Konrad Lorenz (1981) - The Foundations of Ethology

Mr. President, ladies and gentlemen. I first intended to give this speech in English. Then, on second thought, I decided to speak German, which is easier to me and easier for the majority of my audience, but on third thought, I decided to speak English after all. Most of the younger German understand English anyhow, and I know that as a German one understands English particularly well, if it is badly spoken by another German. This is an excuse to my undemocratic procedure. What I am going to talk about is a very banal story. I wanted to talk about ethology, ‘Vergleichende Verhaltensforschung’. Ethology is extremely easy to define. The definition is simply the science in which the same approaches and the same methods are used with behaviour as a subject, as are used in all other kinds of biological sciences since the times of my old friend Charlie Darwin. Charles Darwin knew all about ethology and, you know you can always claim Darwin as the first of your science, in biological sciences, because he started the hare in so very many of them. And if you read Charles Darwin’s book on the expression of emotions, in man and animals, you find practically between the lines, or even clearly outspoken, practically everything we are talking about and puzzling about nowadays. The story of ethology is interesting from the historical point of view because it is so obvious that animal and human behaviour is a life function, a function of life. So it seems to me a matter of course that we approach it as any other life forces. Why did that not happen until about 1910, very late after Darwin. And the reason of this is purely historical. It is the outcome of a contention between two great schools of psychology. The purposive psychologists on one side, McDougall, William McDougall, Edward Chace Tolman being the most important representatives. And by the behaviourists, Wilson, Jörgs and so on on the other side. This contention was a grudge fight on both sides because they fought for their philosophies, for their ‘Weltanschauungen’. And the purposive psychologists contended that instinct was a preternatural factor. Neither in need of, nor accessible to a natural explanation. The instinct told the animal what to do. It taught, and also they equated the purpose, hence the term purposive psychologists, they equated the purpose followed by the animal or by man, following the urges of an instinct, was pursuing the purpose of survival. The species preserving function of the action. And this is an obvious error because the young man courting a girl is certainly not aiming or purposing the baby, nor does any courting animal do the same thing. So the equation of purpose of the teleonomy or the action and the purpose pursued by the subject is obviously wrong. That’s our fight with McDougall. Otherwise the purposive psychologists knew a lot about animals. And particularly McDougall stressed the important fact that animal behaviour was spontaneous. His slogan, the healthy animal is up and doing, is true, that’s indeed a fact. On the other hand they were clearly vitalists and the introduction of an ununderstandable factor is clearly unscientific. Also they were actually averse to experiments. Now, the behaviourists reacted clearly to the error of the purposivists and the opposite of an error is not truth but the opposite error. And both the purposivist and the behaviourists took positions which none of them would ever have taken, if they hadn’t known of the opposite position of the antagonist, like people engaging in a tug-of-war. If the innate, what Eigen would call the instruction of the genome did not exist, was for one side, for the purposivist a preternatural god, ‘instinct’ spelt with a capital I, then the behaviourists simply denied its existence. They were so averse to teleology that they even did not want to concede the species preserving structures and behaviour patterns of animals. The questions ‘what for?’, ‘wozu?’ was an anatoma to them. And this is why they put so much value on the blind experiment, the experiment which was not put with a question but just to see what happens. On the point that they wanted to turn the behaviour study into an experimental animal, in an experimental research, they were absolutely right. And their choice of object was the conditioned response, the learning process, the reflex was very important and taken as the most important explanatory principle in physiology and the conditioned reflex by Pavlov was just new and it was very obvious to choose the conditioning of a reflex, the conditioned reflex as the main object of experiment, because it is indeed very accessible to experimentation. The mistake of which we accuse behaviourists is not what they did. What they did is glorious. The investigation of the conditioned response and particularly by the conditioning by reward, by reinforcement has the greatest possible merit. But what we accuse the behaviourist of is what they do not do. Very many file of animals, independently of each other, have developed a mechanism in which the success of some behaviour is fed back on the present behaviour. Encouraging the animal to repeat that or saying don’t do that, that was bad. This feedback was invented, I say “invented”, by practically all file of animal which achieved a central nervous system. This one apparatus is very much the same, in the pigeon, in an octopus, in a rat, in a man. But investigating nothing else but this apparatus, you leave out of regard everything that makes a man a man, makes a rat a rat, an octopus an octopus and so on. Now, this attitude of extreme experimentalism would never have been assumed by the behaviourists if they hadn’t known about the absolute nonsense the purposivists were talking about with their great preternatural instinct. And it is what characteristic that in this time it was not psychologists, nor people particularly concerned with behaviour, but plain zoologists, “Viechzoologen”, who made the important discovery. And independently of each other, Charles Otis Whitman in America and Oskar Heinroth in Germany. Oskar Heinroth didn’t know of Whitman. At the 1932 Ornithological Congress in Oxford Margaret Morse Nice, an American ornithologist told me about the existence of Whitman and particularly of his pupil Wallace Craig, who became one of my most important teachers. Now, what was the great discovery, which I consider as the archimedic point from which ethology, ‘Vergleichende Verhaltensforschung’ took its origin. These people were systematicists, they were investigating the relationships between closely related species. And in doing this you must collect characters, characters, characteristic of subspecies, species, subspecies and other characters more generalities among genesis. And this is the typical crazy zoologist, the man you find sitting with a microscope and counting filopods or counting the veins in a little fly’s wings, the typical crazy funny people professor. Now, what these people really do is preparing the foundation for building the great bridge from genetics to phylogenetics. From phylogeny, from evolution to genetics. And it was Erwin Stresemann and particularly his pupil Ernst Mayr who used the tremendous material stored up in musea by what we call bird maniacs, “Ornithomanen”, for entirely different purposes, from the urge of collecting, and in particular Stresemann was the first to use this material from the evolutionary point of view. And this search for characters, for more and more characters, the closer that two species are aligned, I have not the time to go into this, the more characters you need to ascertain their blood relationship. And in their avidity, their avid search, their need for characters, they hit on behaviour characters. And they found out that there are motor patterns, particularly motor patterns of courtship, which are as reliable guides to blood relationship, as reliable documents of phylogeny as are the form of teeth, so much preferred by mammalogists or the number of vertebrate in ethology and so on. And if you read the primitive naïve papers of these people, you find that they say ‘this species has, does the species have the grunt-whistle or the head and tail lift or any, as if this grunt-whistle or this head and tail lift was a physiological, a morphological character. This discovery could only have been made by people who were amateurs, who were dilettante. And the man who doesn’t take pleasure in his object, even if he were a Tibetan Lama, he wouldn’t have the patience to sit long enough staring in an aquarium or on a bond with ducks as is necessary to see these things. Once the discovery was made, the use of behaviour patterns as systematic characters began rapidly to spread and today, if you come to an ornithological congress, you hear quite a lot about it. Now, neither Whitman nor Heinroth ever said a word about the physiology of this particular kind of movements. But even without theorising on their physiological causality, they used only a certain kind of motor pattern of which we now know quite something physiologically. One character of this behaviour patterns is that they can differ in intensity. One might say the opposite of an all or nothing rule prevails for them. The same motor pattern can be performed in a slight hint, in a slight suggestion of the motor pattern. And if you measure it, you find that the relationship between phase and amplitudes of excursion remain constant. And that way you recognise the motor pattern, even at the slight suggestion, exactly as you can recognise a melody played very, very softly. From these intention movements there is a continuity of gradations to the motor pattern in which the real teleonomic biological, before a function of the movement is performed. Very many of these motor patterns respond to a highly specific, simple stimulus situation. If you put a shining flat mirror or simply a window pane before a young falcon or young …., it will sit on it and begin to perform bathing movements. Or if you show a stickleback a crude dummy which has nothing in common with another real stickleback but the red on its undersurface, it will start to fight it. As long as this very specific stimulus situation, which sets off the motor pattern was regarded as part of the motor pattern itself, nothing oblige us to pay particular attention to its physiology, it simply was the first of a chain of reflexes. Very many things in very many characters of these behaviour patterns can tell us, could tell us at that early moment that they are not reflexes. And at a very early stage Wallace Craig, who became one of my most important teachers, wrote to me in a letter: but isn’t it nonsense to speak of a reaction to a stimulus not yet received?’ Because all these activities are spontaneous. And if the activity is not released for some time, its threshold decreases, ‘Schwellenerniedrigung’. It decreases more and more and it decreases in the end to the extent of letting the motor pattern explode in vacuum. This was well-known as early as before 1900 to Wallace Craig, who experimented with a blond ringdove, male, which would normally perform its “coo boo and coo boo” only to the female of the species after having been kept in isolation, deprived of stimulation, it would perform to a domestic pigeon, after some more damming up of the reaction, it would respond to the fist of the experimentor and at last it would perform to the corner where the reliance of the box converged. So there is still some point of reference to which the motor pattern is addressed. This is called vacuum activities, we call it ‘Leerlaufreaktion’. The word was invented by my friend Bernhard Hellmann when we were 17 years old and it is called, not from a physical parable but from the motorcycle, ‘auf Leerlauf’, in neutral, letting the motor run in neutral, Now, all of these facts about threshold lowering, running off in vacuum and, last but not least, appetitive behaviour are quite inexplicable on the basis of reflex theory. I have not yet mentioned appetitive behaviour because it’s not my child, it was something to which Wallace Craig drew attention, and that is the fact that if you deprive the animal for some time of the necessary key stimuli, normally releasing this motor pattern, it will not only react to lesser stimuli, but it will become restless and begin to search actively for the stimuli which specifically – this restlessness, this searching in its simple case is simply a random movement which still increases the probability of finding the stimuli, but in more complicated cases it is a very well-directed search and all conditioning, all learning by reward is done in the course of what Wallace Craig has called appetitive behaviour. So it’s interesting to know what the animal is striving for. If you want to investigate its learning processes. So appetitive behaviour, finding of the key stimuli, releasing of an innate releasing mechanism and the following discharge of the consummatory act seem to be the unit of animal behaviour. Heinroth called it ‘arteigene Triebhandlungen’ and very early after I had given a conference on this in the Harnack-Haus in the Max Planck Gesellschaft that was my first debut with Max Planck. Critic: Charlotte Kogon said that it aught not to be ‘Instinkthandlung’ or ‘Triebhandlung’, but ‘Instinktbewegung’, instinctive movement. And shortly after this meeting in Max Planck we had a meeting under Van der Klaauw in Leiden, and there we discussed the elements of what Heinroth called ‘arteigene Triebhandlung’, and at this time we were already so convinced that the spontaneity of this movement was due to internal impulse production which Erich von Holst had demonstrated. That we were very clear about the question that the mechanism receiving stimuli, the filter which only passes very specific symbolition on to the releasing of the action was a process entirely different from the following consummatory act which clearly was akin to Erich von Holst’s endogenously produced and centrally coordinated motor patterns. And we do not know who first, whether it was Tinbergen there or I, who first used the term innate releasing mechanism, it was first called ‘angeborenes Schema’, but then we realised that it was not at all like a diagram but the summation of stimuli, question to which I needn’t go in here. Now, this whole story of elements which constitute innately programmed motor patterns has led to a lot of research and particularly Tinbergen and Tinbergen’s pupils experimented on the releasing mechanism about which we know something. But on the other hand the physiological properties of the endogenous impulses have not been really investigated since Erich von Holst died, and none of his pupils has really – they are so much dependent on what we know about the spontaneity, we know of the spontaneity of complex behaviour but the basis, that’s why my today’s lecture has the title of fundaments of ethology, the physiological basis of endogenous impulse production and of endogenous coordination has not really been experimentally pursued ever since. I want to say two words about man. About our attitude to human beings. We ethologists are very often accused of regarding human beings as animals, of underassessing the difference between man and animals, and that is a very unjust accusation. Man is a very particular creature because man has, and it was already Julian Huxley who drew attention to this fact, man possesses a new hereditary apparatus. It is an error to believe that evolution always proceeds gradually. Evolution can perform very sudden movements, which mostly consist in two pre-existent systems being united in one system. And you can show even on the basis of electronic apparatus that the fusion of two systems may produce a new system with entirely new and hardly predicable systemic properties. And it is my conviction that man has made the jump from the non-reasonable to the reasonable being by such a confluence of pre-existing mechanisms, some of which we know very well. One of them is Gestalt - perception, perception is able of processes of real abstraction, you abstract the colour which you call a shirt, the shirt is white, is really the process of a complicated calculation in which the colour of the illumination is put in reference with the colour actually reflected at the moment by the object. And it is the object’s colour reflecting preferences which we call ‘its colour’. And this abstraction is a thing which, even in this abstractive apparatus, is able to perform, which obviously is evolved in the service of recognising things, individual things, can do more, it goes a lot of way to forming generic concepts. If I see my dog from far, from near, from behind, from in front, in red light, in green light and recognise that it is also the same dog, this is Dingkonstanz, but if I recognise a property of doggishness in this dog, in the name of poodle and so on, then this is already something very close to abstraction. And another function which comes very close to abstraction is the representation of space. Of which the chimpanzee is already so well able, the chimpanzee, which Professor Eccles has described, solves the problem of the banana to be reached by the box without moving, moving only the eyes, he looks to the banana, he looks to the box, he looks from the box to the point under the banana and up to the banana and then he has the solution. And all these - I won’t go into details - but some other faculties which are already present in animals, but in none so much as in human beings. Fuse together gives something like conceptual thought, exploratory behaviour has a very important role and this exploration consists in activity which is independent from any special drive. The raven, exploring a new object, sees immediately when it gets really hungry, but he plays the whole repertoire of his activities on any new object, unknown object which you offer him, and it is not that he wants to eat but he wants to know whether this is in principle edible. And this kind of exploratory behaviour, which is very well developed in rats and in very many non specifically adapted animals, is also a specific property of man. And all these things work together to make conceptual thought and with it syntactic language. And from the first prebiotic life-like living being up to the most recent ancestors of ourselves, the genome was the main apparatus, the hoary method of mutation and selection was the only method of gathering information. And quite suddenly in the late tertiary period they developed something new. A new apparatus which does the same thing but much faster. And if a modern man were asked to define life, I am quite sure that he would pass over in his definition the double helix discovered by Crick and Watson who are both here. And this would pass over, something very specific of human life, this very fast, new method of gathering information. And whether this new apparatus will prove a benefit or curse is something which the young people will have to decide. Thank you.

Konrad Lorenz (1981)

The Foundations of Ethology

Konrad Lorenz (1981)

The Foundations of Ethology

Comment

Konrad Lorenz lectured three times at the Lindau Meetings and the present lecture is the last of the three. Lorenz had a close working relationship with his co-bird watcher Nikolaas Tinbergen, who was in the audience. Together they made up a winning team (with Karl von Frisch working more independently with bees). In this team, Lorenz in some sense played the role of the theoretician and Tinbergen the role of the experimentalist. This does not mean that Lorenz did his research work at his desk with Tinbergen roaming around the countryside. Both spent considerable time performing the classical work of the ethologist, “watching and wondering”. But it was Tinbergen who experimentally tested many of the ideas that Lorenz got by watching animals. In his lecture, Lorenz first presented a concise history of the science that he himself gave the name “ethology”. In his opinion ethology could have become an established research field much earlier, since already Charles Darwin had understood the foundations. But progress was hampered by conflicts between vitalists and mechanists and the deadlock was not broken until the early 1900’s, by men such as Charles Whitman and Oskar Heinroth, before Lorenz himself came onto the scene. In the second part of the lecture, Lorenz decribes some of the basic concepts used in ethology and at the very end he generously hands over some questions to the younger generation.

Anders Bárány

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