7 Jan 2017

Uexküll (3.7) Theoretical Biology, “The Observer and the Animal”, summary


by Corry Shores


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[The following is summary. All boldface and bracketed commentary is my own. Proofreading is incomplete, so please forgive my typos. Page citations refer to the 1928 German edition first and to the 1926 English edition second. Note: German terms are repeatedly inserted to facilitate comparison with translations of other Uexküll texts.]





Summary of


Jakob von Uexküll


Theoretical Biology

[Theoretische Biologie]


Ch.3 The Content-Qualities

[Die Inhaltsqualitäten]


3.7 The Observer and the Animal

[Die Beobachter und das Tier]




Brief summary:

An animal subject’s body can be divided into two main parts: {1} the receptor half, which is involved with the properties of the animal’s world that affect it and potentially trigger it to act, also called the world-as-sensed [Merkwelt], and {2} the effector half, which is involved with the properties of the animal’s world that facilitate its effective action and which can be altered by means of those actions. This half is involved with the world of action [Wirkwelt]. But because each half is so intimately bound up with its respective world, such that even physical resemblance can be detected, we might think that the animal is simply the imprint of its environment. But this is a faulty conclusion. Our sensory organs are spatially distinct but they tell us about objects whose properties are unified. The animal carves up the world of indications in its own way and by means of its own anatomical structures. But when we consider nature more broadly, we see that there is a larger plan at work governing how each species experiences the world. Also, although we cannot know another animal’s sensations, we can, by means of experimentation with their response behaviors and with a study of their anatomies, discern which environmental indications it is sensitive to and also the form of the relations holding between these indications.







[There are two functional halves to an animal subject: {1} the receptor half that is sensitive to affecting influences, and {2} the effector half that reacts and makes changes in the world.]


[We are continuing our study of the animal on the part of an observer. It seems we will consider the parts of the animal itself. Uexküll, instead of simply making that analysis, is fitting this description in with the prior discussion. So we think of the observer looking at the animal. The animal in a sense then has properties that the observer can detect, and thus those properties serve as indications for the observer (see section 3.5 on indications). On the basis of these discerned indications telling the observer about the animal’s body, she would distinguish two halves of the animal: {1} the receptor half, which is involved with the properties of the animal’s world that affect it and potentially trigger it to act, also called the world-as-sensed, and {2} the effector half, which is involved with the properties of the animal’s world that facilitate its effective action and which can be altered by means of those actions.]

The properties of which the animal is composed are likewise indications for the observer. These, after careful study, he will divide into two halves — a receptor half, corresponding to the world-as-sensed [Merkwelt], and an effector half, corresponding to the world of action [Wirkwelt]. The receptor half receives the actions of the surrounding-world [Umwelt], and the effector half reacts thereto.

(68 / 81, bracketed insertions mine)




[Because the animal’s receptor half is so intimately bound up with its world-as-sensed (Merkwelt) and because its effector half is so intimately bound up with its world of action (Wirkungswelt), it would seem that the animal is an imprint of its surrounding world (Umwelt) and is thus moulded from it.]


The animal has both receptor organs and effector organs. [See §§26, 32, 34-43 of Stroll through the Worlds of Animals and Men.] The receptor organs correspond very closely with the animal’s world-as-sensed, and the effector organs correspond very closely with the animal’s world of action. We thus are led to view the animal as being something like an “imprint of its surrounding world”. [In other words, half of the organism is ‘designed’ to sense the important features of its surrounding world, and the other half is ‘designed’ to effectively operate with and in that world. As such, the features of the surrounding world are directly and obviously implied in the receptive and effective features of the animal’s anatomy and behavior. We can look at the animal’s body and actions, and we can infer what sort of environment it thrives in. The resemblance might even be visually and obviously apparent, like the white fur of an animal that lives in a snow-covered region.] This idea forms the basis for theories that regard the animal as like a material moulded in conformity with its environment.

There is an astonishingly close correspondence, on the one hand, between the animal's receptor organs and the world-as-sensed [Merkwelt] and, on the other, between its effector organs and the world of action [Wirkungswelt]. This must strike every observer, and it gives the impression that the animal is merely an imprint of its surrounding-world [Umwelt]. On this impression are based all the theories that see in the living substance of which all animal bodies are made, merely a plastic element, passively moulded, which adjusts itself more or less exactly to external influences.

(68 / 81, bracketed insertions mine)




[Theories saying that the animal’s anatomy and behavior are moulded by the environment are wrong. For, the coherence of the animal’s limited set of features implies a coherence and unity in the environment’s features, when in fact the environment’s features are countless and variable.]


These theories saying that the animal is moulded from its surrounding world forget something. [The idea seems to be the following. The coherence of the features in an animal seem to imply a coherence of the features in the environment. For example, the whiteness and warmth-providing features of the animal fur are coherently given in one part of the animal’s body (the hair), and it makes us think that the snowy environment has a coherence to it, for example, that it has a material, namely snow, which is cold and white. Uexküll claims that the features of the surrounding world are not inherently and coherently unified, and they only are so when they enter into agreement with the animal. Without that bond with the animal’s features, the properties of the world remain disconnected. I am not sure exactly why that is so. I could see the features of the world having a certain coherence based on natural laws, like snow is always cold and white on account of its physical constitution. Perhaps the idea is that the animal’s anatomy and behavior are sensitive to and interact with only a small selection of the environment’s features, and that limited representation makes us think that the environment has a simple composition. However, the environment will have countless variable features. Let me quote, because I am guessing:]

These theories overlook one essential circumstance, namely, that the surrounding-world [Umwelt] of an animal, if considered by itself, is not a unity. On the contrary, the properties of the surrounding-world [Umwelt] become linked up into a unity only when they are in agreement with the properties of the animal; without this bond, they merely flutter about disconnectedly.

(68 / 81, bracketed insertions mine)




[But animals are not moulded by their environments, because their receptive organs are not distributed on the body like they are in the environment; and also, the animal is receptive only to a limited selection of environmental indications, and this varies highly between animals in the same environment. Furthermore, the features of objects responsible for their many sensible properties are unified in the objects, but they are sensed disunifiedly by distinct sensory organs in the animal’s body.]


So Uexküll does not think that the animal is moulded by its environment. To prove this, we need to show that animals have features that could not have been imprinted upon it by the features of the environment. He acknowledges that such a proof cannot be given definitively. [I am not sure why. Perhaps any cited evidence could still be interpreted somehow as being an imprint. But the reason for this I do not know.] [Uexküll then goes on to cite examples anyway. I unfortunately do not understand his point here, so you should skip to the quotation to follow. I will at least guess the idea is the following. Animals have receptive organs responsible for different sense modalities. Were the receptive organs an imprint of the surrounding environment, then the organs would be distributed on the body in the way the environment affects the body. And since the environment affects the body from all directions, we would expect the receptive organs, as imprints, to be distributed all over the body. In fact they are not. They are concentrated in certain specific places. But that interpretation cannot be right, because it does not seem like he has in mind the idea that an animal would have eyes, ears, etc. all over its body. Perhaps instead Uexküll is not referring to the spatial distribution of the organs on the body, but rather to the select range of indications it is sensitive to. So along this line of thinking, we would note that the environment has many different features. Were the animal an imprint of the environment, it would be sensitive to all of those features. However, animals are sensitive only to a very limited number of them, that is, to just the indication-circles that interest it. Also consider the vast diversity of animal types living in the same environment. They can even have very different sensory systems (consider bats, for example, like Nagel does). If animals are imprints of their environments, then how could there be such diversity of anatomies and behaviors in one same environment? I am not sure that this is the correct interpretation, because we might say that the animal is still imprinted by the environment, even though only by a portion of it. Given what he writes later, there is a third interpretation I would offer. Many of the features of one object are given together by means of the same physical properties of that object. For example, the physical motion of a vibrating instrument string gives both its visual appearance and its sonic appearance. But we do not have one organ that detects both of these features. We have eyes and separately we have ears. So the unity of the object’s properties is not reflected in the animal’s body, which separates those features anatomically. Thus it is not an imprint of the environment. I realize that I have probably not conveyed Uexküll’s own reasoning, so please read the quotation to discern it for yourself.]

The convincing proof that the animal body does not owe its form to external influences can only be given by our showing that it displays properties that could not have been imprinted on it from without. And this proof can never be brought quite definitely. In the arrangement of their receptive or sensory organs, all the higher animals show a distribution which has nothing to do with the arrangement | of the surrounding-world [Umwelt], but which, on the other hand, reproduces spatially the distribution of the circles of their indications [Merkmalskreise]. In the eye are assembled all the nervous elements that are directed on the colour-indications of the surrounding-world [die farbigen Merkmale der Umwelt]. The same holds good for the ear, as regards the indications [Merkmale] of sound. In the mouth lie the receptors for taste, and in the nose those for smell.

(69 / 81-82, bracketed insertions mine)




[Those still defending the idea that bodies are imprints of environments despite the fact that unified phenomena are perceived disunifiedly through various sense organs can make the following reply: the properties of the unified phenomena are conveyed disunifiedly through various distinct physical media, thus various distinct receptive organs are needed to perceive those sensible features.]


Uexküll then considers a counterargument that defends the notion that animals are imprints, despite the separation of sense organs. [Again, you will have to discern the reasoning for yourself. I can only guess what the idea is. It might be that the physics of the sensible phenomena involve a separation into physical media, and thus it is fitting that the anatomy, as an imprint, has a separation of the receptive organs. I do not understand what Uexküll means by etheric vibrations, so let us consider our own example of the vibrating string. Yes, it is one same unified phenomenon responsible for its features, namely the string’s structural properties in relation to its particular physical motion. But this movement affects both the motion of the surrounding air and it also affects the patterns of light waves bouncing off of it. So the sensible properties of this motion is conveyed on the one hand through air vibrations and on the other through light wave patterns. Thus our anatomies, as imprints of the environment, should rightly have separate eyes and ears, according to this reply.]

As a rule people try to explain this remarkable anatomical separation of the quality-circles [Qualitätenkreise] within the body by pointing to the uniformity of the method [gemeinsame Wirkungsart] in which the related properties act in the surrounding-world [Umwelt]. The etheric vibrations require specific transformers if they are to be converted into nervous excitation; so do the air-waves. The same may be said of the substances soluble in water that yield the taste stimuli; and the stimuli of the sense of smell are conveyed by air- currents.

(69 / 82, bracketed insertions mine)




[But in the case of fish, the same medium conveys both olfactory and gustatory information, yet the receptive organs are distinct. So the above defense of the imprint theory does not hold in all cases.]


Uexküll rejects this defense [namely, that the distinct physical media conveying the sensible features of unified physical phenomena call for distinct sensory organs.] He notes that for aquatic animals, there is one medium conveying both smell and taste, as they are both “delivered by substances dissolved in water”. However, “fishes, like other vertebrates, have their olfactory and gustatory organs quite distinct from one another” (69 / 82). [So in other words, a shared medium for two sorts of sensible features of things in the world should correspond with one sensory organ sensing them both, according the imprint theory. But in fish, this is not so.]




[What determines the distribution of sense functions in sense organs on an animal’s body is the result of how their quality-circles are arranged, such that when certain types of qualities lie in different circles, they could be received by distinct sensory organs that are found on different spatial locations of the body.]


Thus “the anatomical separation of the receptive organs into quite distinct unities” cannot be explained on the basis of “the chemical or physical connection with the surround world [Umwelt]” (69 / 82, bracketed insertion mine). [I will have to guess again at the next idea, so please consult the quotation. Uexküll might be saying the following. We have mark-signs (Merkzeichen), which are the qualitative differences we perceive (see section 3.4). The next sentence I am not sure how to interpret: “What are responsible are the attention-forms [Aufmerksamkeitsformen] of the mark-signs [Merkzeichen], of which they are the spatial images”. The attention-forms might be the relations between attention thresholds. Perhaps the mark-signs are spatial images of these relations. But I do not think the mark-signs are spatialized. So perhaps somehow the receptive organs are spatial images of these relations, although it is not entirely clear to me how that is so. At any rate, it seems the attention-forms (whatever they are) of the mark-signs are responsible for the anatomical distribution of the receptive organs, with one component being a spatial images of another. Let us consider some possible interpretations. Does he simply mean that because visual mark-signs are not found in the same quality-circles as sonic mark-signs (for some species), that these quality circles can be considered spatially distinct somehow, and thus on the body the eyes and ears are spatially separated? In this case, the “spatial image” would seem to be the spatial distance between the receptive organs which corresponds to the non-overlap of the different quality-circles. Or does he mean that within one sensory organ there is a spatial distribution that reflects the distribution of mark-signs? For example, in the part of the ear where there are hairs that vibrate, perhaps the hairs are arranged in an ascending order that reflects the ascending order of frequencies we are sensitive too. But this does not explain the spatial distinctness of different sensory organs. Please read the following paragraph, and I would be interested in what you think. The interpretation I will go with for now is that because for example sight and sound data are in different qualitative circles, they are received by different organs. Were they somehow in the same circle, they would be received by the same organ. We might consider bats for example. For them, spatial properties and sonic properties might very well fall into the same quality-circle. In other words, it both hears and “sees”  in the same receptive act, using the same receptive organ, because the form of its attention places spatial qualities and sonic qualities in the same circle of quality variation.]

For the anatomical separation of the receptive organs into quite distinct unities, the chemical or physical connection with the surrounding-world [Umwelt] cannot be held responsible. What are responsible are the attention-forms [Aufmerksamkeitsformen] of the mark-signs [Merkzeichen], of which they are the spatial images. And thus the representations we draw in space of the quality-circles [Qualitätskreise] acquire increased significance [Bedeutung]. When we study the sense-organs of animals, we see Nature herself at work, and in the very act of reproducing in extensive forms the laws of intensive magnitudes.

(69 / 82, bracketed insertions mine).




[Indicators are common among all animals, because we live in the same physical world, although each animal groups them differently and is sensitive to a limited range of them. We do not have access to another animal’s sensations, but by conducting experiments, we can determine the indicators they are sensitive to and the ways those indicators are related. This indirectly gives us a picture of the animal’s Umwelt and of its general manner of experiencing that world.]


[Again please consult the quotation, but I will guess at an interpretation of this next paragraph. We cannot know the animal’s sensations, but we still want to know what its inner world is like or at least what its life is like in its own particular Umwelt. To arrive upon this knowledge, w can perhaps conduct experiments to see how animals respond to variations in the indications of their environment, and in addition we can study how their nervous systems work while perceiving these variations. This can tell us what their indications are and thus what their mark-signs might be, even though we do not have access to their own experience of sensations. His next point seems to build from this idea that the way our indications are grouped in our experience is the same as for other animals. This was a point I had trouble with, but as far as I could tell, he is not saying that our indication groupings are identical but rather that the general laws ours follow will also be followed by the animals. For example, he says, an indicator of spatial relations between objects for us would not be able to indicate a different spatial relation between the objects for another animal. The indications, he explained, are thought to be parts of the world rather than parts of inner experience (the mark-signs are more on the side outer of inner experience). He seems to be working with the idea then that while our mark-signs might differ between creatures in how we experience them, the indications that one animal finds in the world are potentially discoverable by other animals, because, it seems he assumes, we share the same physical world, even though we all constitute it differently and subjectively. So perhaps we might say that we each live in our own appearance-world with its own mark-signs, but we also live in a shared world with common indications. Uexküll’s next point seems to be that on the basis of how we determine the indication groupings of higher animals, we then do so for lower animals. They might have sense organs very different from our own, but so long as we can group their indicators, we can better study their Umwelts and manners of experiencing the world. But please consult the quotation to see its meaning:]

This makes our research very much easier, for, in investigating animals, we can never hope to attain to a knowledge of their sensations. All we can determine by experiment | is the number and the nature of the indications [Merkmale] in the sensed world [Merkwelt] to which the animal reacts. Thus far, we have been able to group indications [Merkmale] only according to the forms of our own attention [Formen unserer eigenen Aufmerksamkeit]. But by realising that the sense-organs of the higher animals correspond to this grouping, we are put in a position to go further; by anatomical study of the lower animals, we can now undertake to group their indications [Merkmale] also, and this even when we come across sense-organs that are quite unfamiliar to us.

(69 / 62-83, bracketed insertions mine)




[We can infer that there is a higher order of unified nature on account of the super-subjective cause of how we fashion our appearance-worlds.]



[I again am uncertain what is meant in this next paragraph. Let me quote it first.]

The most important advance, however, lies in the following conclusion. If the laws manifested in the forms of our attention [Formen unserer eigenen Aufmerksamkeit] (which is the deciding factor for the appearance-world [Erscheinungswelt] of our own subject) can be recognised not only in the shape of our own body, but also in the shape of the bodies of other subjects of whose attention-forms [Aufmerksamkeitsformen] we know nothing, then this indicates that the work of fashioning by the mark-signs [Merkzeichen] is not determined purely by our own subject, but is super-subjective. This means that we are on the track of a control by Nature [Naturwalten] pointing to a unity even higher than our own apperception, in which otherwise we must see the final unity.

(70 / 83, bracketed insertions mine)

[So somehow the shape of our bodies expresses the forms of our attention. But we also know that the shapes of the bodies of other animals also expresses the forms of their attention, even though we do not have any knowledge of those forms (except what we learn through objective experimentation on the animals). The part that confuses me is: “the work of fashioning by the mark-signs [Merkzeichen] is not determined purely by our own subject, but is super-subjective”. I do not know what ‘the work of fashioning by the mark-signs’ [die Formgebung der Merkzeichen] is. Let me make a wild guess. The world objectively speaking is full of indications of its very many different properties. Each animal is sensitive only to a limited set of them. So for example, dogs might see fewer colors than we do, but mantis shrimp see more. Our appearance-worlds are made of the mark-signs corresponding to the indications we are receptive to. So perhaps ‘the work of fashioning by the mark-signs’ is the way that our limited range of mark-signs fashions our own unique appearance-world, which varies from creature to creature. They are sort of like the palate we use to paint our world, and each creature paints their world with a different palate. And so he says that because we can see this through the anatomies of other animals, we can say that this fashioning of the phenomenal world is super-subjective and involves the higher unity of Nature. But I do not understand that statement or the reasoning involved in it. I wonder if the idea is the following, but this is an unreliable guess. Each animal’s body carves out its own appearance-world, giving it its own place in relation to the other creatures. But somehow all these different Umwelten interlock and function together coherently. So perhaps the particulars of our own species’ attention-forms is governed by a higher organization of all species in nature. I doubt that is the meaning, but I am not sure what else to propose. See the quotation above.]




[Our inner mental life, which experiences intensive variations, corresponds to the extensive differences of our receptive organs and of the physical stimuli. But there is not a paralelism; rather, there is a factor or law that determines their mutual conformation.]


[The final paragraph also confuses me, so as always, please consult it for yourself. It might be making the following points. The forms of our attention conform to the forms of our anatomy. I am not exactly sure how, but perhaps it is again the idea that certain qualities are grouped in their own circles, and the separation between these circles corresponds with the separation of their respective receptive organs. From this Uexküll says we can infer that there is some factor that determines both our physical and mental activities. I am not sure what that factor is. Perhaps it is his idea of a “plan” that we will return to later. (Now, some might object that there is no factor governing the mind’s and the body’s correlation, but rather that they are simply parallel.) He does not think that we can explain this correlation with a notion of parallelism, because the intensive and the extensive are never parallel. However, there are certain laws that can express themselves both intensively and extensively. (I am not certain exactly how Uexküll makes the intensive/extensive distinction. From before it seemed like a difference between a spatialized sort of variation with a variation by degrees. Here maybe I wonder if there is the element of the extensive being external and a part of the objective world while intensive is a part of the inner world of mental experience. I do not know the laws that express themselves in both. Perhaps we might think of there being an extensive difference between sound waves of different frequencies and an intensive difference between how we experience them as different pitches.]

The fact that the forms of our attention [Formen unserer Aufmerksamkeit] find expression in the conformation of our own frame is sufficient to suggest that there is a factor which uniformly determines the activity both of our consciousness and of our body. It is not enough to speak of a parallelism between mental and bodily processes; an expression like this loses its sense when we are dealing with a comparison between intensive and extensive forms, for such forms are never parallel to one another. On the other hand, we may speak of identical laws, which express themselves both in intensive and extensive forms.

(70 / 83, bracketed insertions mine)




Works cited (in this order):


Uexküll, Jakob von. 1928. Theoretische Biologie, 2. gänzlich neu bearbeitete Auflage. Berlin: Springer.


Uexküll, Jakob von. 1926. Theoretical Biology. Translated by Doris Livingston MacKinnon. London: Kegan Paul, Trench, Trubner & Co. / New York: Harcourt, Brace & Company. PDF available at:





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