The Foundations of Ethology
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Transcript: English(auto-generated)
00:12
Mr. President, ladies and gentlemen, I first intended to give this speech in English.
00:22
Then, on second thought, I decided to speak German, which is easy to me and easy to the majority of my audience. But on third thought, I decided to speak English after all. Most of the younger German understand English
00:42
anyhow. And I know that, as a German, one understands English particularly well if it is badly spoken by another German. So this is an excuse to my undemocratic procedure.
01:02
What I'm going to talk about is a very banal story. I wanted to talk about ethology. Vergleich in der Verheitensforschung. Ethology is extremely easy to define. The definition is simply the science
01:21
in which the same approaches and the same methods are used with behavior as a subject, as are used in all other kinds of biological sciences, since the times of my old friend Charlie with Charlie Darwin.
01:42
Charles Darwin knew all about ethology. And you can always claim Darwin as the first of your science in biological sciences because he started the hair off in so very many of them. And if you read Charles Darwin's book
02:04
on the expression of emotions in man and animals, you find practically between the lines or even clearly outspoken all practically everything we are talking about and puzzling about nowadays.
02:23
The story of ethology is interesting from the historical point of view because it is so obvious that animal and human behavior is a life function, a function of life. So it seems to be a matter, of course,
02:42
that we approach it as any other life process. Why did not that happen until about 1910, very late after Darwin? And the reason of this is purely historical. It is the outcome of a contention
03:03
between two great schools of psychology, the purposive psychologists on one side, MacDougall, William MacDougall, Edward Chase Tolman, Tolman being the most important representatives,
03:20
and by the behaviorists, Wilson, Yerkes, and so on on the other side. This contention lasted, was a grudge fight on both sides because they fought for their philosophies, for their Weltanschauern.
03:41
And the purposive psychologists contended that instinct was a preternatural factor, neither in need of, nor accessibly to a natural explanation. The instinct taught the animal what to do.
04:02
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,
04:22
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 as a baby, rabbe, rabbe, nor does any courting animal
04:44
do the same thing. So the equation of purpose of the teleonomy of the action and the purpose pursued by the subject is obviously wrong. That's our fight with MacDougall. Otherwise, the purposive psychologists
05:02
knew a lot about animals. And particularly, MacDougall stressed the important fact that animal behavior was spontaneous. His slogan, the healthy animal is up and doing, is true. That's indeed the fact.
05:21
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 behaviorists reacted clearly
05:41
to the error of the purposivists. And the opposite of an error is not truth, but the opposite error. And both the purposivists and the behaviorists took positions which none of them would ever have taken.
06:02
If they hadn't known of the opposite position of the antagonist, like people engaging in a terror war, if the innate, what Eigen would call the instruction of the genome, did not exist, was for one side,
06:24
for the purposivists, a preternatural god, instinct spelled with a capital I, then the behaviorists simply denied its existence. They were so averse to teleology
06:44
that they even did not want to concede the species preserving structures and behavior patterns of animals. The questions, what for, what to, was anathema to them.
07:05
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.
07:21
On the point that they wanted to turn the behavior 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.
07:41
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,
08:05
as the main object of experiment, because it is indeed very accessible to experimentation. The mistake of which we excuse behaviorists is not. What they did.
08:20
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 behaviorists of
08:40
is what they do not do. Very many filers of animals have developed independently of each other, have developed a mechanism in which the success of some behavior is fed back on the present behavior, encouraging the animal to repeat that or saying,
09:00
fu, fu, don't do that. That was bad. This feedback was invented, I say, quote, invented, unquote, by practically all filer of animal which achieved a central nervous system. This one apparatus is very much the same in a pigeon and octopus in a rat in a man.
09:24
But in investigating nothing else but this apparatus, you leave out of regard everything that makes a man, makes a rat a rat, an octopus an octopus, and so on. Now, this attitude of extreme experimentalism
09:40
would never have been assumed by the behaviorists if they hadn't known about the absolute nonsense the proposalists were talking about with their great preternatural instinct. And it is quite characteristic that in this time,
10:00
it was not psychologists nor people particularly concerned with behavior, but plain zoologists, speak so long, who made the important discovery and independently of each other, Charles Otis Whitman in America and Oskar Heinrod in Germany.
10:24
Oskar Heinrod didn't know of Whitman. At the 1930s, on his logical Congress in Oxford, 1932, sorry, on his logical Congress in Oxford, Margaret Moore Snice, an American ornithologist,
10:40
told me about the existence of Whitman, and particularly of his pupil, Wallace Greig, who became one of my most important teachers. Now, what was the great discovery which I consider as the archimedic point from which ethology, Vergleiche, and the Weidensforschung
11:00
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,
11:22
and other characters more generally distributed among genuses. And this is the typical crazy zoologist, the man you find sitting at a microscope and counting the kitty of philippodes or counting the veins in a little fly's wing,
11:45
the typical crazy, funny paper professor. Now, what these people really do is preparing the foundation for building the great bridge from genetics to phylogenetics,
12:02
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 musia by what we call bird maniacs,
12:26
Orny Tomainen, for entirely different purposes from the urge of collecting. And particularly Stresemann was the first to use this material from the evolutionary point of view.
12:44
And this search for characters, for more and more characters, the closer two species are allied. I have not the time to go into this. The more characters you need to ascertain their blood relationship. And in the avidity, in the avid search,
13:04
the need for characters, they hit on behavior characters. And they found out that there are motor patterns, particularly motor patterns of courtship, which are as reliable guides to blood relationship,
13:21
as reliable documents of phylogeny, as are the form of teeth, so much preferred by mammalologists, or the number of vertebrates in ecologists, and so on. And if you read the primitive, naive papers of these people,
13:44
you'll find that they say this species has. Does this species have the grunt whistle or the head of tail? As if this grunt whistle or this head of tail was a morphological character.
14:04
This discovery could only have been made by people who were amateurs, who were dilettante. Dilettante means take pleasure in. And a man who doesn't take pleasure in his object,
14:22
even if he were a dependent llama, he wouldn't have the patience to sit long enough, staring into an aquarium or on a pond with ducks, as is necessary to see these things. Once the discovery was made, the use of behavior patterns
14:42
as morphological, as systematic characters, began rapidly to spread. And today, if you come to an ontological Congress, you'll hear quite a lot about it. Now, neither Whitman nor Heinold ever said a word about the physiology
15:02
of these particular kind of movements. But even without theorizing on their physiological causality, they used only a certain kind of motor pattern of which we now know quite something physiologically.
15:25
One character of these behavior 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,
15:45
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.
16:01
And that way, you recognize the motor pattern, even at the slight suggestion, such exactly as you can recognize a melody played very, very softly. From these intention movements, there is a continuity of gradations to the motor pattern
16:22
in which the real teleonomic biological function of the movement is performed. Heinold, most of these motor patterns, not most, very many of these motor patterns respond to a highly specific simple stimulus situation.
16:45
If you put a shining flat mirror or windowpane before a young falcon, a young cernice, it will sit on it and begin to perform bathing movements.
17:03
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,
17:23
which sets off the motor pattern, was regarded as part of the motor pattern itself, nothing obliged us to pay particular attention to its physiology.
17:40
It simply was the first of a chain of reflexes. Very many things in very many characters of these behavior patterns can tell us, could tell us at the early moment that they are not reflexes. And at a very early stage, Wallace Craig,
18:02
who became one of my most important teachers, wrote to me in a letter, now look, you persist in this being chain reflexes, but isn't it nonsense to speak of a reaction to a stimulus not yet received?
18:21
Because all these activities are spontaneous. And if the activity is not released for some time, its threshold decreases. Schwellner niederung. It decreases more and more, and it decreases in the end to the extent of letting the motor pattern explode in vehicle.
18:47
This was well known as early as before 1900 to Wallace Craig, who experimented with a blonde ringed-off male, which would normally
19:00
perform its coo-boo and coo-boo-boo-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 experimenter.
19:24
And at last, it would perform to the corner where the three lines of the box converge. So there's still some point of reference to which the motor pattern is addressed. This is called vacuum activities.
19:43
We call it Leherlauf reaction. The board was invented by my friend, Bernard Hellman, when we were 17 years old. And it is colored not from a physical parable, but from the motorcycle of Leherlauf in neutral, letting the motor run in neutral of Leherlauf.
20:03
That was the origin of the name. Now, all these facts about threshold lowering, running off in a vacuum, and last but not least, appetitive behavior are quite inexplicable on the basis
20:22
of reflex theory. I have not yet mentioned appetitive behavior, 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,
20:44
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
21:02
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.
21:21
And all conditioning, all learning by reward is done in the course of what Wallace Craig has called appetitive behavior. So it's interesting to know what the animal is striving for if you want to investigate
21:42
its learning processes. So appetitive behavior, finding of the key stimuli, releasing of an innate releasing mechanism, and the following discharge of the consummatory act seemed to be the unit of animal behavior.
22:04
Heinrich called it art, agony, tree-pandelman. And very early, after I had given a conference on this in the Harnack House in the Max Planck Institute, that was my first debut with Max Planck,
22:24
critic Charlotte Kogon said that it ought not be instinct-handung or tree-pandelman, but instinct-pewewung, instinctive movement. And shortly after this meeting in Max Planck, we had a meeting under van der Klaue in Leiden.
22:45
And there, we discussed the elements of what Heinrich called art, agony, tree-pandelman. And at this time, we were already so convinced that the spontaneity of this movement
23:01
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 simulation onto the releasing
23:22
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.
23:42
And we do not know who first, whether it was Tinberg there or I, who first used the term innate releasing mechanism. It was first called Angeborn-Shema. But then we realized that it was not at all like a diagram,
24:03
but summation of stimuli, a question into 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.
24:23
And particularly Tinberg's and Tinberg'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
24:42
been really investigated since Erich von Holst died. And none of his pupils has really, they're so much dependent on what we know about the spontaneity. We know of the spontaneity of complex behavior. But the basis, that's why my today's lecture
25:03
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.
25:23
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,
25:41
of under-assessing the difference between men 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
26:00
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
26:22
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
26:44
predictable 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
27:01
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 color at which you call a shirt. The shirt is white.
27:20
It's really the process of a complicated calculation in which the color of the illumination is put in reference with a color actually reflected at the moment by the object. And it is the object's color reflecting preferences, which we call its color.
27:41
And this abstraction is a thing which, even in this abstractive apparatus, is able to perform, which obviously is evolved in the service of recognizing things, individual things, can do more. It goes a lot of way to forming generic concepts.
28:05
If I see my dog from far, from near, from behind, from in front, in red light, in green light, and recognize it is also the same dog, this is Ding Constance. But if I recognize a property of dogishness
28:24
in this dog, in the neighbor's 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
28:41
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
29:01
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. Fused together gives something like conceptual thought.
29:24
Exploratory behavior has a very important role in this. Exploration consists in an activity which is independent from any special drive.
29:40
The raven playing, exploring a new object, sees immediately when he gets really hungry, but he plays the whole repertoire of his activities on any new objects, unknown objects, which you offer him. And it is not that he wants to eat,
30:01
but he wants to know whether this is, in principle, edible. And this kind of exploratory behavior, which is very well developed in rats and in very many non-specifically adapted animals, is also a specific property of man.
30:24
And all these things work together to make conceptual thought, which end with it, syntactic language. And from the first prebiotic, virus-like living being,
30:42
to up to the most recent ancestors of ourselves in Old White Gorge, the genome was the main apparatus, the holy method of mutation and selection, was the only method of gathering information.
31:03
And quite suddenly, in the late tertiary period, these anthroposes developed something new, a new apparatus with just the same thing, but much faster. And if a modern man were asked to define life,
31:24
I'm quite sure that he would pass over, in his definition, the double helix discovered by Crick and Wachsen, who are both here, and this would pass over something very specific of human life, this very fast,
31:43
new method of gathering information. And whether this new apparatus will prove a benefit or a curse is something which the young people will have to decide. Thank you.
32:04
Thank you. Thank you very much. Thank you.
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