Download Synopsis of Organismic Theory

J. Theoret. Biol. (1964) 7, 53-61
Synopsis of Organismic
WALTER
Department
of Geology, Princeton
M.
Theory
ELSASSER
University, Princeton,
(Received 1 July 1963, and in revisedform
New Jersey, U.S.A.
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, assumptions and propositions, with connecting and explanatory text. Organismic
theory is not identical with biological theory; it is a formal scheme prerequisite 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 inhomogeneity 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 simplification 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 astrophysics 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 observations, 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
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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 homogeneous 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
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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.
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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 idealization 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 concerned 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 oneclass. 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 description 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. asymmetrically 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 tremendous 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
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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 concentrates 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 significantly 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 inhomogeneity 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 determination 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 perturbations in the object which become very severe if the measurements are thoroughgoing 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 perturbations 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 quantitative 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:
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AH. All organisms have component processes which can be described
adequately in terms of homogeneous classes. These processes can be reproduced 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 generalizations 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 inhomogeneity 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”
abandoned.
used earlier by the author (Elsasser, 1958) has been
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be reduced to physical laws, nor can a contradiction to the latter be constructed, 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 inhomogeneity 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 introduced by a number of outstanding contemporary biologists on the basis of
empirical observation (Concepts of Biology, 1958; see, for instance, the quotations 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
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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 information by an automaton; this is known as the theory of epigenesis. It is unfortunately 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 determines 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
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M.
ELSASSER
for a complete storage mechanism for innumerable other, especially the more
complicated, morphological features. At this point we must strongly emphasize the essentially empirical character of epigenetic views. If these views mean
anything in quantitative terms, they can only mean that there is not a complete “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 designates 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 viewpoints, 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 functions 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 information after all the redundancy has already been removed.) There has been some
SYNOPSIS
OF ORGANISMIC
67
THEORY
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 characteristic 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 introduction of automata theory can be retained without any restriction in
organismic theory.
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