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diff --git a/old/52134-0.txt b/old/52134-0.txt deleted file mode 100644 index 706979a..0000000 --- a/old/52134-0.txt +++ /dev/null @@ -1,2845 +0,0 @@ -The Project Gutenberg EBook of Ants and Some Other Insects: An Inquiry -Into the Psychic Powers of These Animal, by Auguste Forel - -This eBook is for the use of anyone anywhere in the United States and most -other parts of the world at no cost and with almost no restrictions -whatsoever. You may copy it, give it away or re-use it under the terms of -the Project Gutenberg License included with this eBook or online at -www.gutenberg.org. If you are not located in the United States, you'll have -to check the laws of the country where you are located before using this ebook. - -Title: Ants and Some Other Insects: An Inquiry Into the Psychic Powers of These Animals - -Author: Auguste Forel - -Translator: William Morton Wheeler - -Release Date: May 23, 2016 [EBook #52134] - -Language: English - -Character set encoding: UTF-8 - -*** START OF THIS PROJECT GUTENBERG EBOOK ANTS, OTHER INSECTS: PSYCHIC POWERS *** - - - - -Produced by Wayne Hammond, MWS, Bryan Ness and the Online -Distributed Proofreading Team at http://www.pgdp.net (This -file was produced from images generously made available -by The Internet Archive) - - - - - - - - - - _Ants and Some Other Insects_ - - - _An Inquiry into_ - - _The Psychic Powers of these Animals_ - - _With an Appendix on_ - - _The Peculiarities of Their Olfactory Sense_ - - - _By_ - - _Dr. August Forel_ - - _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, Trübner & Co. Ltd._ - - _1904_ - - COPYRIGHT, 1904 - THE OPEN COURT PUBLISHING CO. - CHICAGO - - - - -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 -organisation, but on the other hand, have come, through so-called -convergence, to possess in the form of a social commonwealth 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 Suisse_, and many others. Since -the functions of 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 Generale_, 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 recognise 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. - -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 conscious. 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 automatised 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 -organisation 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 psycho-physical -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 symbolised 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 -symbolised 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 conscious 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 may be forthwith discredited. -Energy can only be transformed qualitatively, not quantitatively. A -mind conceived as dualistic, if supposed 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 itself 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 sense-impressions by the brain; the -will represents the psycho- or cerebrofugal resultants of cognition -and the feelings together with their final transmission to the -muscles. 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 refined by the -former, but have profound hereditary and phylogenetic origins and are -relatively independent. There is a continual interaction 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 pleasure 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 -onomatopœas, 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 symbolised by words, as a -higher stage in generalisation. All these things give man a colossal -advantage, since he is thereby enabled to stand on the shoulders of -the written encyclopædia 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 symbolises nothing. On the other hand, some animals -possess phyletic, i. e., hereditarily and instinctively fixed cries -and gestures, 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 -utilising 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_. - -The so-called reflexes and their temporary, purposefully adaptive, but -hereditarily 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 reproduces 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 living protoplasm. Everything points to -the conclusion that the instinctive automatisms have been gradually -acquired and hereditarily 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 especially 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 -crystalised 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 Amœba, 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. They grade over into each other, 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 Amœba belong therefore rather to the -plasticity of living molecules, but not as yet to that of coöperating -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 considerable 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. - -[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. - -Spoken and especially written language, moreover, enable man to exploit -his brain to a wonderful extent. This leads us to underestimate -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. - -I beg your indulgence for this lengthy introduction to my subject, -but it seemed necessary that we should come to some understanding -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-fibres 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 supra-oesophageal ganglion and enable -them to possess themselves of the females during the nuptial flight. -No muscles are innervated by the supra-oesophageal ganglion. These -conditions greatly facilitate the comparison of the perceptive organs, -i. e., of the brain (_corpora pedunculata_) in the three sexes. This -is very large in the worker, much smaller in the female, 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 intended to say. - -In ants, injury to the cerebrum leads to the same results as injury 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 facetted 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 localise large spatial areas -admirably, but must show less definite contours of the objects than -our eyes. The compound eye yields only a single upright 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 insects are easily captured when very slowly -approached. In flight insects orient themselves in space by means -of their compound eyes. 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 skilful -workmanship. This may be due, to admixtures of the ultra-violet rays -which are invisible to our eyes. - -[Illustration: Fig. _W._] - -[Illustration: Fig. _F._] - -[Illustration: Fig. _M._] - - -EXPLANATION OF THE FIGURES. - -Brain (supra-œsophageal 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 olfactorius sive antennalis (olfactory lobe). _N._ = Facetted -eye. _N. olf._ = Nervus olfactorius 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. - -The ocelli (simple eyes) play a subordinate rôle, 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 antennæ, usually in the -club-shaped flagellum, or rather in the pore-plates and olfactory -rods of these portions of the antennæ. On account of its external -and moveable 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 itself 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 antennæ -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 polarisation. -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 (Dugès). - -The tactile sense is everywhere represented by tactile hairs and -papillæ. 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 psychology. -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 -antennæ 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 recognise 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 cannot 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 -antenna, seize a companion who forthwith rolls herself 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 suitable 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_) undertake 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 pupæ of the _Formica 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 contains pupæ, the robbers return on the same or following days -and carry off the remainder, but if there are no pupæ left they do not -return. How do the Polyergus know whether there are pupæ remaining? -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 pupæ still -remain behind in the plundered nest can induce them to return. 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 shovelling 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 recognise friends even after the expiration -of months. In ants and bees there are very complex combinations and -mixtures of odors, which Von Buttel 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 -rôle 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 execution 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 pre-existing -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 instinct-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 attempted 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 niger_. In Algiers I always saw the -nest-opening wide open. There are many similar examples which go to -show that these tiny animals can utilise 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 antennæ 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! 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 antennæ, stridulatory -movement of the abdomen, etc.). Moreover, imitation plays a great -rôle. 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, without 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 10th, 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. - -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 (_c_) continued to be visited like those -completely visible. The bees often flew to Dahlias (_b_) 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 entrance to a covered -Dahlia from the side or from below. Thenceforth 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 insects 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, (usually 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 13th 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 artefacts 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 artefacts, 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 insects. -I placed artefacts 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. 19th, the following rather crude paper-flowers: - -α. A red flower; - -β. A white flower; - -γ. A blue flower; - -δ A blue flower, with a yellow center made from a dead leaf; - -ε. A rose-colored piece of paper with a dry Dahlia disc; - -ζ. A green Dahlia leaf (unchanged). - -It was nine o’clock in the morning. I placed a drop of honey on each of -the six artefacts mounted among the Dahlias. For a quarter of an hour -many bees flew past, very close to my artefacts but without perceiving -and hence without smelling the honey. I went away for an hour. On my -return artefact δ was without honey, and must therefore have been -discovered by the bees. All the others had remained quite untouched and -unnoticed. - -With some difficulty I next undertook to bring artefact α 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 up the honey from the -paper-flower. I marked the bee’s back with blue paint so that I might -be able to recognise her, and repeated the experiment with β and ε. 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 α, first hovering about it dubiously, then to -δ, where she fed, then again to α, but not to the Dahlias. Later the -yellow bee returned to β and fed, and flew to α and δ where she again -fed, but gave as little heed to the Dahlias as did the blue bee. - -Thereupon the white bee returned seeking ε, 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 _idée fixe_ of honey. She -returned to the artefacts, 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 ε, which happened to be turned down -somewhat behind, and began lapping up the honey. - -Thenceforth the three painted bees, and these alone, returned regularly -to the artefacts and no longer visited the Dahlias. The fact is of -great importance that the painted bees entirely of their own accord, -undoubtedly through an instinctive inference from analogy, discovered -the other artefacts as soon as their attention had been attracted by -the honey on one of them, notwithstanding the fact that the artefacts -were some distance from one another and of different colors. For were -not the Dahlias, too, which they had previously visited, of different -colors? Thus the blue bee flew to α, β, γ, and δ, the yellow to β, α, -δ, and γ, the white ε, α, β, and δ. Matters continued thus for half -an hour. The hidden green ζ was not found, evidently because it was -indistinguishable from the green foliage. - -Finally one bee, by herself, having had in all probability her -attention attracted by the three others, came to δ and fed. I marked -her with carmine. Thereupon she flew to α and drove the blue bee away. -Another bee was attracted to ε of her own accord and was painted with -cinnobar. Still another bee came by herself to β and was painted green. -It was now 12.30 o’clock. The experiment had therefore lasted more than -three hours, and during this time only six bees had come to know the -artefacts, while the great majority still kept on visiting the Dahlias. -But now the other bees began to have their attention attracted by the -visitors to the artefacts. One, then two, then three, and finally more -new ones followed, and I had not sufficient colors with which to mark -them. Every moment I was obliged to replenish the honey. Then I went to -dinner and returned at 1.25. At this moment seven bees were feeding on -β, two on α, one on γ, three on δ, the white one alone on ε. More than -half of all these were new, unpainted followers. Now a veritable swarm -of bees threw themselves on the artefacts and licked up the last traces -of the honey. Then for the first time, after more than four hours, a -bee from the swarm discovered the honey on the artefact ζ, which on -account of its color had remained concealed up to this time! - -As a pack of hounds throws itself on an empty skeleton, the swarm of -bees, now completely diverted from the Dahlias, cast themselves on the -completely empty artefacts and vainly searched every corner of them for -honey. It was 1.55 P. M. The bees began to scatter and return to the -Dahlias. Then I replaced α and β by a red and white paper respectively, -which had never come in contact with honey and could not therefore -smell of the substance. These pieces of paper, nevertheless, were -visited and examined by various bees, whose brains were still possessed -with the fixed idea of the flavor of honey. The white bee, e. g., -investigated the white paper very carefully for a period of three to -four minutes. There could, of course, be no such thing as an unknown -force or attraction of odor, or brilliancy of floral colors. This fact -can only be explained by an association of space, form, and color -memories with memories of taste. - -Thereupon I took all the artefacts in my left hand for the purpose of -carrying them away. Two or three bees followed me, hovering about my -left hand, and tried to alight on the empty artefacts. The space-image -had changed and only the color and form could any longer be of service -to the bees in their recognition of these objects. - -This experiment is so clear and unequivocal that I mention it here -among many others. It demonstrates: - -1. The space, form, and color perceptions of the honey-bee. That these -are possible only through the agency of the compound eyes is proved by -other experiments (varnishing the eyes, extirpation of the antennæ, -mouth-parts, etc.). - -2. The memory of the honey-bee, in particular her visual and gustatory -memory. - -3. Her power of associating gustatory with visual memories. - -4. Her ability instinctively to draw inferences from analogy: If she -has once been offered honey in an artefact, she will investigate -others, even those of a different color and hitherto unnoticed. These -she compares by means of the visual sense, since they are relatively -similar, and recognises them as similar though such objects are most -unusual in the bee’s experience. - -5. Her poor olfactory sense, which is useful only at very close range. - -6. The onesidedness and narrow circle of her attention. - -7. The rapid formation of habits. - -8. The limits of imitation of bees by one another. - -Of course, I should not allow myself to draw these conclusions from -a single experiment, if they had not been confirmed by innumerable -observations by the ablest investigators in this field. Lubbock showed -clearly that it is necessary to train a bee for some time to go to -a particular color if one wishes to compel her to pay no attention -to other colors. This is the only way in which it is possible to -demonstrate her ability to distinguish colors. My bees, on the -contrary, had been trained on differently colored objects (Dahlias and -artefacts) and therefore paid no attention to differences in color. It -would be a fallacy to conclude from this that they do not distinguish -colors. On the contrary, by means of other experiments I have fully -confirmed Lubbock’s results. - -By 2.20 P. M. all of my bees, even the painted ones, had returned to -the Dahlias. - -On September 27, a week later, I wished to perform a fresh experiment -with the same bees. I intended to make them distinguish between -differently colored discs, placed at different points on a long scale, -representing on a great sheet of paper, varying intensities of light -from white through gray to black. First, I wished to train a bee to -a single color. But I had calculated without the bee’s memory, which -rendered the whole experiment impracticable. Scarcely had I placed -my paper with the discs on the lawn near the Dahlia bed, and placed -one or two bees on the blue discs and marked them with colors, when -they began to investigate all the red, blue, white, black and other -discs with or without honey. After a few moments had elapsed, other -bees came from the Dahlia bed and in a short time a whole swarm threw -itself on the paper discs. Of course, those that had been provided with -honey were most visited, because they detained the bees, but even the -discs without honey were stormed and scrutinised by bees following one -another in their flight. The bees besieged even the paint-box. Among -these there was one that I had previously deprived of her antennæ. She -had previously partaken of the honey on the blue discs and had returned -to the hive. This bee examined the blue piece of paint in the color-box. - -In brief, my experiment was impossible, because all the bees still -remembered from a former occasion the many-colored artefacts provided -with honey, and therefore examined all the paper discs no matter of -what color. The association between the taste of the honey and the -paper discs had been again aroused by the sight-perception of the -latter, and had acquired both consistency and rapid and powerful -imitation, because honey happened to be actually found on some of the -discs. - -Together with the perceptive and associative powers, the power of -drawing simple, instinctive inferences from analogy is also apparent. -Without this, indeed, the operation of perception and memory would be -inconceivable! We have just given an example. I have shown on a former -occasion that humble-bees, whose nest I had transferred to my window, -when they returned home often confounded other windows of the same -façade and examined them for a long time before they discovered the -right one. Lubbock reports similar facts. Von Buttel shows that bees -that are accustomed to rooms and windows, learn to examine the rooms -and windows in other places, i. e., other houses. When Pissot suspended -wire netting with meshes twenty-two mm. in diameter in front of a wasp -nest, the wasps hesitated at first, then went around the netting by -crawling along the ground or avoided it in some other way. But they -soon learned to fly directly through the meshes. The sense of sight, -observed during flight, is particularly well adapted to experiments -of this kind, which cannot therefore be performed with ants. But the -latter undoubtedly draw similar inferences from the data derived from -their topochemical antennal sense. The discovery of prey or other food -on a plant or an object induces these insects to examine similar plants -or objects and to perform other actions of a like nature. - -There are, on the other hand, certain very stupid insects, like -the males of ants, the Diptera and may-flies (Ephemerids) with -rudimental brains, incapable of learning anything or of combining -sense-impressions to any higher degree than as simple automatisms, and -without any demonstrable retention of memory-images. Such insects lead -a life almost exclusively dominated by sensory stimuli; but their lives -are adapted to extremely simple conditions. In these very instances the -difference is most striking, and they demonstrate most clearly through -comparison and contrast the _plus_ possessed by more intelligent -insects. - - -THE REALM OF WILL. - -The notion of volition, in contradistinction to the notion of reflex -action, presupposes the expiration of a certain time interval and -the operation of mediating and complex brain-activities between the -sense-impression and the movement which it conditions. In the operation -of the purposeful automatisms of instinct which arouse one another -into activity in certain sequences, there is also a time interval, -filled out by internal, dynamic brain-processes as in the case of -the will. Hence these are not pure reflexes. They may for a time -suffer interruption and then be again continued. But their operation -is brought about in great measure by a concatenation of complicated -reflexes which follow one another in a compulsory order. On this -account the term automatism or instinct is justifiable. - -If we are to speak of will in the narrower sense, we must be able to -establish the existence of individual decisions, which can be directed -according to circumstances, i. e., are modifiable, and may, for a -certain period, remain dormant in the brain to be still performed -notwithstanding. Such volition may be very different from the complex -volition of man, which consists of the resultants of prodigiously -manifold components that have been long preparing and combining. The -ants exhibit positive and negative volitional phenomena, which cannot -be mistaken. The ants of the genus Formica Linné are particularly -brilliant in this respect, and they also illustrate the individual -psychical activities most clearly. The above-mentioned migrations from -nest to nest show very beautifully the individual plans of single -workers carried out with great tenacity. For hours at a time an ant -may try to overcome a multitude of difficulties for the purpose of -attaining an aim which she has set herself. This aim is not accurately -prescribed by instinct, as the insect may be confronted with several -possibilities, so that it often happens that two ants may be working in -opposition to each other. This looks like stupidity to the superficial -observer. But it is just here that the ant’s plasticity reveals itself. -For a time the two little animals interfere with each other, but -finally they notice the fact, and one of them gives in, goes away, or -assists the other. - -These conditions are best observed during the building of nests or -roads, e. g., in the horse-ant (_Formica rufa_) and still better in _F. -pratensis_. It is necessary, however, to follow the behavior of a few -ants for hours, if one would have a clear conception of this matter, -and for this much patience and much time are necessary. The combats -between ants, too, show certain very consistent aims of behavior, -especially the struggles which I have called chronic combats (_combats -à froid_). After two parties (two colonies brought together) have made -peace with each other, one often sees a few individuals persecuting and -maltreating certain individuals of the opposite party. They often carry -their victims a long distance off, for the purpose of excluding them -from the nest. If the ant that has been borne away returns to the nest -and is found by her persecutrix, she is again seized and carried away -to a still greater distance. In one such case in an artificial nest -of a small species of Leptothorax, the persecuting ant succeeded in -dragging her victim to the edge of my table. She then stretched out her -head and allowed her burden to fall on the floor. This was not chance, -for she repeated the performance twice in succession after I had again -placed the victim on the table. Among the different individuals of the -previously hostile, but now pacified opposition, she had concentrated -her antipathy on this particular ant and had tried to make her return -to the nest impossible. One must have very strong preconceived opinions -if in such and many similar cases one would maintain that ants are -lacking in individual decision and execution. Of course, all these -things happen within the confines of the instinct-precincts of the -species, and the different stages in the execution of a project are -instinctive. Moreover, I expressly defend myself against the imputation -that I am importing human reflection and abstract concepts into this -volition of the ant, though we must honestly admit, nevertheless, -that in the accomplishment of our human decisions both hereditary and -secondary automatisms are permitted to pass unnoticed. While I am -writing these words, my eyes operate with partially hereditary, and my -hand with secondary automatisms. But it goes without saying that only -a human brain is capable of carrying out my complex innervations and -my concomitant abstract reflections. But the ant must, nevertheless, -associate and consider somewhat in a concrete way after the manner of -an ant, when it pursues one of the above-mentioned aims and combines -its instincts with this special object in view. While, however, -the instinct of the ant can be combined for only a few slightly -different purposes, by means of a small number of plastic adaptations -or associations, individually interrupted in their concatenation or -_vice versa_, in the thinking human being both inherited and secondary -automatisms are only fragments or instruments in the service of an -overwhelming, all-controlling, plastic brain-activity. It may be said -incidentally that the relative independence of the spinal chord and of -subordinate brain-centers in the lower animals (and even in the lower -mammals) as compared with the cerebrum, may be explained in a similar -manner if they are compared with the profound dependence of these -organs and their functions on the massive cerebrum in man and even -to some extent in the apes. The cerebrum splits up and controls its -automatisms (_divide et impera_). - -While success visibly heightens both the audacity and tenacity of -the ant-will, it is possible to observe after repeated failure or in -consequence of the sudden and unexpected attacks of powerful enemies -a form of abulic dejection, which may lead to a neglect of the most -important instincts, to cowardly flight, to the devouring or casting -away of offspring, to neglect of work, and similar conditions. There is -a chronically cumulative discouragement in degenerate ant-colonies and -an acute discouragement when a combat is lost. In the latter case one -may see troops of large powerful ants fleeing before a single enemy, -without even attempting to defend themselves, whereas the latter a -few moments previously would have been killed by a few bites from the -fleeing individuals. It is remarkable how soon the victor notices and -utilises this abulic discouragement. The dejected ants usually rally -after the flight and soon take heart and initiative again. But they -offer but feeble resistance, e. g., to a renewed attack from the same -enemy on the following day. Even an ant’s brain does not so soon forget -the defeats which it has suffered. - -In bitter conflicts between two colonies of nearly equal strength the -tenacity of the struggle and with it the will to conquer increases till -one of the parties is definitively overpowered. In the realm of will -imitation plays a great rôle. Even among ants protervity and dejection -are singularly contagious. - - -THE REALM OF FEELING. - -It may perhaps sound ludicrous to speak of feelings in insects. But -when we stop to consider how profoundly instinctive and fixed is our -human life of feeling, how pronounced are the emotions in our domestic -animals, and how closely interwoven with the impulses, we should expect -to encounter emotions and feelings in animal psychology. And these -may indeed be recognised so clearly that even Uexkuell would have to -capitulate if he should come to know them more accurately. We find -them already interwoven with the will as we have described it. Most -of the emotions of insects are profoundly united to the instincts. -Of such a nature is the jealousy of the queen bee when she kills the -rival princesses, and the terror of the latter while they are still -within their cells; such is the rage of fighting ants, wasps, and -bees, the above-mentioned discouragement, the love of the brood, the -self-devotion of the worker honey-bees, when they die of hunger while -feeding their queen, and many other cases of a similar description. But -there are also individual emotions that are not compelled altogether -by instinct, e. g., the above-mentioned mania of certain ants for -maltreating some of their antagonists. On the other hand, as I have -shown, friendly services (feeding), under exceptional circumstances, -may call forth feelings of sympathy and finally of partnership, -even between ants of different species. Further than this, feelings -of sympathy, antipathy, and anger among ants may be intensified by -repetition and by the corresponding activities, just as in other -animals and man. - -The social sense of duty is instinctive in ants, though they exhibit -great individual, temporary, and occasional deviations, which betray a -certain amount of plasticity. - - -PSYCHIC CORRELATIONS. - -I have rapidly reviewed the three main realms of ant-psychology. It -is self-evident that in this matter they no more admit of sharp -demarcation from one another than elsewhere. The will consists of -centrifugal resultants of sense-impressions and feelings and in turn -reacts powerfully on both of these. - -It is of considerable interest to observe the antagonism between -different perceptions, feelings, and volitions in ants and bees, and -the manner in which in these animals the intensely fixed (obsessional) -attention may be finally diverted from one thing to another. Here -experiment is able to teach us much. While bees are busy foraging on -only one species of flower, they overlook everything else, even other -flowers. If their attention is diverted by honey offered them directly, -although previously overlooked, they have eyes only for the honey. An -intense emotion, like the swarming of honey-bees (von Buttel) compels -these insects to forget all animosities and even the old maternal -hive to which they no longer return. But if the latter happens to be -painted blue, and if the swarming is interrupted by taking away the -queen, the bees recollect the blue color of their old hive and fly to -hives that are painted blue. Two feelings often struggle with each -other in bees that are “crying” and without a queen: that of animosity -towards strange bees and the desire for a queen. Now if they be given a -strange queen by artificial means, they kill or maltreat her, because -the former feeling at first predominates. For this reason the apiarist -encloses the strange queen in a wire cage. Then the foreign odor -annoys the bees less because it is further away and they are unable to -persecute the queen. Still they recognise the specific queen-odor and -are able to feed her through the bars of the cage. This suffices to -pacify the hive. Then the second feeling quickly comes to the front; -the workers become rapidly inured to the new odor and after three or -four days have elapsed, the queen may be liberated without peril. - -It is possible in ants to make the love of sweets struggle with the -sense of duty, when enemies are made to attack a colony and honey is -placed before the ants streaming forth to defend their nest. I have -done this with _Formica pratensis_. At first the ants partook of the -honey, but only for an instant. The sense of duty conquered and all of -them without exception, hurried forth to battle and most of them to -death. In this case a higher decision of instinct was victorious over -the lower impulse. - -In _résumé_ I would lay stress on the following general conclusions: - -1. From the standpoint of natural science we are bound to hold -fast to the psychophysiological theory of identity (Monism) in -contradistinction to dualism, because it alone is in harmony with the -facts and with the law of the conservation of energy. - -Our mind must be studied simultaneously both directly from within and -indirectly from without, through biology and the conditions of its -origin. Hence there is such a thing as comparative psychology of other -individuals in addition to that of self, and in like manner we are -led to a psychology of animals. Inference from analogy, applied with -caution, is not only permissible in this science, but obligatory. - -2. The senses of insects are our own. Only the auditory sense still -remains doubtful, so far as its location and interpretation are -concerned. A sixth sense has not yet been shown to exist, and a special -sense of direction and orientation is certainly lacking. The vestibular -apparatus of vertebrates is merely an organ of equilibration and -mediates internal sensations of acceleration, but gives no orientation -in space outside of the body. On the other hand the visual and -olfactory senses of insects present varieties in the range of their -competency and in their specific energies (vision of ultra-violet, -functional peculiarities of the facetted eye, topochemical antennal -sense and contact-odor). - -3. Reflexes, instincts, and plastic, individually adaptive, central -nervous activities pass over into one another by gradations. Higher -complications of these central or psychic functions correspond to -a more complicated apparatus of superordinated neuron-complexes -(cerebrum). - -4. Without becoming antagonistic, the central nervous activity in the -different groups and species of animals complicates itself in two -directions: (_a_) through inheritance (natural selection, etc.) of -the complex, purposeful automatisms, or instincts; (_b_) through the -increasingly manifold possibilities of plastic, individually adaptive -activities, in combination with the faculty of gradually developing -secondary individual automatisms (habits). - -The latter mode requires many more nerve-elements. Through hereditary -predispositions (imperfect instincts) of greater or less stability, it -presents transitions to the former mode. - -5. In social insects the correlation of more developed psychic powers -with the volume of the brain may be directly observed. - -6. In these animals it is possible to demonstrate the existence of -memory, associations of sensory images, perceptions, attention, habits, -simple powers of inference from analogy, the utilisation of individual -experiences and hence distinct, though feeble, plastic, individual -deliberations or adaptations. - -7. It is also possible to detect a corresponding, simpler form of -volition, i. e., the carrying out of individual decisions in a more -or less protracted time-sequence, through different concatenations -of instincts; furthermore different kinds of discomfort and pleasure -emotions, as well as interactions and antagonisms between these diverse -psychic powers. - -8. In insect behavior the activity of the attention is one-sided -and occupies a prominent place. It narrows the scope of behavior -and renders the animal temporarily blind (inattentive) to other -sense-impressions. - -Thus, however different may be the development of the automatic and -plastic, central neurocyme activities in the brains of different -animals, it is surely possible, nevertheless, to recognise certain -generally valid series of phenomena and their fundamental laws. - -Even to-day I am compelled to uphold the seventh thesis which I -established in 1877 in my habilitation as _privat-docent_ in the -University of Munich: - -“All the properties of the human mind may be derived from the -properties of the animal mind.” - -I would merely add to this: - -“And all the mental attributes of higher animals may be derived from -those of lower animals.” In other words: The doctrine of evolution -is quite as valid in the province of psychology as it is in all the -other provinces of organic life. Notwithstanding all the differences -presented by animal organisms and the conditions of their existence, -the psychic functions of the nerve-elements seem nevertheless, -everywhere to be in accord with certain fundamental laws, even in the -cases where this would be least expected on account of the magnitude of -the differences. - - -APPENDIX. - -THE PECULIARITIES OF THE OLFACTORY SENSE IN INSECTS. - -Our sense of smell, like our sense of taste, is a chemical sense. But -while the latter reacts only to substances dissolved in liquids and -with but few (about five) different principal qualities, the olfactory -sense reacts with innumerable qualities to particles of the most -diverse substances dissolved in the atmosphere. Even to our relatively -degenerate human olfactories, the number of these odor-qualities seems -to be almost infinite. - -In insects that live in the air and on the earth the sense of taste -seems to be located, not only like our own, in the mouth-parts, but -also to exhibit the same qualities and the corresponding reactions. At -any rate it is easy to show that these animals are usually very fond -of sweet, and dislike bitter things, and that they perceive these two -properties only after having tasted of the respective substances. F. -Will, in particular, has published good experiments on this subject. - -In aquatic insects the conditions are more complicated. Nagel, who -studied them more closely, shows how difficult it is in these cases -to distinguish smell from taste, since substances dissolved in water -are more or less clearly perceived or discerned from a distance by -both senses and sought or avoided in consequence. Nagel, at any rate, -succeeded in showing that the palpi, which are of less importance in -terrestrial insects, have an important function in aquatic forms. - -In this place we are concerned with an investigation of the sense of -smell in terrestrial insects. Its seat has been proved to be in the -antennæ. A less important adjunct to these organs is located, as Nagel -and Wasmann have shown, in the palpi. In the antennæ it is usually the -club or foliaceous or otherwise formed dilatations which accommodate -the cellular ganglion of the antennary nerve. I shall not discuss the -histological structure of the nerve-terminations but refer instead to -Hicks, Leydig, Hauser, my own investigations and the other pertinent -literature, especially to K. Kraepelin’s excellent work. I would merely -emphasise the following points: - -1. All the olfactory papillæ of the antennæ are transformed, hair-like -pore-canals. - -2. All of these present a cellular dilatation just in front of the -nerve-termination. - -3. Tactile hairs are found on the antennæ together with the olfactory -papillæ. - -4. The character and form of the nerve-terminations are highly -variable, but they may be reduced to three principal types: -pore-plates, olfactory rods, and olfactory hairs. The two latter -are often nearly or quite indistinguishable from each other. The -nerve-termination is always covered with a cuticula which may be never -so delicate. - -Other end-organs of the Hymenopteran antenna described by Hicks and -myself, are still entirely obscure, so far as their function is -concerned, but they can have nothing to do with the sense of smell, -since they are absent in insects with a delicate sense of smell (wasps) -and occur in great numbers in the honey-bees, which have obtuse -olfactories. - -That the antennæ and not the nerve-terminations of the mouth and palate -function, as organs of smell, has been demonstrated by my control -experiments, which leave absolutely no grounds for doubt and have, -moreover, been corroborated on all sides. Terrestrial insects can -discern chemical substances at a distance by means of their antennæ -only. But in touch, too, these organs are most important and the palpi -only to a subordinate extent, namely in mastication. The antennæ enable -the insect to perceive the chemical nature of bodies and in particular, -to recognise and distinguish plants, other animals and food, except -in so far as the visual and gustatory senses are concerned in these -activities. These two senses may be readily eliminated, however, since -the latter functions only during feeding and the former can be removed -by varnishing the eyes or by other means. Many insects, too, are blind -and find their way about exclusively by means of their antennæ. This is -the case, e. g., with many predatory ants of the genus Eciton. - -But I will here assume these questions to be known and answered, nor -will I indulge in polemics with Bethe and his associates concerning -the propriety of designating the chemical antennal sense as “smell.” -I have discussed this matter elsewhere.[2] What I wish to investigate -in this place is the psychological quality of the antennal olfactory -sense, how it results in part from observation and in part from the -too little heeded correlative laws of the psychological exploitation -of each sense in accordance with its structure. I assume as known the -doctrines of specific energies and adequate stimuli, together with the -more recent investigations on the still undifferentiated senses, like -photodermatism and the like, and would refer, moreover, to Helmholtz’s -_Die Thatsachen in der Wahrnehmung_, 1879. Hirschwald, Berlin. - -[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. - -When in our own human subjective psychology, which alone is known -to us directly, we investigate the manner in which we interpret our -sensations, we happen upon a peculiar fact to which especially Herbert -Spencer has called attention. We find that so-called perceptions -consist, as is well known, of sensations which are bound together -sometimes firmly, sometimes more loosely. The more intimately the -sensations are bound together to form a whole, the easier it is for us -to recall in our memory the whole from a part. Thus, e. g., it is easy -for me to form an idea from the thought of the head of an acquaintance -as to the remainder of his body. In the same manner the first note of -a melody or the first verse of a poem brings back the remainder of -either. But the thought of an odor of violets, a sensation of hunger, -or a stomach-ache, are incapable of recalling in me either simultaneous -or subsequent odors or feelings. - -These latter conditions call up in my consciousness much more easily -certain associated visual, tactile, or auditory images (e. g., the -visual image of a violet, a table set for a meal). As ideas they are -commonly to be represented in consciousness only with considerable -difficulty, and sometimes not at all, and they are scarcely capable of -association among themselves. We readily observe, moreover, that visual -images furnish us mainly with space recollections, auditory images with -sequences in time, and tactile images with both, but less perfectly. -These are indubitable and well-known facts. - -But when we seek for the wherefore of these phenomena, we find the -answer in the structure of the particular sense-organ and in its manner -of functioning. - -It is well known that the eye gives us a very accurate image of the -external world on our retina. Colors and forms are there depicted in -the most delicate detail, and both the convergence of our two eyes and -their movement and accommodation gives us besides the dimensions of -depth through stereoscopic vision. Whatever may be still lacking or -disturbing is supplied by instinctive inferences acquired by practice, -both in memory and direct perception (like the lacunæ of the visual -field), or ignored (like the turbidity of the corpus vitreum). But the -basis of the visual image is given in the coördinated _tout ensemble_ -of the retinal stimuli, namely the retinal image.[3] Hence, since the -retina furnishes us with such spatial projections, and these in sharp -details, or relations, definitely coöordinated with one another, the -sense of sight gives us knowledge of space. For this reason, also, and -solely on this account, we find it so easy to supply through memory by -association the missing remnant of a visual spatial image. For this -reason, too, the visual sensations are preëminently associative or -relational in space, to use Spencer’s expression. For the same reason -the insane person so readily exhibits hallucinations of complicated -spatial images in the visual sphere. This would be impossible in the -case of the olfactory sense. - -[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. - -Similarly, the organ of Corti in the ear gives us tone or sound scales -in accurate time-sequence, and hence also associations of sequence much -more perfectly than the other senses. Its associations are thus in the -main associations of sequence, because the end-apparatus registers -time-sequences in time-intervals and not as space images. - -The corresponding cortical receptive areas are capable, in the first -instance, merely of registering what is brought to them by the -sense-stimuli and these are mainly associated spatial images for sight -and tone or sound-sequences for hearing. - -Let us consider for a moment how odors strike the mucous membranes of -our choanæ. They are wafted towards us as wild mixtures in an airy -maelstrom, which brings them to the olfactory terminations without -order in the inhaled air or in the mucous of the palate. They come in -such a way that there cannot possibly be any spatial association of the -different odors in definite relationships. In time they succeed one -another slowly and without order, according to the law of the stronger -element in the mixture, but without any definite combination. If, after -one has been inhaling the odor of violets, the atmosphere gradually -becomes charged with more roast meat than violet particles, the odor -of roast succeeds that of violet. But nowhere can we perceive anything -like a definitely associated sequence, so that neither our ideas of -time nor those of space comprise odors that revive one another through -association. By much sniffing of the surface of objects we could at -most finally succeed in forming a kind of spatial image, but this would -be very difficult owing to man’s upright posture. Nevertheless it is -probable that dogs, hedge-hogs, and similar animals acquire a certain -olfactory image by means of sniffing. The same conditions obtain in -the sphere of taste and the visceral sensations for the same reasons. -None of these senses furnish us with any sharply defined qualitative -relations either in space or time. On this account they furnish by -themselves no associations, no true perceptions, no memory images, -but merely sensations, and these often as mixed sensations, which are -vague and capable of being associated only with associative senses. -The hallucinations of smell, taste, and of the splanchnic sensations, -are not deceptive perceptions, since they cannot have a deceptive -resemblance to objects. They are simply paræsthesias or hyperæsthesias, -i. e., pathological sensations of an elementary character either -without adequate stimulus or inadequate to the stimulus. - -The tactile sense furnishes us with a gross perception of space and of -definite relations, and may, therefore, give rise to hallucinations, or -false perceptions of objects. By better training its associative powers -in the blind may be intensified. The visual sensations are usually -associated with tactile localisations. - -Thus we see that there is a law according to which the psychology -of a sense depends not only on its specific energy but also on the -manner in which it is able to transmit to the brain the relations -of its qualities in space and time. On this depends the knowledge -we acquire concerning time and space relations through a particular -sense and hence also its ability to form perceptions and associations -in the brain. More or less experience is, of course, to be added or -subtracted, but this is merely capable of enriching the knowledge -of its possessor according to the measure of the relations of the -particular sense-stimuli in space and time. - -I would beg you to hold fast to what I have said and then to picture to -yourselves an olfactory sense, i. e., a chemical sense effective at a -distance and like our sense of smell, capable of receiving impressions -from particles of the most diverse substances diffused through the -atmosphere, located not in your nostrils, but on your hands. For of -such a nature is the position of the olfactory sense on the antennal -club of the ant. - -Now imagine your olfactory hands in continual vibration, touching all -objects to the right and to the left as you walk along, thereby rapidly -locating the position of all odoriferous objects as you approach or -recede from them, and perceiving the surfaces both simultaneously and -successively as parts of objects differing in odor and position. It is -clear from the very outset that such sense-organs would enable you to -construct a veritable odor-chart of the path you had traversed and one -of double significance: - -1. A clear contact-odor chart, restricted, to be sure, to the immediate -environment and giving the accurate odor-form of the objects touched -(round odors, rectangular odors, elongate odors, etc.) and further hard -and soft odors in combination with the tactile sensations. - -2. A less definite chart which, however, has orienting value for a -certain distance, and produces emanations which we may picture to -ourselves like the red gas of bromine which we can actually see. - -If we have demonstrated that ants perceive chemical qualities through -their antennæ both from contact and from a distance, then the antennæ -must give them knowledge of space, if the above formulated law is true, -and concerning this there can be little doubt. This must be true even -from the fact that the two antennæ simultaneously perceive different -and differently odoriferous portions of space.[4] - -[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. - -They must therefore also transmit perceptions and topographically -associated memories concerning a path thus touched and smelled. Both -the trail of the ants themselves and the surrounding objects must leave -in their brains a chemical (odor-) space-form with different, more -or less definitely circumscribed qualities, i. e., an odor-image of -immediate space, and this must render associated memories possible. -Thus an ant must perceive the forms of its trail by means of smell. -This is impossible, at least for the majority of the species, by means -of the eyes. If this is true, an ant will always be able, no matter -where she may be placed on her trail, to perceive what is to the -right, left, behind or before her, and consequently what direction -she is to take, according to whether she is bound for home, or in the -opposite direction to a tree infested with Aphides, or the like. - -Singularly enough, I had established this latter fact in my “Études -Myrmécologiques en 1886” (_Annales de la Societé Entomologique de -Belgique_) before I had arrived at its theoretical interpretation. -But I was at once led by this discovery in the same work to the -interpretation just given. Without knowing of my work in this -connection, A. Bethe has recently established (discovered, as he -supposes) this same fact, and has designated it as “polarisation of -the ant-trail.” He regards this as the expression of a mysterious, -inexplicable force, or polarisation. As we have seen, the matter is -not only no enigma, but on the contrary, a necessary psychological -postulate. We should rather find the absence of this faculty -incomprehensible. - -But everything I have just said presupposes a receptive brain. The -formation of lasting perceptions and associations cannot take place -without an organ capable of fixing the sense-impressions and of -combining them among themselves. Experience shows that the immediate -sensory centers are inadequate to the performance of this task. Though -undoubtedly receptive, they are, nevertheless, incapable of utilising -what has been received in the development of more complex instincts -and can turn it to account only in the grosser, simpler reflexes and -automatisms. To be sure, a male ant has better eyes than a worker ant, -and probably quite as good antennæ, but he is unable to remember what -he has seen and is especially incapable of associating it in the form -of a trail-image, because he is almost devoid of a brain. For this -reason he is unable to find his way back to the nest. On the other -hand, it is well known that the brain of a man who has lost a limb or -whose hearing is defective, will enable him to paint pictures with his -foot, write with the stump of an arm or construct grand combinations -from the images of defective senses. - -I venture, therefore, to designate as topochemical the olfactory -antennal sense of honey-bees, humble-bees, wasps, etc. - -Can we generalise to such an extent as to apply this term without -further investigation to all arthropods? To a considerable extent this -must be denied. - -In fact, the multiformity in the structure and development of the -arthropod sense-organs is enormous, and we must exercise caution in -making premature generalisations. - -It is certain that in some aerial insects the olfactory sense has -dwindled to a minimum, e. g., in those species in which the male -recognises and follows the female exclusively by means of the eyes, as -in the Odonata (dragon-flies). To insects with such habits an olfactory -sense would be almost superfluous. Here, too, the antennæ have dwindled -to diminutive dimensions. - -But there are insects whose antennæ are immovable and quite unable to -touch objects. This is the case in most Diptera (flies). Still these -antennæ are often highly developed and present striking dilatations -densely beset with olfactory papillæ. By experiment I have demonstrated -the existence of an olfactory sense in such Dipteran antennæ, and I -have been able to show that, e. g., in _Sarcophaga vivipara_ and other -carrion flies, the egg-laying instinct is absolutely dependent on the -sensation of the odor of carrion and the presence of the antennæ. In -these cases the contact-odor sense is undoubtedly absent. More or -less of a topochemical odor-sense at long range must, of course, be -present, since the antennæ are external, but the precision of the -spatial image must be very imperfect, owing to the immobility of the -antennæ. Nevertheless, flies move about so rapidly in the air that they -must be able by means of their antennæ to distinguish very quickly -the direction from which odors are being wafted. These insects do, in -fact, find the concealed source of odors with great assurance. But -this is no great art, for even we ourselves are able to do the same by -sniffing or going to and fro. But the flies find their way through the -air with their eyes and not at all by means of their sense of smell. -Hence their olfactory powers probably constitute a closer psychological -approximation to those of mammals than to the topochemical odor-sense -of ants, for they can hardly furnish any constant and definite -space-relations. - -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. 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