Psychic powers of insects and ants
Psychic powers of insects and ants, when I came across this old document created by Forel, August, 1848-1931, I had to take the time to format (not easy!) and put it up on the site. It is indeed very old, and some paragraphs may be fragmented but I’ve tried to fix such errors as I find them.
I have included a link to the pdf of the original and for ease of reading, the text is included below.
An Inquiry into The Psychic Powers of these Animals With an Appendix on The Peculiarities of Their Olfactory Sense
By Dr. August For el
Late Professor of Psychiatry at the University of Zurich
Translated from the German By Prof. William Morton Wheeler
American Museum of Natural History, New York
Chicago
The Open Court Publishing Company
London
Kegan Paul, Trench, Trubner &* Co. Ltd.
COPYRIGHT, 1904
THE OPEN COURT PUBLISHING CO.
CHICAGO
/Of
ANTS AND SOME OTHER INSECTS.
WHEN discussing the ant-mind, we must consider that these small animals, on the one hand, differ very widely from ourselves in organization, but on the other hand, have come, through so-called convergence, to possess in the form of a social common wealth a peculiar relationship to us. My subject, however, requires the discussion of so many complicated questions that I am compelled to assume acquaintance with the work of others, especially the elements of psychology, and in addition the works of P. Huber, Wasmann, von Buttel-Reepen, Darwin, Romanes, Lubbock, my Fourmis de la Sutsse, and many others. Since the functions ot the sense-organs constitute the basis of comparative psychology, I must also refer to a series of articles entitled “Sensations des Insectes” which I have recently published (1900-1901) in the Rivista de Biologia Generate, edited by Dr. P. Celesia. In these papers I have defined my position with respect to various authors, especially Plateau and Bethe.
Very recently Bethe, Uexkull, and others have denied the existence of psychic powers in invertebrate animals. They explain the latter as reflex-machines, and take their stand on the ground of the so-called psycho-physical parallelism for the purpose of demonstrating our inability to recognize mental qualities in these animals.
They believe, however, that they can prove the mechanical regularity of behavior, but assume unknown forces whenever they are left in the lurch in their explanations. They regard the mind as first making its appearance in the vertebrates, whereas the old Cartesians regarded all animals, in contradistinction to man, as mindless (unconscious) machines.
2 ANTS AND SOME OTHER INSECTS.
The Jesuit father E. Wasmann and von Buttel-Reepen are willing, on the other hand, to accept the inductive inference from analogy as a valid scientific method. Like Lubbock, the lecturer and others, they advocate a comparative psychology of the invertebrates and convincingly demonstrate the existence of psychic faculties in these animals. Wasmann, however, puts a very low estimate on the mental powers of the higher vertebrates and, in my opinion, improperly, denies to them any ability of drawing inferences from experience when in the presence of new conditions (this alone he designates as intelligence); he believes that man alone possesses an immortal soul (independent of natural laws?) in addition to the animal mind.
It is necessary, first of all, to arrive at some common understanding concerning the obscure notion “psychic” in order that we may avoid logomachy, and carrying on theology in the sense of Goethe’s Mephistopheles. Two concepts are confounded in an obscure manner in the word “psychic” : first, the abstract concept of introspection, or subjectivism, i. e. , observation from within, which every person knows only, and can know only, in and by himself. For this let us reserve the term “consciousness.” Second, the “activity” of the mind or that which determines the contents of the field of consciousness. This has been included without further ado with consciousness in the wider sense, and thence has arisen the confusion of regarding consciousness as an attribute of the mind. In another place I have designated the molecular wave of activity of the neural elements as “neurocyme.”
We cannot speak of the consciousness of human beings other than ourselves without drawing an inference from analogy ; quite as little ought we to speak of a consciousness of forgotten things. The field of our consciousness is constantly changing. Things appear in it and disappear from it. Memory, through association, enables us to recall, more or less directly and with more or less difficulty, things which appear to be momentarily absent from consciousness. Moreover, both the experience of self-observation and the phenomena of hypnotism teach us experimentally that many things of which we seem to be unconscious, are nevertheless present in consciousness or have been. Indeed, certain sense-impressions remain, at the moment of their occurrence, unconscious so far as our ordinary consciousness or superconsciousness is concerned, although they can be subsequently recalled into consciousness by suggestion. Whole chains of brain-activities, (dreams, somnambulism, or secondary consciousness) seem ordinarily to be excluded from the superconsciousness, but may subsequently be associated by suggestion with the remembered contents of consciousness. In all these cases, therefore, what seems to be unconscious is after all proved to be onscious.
The above-mentioned phenomena have frequently led to mystical interpretations, but they are explainable on a very simple assumption. Let us assume and this is quite in harmony with observation that the fields of the introspectively conscious brain-activities are limited by so-called association or dissociation processes, i. e., that we are unable actively to bring them all into connection at the same time, and that therefore all that seems to us unconscious has also in reality a consciousness, in other words, a subjective reflex, then the following results : Our ordinary waking consciousness or superconsciousness is merely an inner subjective reflex of those activities of attention which are most intimately connected with one another, i. e., of the more intensively concentrated maxima of our cerebral activities during waking. There exist, however, other consciousnesses, partly forgotten, partly only loosely or indirectly connected with the contents of the superconsciousness, in contradistinction to which these may be designated as subconsciousness. They correspond to other less concentrated or otherwise associated cerebral activities. We are bound to assume the existence of still more remotely interconnected subconsciousnesses for the infra-cortical (lower) brain centers, and so on.
It is easy to establish the fact that the maximum of our psychic activity, namely, attention, passes every moment from one perception or thought to another. These objects of attention, as visual or auditory images, will-impulses, feelings or abstract thoughts, come into play and of this there is no doubt in different brain regions or neuron-complexes. We can therefore compare attention to a functional macula lutea wandering in the brain, or with a wandering maximal intensity of neurocymic activity. But it is quite as satisfactorily established that other psychic phenomena external to attention are likewise present in consciousness, though in a feebler condition. Finally, it is well known that all that has been in consciousness even that which is now more, now less, forgotten is included in the psychic, i. e., in the contents of consciousness. On superficial consideration this appears to satisfy theoretical requirements. But in fact and in truth there are innumerable processes of which we are feebly conscious for only a scarcely appreciable instant and which anon disappear from consciousness. Here and not in the strong and repeated “psychomes” I beg your indulgence for this word, with which I would for the sake of brevity designate each and every psychic unit are we to seek the transition from the conscious to the apparently unconscious. Even in this case, however, the feeble condition of consciousness is only apparent, because the inner reflex of these processes can merely echo faintly in the field of a strongly diverted attention. This, therefore, in no wise proves that such half conscious processes are in and for themselves so feebly represented in consciousness, since a flash of attention is sufficient subsequently to give them definite shape in consciousness. Only in consequence of the diversion of the attention do they lose more and more their connection with the chain of intensity-maxima which, under ordinary circumstances, constitute the remembered contents of our superconsciousness.
The more feebly, however, they are bound to the latter, with the more difficulty are such half-conscious processes later associated anew through memory with the dominant chain. Of such a nature are all dreams, all the subordinate circumstances of our lives, all automated habits, all instincts. But if there exists between the clearly conscious and the unconscious, a half-conscious brain-life, whose consciousness appears to us so feeble merely on account of the deviation of our ordinary train of memories, this is an unequivocal indication that a step further on the remaining connection would be completely severed, so that we should no longer have the right to say that the brain-activities thus fading away nebulously from our superconsciousness do not have consciousness in and for themselves. For the sake of brevity and simplicity we will ascribe subconsciousness to these so-called unconscious brain -processes.
If this assumption is correct and all things point in this direction we are not further concerned with consciousness. It does not at all exist as such, but only through the brain-activity of which it is the inner reflex. With the disappearance of this activity, consciousness disappears. When the one is complicated, the other, too, is complicated. When the one is simple, the other is correspondingly simple. If the brain-activity be dissociated, consciousness also becomes dissociated. Consciousness is only an abstract concept, which loses all its substance with the falling away of “conscious” brain-activity. The brain-activity reflected in the mirror of consciousness appears therein subjectively as a summary synthesis, and the synthetical summation grows with the higher complications and abstractions acquired through habit and practice, so that details previously conscious (e. g., those involved in the act of reading) later become subconscious, and the whole takes on the semblance of a psychical unit.
Psychology, therefore, cannot restrict itself merely to a study of the phenomena of our superconsciousness by means of introspection, for the science would be impossible under such circumstances. Everybody would have only his own subjective psychology, after the manner of the old scholastic spiritualists, and would therefore be compelled to doubt the very existence of the external world and his fellow-men. Inference from analogy, scientific induction, the comparison of the experiences of our five senses, prove to us the existence of the outer world, our fellow-men and the psychology of the latter. They also prove to us that there is such a thing as comparative psychology, a psychology of animals. Finally our own psychology, without reference to our brain–activity, is an incomprehensible patchwork full of contradictions, a patchwork which above all things seems to contradict the law of the conservation of energy.
It follows, furthermore, from these really very simple reflections that a psychology that would ignore brain-activity, is a monstrous impossibility. The contents of our superconsciousness are continually influenced and conditioned by subconscious brain-activities. Without these latter it can never be understood. On the other hand, we understand the full value and the ground of the complex organization of our brain only when we observe it in the inner light of consciousness, and when this observation is supplemented by a comparison of the consciousness of our fellow-men as this is rendered possible for us through spoken and written language by means of very detailed inferences from analogy. The mind must therefore be studied simultaneously from within and from without. Outside ourselves the mind can, to be sure, be studied only through analogy, but we are compelled to make use of this the only method which we possess.
Some one has said that language was given to man not so much for the expression as for the concealment of his thoughts. It is also well known that different men in all honesty attribute very different meanings to the same words. A savant, an artist, a peasant, a woman, a wild Wedda from Ceylon, interpret the same words very differently. Even the same individual interprets them differently according to his moods and their context. Hence it follows that to the psychologist and especially to the psychiatrist and as such I am here speaking the mimetic expression, glances and acts of a man often betray his true inner being better than his spoken language. Hence also the attitudes and behavior of animals have for us the value of a “language,” the psychological importance of which must not be underestimated. Moreover, the anatomy, physiology and pathology of the animal and human brain have yielded irrefutable proof that our mental faculties depend on the quality, quantity, and integrity of the living brain and are one with the same. It is just as impossible that there should exist a human brain without a mind, as a mind without a brain, and to every normal or pathological change in the mental activity, there corresponds a normal or pathological change of the neurocymic activity of the brain, i. e., of its nervous elements. Hence what we perceive introspectively in consciousness is cerebral activity.
As regards the relation of pure psychology (introspection”) to the physiology of the brain (observation of brain-activity from without), we shall take the theory of identity for granted so long as it is in harmony with the facts. The word identity, or monism, implies that every psychic phenomenon is the same real thing as the molecular or neurocymic activity of the brain-cortex coinciding with it, but that this may be viewed from two standpoints. The phenomenon alone is dualistic, the thing itself is monistic. If this were otherwise there would result from the accession of the purely psychical to the physical, or cerebral, an excess of energy which would necessarily contradict the law of the conservation of energy.
Such a contradiction, however, has never been demonstrated and would hold up to derision all scientific experience. In the manifestations of our brain-life, wonderful as they undoubtedly are, there is absolutely nothing which contradicts natural laws and justifies us in postulating the existence of a mythical, supernatural “.psyche.”
On this account I speak of monistic identity and not of psychophysical parallelism. A thing cannot be parallel with itself. Of course, psychologists of the modern school, when they make use of this term, desire merely to designate a supposed parallelism of phenomena without prejudice either to monism or dualism. Since, however, many central nervous processes are accessible neither to physiological nor to psychological observation, the phenomena accessible to us through these two methods of investigation are not in the least parallel, but separated from one another. very unequally by intermediate processes. Moreover, inasmuch as the dualistic hypothesis is scientifically untenable, it is altogether proper to start out from the hypothesis of identity.
It is as clear as day that the same activity in the nervous system of an animal, or even in my own nervous system, observed by myself, first by means of physiological methods from without, and second, as reflecting itself in my consciousness, must appear to me to be totally different, and it would indeed be labor lost to try to convert the physiological into psychological qualities or vice versa. We cannot even convert one psychological quality into another, so far as the reality symbolized by both is concerned ; e. g., the tone, the visual and tactile sensation, which a uniform, low, tuning-fork vibration produces on our three corresponding senses. Nevertheless, we may infer inductively that it is the same reality, the same vibration which is symbolized for us in these three qualitatively and totally different modes; i. e., produces in us these three different psychical impressions which cannot be transformed into one another. These impressions depend on activities in different parts of the brain and are, of course, as such actually different from one another in the brain. We speak of psycho-physiological identity only when we mean, on the one hand, the cortical neurocyme which directly conditions the tonscious phenomena known to us, on the other hand, the corresponding phenomena of consciousness.
And, in fact, a mind conceived as dualistic could only be devoid of energy or energy-containing. If it be conceived as devoid of energy (Wasmann), i. e., independent of the laws of energy, we have arrived at a belief in the miraculous, a belief which countenances the interference with and arbitrary suspension of the laws of nature. If it be conceived as energy-containing, one is merely playing upon words, for a mind which obeys the law of energy is only a portion of the cerebral activities arbitrarily severed from its connections and dubbed “psychic essence,” only that this maybe forthwith discredited. Energy can only be transformed qualitatively, not quantitatively. A mind conceived as dualistic, if sup- posed to obey the law of energy, would have to be transformed completely into some other form of energy. But then it would no longer be dualistic, i. e., no longer essentially different from the brain-activities.
Bethe, Uexkull, and others would require us to hold fast to the physiological method, because it alone is exact and restricts it- self to what can be weighed and measured. This, too, is an error which has been refuted from time immemorial. Only pure mathematics is exact, because in its operations it makes use solely of equations of abstract numbers. The concrete natural sciences can never be exact and are as unable to subsist without the inductive method of inference from analogy as a tree without its roots. Bethe and Uexkull do not seem to know that knowledge is merely relative. They demand absolute exactitude and cannot understand that such a thing is impossible. Besides, physiology has no reason to pride itself upon the peculiar exactitude of its methods and results.
Although we know that our whole psychology appears as the activity of our cerebrum in connection with the activities of more subordinate nerve-centers, the senses and the muscles, nevertheless for didactic purposes it may be divided into the psychology of cognition, of feeling and Volition. Relatively speaking, this subdivision has an anatomico-physiological basis. Cognition depends, in the first instance, on the elaboration of sen^e- impressions by the brain ; the will represents the psychooreerebrofugal resultants of cognition and the feelings together with their final transmission to the musclers. The feelings represent general conditions of excitation of a central nature united with elements of cognition and with cerebrofugal impulses, which are relatively differentiated and re- fined by the former, but have profound hereditary and phylogenetic origins and are relatively independent. There is a continual inter- action of these three groups of brain-activities upon one another. Sense-impressions arouse the attention ; this necessitates movements ; the latter produce new sense-impressions and call for an active selection among themselves. Both occasion feelings of pleas- ure and pain and these again call forth movements of defense, flight, or desire, and bring about fresh sense-impressions, etc. Anatomically, at least, the sensory pathways to the brain and their cortical centers are sharply separated from the centers belonging to the volitional pathways to the muscles. Further on in the cerebrum, however, all three regions merge together in many neurons of the cortex.
Within ourselves, moreover, we are able to observe in the three above-mentioned regions all varieties and degrees of so-called psychic dignity, from the simplest reflex to the highest mental manifestations. The feelings and impulses connected with self- preservation (hunger, thirst, fear) and with reproduction (sexual love and its concomitants) represent within us the region of long- inherited, profoundly phyletic, fixed, instinct-life. These instincts are nevertheless partially modified and partly kept within due bounds through the interference of the higher cerebral activities. The enormous mass of brain-substance, which in man stands in no direct relation to the senses and musculature, admits not only of an enormous storing up of impressions and of an infinite variety of motor innervations, but above all, of prodigious combinations of these energies among themselves through their reciprocal activities and the awakening of old, so-called memory images through the agency of new impressions. In contradistinction to the compulsory, regular activities of the profoundly phyletic automatisms, I have used the term “plastic” to designate those combinations and individual adaptations which depend on actual interaction in the activities of the cerebrum. Its loftiest and finest expression is the plastic imagination, both in the province of cognition and in the province of feeling, or in both combined. In the province of the will the finest plastic adaptability, wedded to perseverance and firmness, and especially when united with the imagination, yields that loftiest mental condition which gradually brings to a conclusion during the course of many years decisions that have been long and carefully planned and deeply contemplated. Hence the plastic gift of combination peculiar to genius ranks much higher than any simpler plastic adaptability.
The distinction between automatism and plasticity in brain- activity is, however, only a relative one and one of degree. In the most different instincts which we are able to influence through our cerebrum, i. e., more or less voluntarily, like deglutition, respiration, eating, drinking, the sexual impulse, maternal affection, jealousy, we observe gradations between compulsory heredity and plastic adaptability, yes, even great individual fluctuations according to the intensity of the corresponding hereditary predispositions.
Now it is indisputable that the individual Pithecanthropus or allied being, whose cerebrum was large enough gradually to construct from onomatopceas, interjections and the like, the elements of articulate speech, must thereby have acquired a potent means of exploiting his brain. Man first fully acquired this power through written language. Both developed the abstract concept symbolized by words, as a higher stage in generalization. All these things give man a colossal advantage, since he is thereby enabled to stand on the shoulders of the written encyclopedia of his predecessors. This is lacking in all animals living at the present time. Hence, if we would compare the human mind with the animal mind, we must turn, not to the poet or the savant, but to the Wedda or at any rate to the illiterate. These people, like children and animals, are very simple and extremely concrete in their thinking. The fact that it is impossible to teach a chimpanzee brain the symbols of language proves only that it is not sufficiently developed for this purpose. But the rudiments are present nevertheless. Of course the “language” of parrots is no language, since it symbolizes nothing. On the other hand, some animals possess phyletic, i. e., hereditarily and instinctively fixed cries and gesture, which are as instinctively understood. Such instinctive animal languages are also very widely distributed and highly developed among insects, and have been fixed by heredity for each species. Finally it is possible to develop by training in higher animals a certain mimetic and acoustic conventional language-symbolism, by utilizing for this purpose the peculiar dispositions of such species. Thus it is possible to teach a dog to react in a particular manner to certain sounds or signs, but it is impossible to teach a fish or an ant these things. The dog comprehends the sign, not, of course, with the reflections of human understanding, but with the capacity of a dog’s brain. And it is, to be sure, even more impossible to teach its young an accomplishment so lofty for its own brain as one which had to be acquired by training, than for the Wedda or even the negro to transmit his acquired culture by his own impulse. Even the impulse to do this is entirely lacking. Nevertheless, every brain that is trained by man is capable of learning and profiting much from the experience of its own individual life. And one discovers on closer examination that even lower animals may become accustomed to some extent to one thing or another, and hence trained, although this does not amount to an understanding of conventional symbols.
In general we may say, therefore, that the central nervous system operates in two ways : automatically and plastically.
12 ANTS AND SOME OTHER INSECTS.
The so-called reflexes and their temporary, purposefully adaptive, but hereditary stereotyped combinations, which respond always more or less in the same manner to the same stimuli, constitute the paradigm of automatic activities. These have the deceptive appearance of a “machine” owing to the regularity of their operations. But a machine which maintains, constructs, and re- produces itself is not a machine. In order to build such a machine we should have to possess the key of life, i. e., the understanding of the supposed, but by no means demonstrated, mechanics of liv- ing protoplasm. Everything points to the conclusion that the instinctive automatisms have been gradually acquired and heredita- rily fixed by natural selection and other factors of inheritance. But there are also secondary automatisms or habits which arise through the frequent repetition of plastic activities and are therefore espe- cially characteristic of man’s enormous brain-development.
In all the psychic provinces of intellect, feeling, and will, habits follow the constant law of perfection through repetition. Through practice every repeated plastic brain-activity gradually becomes automatic, becomes “second nature,” i. e., similar to instinct. Nevertheless instinct is not inherited habit, but phylogenetically inherited intelligence which has gradually become adapted and crystallized by natural selection or by some other means.
Plastic activity manifests itself, in general, in the ability of the nervous system to conform or adapt itself to new and unexpected conditions and also through its faculty of bringing about internally new combinations of neurocyme. Bethe calls this the power of modification. But since, notwithstanding his pretended issue with anthropomorphism, he himself continually proceeds in an anthropomorphic spirit and demands human ratiocination of animals, if they are to be credited with plasticity (power of modification), he naturally overlooks the fact that the beginnings of plasticity are primordial, that they are in fact already present in the Amoeba, which adapts itself to its environment. Nor is this fact to be conjured out of the world by Loeb’s word “tropisms.”
Automatic and plastic activities, whether simple or complex, are merely relative antitheses.
ANTS AND SOME OTHER INSECTS. 13
e. g., in the formation of habits but also in instincts. In their extreme forms they resemble two terminal branches of a tree, but they may lead to similar results through so-called convergence of the conditions of life (slavery and cattle-keeping among ants and men). The automatic may be more easily derived from the plastic activities than vice versa. One thing is established, however: since a tolerably complicated plastic activity admits of many possibilities of adaptation in the individual brain, it requires much more nervous substance, many more neurons, but has more resistances to overcome in order to attain a complicated result. The activities of an Amoeba belong therefore rather to the plasticity of living molecules, but not as yet to that of cooperating nerve-elements; as cell-plasticity it should really be designated as “undifferentiated.” 1 There are formed in certain animals specially complex automatisms, or instincts, which require relatively little plasticity and few neurons. In others, on the contrary, there remains relatively consider- able nerve-substance for individual plasticity, while the instincts are less complicated. Other animals, again, have little besides the lower reflex centers and are extremely poor in both kinds of complex activities. Still others, finally, are rich in both. Strong so- called “hereditary predispositions” or unfinished instincts constitute the phylogenetic transitions between both kinds of activity and are of extraordinarily high development in man.
Spoken and especially written language, moreover, enable man to exploit his brain to a wonderful extent. This leads us to under- estimate animals. Both in animals and man the true value of the brain is falsified by training, i. e., artificially heightened. We overestimate the powers of the educated negro and the trained dog and underestimate the powers of the illiterate individual and the wild animal.
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.
14 ANTS AND SOME OTHER INSECTS.
You could oject, but it seemed necessary that we should come to some under- standing concerning the validity of comparative psychology. My further task now consists in demonstrating to you what manner of psychical faculties may be detected in insects. Of course, I shall select in the first place the ants as the insects with which I am most familiar. Let us first examine the brain of these animals.
In order to determine the psychical value of a central nervous system it is necessary, first, to eliminate all the nerve-centers which subserve the lower functions, above the immediate innervation of the muscles and sense-organs as first centers. The volume of such neuron-complexes does not depend on the intricacy of mental work but on the number of muscle-fibers concerned in it, the sensory surfaces, and the reflex apparatus, hence above all things on the size of the animals. Complex instincts already require the intervention of much more plastic work and for this purpose such nerve- centers alone would be inadequate.
A beautiful example of the fact that complex mental combinations require a large nerve-center dominating the sensory and muscular centers is furnished by the brain of the ant. The ant-colony commonly consists of three kinds of individuals : the queen, or female (largest), the workers which are smaller, and the males which are usually larger than the workers. The workers excel in complex instincts and in clearly demonstrable mental powers (memory, plasticity, etc.). These are much less developed in the queens. The males are incredibly stupid, unable to distinguish friends from enemies and incapable of finding their way back to their nest. Nevertheless the latter have very highly developed eyes and antennae, i. e., the two sense-organs which alone are connected with the brain, or supraoesophageal ganglion and enable them to possess themselves of the females during the nuptial flight. No muscles are innervated by the supraoesophageal ganglion. These conditions greatly facilitate the comparison of the perceptive organs, i. e., of the brain (corpora pedunculate?) in the three sexes. This is very large in the worker, much smaller in the fe- male, and almost vestigial in the male, whereas the optic and olfactory lobes are very large in the latter. The cortical portion of the large worker brain is, moreover, extremely rich in cellular elements. In this connection I would request you to glance at the figures and their explanation.
Very recently, to be sure, it has come to be the fashion to underestimate the importance of brain-morphology in psychology and even in nerve-physiology. But fashions, especially such absurd ones as this, should have no influence on true investigation. Of course, we should not expect anatomy to say what it was never in- tended to say.
In ants, injury to the cerebrum leads to the same results as in- jury to the brain of the pigeon.
In this place I would refer you for a fuller account of the details of sensation and the psychic peculiarities of insects to my more extended work above mentioned : Sensations des Insectes.
It can be demonstrated that insects possess the senses of sight, smell, taste, and touch. The auditory sense is doubtful. Perhaps a sense of touch modified for the perception of delicate vibrations may bear a deceptive resemblance to hearing. A sixth sense has nowhere been shown to occur. A photodermatic sense, modified for light-sensation, must be regarded as a form of the tactile sense. It occurs in many insects. This sense is in no respect of an optic nature. In aquatic insects the olfactory and gustatory senses perhaps grade over into each other somewhat (Nagel), since both perceive chemical substances dissolved in the water.
The visual sense of the faceted eyes is especially adapted for seeing movements, i. e. , for perceiving relative changes of position in the retinal image. In flight it is able to localize large spatial areas admirably, but must show less definite contours of the objects than our eyes. The compound eye yields only a single up- right image (Exner), the clearness of which increases with the number of facets and the convexity of the eye. Exner succeeded in photographing this image in the fire-fly (Lampyris). As the eyes are immovable the sight of resting objects soon disappears so far as the resting insect is concerned. For this reason resting in- sects are easily captured when very slowly approached. In flight insects orient themselves in space by means of their compound
EXPLANATION OF THE FIGURES.
Brain (supra-oesophageal ganglion) of an ant (Lasius fuliginosus), magnified 60 diameters, seen from above.
Fig. W. Brain of the Worker.
Fig. F. Brain of the Queen (Female)
Fig. M. Brain of the Male.
St. = Brain trunk. L. op. =Lobus opticus (optic lobe). L. olf. = Lobus olfac- torius si ve antennalis (olfactory lobe). N. = Facetted eye. N. olf. =Nervus olfac- torius sive antennalis (olfactory nerve). O.= Ocelli, or simple eyes with their nerves (present only in the male and queen). H. = Cellular brain cortex (developed only in the worker and queen). C. p. = Corpora pedunculata, or fungiform bodies (developed only in the worker and queen). R. =Rudimental cortex of male.
The length of the whole ant is :
in the worker 4.5 mm ;
in the queen 6.0 mm ;
in the male 4.5 mm.
N. B. The striation of the corpora pedunculata and their stems is represented diagrammatically, for the purpose of indicating rather coarsely their extremely delicate fibrillar structure.
l6 ANTS AND SOME OTHER INSECTS.
Odor, when perceived, merely draws these animals in a particular direction. When the compound eyes are covered, all powers of orientation in the air are lost. Many insects can adapt their eyes for the day or night by a shifting of the pigment. Ants see the ultra-violet with their eyes. Honey-bees and humble-bees can distinguish colors, but obviously in other tones than we do, since they cannot be deceived by artificial flowers of the most skillful workmanship. This may be due to admixtures of the ultra-violet rays which are invisible to our eyes.
The ocelli (simple eyes) play a subordinate role, and probably serve as organs of sight for objects situated in the immediate vicinity and in dark cavities.
The olfactory sense has its seat in the antennae, usually in the club-shaped flagellum, or rather in the pore-plates and olfactory rods of these portions of the antennas. On account of its external and movable position at the tip of the antenna, the olfactory organ possesses two properties which are lacking in the vertebrates, and particularly in man. These are :
1. The power of perceiving the chemical nature of a body by direct contact (contact-odor);
2. The power of space-perception and of perceiving the form of objects and that of the animal’s own trail by means of odor, and the additional property of leaving associated memories.
The olfactory sense of insects, therefore, gives these animals definite and clear-cut perceptions of space-relations, and enables the animal while moving on the surface of the ground to orient it- self with facility. I have designated this sense, which is thus qualitatively, i. e., in its specific energy, very different from our olfactory sense, as the topochemical (olfactory) sense. Probably the pore-plates are used for perceiving odor at a distance and the olfactory rods for contact-odor, but this is pure conjecture. Extirpation of the antennae destroys the power of distinguishing friends from enemies and deprives the ant of the faculty of orienting itself on the ground and of finding its way, whereas it is possible to cut off three legs and an antenna without seriously impairing these powers. The topochemical sense always permits the ant to distinguish between the directions of its trail, a faculty which Bethe attributes to a mysterious polarization. The ability to sense different odors varies enormously in different insects. An object possessing odor for one species is often odorless for other species (and for ourselves) and vice versa.
The gustatory organs are situated on the mouth-parts. Among insects the reactions of this sense are very similar to our own. Will accustomed some wasps to look in a particular place for honey, which he afterwards mixed with quinine. The wasps detected the substance at once, made gestures of disgust, and never returned to the honey. Mixing the honey with alum had the same result. At first they returned, but after the disagreeable gustatory experience they failed to reappear. Incidentally this is also a proof of their gustatory memory and of their powers of association.
Several organs have been found and described as auditory. But after their removal the supposed reaction to sounds persists. This would seem to indicate that a deceptive resemblance to hearing may be produced by the perception of delicate vibrations through the tactile sense (Duges).
The tactile sense is everywhere represented by tactile hairs and papillae. It reacts more especially to delicate tremors of the atmosphere or soil. Certain arthropods, especially the spiders, orient themselves mainly by means of this sense.
It may be demonstrated that insects, according to the species and conditions of life, use their different senses in combination for purposes of orienting themselves and for perceiving the external world. Many species lack eyes and hence also the sense of sight. In others, again, the olfactory sense is obtuse ; certain other forms lack the contact-odor sense (e. g., most Diptera).
It has been shown that the superb powers of orientation exhibited by certain aerial animals, like birds (carrier-pigeons), bees, etc., depend on vision and its memories. Movement in the air gives this sense enormous and manifold values. The semi-circular canals of the auditory organ are an apparatus of equilibrium in vertebrates and mediate sensations of acceleration and rotation (Mach-Breuer), but do not give external orientation. For the demonstration of these matters I must refer you to my work above- cited. A specific, magnetic, or other mode of orientation, independent of the known senses, does not exist.
The facts above presented constitute the basis of insect psycholog)’. The social insects are especially favorable objects for study on account of their manifold reciprocal relationships. If in speaking of their behavior I use terms borrowed from human life, I request you, once for all, to bear in mind that these are not to be interpreted in an anthropomorphic but in an analogous sense.
THE PROVINCE OF COGNITION.
Many insects (perhaps all, in a more rudimental condition) possess memory, i. e., they are able to store up sense-impressions in their brains for subsequent use. Insects are not merely attracted directly by sensory stimuli, as Bethe imagines. Huber, myself, Fabre, Lubbock, Wasmann, Von Buttel-Reepen, have demonstrated this fact experimentally. That bees, wasps, etc., can find their way in flight through the air, notwithstanding wind and rain (and hence under circumstances precluding the existence of any possible odoriferous trail), and even after the antennae have been cut off, to a concealed place where they have found what they desired, though this place may be quite invisible from their nest, and this even after the expiration of days and weeks, is a fact of special importance as proof of the above assertion. It can be shown that these insects recognize objects by means of their colors, their forms, and especially by their position in space. Position they perceive through the mutual relations and succession of the large objects in space, as these are revealed to them in their rapid change of place during flight in their compound eyes (shifting of retinal images). Especially the experiments performed by Von Buttel-Reepen and myself leave no doubt concerning this fact. Additional proof of a different nature is furnished by Von Buttel, who found that ether or chloroform narcosis deprives bees of all memory. By this means enemies can be converted into friends. Under these circumstances, too, all memory of locality is lost and must be reacquired by means of a new flight of orientation. An animal, however, certainly can- not forget without having remembered.
The topochemical antennal sense also furnishes splendid proofs of memory in ants, bees, etc. An ant may perform an arduous journey of thirty meters from her ruined nest, there find a place suitable for building another nest, return, orienting herself by means of her antennae, seize a companion who forthwith rolls her- self about her abductrix, and is carried to the newly selected spot. The latter then also finds her way to the original nest, and both each carry back another companion, etc. The memory of the suit- able nature of the locality for establishing a new nest must exist in the brain of the first ant or she would not return, laden with a companion, to this very spot. The slave-making ants {Polyergus} un- dertake predatory expeditions, led by a few workers, who for days and weeks previously have been searching the neighborhood for nests of Formica fusca. The ants often lose their way, remain standing and hunt about for a long time till one or the other finds the topochemical trail and indicates to the others the direction to be followed by rapidly pushing ahead. Then the pupae of the For- mica fusca nest, which they have found, are brought up from the depths of the galleries, appropriated and dragged home, often a distance of forty meters or more. If the plundered nest still con- tains pupae, the robbers return on the same or following days and carry off the remainder, but if there are no pupae left they do not return. How do the Polyergus know whether there are pupae re- maining? It can be demonstrated that smell could not attract them from such a distance, and this is even less possible for sight or any other sense. Memory alone, i. e., the recollection that many pupae still remain behind in the plundered nest can induce them to re- turn. I have carefully followed a great number of these predatory expeditions.
While Formica species follow their topochemical trail with great difficulty over new roads, they nevertheless know the immediate surroundings of their nest so well that even shoveling away the earth can scarcely disconcert them, and they find their way at once, as Wasmann emphatically states and as I myself have often observed. That this cannot be due to smelling at long range can be demonstrated in another manner, for the olfactory powers of the genus Formica, like those of honey-bees, are not sufficiently acute for this purpose, as has been shown in innumerable experiments by all connoisseurs of these animals. Certain ants can recognize friends even after the expiration of months. In ants and bees there are very complex combinations and mixtures of odors, which Von But- tel has very aptly distinguished as nest-odor, colony- (family-) odor, and individual odor. In ants we have in addition a species-odor, while the queen-odor does not play the same role as among bees.
It follows from these and many other considerations that the social Hymenoptera can store up in their brains visual images and topochemical odor-images and combine these to form perceptions or something of a similar nature, and that they can associate such perceptions, even those of different senses, especially sight, odor, and taste, with one another and thereby acquire spatial images.
Huber as well as Von Buttel, Wasmann, and myself have always found that these animals, through frequent repetition of an activity, journey, etc., gain in the certainty and rapidity of the ex- ecution of their instincts. Hence they form, very rapidly to be sure, habits. Von Buttel gives splendid examples of these in the robber-bees, i. e., in some of the common honey-bees that have acquired the habit of stealing the honey from the hives of strangers. At first the robbers display some hesitation, though later they become more and more impudent. But he who uses the term habit, must imply secondary automatism and a preexisting plastic adaptability. Von Buttel adduces an admirable proof of this whole matter and at the same time one of the clearest and simplest refutations of Bethe’s innumerable blunders, when he shows that bees that have never flown from the hive, even though they may be older than others that have already flown, are unable to find their way back even from a distance of a few meters, when they are unable to see the hive, whereas old bees know the whole environment, often to a distance of six or seven kilometers.
It results, therefore, from the unanimous observations of all the connoisseurs that sensation, perception, and association, inference, memory and habit follow in the social insects on the whole the same fundamental laws as in the vertebrates and ourselves. Furthermore, attention is surprisingly developed in insects, often taking on an obsessional character and being difficult to divert.
On the other hand, inherited automatism exhibits a colossal preponderance. The above-mentioned faculties are manifested only in an extremely feeble form beyond the confines of the in- stinct-automatism stereotyped in the species.
An insect is extraordinarily stupid and inadaptable to all things not related to its instincts. Nevertheless I succeeded in teaching a water-beetle {Dytiscus marginalis} which in nature feeds only in the water, to eat on my table. While thus feeding, it always executed a clumsy flexor-movement with its fore-legs which brought it over on its back. The insect learned to keep on feeding while on its back, but it would not dispense with this movement, which is adapted to feeding in the water. On the other hand, it always at- tempted to leap out of the water (no longer fleeing to the bottom of the vessel) when I entered the room, and nibbled at the tip of my finger in the most familiar manner. Now these are certainly plastic variations of instinct. In a similar manner some large Algerian ants which I transplanted to Zurich, learned during the course of the summer months to close the entrance of their nest with pellets of earth, because they were being persecuted and annoyed by our little Lasius. In Algiers I always saw the nest- opening wide open. There are many similar examples which go to show that these tiny animals can utilize some few of their experiences even when this requires a departure from the usual instincts.
That ants, bees, and wasps are able to exchange communications that are understood, and that they do not merely titillate one another with their antennae as Bethe maintains, has been demonstrated in so many hundred instances, that it is unnecessary to waste many words on this subject. The observations of a single predatory expedition of Polyergus, with a standing still of the whole army and a seeking for the lost trail, is proof sufficient of the above statement. But, of course, this is not language in the human sense !
22 ANTS AND SOME OTHER INSECTS.
There are no abstract concepts corresponding to the signs. We are here concerned only with hereditary, instinctively automatic signs. The same is true of their comprehension (pushing with the head, rushing at one another with wide-open mandibles, titillation with the antennae, stridulatory movement of the abdomen, etc.). Moreover, imitation plays a great role. Ants, bees, etc., imitate and follow their companions. Hence it is decidedly erroneous (and in this matter Wasmann, Von Buttel, and myself are of but one opinion) to inject human thought-conception and human ratiocination into this instinct language, as has been done to some extent, at least, even by Pierre Huber, not to mention others. It is even very doubtful whether a so-called general sensory idea (i. e. , a general idea of an object, like the idea “ant,” “enemy,” “nest,” “pupa”) can arise in the emmet brain. This is hardly capable of demonstration. Undoubtedly perception and association can be carried on in a very simple way, after the manner of insects, with- out ever rising to such complex results. At any rate proofs of such an assumption are lacking. But what exists is surely in itself sufficiently interesting and important. It gives us at least an insight into the brain-life of these animals.
Better than any generalisations, a good example will show what I mean.
Plateau had maintained that when Dahlia blossoms are covered with green leaves, bees nevertheless return to them at once. At first he concealed his Dahlias incompletely (i. e., only their ray- florets), afterwards completely, but still in an unsatisfactory manner, and inferred from the results that bees are attracted by odor and not by sight.
a. In a Dahlia bed visited by many bees and comprising about forty-three floral heads of different colors, I covered first seventeen and then eight at 2.15 P. M., September loth, with grape-leaves bent around them and fastened with pins.
b. Of four I covered only the yellow disc ;
c. Of one, on the other hand, I covered only the outer ray- florets, leaving the disc visible.
ANTS AND SOME OTHER INSECTS. 23
So many bees were visiting the Dahlias that at times there were two or three to a flower.
Result : Immediately all the completely covered flowers ceased to be visited by the bees. Dahlia (V) continued to be visited like those completely visible. The bees often flew to Dahlias (<5) but at once abandoned them ; a few, however, succeeded in finding the disc beneath the leaves.
Then as soon as I removed the covering from a red Dahlia the bees at once flew to it ; and soon a poorly concealed specimen was detected and visited. Later an inquisitive bee discovered the en- trance to a covered Dahlia from the side or from below. Thence- forth this bee, but only this one, returned to this same covered flower.
Nevertheless several bees seemed to be seeking the Dahlias which had so suddenly disappeared. Towards 5.30 o’clock some of them had detected the covered flowers. Thenceforth these in- sects were rapidly imitated by the other bees, and in a short time the hidden flowers were again being visited. As soon as a bee had discovered my imposition and found the entrance to a hidden flower, she flew in her subsequent journeys, without hesitation to the concealed opening of the grape-leaf. As long as a bee had merely made the discovery by herself, she remained unnoticed by the others. When this was accomplished by several, however, (usu- ally by four or five,) the others followed their example.
Plateau, therefore, conducted his experiments in a faulty manner and obtained erroneous results. The bees still saw the Dahlias which he at first incompletely concealed. Then, by the time he had covered them up completely, but only from above, they had already detected the fraud and saw the Dahlias also from the side. Plateau had failed to take into consideration the bee’s memory and attention.
September i3th I made some crude imitations of Dahlias by sticking the yellow heads of Hieracium (hawkweed) each in a Petunia flower, and placed them among the Dahlias. Neither the Petunias nor the Hieracium had been visited by the bees. Nevertheless many of the honey and humble-bees flew at first to the artefact’s in almost as great numbers as to the Dahlias, but at once abandoned the flowers when they had detected the error, obviously by means of their sense of smell. The same results were produced by a Dahlia, the disc of which had been replaced by the disc of a Hieracium.
As a control experiment I had placed a beautiful, odorous Dahlia disc among the white and yellow Chrysanthemums which had been neglected by the bees. For a whole half hour the bees flew by only a few centimeters above the disc without noticing it ; not till then was it visited by a bee that happened to be followed by a second. From this moment the Dahlia disc which lay in the path of flight was visited like the others, whereas on the other hand the Petunia-Hieracium artifacts, now known to be fraudulent, were no longer noticed.
Plateau has demonstrated that artificial flowers, no matter how carefully copied from the human standpoint, are not noticed by in- sects. I placed artifacts of this description among the Dahlias. They remained in fact entirely neglected. Perhaps, as above suggested, the bees are able to distinguish the chlorophyll colors from other artificial hues, owing to admixtures of the ultra-violet rays, or by some other means. But since Plateau imagines that the artificial flowers repel insects, I cut out, Sept. igth, the following rather crude paper-flowers :
a. A red flower ;
b. A white flower ;
c. A blue flower ;
d A blue flower, with a yellow center made from a dead leaf;
e. A rose-colored piece of paper with a dry Dahlia disc ;
f. A green Dahlia leaf (unchanged).
It was nine o’clock in the morning. I placed a drop of honey on each of the six artifacts mounted among the Dahlias. For a quarter of an hour many bees flew past, very close to my artifacts but without perceiving and hence without smelling the honey. I went away for an hour. On my return artefact 8 was without honey, and must therefore have been discovered by the bees. All the others had remained quite untouched and unnoticed.
ANTS AND SOME OTHER INSECTS. 25
With some difficulty I next undertook to bring artifact a very close to a bee resting on a Dahlia. But the attention of the bee was so deeply engrossed by the Dahlia that I had to repeat the experiment four or five times till I succeeded in bringing the honey within reach of her proboscis. The insect at once began to suck tip the honey from the paper-flower. I marked the bee’s back with blue paint so that I might be able to recognize her, and repeated the experiment with /? and e. In these cases one of the bees was painted yellow, the other white.
Soon the blue bee, which had in the meantime gone to the hive, returned, flew at once to a, first hovering about it dubiously, then to 8, where she fed, then again to a, but not to the Dahlias. Later the yellow bee returned to (3 and fed, and flew to a and 8 where she again fed, but gave as little heed to the Dahlias as did the blue bee.
Thereupon the white bee returned seeking e, but failing to find it, at once went to feeding on some of the Dahlias. But she tarried only a moment on each Dahlia as if tortured by the idea of honey. She returned to the artifacts, the perception of which, however, she was not quite able to associate with the memory of the honey flavor. At last she found a separate piece of e, which happened to be turned down somewhat behind, and began lapping up the honey.
This is part one of two, part two can be found at Psychic powers of insects and ants.
