapted to perceiving very different substances. All biological observations favor this view, and our psycho-chemical theories will have to make due allowance for the fact.]
1
If I expressly refrain from accepting the premature and unjustifiable identification of cell-life with a “machine,” I nevertheless do not share the so-called vitalistic views. It is quite possible that science may sometime be able to produce living protoplasm from inorganic matter. The vital forces have undoubtedly originated from physico-chemical forces. But the ultimate nature of the latter and of the assumed material atoms is, of course, metaphysical, i. e., unknowable.
2
“Sensations des Insectes,” Rivista di Biologia Generale. Como, 1900-1901. For the remainder see also A. Forel, Mitth. des Münchener entom. Vereins, 1878, and Recueil. Zool. Suisse, 1886-1887.
3
It is well known that in this matter the movements of the eyes, the movements of the body and of external objects play an essential part, so that without these the eye would fail to give us any knowledge of space. But I need not discuss this further, since the antennæ of ants are at least quite as moveable and their olfactory sense is even more easily educated in unison with the tactile sense.
4
It is not without interest to compare these facts with Condillac’s discussion (Treatise on the Sensations) concerning his hypothetical statue. Condillac shows that our sense of smell is of itself incapable of giving us space knowledge. But it is different in the case of the topochemical sense of smell in combination with the antennary movements. Here Condillac’s conditions of the gustatory sense are fulfilled.