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ANIMAL BEHAVIOUR
ANIMAL BEHAVIOUR
BY C. LLOYD MORGAN, F.R.S. AUTHOR OF “THE SPRINGS OF CONDUCT,” “HABIT AND INSTINCT,” “PSYCHOLOGY FOR TEACHERS,” ETC. ETC. ILLUSTRATED SECOND EDITION THIRD IMPRESSION LONDON EDWARD ARNOLD 1920 ( All rights reserved ) My book on “Animal Life and Intelligence” being out of print, I undertook to revise it for a new Edition. As the work of revision proceeded, however, it appeared that the amended treatment would not fall conveniently under the previous scheme of arrangement. I therefore decided to wr
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I.—Behaviour in General
I.—Behaviour in General
We commonly use the word “behaviour” with a wide range of meaning. We speak of the behaviour of troops in the field, of the prisoner at the bar, of a dandy in the ball-room. But the chemist and the physicist often speak of the behaviour of atoms and molecules, or that of a gas under changing conditions of temperature and pressure. The geologist tells us that a glacier behaves in many respects like a river, and discusses how the crust of the earth behaves under the stresses to which it is subject
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II.—Behaviour of Cells
II.—Behaviour of Cells
From what has already been said it may be inferred that our use of the term “behaviour” neither implies nor excludes the presence of consciousness. Few are prepared to contend that the iron filings in a magnetic field consciously group themselves in definite and symmetrical patterns, or that sand grains on a vibrating plate assemble along certain nodal lines because they are conscious of the effects of the bow by which the plate is set in sounding vibration. But where organic response falls unde
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III. Corporate Behaviour
III. Corporate Behaviour
The word “corporate” is here applied to the organic behaviour of cells when they are not independent and free, but are incorporated in the animal body, and act in relation to each other. If the behaviour of the individual cell during division impresses us with the subtle intricacy of organic processes, the behaviour of the growing cell-republic during the early stages of organic development must impress us no less forcibly. We place the fertilized egg of a hen in an incubator, and supply the req
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IV.—The Behaviour of Plants
IV.—The Behaviour of Plants
A short parenthetic section on the behaviour of plants may serve further to illustrate the nature of organic behaviour. We have seen that Paramecium is apparently attracted by faintly acid solutions, and have briefly considered Dr. Jennings’s interpretation of the facts disclosed by careful observation. In the ferns the female element, or ovum, is contained in a minute flask-shaped structure (archegonium), in the neck and mouth of which mucilaginous matter, with a slightly acid reaction, is deve
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V.—Reflex Action
V.—Reflex Action
It is sometimes said that the tentacles of the Sun-dew leaf indicate a primitive kind of reflex action in plants, and that they afford evidence of discrimination. “It is,” says Romanes, “the stimulus supplied by continuous pressure that is so delicately perceived, while the stimulus supplied by impact is disregarded.” [11] And, comparing this with what is observed in the Venus’s Fly Trap, he says: “In these two plants the power of discriminating between these two kinds of stimuli has been develo
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VI.—The Evolution of Organic Behaviour
VI.—The Evolution of Organic Behaviour
The interpretation of organic behaviour in terms of evolution mainly depends on the answer we give to the question: Are acquired modes of behaviour inherited? A negative answer to this question is here provisionally accepted. But the premisses from which this conclusion is drawn are too technical for discussion in these pages. It must suffice to state as briefly as possible what this conclusion amounts to, and to indicate some of the consequences which follow from its acceptance. The fertilized
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I.—The Conscious Accompaniments of Certain Organic Changes
I.—The Conscious Accompaniments of Certain Organic Changes
It is possible that all organic behaviour is accompanied by consciousness. But there is no direct means of ascertaining whether it is so or not. This is, and must remain, a matter of more or less plausible conjecture. We have, indeed, no direct knowledge of any consciousness save our own. Undue stress should not, however, be laid on this fundamental isolation of the individual mind. We confidently infer that our fellow-men are conscious, because they are in all essential respects like us, and be
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II.—The Early Stages of Mental Development
II.—The Early Stages of Mental Development
We use the phrase “mental development” in its broadest acceptation as inclusive of, and applicable to, all phases of effective consciousness. We shall assume that throughout this development there is a concomitant development of nerve-centres and of their organic connections. And we shall further assume that experience, as such, is not inherited. The nature of the grounds on which the latter assumption is based must first be briefly indicated. It is commonly asserted that fear of man, the invete
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III.—Later Phases in Mental Development
III.—Later Phases in Mental Development
Some surprise may be felt that in our brief discussion of the early stages of mental development nothing has been said of percepts and concepts, nothing of abstraction or generalization. The omission is not only due to a desire to avoid the subtle technicalities of psychological nomenclature. It is partly due to the wish not to forejudge a difficult question of interpretation. Spirited passages of arms from time to time take place between psychologists in opposing camps, as to whether animals ar
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IV.—The Evolution of Consciousness
IV.—The Evolution of Consciousness
The origin of consciousness, like that of matter or energy, appears to be beyond the pale of scientific discussion. The appearance of effective consciousness on the scene of life does indeed seem to justify the belief in the prior existence of sentience as the mere accompaniment of organic behaviour. Ex nihilo nihil fit. And since effective consciousness must, on this principle, be developed from something, it is reasonable to assume that this something is pre-existing sentience. Again, we may a
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I.—Definition of Instinctive Behaviour
I.—Definition of Instinctive Behaviour
There are probably few subjects which have afforded more material for wonder and pious admiration than the instinctive endowments of animals. “I look upon instinct,” wrote Addison in one of his graceful essays, “as upon the principle of gravitation in bodies, which is not to be explained by any known qualities inherent in the bodies themselves, nor from any laws of mechanism, but as an immediate impression from the first Mover and the Divine Energy acting in the creatures.” [21] In like manner S
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II.—Instinctive Behaviour in Insects
II.—Instinctive Behaviour in Insects
Since instinctive behaviour is, by definition, independent of experience, and since the animals which act instinctively are also, in many cases, able to act intelligently, it is clear that, apart from hereditary variations, we must expect to find acquired modifications of instinct. As Huber said of bees, their instinctive procedure often indicates “a little dose of judgment.” It is, indeed, exceedingly difficult, as a matter of observation, to distinguish between hereditary variation and acquire
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III.—The Instinctive Behaviour of Young Birds
III.—The Instinctive Behaviour of Young Birds
Since it is easy to hatch birds of many species in an incubator, and to rear them under conditions which not only afford facilities for observation but exclude parental influence, their study has special advantages. One can with some approach to accuracy distinguish the instinctive from the acquired factors in their behaviour. [34] Fig. 15. —Newly-hatched Chick swimming. The callow young of such birds as pigeons, jays, and thrushes are hatched in a helpless condition, and require constant and as
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IV.—The Conscious Aspect of Instinctive Behaviour
IV.—The Conscious Aspect of Instinctive Behaviour
In our definition of instinctive behaviour all positive reference to the presence of conscious states was omitted. By some writers, however, the fact that it is accompanied by consciousness is regarded as a distinguishing feature of instinct. Romanes introduced his definition with the words: [38] “Instinct is reflex action into which there is imported the element of consciousness.” And he emphasized the conscious aspect when he said: “The term comprises all those faculties of mind which are conc
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V.—The Evolution of Instinctive Behaviour
V.—The Evolution of Instinctive Behaviour
It may be assumed that the fact of evolution is generally admitted. The question of its method is, however, still open to discussion. It is possible that, as some biologists contend, there is an inherent tendency in organic beings to evolve in certain definite directions independently of their relation to the environment. But it is scarcely probable that instinctive behaviour is mainly due to any such inherent tendency—of the nature of which in any case we know but little. Setting this on one si
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I.—The Nature of Intelligent Behaviour
I.—The Nature of Intelligent Behaviour
Such an animal as a newly hatched bird or an insect just set free from the chrysalis is a going concern, a living creature. It is the bearer of wonderfully complex automatic machinery, capable, under the initiating influence of stimuli, of performing instinctive acts. But if this were all we should have no more than a cunningly wrought and self-developing automatic machine. What the creature does instinctively at first it would do always, perhaps a little more smoothly as the organic mechanism s
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II.—Intelligent Behaviour in Insects
II.—Intelligent Behaviour in Insects
It is, as we have seen, among the higher invertebrates, especially in insects, that some of the most remarkable and complex instincts may be found. There is, [50] however, a tendency to ascribe the behaviour of insects entirely to instinct, without sufficient evidence that neither imitation, instruction, nor intelligent learning play any part. This is, perhaps, a survival of the old-fashioned view that all the acts of the lower animals are performed from instinct, whereas those of human beings a
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III.—Some Results of Experiment
III.—Some Results of Experiment
It is unnecessary to give a resumé of entertaining anecdotes illustrative of intelligent behaviour in the higher animals. Such anecdotes are too often the outcome of casual observations by untrained observers; and the interpretation put upon the facts frequently shows a want of psychological discrimination. Such is not the material of which science is constituted. What is needed is systematic observation conducted, so far as possible, under controlled conditions. Two things are necessary: first,
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IV.—The Evolution of Intelligent Behaviour
IV.—The Evolution of Intelligent Behaviour
No attempt can be made in this section to trace the successive stages of the evolutionary progress of intelligence from its lower to its higher developments. It is indeed questionable whether comparative psychology has, as yet, accumulated a sufficient body of data to render such a task profitable or even possible. And the lower the level of intelligence with which we have to deal, the less reliable are the scanty psychological data which we can obtain. To interpret the mental processes which ac
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V.—The Influence of Intelligence on Instinct
V.—The Influence of Intelligence on Instinct
We have seen that the relation of instinct to intelligence is essentially that of congenital to acquired behaviour. We have seen, too, that in the Lamarckian interpretation what is acquired in the course of life may be transmitted through inheritance, and thus the intelligent behaviour of one generation may become instinctive and congenital in the next. But serious biological difficulties stand in the way of the acceptance of this interpretation; there is, moreover, little or no evidence of the
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I.—Imitation
I.—Imitation
The characteristic feature of social behaviour is that it is in large degree determined by the behaviour of other members of the social community. In all animals which mate there is a temporary or more lasting influence on each other of the individuals which unite to procreate their kind; and in those which foster their young there is a social relation of parents and offspring. Some of these mutual relationships will be discussed, in their emotional aspects, in the next chapter. Here we will con
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II.—Intercommunication
II.—Intercommunication
The foundations of intercommunication, like those of imitation, are laid in certain instinctive modes of response, which are stimulated by the acts of other animals of the same social group. These have been fostered by natural selection as a means of social linkage furthering the preservation, both of the individual and of the group. Some account has already been given of the sounds made by young birds, which seem to be instinctive and to afford an index of the emotional state at the time of utt
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III.—Social Communities of Bees and Ants
III.—Social Communities of Bees and Ants
Apart from human societies the most noteworthy social communities of animals are found among insects, especially in ants, bees, wasps, and termites. It is true that in the mammalia we find such communities as the troop of apes, the herd of cattle, the pack of wolves, the school of porpoises, the so-called “rookeries” of seals, and the colonies of “prairie dogs” and of beavers; and that among birds there are analogous communities. Undoubtedly the temporary or permanent association of many individ
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IV.—Animal Tradition
IV.—Animal Tradition
In that interaction between instinct and intelligence which, when further detailed work has sifted and purified our knowledge of the psychology of animal communities, may prove sufficient to account for the well-established facts, animal tradition will probably have to be recognized as of no little importance. When a newly emerging ant or bee, or a young bird or mammal is born into a community where certain modes of behaviour are already in full swing, an imitative tendency of the follow-my-lead
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V.—The Evolution of Social Behaviour
V.—The Evolution of Social Behaviour
“Animals of many kinds,” said Darwin, [108] “are social; every one must have noticed how miserable horses, dogs, sheep, etc., are when separated from their companions. The most common mutual service in the higher animals is to warn one another of danger. Every sportsman knows how difficult it is to approach animals in a herd or troop. Wild horses and cattle do not, I believe, make any danger-signal; but the attitude of any one of them who first discovers an enemy warns the others. Rabbits stamp
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I.—Impulse, Interest, and Emotion
I.—Impulse, Interest, and Emotion
Any discussion of animal behaviour must deal largely with what is termed the conative aspect of consciousness. “The states designated by such words as craving , longing , yearning , endeavour , effort , desire , wish , and will ,” says Dr. Stout, in his admirable “Manual of Psychology,” [117] “have one characteristic in common. In all of these there is an inherent tendency to pass beyond themselves and become something different. This tendency is not only a fact, but an experience; and the pecul
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II.—Play
II.—Play
“There are two quite different popular ideas of play,” says Professor Groos, in his admirable work on “The Play of Animals.” “The first is that the animal (or man) begins to play when he feels particularly cheerful, healthy, and strong; the second that the play of young animals serves to fit them for the tasks of later life.” The former view, in which the latter may be included incidentally as a result, is closely associated with the names of Schiller, [123] who suggested it, and of Mr. Herbert
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III.—Courtship
III.—Courtship
We have seen that Professor Groos regards play as the practice and preparation for the serious business of animal life. Founded on instinctive tendencies, it has its biological value in the acquisition of practical acquaintance with the environment, and of skill in dealing with it effectually. It is an education in behaviour of the utmost service in view of the struggle for existence. It is full of the pleasure derived from the satisfaction of innate impulse, the success of conative effort, and
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IV.—Animal “Æsthetics” and “Ethics”
IV.—Animal “Æsthetics” and “Ethics”
In this section we shall consider some types of behaviour which suggest situations that contain the germs of æsthetics and ethics, with a view to determining, so far as possible, the principles on which they should be interpreted. This is a peculiarly difficult subject; for we are endeavouring to get behind the behaviour, and to infer the mental conditions which accompany it, and through which it assumes its distinctive character. The difficulty is twofold: first, because, as Dr. Stout puts it,
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V.—The Evolution of Feeling and Emotion
V.—The Evolution of Feeling and Emotion
“Whatever conditions,” says Dr. Stout, [175] “further and favour conation in the attainment of its end, yield pleasure. Whatever conditions obstruct conation in the attainment of its end, are sources of displeasure. This is the widest generalization which we can frame, from a purely psychological point of view, as regards the conditions of pleasure and displeasure respectively.” Here Dr. Stout seems carefully to avoid the commonly accepted and much advertised conclusion, that pleasure and pain (
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I.—The Physiological Aspect
I.—The Physiological Aspect
At the outset of our inquiry, we used the word “behaviour” in a wide and comprehensive sense. Thus broadly used, I said, the term in all cases indicates and draws attention to the reaction of that which we speak of as behaving in response to certain surrounding forces or circumstances which evoke the behaviour. The behaviour of living cells is dependent on changes in their environment; that of deciduous trees, as they put forth their leaves in the spring or shed them in the autumn, is related to
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II.—The Biological Aspect
II.—The Biological Aspect
The biological aspect of behaviour—its relation to biological ends—has so often come under our consideration in the foregoing chapters that little need be added in this section: and that little may be most appropriately devoted, first to the question whether consciousness does influence behaviour; and secondly, this being accepted, to the importance of the rôle that is played by the development of conscious situations in securing, in the higher animals, the biological end of racial preservation.
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III.—The Psychological Aspect
III.—The Psychological Aspect
On the hypothesis of monism, the nature of which, so far as it bears on our inquiry, was briefly indicated in the foregoing section, the conscious situation is the psychical or mental expression of that which for the physiologist is what we may term a neural situation. As such it does not enter into the chain of physical causation; nor do physical events as such—that is to say, save as experienced—enter into the chain of mental causation. For mental development they have no independent existence
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IV.—Continuity in Evolution
IV.—Continuity in Evolution
Under the head of organic behaviour, in the widest acceptation of the term, fall the whole of physiology, the whole of embryological development, nay, more, the whole of organic evolution; while mental evolution, in all its stages, may be regarded as the psychological aspect of that which, from the physiological aspect, is the evolution of nervous systems. Life itself is the behaviour of a particular kind of substance which is found more or less abundantly under natural conditions. No other know
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