J. Theoret. Biol. (1964) 7, 53-61

Synopsis of Organismic Theory

WALTER M. ELSASSER

Department of Geology, Princeton University, Princeton, New Jersey, U.S.A.

(Received 1 July 1963, and in revisedform 10 February 1964)

The theory developed over a number of years by this author is here surveyed in a concise form, there being a sequence of definitions, assump- tions and propositions, with connecting and explanatory text. Organismic theory is not identical with biological theory; it is a formal scheme pre- requisite to any such theory. It inquires into the possible modes of behavior of inhomogeneous systems insofar as they may differ from the behavior of the macroscopic homogeneous systems usually considered in physics and chemistry. While assuming the unqualified validity of the laws of quantum physics in the organism, it deals with the novel methods of analysis required to take into account the tremendous complexity and inhomo- geneity which is found in organisms but in no comparable degree in the inorganic world. In the course of the preparation of this summary we believe, moreover, to have achieved a substantial clarification and simpli- fication of some of the concepts developed earlier.

Organismic theory as we understand it here is not, properly speaking, one of the branches of theoretical biology, neither is it co-extensive with the latter. By way of introduction it might be useful to approach its nature in terms of an analogy: if one studies the universe at large there is at one’s disposal as a tool, the theory of general relativity. This theory is not a form of astro- physics nor is it even equivalent to cosmology. Instead, it is merely an abstract scheme obtained by comparatively simple generalization of a body of theory which is known to be valid in our small fraction of the universe. It imposes certain sharp restrictions upon models of the universe at large while at the same time admitting of a considerable and satisfying variety of such models. The actual construction of a cosmological scheme must be based upon observation. But in the absence of the relativistic formalism one would be deprived of any theoretical guidance in cosmological questions and would be reduced to a rather crude form of incoherent empiricism.

Organismic theory as we have tried to develop it over a number of years starts by assuming the unqualified validity of the laws of ordinary quantum mechanics for the physical and chemical processes going on in organisms (moreover, we assume the validity of the basic statistical postulates from which the second law of thermodynamics follows). The main subject of

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inquiry in organismic theory is the manner in which the basic state functions of quantum mechanics must be combined so as to form representations of that extreme complexity and inhomogeneity which characterizes organisms. We claim that the uncritical extrapolation of some of the rules of statistical mechanics as commonly practised leads to a disastrous undervaluation of the potentialities of such utterly complex systems. While leading up to a logical and statistical analysis of the consequences of extreme complexity and inhomogeneity this theory, owing to its formal character, prevents at the same time the introduction of speculative hypotheses such as vitalistic concepts, or the abrogation of the second law of thermodynamics.

Since ideas of the latter type are not particularly fashionable these days, the chief interest of organismic theory at present lies in a more thorough evaluation of the potential of modern quantum physics to serve as a basis for more extensive and varied representations of natural systems than have heretofore been considered available. Thus, to repeat once more, organismic theory is concerned with the logic and analysis pertaining to the description of systems of extraordinary complexity and inhomogeneity of structure and dynamics, whereas conventional statistical mechanics has almost invariably been limited to relatively homogeneous systems on dealing with bodies whose dimensions are well beyond those of the simpler molecules.

Notwithstanding the conservative character of the theory which is implied in a refusal to modify the laws of quantum physics or the second law, it would be very surprising if in going from physical theory to the foundations of biological theory one would not encounter, at some places, radical changes of a logical and mathematical nature. These must be expressed in the way in which the state functions of quantum mechanics are being combined to form representations of natural systems; the principles involved are outlined in the next two sections. (In the language of statistical mechanics, it is necessary to re-investigate the meaning of ergodic theory. To revert to our previous example of relativity, ergodic assumptions play here a role similar to that of Euclid’s axiom of the parallels in geometry. Fortunately, it will not be necessary to enter into the intricate subject of ergodic theory in this paper.)

Although the present outline cannot as yet have the rigor and consistency of a fully developed theory we have tried to cast the presentation into a fairly succinct form. We shall designate by A an assumption (which would correspond to a postulate in a more formal scheme), by D a definition, and by P a derived proposition, which latter may correspond either to a deduction following from the A’s, or to an empirical law derived from adequate observa- tions, or to a mixture of both, as the case may be. More refined discriminations do not seem necessary at the present stage of the inquiry. These various types of statements will be numbered through in the order of their appearance. To

SYNOPSIS OF ORGANISMIC THEORY 55

avoid italicizing them all, their end in each case is indicated by the end of the corresponding paragraph.

I

Al. The laws of quantum theory hold in the organism without any modifications, as they do in theoretical chemistry, in the physics of solids, etc.

A2. The postulate of equal statistical weight of non-degenerate quantum states as well as the assumption of randomness of phases of the corresponding wave functions hold in the organism.

It is known from statistical mechanics that assumptions A2 insure the general validity of the second law of thermodynamics. But it is perhaps no accident that so many biologists have questioned the application of this law to organisms. This is the result of the widespread misconception that there is only one kind of order, which may be measured by the negative of the entropy. Actually there can be types of order at different levels of organization which need not be related to each other. If all types of order are treated on the same footing there will arise many logical inconsistencies; some of them have been discussed in detail but without an identification of their origin, by Brillouin (1962; it is unfortunate that in this remarkable work the logical distinction of various relatively independent levels of order is not made). Take as an example an automatic device which might have a simple homo- geneous input of raw material and which generates from it bodies with highly complicated, ordered shapes. It is clear that such an automaton while creating order at a higher level does not thereby violate the second law. The logical discrimination between order at various levels of organization is of fundamental importance in biology (see Concepts ~fSi&g~, 1958). Therefore the implication of A2 is that the order which is so conspicuous in organisms pertains to a higher level of organization and that the existence of such order is altogether compatible with the disorder at the molecular level which finds its expression in the second law. (The order of a crystal on the other hand is clearly of the molecular type and can be understood in detail on considerations of statistical thermodynamics.)

We proceed now at once to some of the chief problems involved in the description of complex systems. In statistical mechanics one considers the number of ways in which systems can be assigned to available states. The simplest combinatorial problem of this type (which it may suffice to mention here) consists in determining the number of ways in which N different objects can be ordered, this number being, say,

Z=N!-JP

56 WALTER M. ELSASSER

and thus log 2 = N log N. If here N is taken as of the order of the number of molecules in a body the size of a small cell, say, N will be a very large number, and so will be log Z.

D3. A number will be called immense if its (decadic) logarithm is a large number, Similarly, if the ratio a/b is immense we shall say that a is immensely large compared to b or, conversely, that b is immensely small compared to a. The term “immensely rare event” has a corresponding meaning.

Thus Z, above, is an immense number. There exists no common convention as to when a number is to be considered large, this being usually a matter of expediency. D3 reduces the definition of an immense number to that of a large number and hence is subject to the same ambiguity. Just to fix our ideas, we may, for instance, assume that a number is large if it is of the order of a few hundred.

D4. The totality of quantum states accessible to a system in a given temperature range will be designated as the phase space corresponding to this range. In the sequel the temperature range will be taken as that in which organisms are viable.

[This definition is something of a “mixed metaphor” in that the phase space is a construct of classical theory not immediately applicable to quantum physics. For purposes of our conceptual schemes this loose usage will be sufficient; and since the mathematical apparatus involved is well explored it should not be difficult to substitute more precise terms if desired. Moreover, on later using the term “phase space of a class of organisms”, we are glossing over certain more serious mathematical complexities (distinction between the canonical and the grand ensemble). To be highly precise here would require extensive mathematical elaboration; we feel confident, however, that the following conceptual analysis would not be radically modified thereby.]

P5. The number of quantum states in the phase space of any system as large as an organism is immense.

This is a well-known result of standard statistical mechanics. In the present context it assumes its main interest by virtue of the fact that sets of real objects do not contain immense numbers. This is most readily made clear by referring to the fact that astronomical observations show our universe to be finite. An age of 30 billion years and a radius of 30 billion light years are probably rather generous upper limits for the dimensions of the universe. We shall presently indicate that in a world of this magnitude the number of biological occurrences of any one type will not be immense.

D6. A class is a set of objects having certain properties in common. Biological classes may refer to any sub-divisions of taxonomy. The limiting case of a class having one and only one member will be designated as a one-class.

SYNOPSIS OF ORGANISMIC THEORY 57

Note that the class concept has here been specifically applied to designate objects of experience. Actually we are dealing with abstract symbols which are representations of the real objects in our scientific language. Thus, while the concept of a class with an infinitely large membership may have no pragmatic meaning in actual experience, it is of course a legitimate idealiza- tion of theory. Again, in dealing with purely mathematical abstractions not representing objects of experience, it is convenient to use the alternate term set.

The number of organisms, or cells, of any class existing on the earth while it may be extremely large, is not immense in the sense of our definition, as even the simplest of estimates will show. We are, however, not so much con- cerned with the number of organisms as such as we are with another quantity : every organism changes its internal chemical and electrochemical configuration all the time, and we shall be interested in the variety of such configurations that a class of organisms can assume. To obtain a quantitative measure of this, let us for instance assume that a cell changes its internal configuration, say, every microsecond (although some other unit of time would do as well). We shall designate a configuration defined in this sense as a “system event”.

D7. The number of system events of an object is equal to the lifetime of the object measured in an appropriately small unit. The number of system events of a class is obtained from this by summing over all the individuals in this class.

Note, however, that the term “event” here is purely abstract in the sense that it does not as yet denote an operationally defined internal state, a quantum state, say, or some other well-defined internal configuration. We know that these configurations occur but we might be unable to analyze them exhaustively for individual cases (see below).

P8. For a universe of finite size comparable to the known one, the number of system events in any class of organisms may be very large but is finite; one may estimate a number between 106’ and 10” as a rather safe upper limit. This number, however, is immensely small compared to the number of quantum states (and possibly of other chemical or electrochemical configurations) in the phase space of the class.

The numbers given here differ somewhat from those given earlier in the author’s book (Elsasser, 1958) but the general result as expressed in the second sentence of P8 remains the same. PS may be cast in the following alternate form which perhaps brings out better its significance in organismic theory.

P9. Each system event of an organism represents an immensely rare occurrence in the phase space of any class to which the organism may be

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assigned. This may be stated by saying that in a finite universe each system event of a complex system such as an organism represents effectively a one- class. The actual system events occupy only an immensely small fraction of the available phase space. Alternately, since the assignment of a specific quantum state to a macroscopic system can only be made on a statistical basis the probability of any specific quantum state ever being realized is immensely small.

In discussing the axiomatics of quantum theory one usually assumes implicitly, or else postulates explicitly (e.g. Tisza, 1963) that any object of physics and chemistry such as an atom, molecule, etc., can be procured in an unlimited number of fundamentally indistinguishable copies. Hence there is no limit to the number of measurements that can be made on such a class of objects. This assumption must be abandoned in organismic theory. The descrip- tion of nature can therefore no longer be carried out in terms of an idealized picture in which classes of infinite membership are admitted.

AlO. In organismic theory the mathematical tool of description is an abstract structure in which all sets of mathematical entities representing actual objects of experience have finite membership. Such a structure will be designated as afinite universe of discourse (abbreviated FUD).

These notions are in sharp contrast to the accepted methodology of physics which is based on analysis, in the purely mathematical sense of the term. Analysis cannot even be formulated without introducing the concept of infinite sets. In a FUD, individuality has a rather immediate but entirely abstract meaning. In the simplest case it expresses the existence of one-classes; a somewhat more complicated case is that of finite classes which have individuality, such as the classes of taxonomy.

We may ask whether the distinction between mathematical analysis with its infinite sets and a FUD does not, in practice, become trivial provided only the FUD is made large enough. This will only be true, however, if the FUD is sufficiently homogeneous. Now in a FUD there may exist inhomogeneous classes (i.e. classes whose members may have many properties in common but who differ from each other in certain other properties). We now find a radical difference between a FUD and a universe of classes with infinite membership: suppose a class of the latter type, an infinite class for short, is inhomogeneous to begin with. By a simple process of selection continued indefinitely the class can then be decomposed into two or several sub-classes each of which is more homogeneous than the original class. This is an operation wholly familiar in its mathematical aspects from set theory. In this decomposition, the more homogeneous sub-classes will in general again be infinite. On repeating this process of selection a sufficient number of times one obtains classes which can be made as homogeneous as one chooses while still having infinite

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membership. In quantum mechanics there exists, moreover, the concept of a perfectly homogeneous class (known as a “pure state”) namely, a set of identical systems, all in the same quantum state.

II

Pl 1. In a FUD there may exist classes that cannot be homogenized by selection. The process of the selection of sub-classes with respect to some given set of characteristics terminates, instead, in a collection of one-classes with respect to these characteristics.

The intrinsic inhomogeneity of classes of this type may be of two kinds: it might either correspond to directly observable characteristics as is the case for macroscopic morphology; or else it might be potential. The latter means that the admissible measurements (see below) inform us of the radical inhomogeneity of the class in the chemical realm without, however, permitting us to ascertain the exact molecular state of each sample. In this second case the inhomogeneity of the class is equivalent to the fact that each system event pertaining to the class represents an immensely rare occurrence in its phase space.

A12. All classes of living systems or collections thereof are radically inhomogeneous in the sense of Pl 1. We designate this assumption as the principle offinite classes (Elsasser, 1958, 1961).

The difference between the formal methodology of physics and that of biology is now readily apparent. In brief, physics deals essentially with homogeneous classes (which may be assumed infinite for mathematical convenience)-biology is the science of inhomogeneous classes.

We should note here that if we speak of inhomogeneity we are first of all concerned with features of a chemical and electrochemical character. This is the basic inhomogeneity of organisms at the lowest level of their organization. As we proceed from there toward the macroscopic realm, we find other inhomogeneities at various levels of morphological organization.

An important property of finite classes, including of course inhomogeneous classes, is the following.

P13. For finite classes, there frequently exist questions which cannot be answered, whereas the corresponding questions for infinite classes may have quantitative answers.

Let us give an extremely primitive example for this proposition: suppose a coin is tossed five times and then destroyed. In all five throws “head” appears. This might give rise to the suspicion that the coin was “loaded”, i.e. asym- metrically constructed. The question can, however, not be answered with any degree of certainty since the probability of this being a random occurrence is l/32, by no means very small. The question could readily be answered

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observationally if instead of doing away with the coin we kept tossing it a few hundred or thousand times.

This example refers to a class of occurrences which is homogeneous. One may assume a fortiori that in inhomogeneous finite classes there exist many unanswerable questions. Also, the example is of an exaggerated simplicity, but to give more complicated examples would require the introduction of some quite involved mathematical apparatus. The existence of many unanswerable questions in the finite classes of organisms with their tremen- dous structural complexity must appear highly plausible a fortiori. In view of A12, therefore, organismic theory can be said to establish limitations on what is often simply called the experimental method but what is realIy an idealization, namely, the more or less uncritical extrapolation of experimental results to the hypothetical case of unlimited repeatability. From the viewpoint of organismic theory, the assumption of an immense number of experiments, in particular, is an operationally meaningless concept.

We come now to the next main question, that of prediction. The value of a theory can usually be measured in terms of its predictive power. Quantum mechanics is a tool of prediction although, even if we start from a single quantum state, the predictions are as a rule only statistical. In homogeneous systems having many equal components the statistical features often average out, and then unique prediction becomes possible.

D14. The predictive process whereby variables of a system or their probabilities are ascertained for future times by integrating the equations of quantum mechanics, will be designated as physicalprediction.

Let us say parenthetically that an alternate form of prediction is one which is based upon organismic properties related to the radical inhomogeneity of classes. This will be taken up again in section III.

Physical prediction depends on our being able to assign a definite state, or in the more general case a set of states, to a system at an initial time; this assignment is, of course, based on suitable measurements. In the conventional treatment of quantum mechanics it is assumed, implicitly or explicitly, that any system can be prepared so as to be in a single quantum state. For systems as complex as organisms, the reduction to a pure state would involve the combination of a tremendously large number of simple measuring processes. In a FUD this is not necessarily an operationally meaningful concept; the description can as a rule not be pushed beyond assigning to a system probabilities for an immense bundle of quantum states. The relative probabilities of these states are inductive probabilities; they differ in this from the probabilities of physical prediction based on given quantum states, which latter probabilities are deductive.

In a manner of speaking, quantum mechanics as usually understood is a

SYNOPSIS OF ORGANISMIC THEORY 61

statistical halfway-house. On admitting the idealized notion that any system can be prepared so as to be in a well-defined quantum state it ignores the inductive aspects of probability theory as applied to basic physics, and con- centrates solely on deductive probabilities. The introduction of a FUD as the basic tool of description implies a radical change. The quantum states upon which the deductive probabilities of physical prediction are based, are themselves only determined probabilistically, by induction from the results of measurements which latter, for large enough systems, are compatible with an immense number of quantum states (for details see Elsasser, 1962b).

Now the preceding sentences pertain to physics, purely, and have little apparent connection with biology. However, they do not lead to any sig- nificantly novel results when applied to the conventional homogeneous, macroscopic systems usually dealt with by the physicist and chemist. They do lead to such novelty, in a high degree, if the classes of systems considered are of the radically inhomogeneous type that we consider characteristic of organisms, by A12. We then have the second of the two cases distinguished in the paragraph preceding A12, the case where a large part of the inhomo- geneity of the organisms is, as it were, submerged in statistical indeterminacy. The latter is just another expression of the fact that in a FUD the determina- tion of the exact quantum state of a sufficiently large system is by inductive inferences only, there being as a rule an immense number of quantum states compatible with the data of the description.

Again, the conditions which limit pushing the description beyond the level of purely inductive inferences are found to be of two kinds:

(1) As Niels Bohr has pointed out long ago (Bohr, 1933, 1958) the process of multiple measurements required to determine the instantaneous state of a system will (by virtue of the quantum uncertainty relations) produce perturba- tions in the object which become very severe if the measurements are thorough- going enough. In our language, we may transcribe Bohr’s chief conclusion as follows :

P15. A set of measurements thorough-going enough so that they confine the description of the organism to within an immensely small fraction (or perhaps only to a very small fraction) of its phase space produces perturba- tions which disrupt the organism so severely that it becomes non-viable. After the measurement it will have ceased to be a member of any class of organisms to which it belonged before.

(For a more detailed explanation of the meaning of classes of organisms see A17, below.) Owing to the inevitability of such perturbations it becomes necessary to balance the permissible measurements against the perturbations. In this way, Bohr’s principle is capable of interpretation in a more quantita- tive fashion. The subject which has been dealt with elsewhere (Elsasser, 1962b)

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need not now detain us. We need only to remember that we are dealing here, essentially, with a limitation of physical prediction, since such prediction can only be based upon the knowledge of the state of a system.

(2) There exists a second type of prediction in physical science, which is mathematically formulated in quantum theory. It may be designated as prediction by sampling of classes. The simplest case is that of a single quantum state, e.g. the ground state of an atom or molecule. Once the system and its state are identified, which often may be done with small perturbation for systems of great simplicity, prediction becomes possible by virtue of the principle of quantum mechanics that two systems of the same composition and in the same state are strictly indistinguishable. If the systems considered are not all in one quantum state but are scattered over a number of states, this type of prediction is still possible but with correspondingly less determinate results.

We can now interpret more clearly the principle of finite classes, A12. This principle imposes upon prediction based on the sampling of classes of organisms intrinsic limitations which are altogether analogous to those limitations which Bohr’s principle, P15, imposes on predictions based upon measurements made on a single organism. Al2 is a logically independent postulate. It is based upon a tremendous amount of empirical evidence (for instance, Concepts of Biology, 1958, from which some salient statements are quoted by Elsasser, 1963). We could not even begin to exhibit this evidence in the present review. Let us merely summarize our basic result:

P16. The postulate Al2 limits physical prediction based on sampling of classes of organisms, as P15 limits physical prediction based on measurements of an individual. In both cases the number of states which can compete in the description of the system by virtue of the use of inductive probabilities is immense; it is moreover immensely large compared to the number of system events in any class of organisms involved.

III

A17. There exists no formal a priori criterion which would permit US to distinguish between living and non-living matter. The identification of an object as being alive is ex post facto, being based on observation extending over a finite span of time and showing that during this time the object has exhibited features which we consider as adequate criteria of its being alive. The concept of life is thus purely empirical and cannot be anchored in any assumptions regarding a quantitative discontinuity between life and non-life in the laws of nature.

Organismic theory has Al7 in common with the so-called reductionist philosophy. Moreover:

SYNOPSIS OF ORGANISMIC THEORY 63

AH. All organisms have component processes which can be described adequately in terms of homogeneous classes. These processes can be repro- duced in vitro, and all regularities of their behavior can be ascertained in terms of physical prediction. Such processes are designated as mechanistic.

Note, however, that according to our previous assumption, A12, radical inhomogeneity is also characteristic of all organisms. Therefore, organisms have a dual aspect which lies in their own structure and dynamics and is then reflected in controversies which arise among theories. The situation has a very close analog in the historical controversy of physics regarding the wave nature or corpuscular nature of light. In physics it was finally recognized that the controversy is not accidental but is founded upon a dualism residing in the phenomena themselves. In biology, again, it is necessary to acknowledge the dualism inherent in the phenomena rather than to try to eliminate it at all costs by a one-sided theory. Organismic theory offers a broad enough formal framework to combine these dual and complementary aspects of the organism.

To return now to that theory: in a FUD populated by intrinsically inhomogeneous classes it cannot be asserted that all processes can be analyzed and predicted mechanistically. Finding out what happens in inhomogeneous classes is entirely a matter of empirical investigation. We shall, however, try to assert some general principles, based largely on such biological generaliza- tions as already exist. The basic assumption is:

A19. There exist relationships among observed phenomena which cannot be evaluated entirely in terms of mechanistic models and their attendant physical predictions. These will be designated as organismic.* No organism is without them.

Notice that Al9 says more than A12. The latter removes the basic logical obstacles against a theory in which a certain dualism is inherent in organisms themselves (after the manner in which the wave-particle dualism is inherent in quantum mechanics); A19 asserts beyond this that the radical inhomo- geneity of classes of organisms gives rise to novel phenomena which are not completely random but involve relationships not identical with those deduced from physics. If such relationships do involve regularities in a time sequence, one may be able to make predictions which are not based entirely on physical law but in part on empirical organismic relationships. This in turn justifies the separate definition of a purely physical prediction as given in D 14, above. We may now elaborate Al9 somewhat more:

P20. The organismic components of observed biological regularities cannot

* The equivalent term “biotonic” used earlier by the author (Elsasser, 1958) has been abandoned.

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be reduced to physical laws, nor can a contradiction to the latter be con- structed, owing to the combined restrictions expressed in Bohr’s principle and the principle of finite classes. Any operational effort to achieve the desired reduction leads to unanswerable questions (P13).

It must at first sight seem peculiar and self-contradictory that the inhomo- geneity of classes which we have postulated as the only admissible deviation from conventional physics should now give rise to regularities (seemingly the very opposite of inhomogeneity). The contradiction is resolved by noting that these two phenomena may occur at different levels of organization. The idea that order at a higher level of organization can be and is superposed upon radical inhomogeneity at a lower-lying level of organization has been intro- duced by a number of outstanding contemporary biologists on the basis of empirical observation (Concepts of Biology, 1958; see, for instance, the quota- tions in Elsasser, 1963). To explain the order at the higher level it would be necessary to study the numerous couplings which exist in an organism between the different levels of organization; it would then appear that what happens at the higher level cannot be explained without relating it to the events at the lower level. At the latter we find radical inhomogeneity which, in the molecular case, corresponds to a set of immensely rare events in the phase space of the corresponding class. We summarize this :

P21. The existence of organismic regularities or organismic order is based upon couplings between different levels of organization. The effort to explain organismic regularities wholly in terms of physical laws is ultimately stopped by radical inhomogeneity at some level of organization. The lowest such level is the chemical one where inhomogeneity appears as a set of immensely rare events in the phase space of the corresponding classes.

[As we have pointed out earlier (Elsasser, 1963) the various levels do not always seem to represent a hierarchy corresponding, say, to geometrical size. Thus in the case of evolutionary processes the environment sometimes appears to take the place of the “lower” level of the above discussion.]

So far we have insisted on the dual structure of the organism, as having mechanistic and organismic aspects. It will hardly suffice, however, to assume that the radical inhomogeneity of organisms just “happens”. Instead, it appears necessary to assume that just as organisms have in the course of evolution developed highly complicated and highly efficient mechanistic processes-so organisms have also developed very eficient processes which subserve the appearance and maintenance of radical inhomogeneity (Elsasser, 1962a). We have termed this type of process, ergodization; we have, moreover, pointed out that at the lowest level of organization, the molecular one, the ergodizing processes seem to be intimately connected with the powerful electrical activity of protein molecules. The study of ergodization is one of

SYNOPSIS OF ORGANISMIC THEORY 65

the most urgent wide-open problems of biochemistry and biophysics. Lack of space prevents us from expanding the subject; let us only summarize:

P22. The dynamics of the organism has a dual aspect. It consists of an intricately interwoven combination of mechanistic, physically predictable processes and of ergodizing processes. The latter subserve the generation and maintenance of that radical inhomogeneity which can lead to the existence of organismic regularities at other levels of organization. On the chemical level, the radical inhomogeneity corresponds to the fact that by P9 any system event is an immensely rare occurrence in the phase space of the class.

The concept of organismic behavior and organismic regularity is clearly so general that it would be naive to attempt even a survey or tabulation of organismic phenomena at this place. Instead, we shall confine ourselves to a very brief discussion of just one exceptionally important case; that is the relation of mechanistic and organismic components in heredity.

The reductionist view of heredity is well-known. It claims that there is complete “information storage” amounting to a description of the adult organism, in the germ cell, after the manner of an automaton. This view goes back to the beginning of the eighteenth century when it was known as preformationism. Equally old is the view that developmental processes are autonomous and cannot be reduced to the “reading out” of stored informa- tion by an automaton; this is known as the theory of epigenesis. It is unfortu- nately true that the latter theory has never been given a quantitative form. Instead, we have the consistent statement of the overwhelming majority of those who have studied or are studying various aspects of developmental processes (growth, embryology, transplantation experiments, etology, etc.) that developmental processes are altogether sui generis and cannot be reduced to mechanistic functions.

Remarkably enough, the controversy between epigenesis and preformation is just about as old as the controversy between the wave or corpuscular nature of light; the former pervades the history of biology as the latter pervades the history of physics. But whereas in physics a synthesis of these opposing concepts on a higher level of abstraction has been achieved by quantum mechanics, a similar advance has not yet been accomplished in biology. It is the purpose of organismic theory to initiate such a synthesis.

The tremendous recent progress in our understanding of the mechanistic aspects of these problems comes, of course, to mind. The well-known DNA-RNA-protein system constitutes an intricate mechanism which deter- mines the structure of newly formed protein. This mechanism guarantees the constancy of the species-specific proteins in mitosis and hence also in developmental processes. These things are so well-known that we do not need to enter into them more closely. On the other hand, there is no evidence

T.B. 5

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for a complete storage mechanism for innumerable other, especially the more complicated, morphological features. At this point we must strongly empha- size the essentially empirical character of epigenetic views. If these views mean anything in quantitative terms, they can only mean that there is not a com- plete “message” in the germ cell, spelling out every detail of the future organism. On the other hand, genetics shows a very clearcut relationship between changes induced in the chromosomes and changes in the morphology or physiology of the adult organisms. Thus, once more, if epigenesis desig- nates anything specific quantitatively, it does mean that information storage is only partial; perhaps, and especially in the case of more complicated organisms, only very fractional (Elsasser, 1958).

To recapitulate, in empirical biology the chief inconsistencies at this time lie in the contradictions between the reductionist and the epigenetic view- points, both based on observational data. Organismic theory furnishes the basic abstract tools for the synthesis. It does this by the introduction of the concept of a FUD and by the assumption that organisms form radically inhomogeneous classes within such a FUD. The theory.withdraws the basis from exaggerated reductionism by denying the unlimited repeatability of sets of experiments; at the same time the structure of the theory guarantees that the laws of quantum physics and the second law of thermodynamics hold in any possible sets of laboratory experiments.

Let us sum up here two areas of contact with rather concrete problems. The first concerns the existence of processes or mechanisms of ergodization. Their function is to reduce physical predictability with respect to certain aspects of the living tissue (the organismic ones) while at the same time other functions (the mechanistic ones) remain highly predictable over some periods of time. The predictability of the mechanistic processes may in many cases ultimately be limited only by their inevitable coupling with the ergodizing functions. The more detailed experimental demonstration of the latter func- tions should be a difficult and time-consuming but by no means an impossible task of biochemical and biophysical investigation (see Elsasser, 1962a).

A second area of contact with the phenomena lies in the distinction between total and partial storage of information. Since this distinction has hardly been made by biologists in a clearcut fashion in the past, a renewed approach to biological interpretation from this viewpoint will yield new and valuable results. Here, we wish to dwell on a point of great theoretical interest, namely, that the reproduction of a structure from partial information is one task which is essentially beyond the capacity of any automaton. (In speaking of partial information we do not mean, of course, merely the elimination of redundancy; we mean, instead, a further significant reduction of the informa- tion after all the redundancy has already been removed.) There has been some

SYNOPSIS OF ORGANISMIC THEORY 67

argument as to whether automata can be creative, that is, generate novel information by trial-and-error methods. This may or may not be true; but there can be no doubt about the fact that an automaton cannot reproduce a structure on a sharply fixed time schedule (characteristic of developmental processes) and with a rather modest rate of incidence of error (also charac- teristic of normal development) unless the automaton has all the requisite information available in stored form. Thus if the epigenetic view is taken seriously, the inadequacy of the reductionist view and the importance of organismic components in development are undeniable. These questions, which are here only hinted at, deserve much further elaboration; they have to a limited extent been dealt with previously by the author (Elsasser, 1958, 1961) as has been the related and equally intriguing problem of the possibility of only partial information storage in the case of conventional (cerebral) memory.

In conclusion, we shall comment again about the relationship between organisms and automata. Certainly, automata theory has greatly advanced biological thinking; disconnected observations and theories have fallen into place; phenomena like homeostasis and nervous reflexes have been subsumed under the common formal principle of feedback, and so on. Now according to Al7 there is no sharply delineated difference between automata on the one hand and organisms on the other; such differences as exist arise essentially out of the tremendously higher degree of complexity and inhomogeneity which organisms can possess as compared to man-made automata, and out of the possible consequences of this inhomogeneity in a FUD. The latter amounts to a denial of the unbounded repeatability of experimentation with any physical system whatever. While this, as we have seen, introduces some radical changes into the interpretation of natural phenomena so far as they pertain to biology, the great advances which have been made by the intro- duction of automata theory can be retained without any restriction in organismic theory.

REFERENCES BRILLOUIN, L. (1962). “Science and Information Theory”, Second ed. New York: Academic

Press. BOHR, N. (1933). Nature, 131, 421, 457. BOHR, N. (1958). “Atomic Physics and Human Knowledge”. New York: John Wiley. Concep@ of Biology (1958). R. W. Gerard, ed., J. Behavioral Sci. 3, 93-215; also available

as Publication 560, Nat. Acad. of %.-Nat. Research Council, Washington 25, D.C. ELSASSER, W. M. (1958). “The Physical Foundation of Biology”. London and New York:

Pergamon Press. ELSASSER, W. M. (1961). J. Theoret. Biol. 1, 27. ELSASSER, W. M. (1962a). J. Theoret. Biol. 3, 164. ELSASSER, W. M. (1962b). 2.f. Phys. 171,66. ELSASSER, W. M. (1963). J. Theoret. Biol. 4, 166. TISZA, L. (1963). Revs. Mod. Phys. 35, 151.

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