Biological Causation

Biological Causation
Author(s): Ralph S. Lillie

Source: Philosophy of Science, Vol. 7, No. 3 (Jul., 1940), pp. 314-336
Published by: The University of Chicago Press on behalf of the Philosophy of Science Association
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Biological Causation
BY
RALPH S. LILLIE
T WOULD appear that among scientific men
discussion of the general principles of natural
[ [k. science (including the problem of causation) has,
on the whole, proved more congenial to mathe-
maticians (with their brothers-in-arms the logi-
cians) and physicists than to biologists. Just
why this should be so might be difficult to explain or justify.
But one reason seems to lie in the comparative ambiguity of the
concept of causation in biology. In general, the term causation
has been used in science to designate the special role of active
factors, rather than of passive or stable factors (more or less
permanent "conditions"), in the determination of single events.
By active factors we mean those which involve physical change,
typically associated with transfer of energy; these are distin-
guished from stable factors or invariants which persist unchanged
throughout the process under consideration. Thus in the classi-
cal isolated system, the total energy represents a stable factor
which remains the same through all transformations of the system.
The energy changes its form, potential or distribution, but not
its total quantity. This rule of conservation defines a static
condition persisting as a limiting factor through any case of
change. Similarly, the permanent or unchanging factor in a
machine consists in the stable properties and structural intercon-
nections of its parts; together those constitute an invariant which
fixes definitely the possible range of activity. Activity itself
requires flow of energy; strictly speaking, a causal factor is not a
314

R. S. Lillie 315
static factor but is a change of some kind; releasing events ("trig-
ger action") are included under causal events since they introduce
factors without which change does not occur.
Observation shows that similar events, acting as "causes,"
may have very different effects under different static "condi-
tions." In many natural processes, however, active and static
factors cannot be sharply distinguished, but shade into one an-
other imperceptibly; and this is especially true of living organisms.
Here structural complexity (organization) becomes a factor of
crucial importance in the special activity exhibited by the system.
Thus the causes of a man's behavior are often said to lie in his
"character," meaning by this term a set of constant determinative
factors inherent in his special personality; these presuppose spe-
cial structural and physiological factors, including (e.g.) features
of nervous organization. But investigation shows that these con-
ditions are not static; they fall rather in the "dynamic" class;
i.e., they are kept at a stationary level by the continual activity
of the organism itself. It is a case of a "steady state,"' as dis-
tinguished from a static equilibrium; there is a balance of opposed
processes. Nevertheless, this steady state, although actively
maintained, is itself a constant factor in the determination of
activity; it presupposes a chain of active causation leading back
to elementary processes of metabolism or beyond. Active causa-
tion and static conditions thus become merged and indistin-
guishable.
This is one of many sources of ambiguity. We ask the ques-
tion, "why" does an animal behave thus and so? Quite evidently
to the naturalist the living organism is a physical system, subject
to all the conditions of physical existence. Yet to ascribe its
behavior to physical conditions alone appears insufficient; it is
more realistic to say that the physical conditions limit but do not
completely determine the special activity. The main reason for
this reservation is that in higher animals (perhaps in all) there is
1 For a recent discussion of the steady state, with special reference to biological con-
ditions, cf. the recent paper of Alan C. Burton, "The Properties of the Steady State
Compared to Those of Equilibrium...." Journal of Cellular and Comparative Physi-
ology, I939, vol. 14, p. 327.

316 Biological Causation
the additional factor of inner or psychological(volitional) deter-
mination. Although this is not physically (externally) observ-
able, its presence is an inescapablenatural fact, in ourselves at
least. Accordingly,animal behavior, while unavoidably bound
up with physical (physiological)conditions, is seen to have its
own special factorsof determinationcomingunder categoriesof a
differenttype, the psychological. Behaviorismgives only a par-
tial account of animal activity, and at present general concepts
of the living organismtend to be based quite as fully on psycho-
logical as on physiological considerations.2 The higher animal
is more realistically described as a psycho-physicalentity. To
certain features of activity we assign a predominantlyphysical,
to others a predominantlypsychical determination;but there is
always the difficulty of isolating the two. While making the
best of different types of abstractionism,we need be under no
delusionas to theiressentiallypartialnature. Apparentlya more
comprehensiveconceptionof biologicalcausationis required.
Causationis a less ambiguousconcept in the physical sciences,
althougheven hereits workingvalue is often questioned. Never-
theless its complete rejection is scarcely possible, since the natural
fact of causal interconnectionbetween events is always present.
Apparently complete isolation is not possible to single natural
events; thereis always the phenomenonof fringesor raggededges,
in the temporal as well as in the spatial sphere. Since events
mergeinto one another,in this sense, it seems necessaryto ascribe
coherenceor unitary character to nature as a whole (monism).
However this may be, it remains characteristicof the physical
sciences, as comparedwith biology, that clarity and unambiguity
of statement and formulationare on the whole easier to attain.
Part of this advantage comes from the fact that physics deals
exclusivelywith the phenomenaof an externalizedor publicworld.
Whena physicalevent happens,it can usually be definitelyrelated
to other observableevents or conditionsin its immediateenviron-
ment. It is "dependent" (in part) on these events and condi-
tions. At least only potentially observable events and condi-
' Cf. e.g. the recent book of K. Goldstein,"The Organism,"New York, American
Book Co., 1939.

R. S. Lillie 3 17
tions are usually taken into account.3 Unequivocality of state-
ment is attainable within a relatively narrow margin of uncer-
tainty. Accordingly, the correspondence between the observed
features of physical events and their pictorial or symbolic repre-
sentation (e.g., in the sign language of mathematics) may be
highly exact, or at least exact enough for most descriptive or
practical purposes.
If we confine our attention to the external world, we observe
in general that events which are spatially and temporally con-
tiguous are interdependent in the sense that the preceding event
determines in an unequivocal way certain definite features of the
immediately succeeding event. When a bat hits a ball, there is
"causal" interconnection; and such interconnection of events is a
constant feature of the physical world. The interconnection
involves change; i.e., it is in time as well as in space; one event
precedes and determines the other and is influenced by it; the
influence is reciprocal. It is important to note that spatial and
temporal contiguity between cause and effect is always required.3
Things act here and now; i.e., in the geometrical representation
there is intersection of the space-time coordinates of the two
events. If a distant event influences one near at hand, it is only
through the intermediacy of radiation or some other type of
physical connection (mechanical, electrical, gravitational) which
establishes continuity between the events.4 Events may occur
at the same time in different spaces, or in the same space at dif-
ferent times, without influencing one another; but when both spaces
and times coincide, the interrelation termed causal inevitably
enters. The preceding event then determines in a definite way
certain characters (not all) of the immediately succeeding event.
s It should be noted, however, in consideringthe totality of the conditionsdeter-
miningan event, that part of these conditionsare not environmentalbut are internalto
the event itself. Also in their very nature these may not all be open to external ob.
servation. This is a fact of special importancein biologicalcausation(see below).
A requirementemphasizedby Hume. Cf. the recent article in this Journal by
R. B. Winn, "The Nature of Causation," vol. 7, I940, p. I92.
4This is the conditionwhichNewton perceivedclearlyin his requirementof a medium
for gravitationalaction.

Interconnection of this type pervades nature and may be de-
scribed as its causal structure.4a
Now living organisms are pre-eminently causal systems; their
continued existence and the stability of their properties and activi-
ties depend upon the precision with which definite "causes" pro-
duce their constant or invariant "effects." It is by virtue of this
causal interconnection of its component events that any organism
preserves its stability or "survives" in its natural environment.
We may note here (incidentally) that the rule of contiguity
imposes definite upper limits on the size of living organisms, for
the reason that stability requires the causal interlocking of a
multiplicity of diverse events, and this is possible only within
certain limits of dimensions. In any single organism the causal
cross-connections are most complex; yet this complexity is per-
fectly compatible with stability, as is shown by the continuance
of life and its activities. Accordingly, living organisms combine
with the maximum complexity of physical constitution an equally
great complexity of activity. This complexity is conjoined with
an equal degree of regularity; constancy and reproducibility are
as characteristic of vital as of non-vital activities. But this fact
is in itself not surprising, for it is just this constancy which is the
essential feature of causation as a universal natural factor; other-
wise the existence of such complex causal systems as living organ-
isms would not be possible. It seems fair to say that in living
organisms causation exhibits the greatest precision of which it is
capable; any complex acquired skill, like the art of the violinist, is
an instance of this precision; in the field of heredity the resem-
blances between identical twins (or identical quintuplets) give
illustrations of the exactness with which complex chains and net-
works of causation may be repeated. The regular features of
causation are thus responsible for the stable side of organic life.
The spontaneous or apparently indeterminate actions of higher
animals undoubtedly require a different reference, as will be indi-
cated later; but at present we are concerned with causation in its
regular or routine aspects only.
4^This phrase, "causal structure," is also used by H. J. Jordan, Acta Biotheoretica,
I935, vol. I, p. I00.

R. S. Lillie 319
The biologicalquestion is: what gives coherenceto the multi-
plicity of events in the system and confersunity on the organismic
sequence,consideredas a whole? This is the problemof integra-
tion, whichmust be regardedas the essentialproblemof biological
causation.
In biological causation we see natural activity as conditioned
by the maximumof structuralcomplexity. Structureis so diver-
sified and complex, and extends so far into the microscopicand
ultramicroscopicfields, that continual expenditureof energy is
required to maintain it. In general we may say that thermo-
dynamic conditionsresist fine-graineddiversificationin predomi-
nately fluid systems, i.e., the automatic tendency is toward uni-
form distribution of energy and components. Yet the fluid
protoplasm,instead of becomingmore homogeneous,retains its
diversity or even exhibits (especiallyin development)a tendency
to increase of structural and other complexity. Within the
frameworkof this structure, there is correspondinglycomplex
activity, directed, controlled and unified in a manner largely
determinedby the special peculiaritiesof the structure.
The general realistic or naturalisticsignificanceto be ascribed
to the term structure should first be considered. Structure
implies complicationor compositecharacter,combinedwith sta-
bility. A structuralentity is one which is internally diversified
(differentiated),and its type of inner diversity has a certain
degree of persistence. Usually by the term "structure," as ap-
plied to objects in the external world, we mean stable spatial
arrangementsof "matter"; and biologists use the term "proto-
plasm"in referenceto the structuralsubstratumof vital processes.
This substratumis stable in the sense that it preservesa "steady
state"; actually it is the seat of continual chemical change
(metabolism);duringlife opposedchemicalprocesses(anabolism
and catabolism) are so balanced (normally) as to maintain a
nearly constant condition (the steady state) of the system, to
whichcorrespondsa constantor specificstructure,i.e., distribution
of components;with this are associatedconstant propertiesand
activities. Living protoplasmis thus conceived as a flux whose
stability dependson the constancy not only of its chemicalcon-

stituents, but also of their special modes of activity and inter-
action; i.e., kinetic as well as static constants are concerned. An
important part of the problemof biologicalcausation has refer-
ence to the conditionsdeterminingthe constant physical proper-
ties of the flux.
First we note that this constancy presupposes constant proper-
ties in the elementary units (atoms and molecules) composing the
system. Hence, any kind of matter, living or non-living, owes
to its atomic constitution the fact that it is itself stable and also
permitsof stable arrangements.4bThe "definition"of matter, in
fact, is just this; material systems are those entities which are
localized in external space and persist in time for a longer or
shorter period, during which their essential properties remain
unchanged. According to the prevailing physical conceptions,
matter as well as energy is subject to the rule of conservation.
The changes in propertieswhich we observe are referred,scien-
tifically, to changes in the arrangementof these stable atomic
components. These changes are associated with transfers of
energy, either between the atomic units themselves (involving,
e.g., changesof chemicalcombination)or to the surroundingsas
radiation. The transfersof energyoccurin units (quanta)having
certainconstant or stable characteristics,e.g., a constant relation
to wave-length.
In the foregoingbrief discussionemphasis has been placed on
factors of stability. Such reference to stability is inseparable
fromany kind of scientificconsideration. This is seen especially
in the mathematicalcharacterwhich all highly developedsciences
tend to assume. Any mathematical formulation presupposes
stability or constancy in the significanceattached to terms and
symbols;and a correspondingconstancyis ascribedto the natural
entities or processesthus symbolized. Mathematicalrepresenta-
bility is a correlateof regularity,i.e., of formalstability. Another
considerationof broadphilosophicalbearingalso requiresempha-
sis at this point, in referenceto the highly individualizedcharacter
which biologicalorganismsexhibit, especially in their higher de-
b Note that underlyingthe atomicstabilityareother kindsof stability,e.g., electronic
charge,quantionsconstant, and so on.

R. S. Lillie 321
velopments. Any single stable entity, say an atom, illustrates

what we call "individuality"; i.e., it persists as a separate unit and
has its own singularity or uniqueness of properties, position and
activity. Individuality* thus implies a certain degree of isolation;
each individual entity has a special character of its own, shown
in its separateness from other entities and in its persistence; thus
at any time each atom has the exclusive occupancy of a certain
region of space and resists encroachment on that region, i.e., it
displaces or excludes other similar entities. In physics this kind
of interaction is conceived as a manifestation of "force"; and in
modern physics energy has come to be regarded as the physical
ultimate; each material particle represents a stable local concen-
tration of energy, and any final distinctions between matter and
radiation disappear. Energy, capability of effecting change, is
the essential; the chemical atoms of the periodic system are
regarded as structured systems, consisting of subatomic energetic
units (protons, electrons, neutrons, etc.), each with stable and
characteristic properties, both static and active. Each unit is an
"endurant",6 having its definite mass, and to a definite mass
corresponds a definite energy, as represented by Einstein's rela-
tivistic equation7 of the inertial value of energy, E = mc2.
Energy (equivalent to actual or potential activity), mainly con-
centrated in atomic units of definite pattern and transmitted
according to definite laws, thus appears as the fundamental natu-
ral reality. At any time this energy is chiefly intra-atomic and
manifests itself as mass, although a part may be in process of
"The word is etymologicallyequivalent to non-divisibility,or non-disintegrability,

correspondingto a certain kind of stability. Note also the psychologicalanalogy of
selfhood (psychic isolation). Obviouslythere is a great gap between the individuality
of an atom and that of a human being, but both types illustrate, in their close com-
binationor interfusionof active and static (= structuralor formal)characters,as well
as in their isolation,the same generalprincipleof naturalindividuation.
Or "continuant." Cf. R. W. Sellars, "The Philosophyof Physical Realism," pp.
301 ff.
7 Cf. (e.g.) Lindsayand Margenau,"Foundationsof Physics,"New York, 1936,p. 35I.
E is total energy,m stationary mass,c velocityof light. In a moving body E - mcs+
1/2 mv2,where v is the velocity of motionof the body. A simplecalculationshows that
for a gram mass moving at high projectilevelocity, one kilometerper second, the ratio
of the intrinsicenergyto the ordinarykineticenergywould be i810to i.

transfer between separate atoms, by radiation or impact. It is
important to note that of the total energy in any material system,
by far the greater part is intrinsic and non-interchangeable; i.e.,
is concentrated in the stable atomic units and their aggregates.
Thermal or other interchange of energy between atomic units is
in quanta, the energy value of which is relatively very small.
Such interchange is typically reversible and does not alter the
essential character of the atoms concerned. Hence the stability
of an atomic system is typically very great, even if not absolute.
As a rule, a large addition of energy is required to disrupt or
transform an atom, as in induced radio-activity or the artificial
formation of isotopes; and relatively few naturally occurring
atoms show appreciable spontaneous disintegration.
According to the subdivisional analysis of physics, all naturally
occurring material systems, including living organisms, consist of
innumerable such localized centers of activity; these are asso-
ciated in various ways to form a hierarchical series of unified and
stable entities, each higher or more complex entity being built
up of simpler entities, with living organisms at the summit of the
hierarchy. General characterizations of this kind, i.e., of differ-
ent "levels of organization," are familiar to most naturalists.
What is particularly to be noted at present is that just as it is the
inner (or intrinsic) activity of the atomic or sub-atomic units
which determines their special properties, so, derivatively (since
the higher units are composites of the lower), an ultimate reference
to intra-atomic activities is implied in any analytical attempt to
assign final physical grounds to vital phenomena.8 But the im-
portant fact for biology is that these activities, whose ultimate
source of energy we thus recognize as intra-atomic, show in their
aggregate, as embodied in individual living beings, such an amaz-
ingly high degree of unification or integration. Each living
organism, even if analyzable into sub-atomic units and their inter-
connections, preserves while living its special structural unity and
is the center and condition of a special kind of unified activity.
To the human observer, this activity appears as having both
physical (external) and psychical (internal) aspects.
8
Compare the remarks of Niels Bohr, "Causality and Complementarity," Philosophy
of Science, 1937, vol. 4, p. 289; cf. p. 295.

R. S. Lillie 323
Analysis reveals the composite character of the living system,
but gives an unsatisfactory account of its unity. In general, the
insufficiency of analytic method as a means of scientific charac-
terization is more evident in biology than in the physical sciences.
Our present question is: How much light on the problem of bio-
logical causation and integration may be expected from analysis,
considered alone? and at what point does synthesis, both in rea-
soning and in experimentation, become necessary? Since the
living organism is the most complex and highly unified product of
synthesis found in nature, it would seem inevitable that biology,
as it advances, must become more and more completely a science
of synthesis. Already a large part of biological science, especially
on its theoretical side, consists of reconstruction or attempts at
reconstruction. Synthetic procedures are also highly developed
in other sciences, especially chemistry and the technologies. But
what gives the living organism its special place among natural
systems is that its very existence depends upon the continual
activity of synthesis. Always in its chemistry we find the same
combination of analytic and synthetic reaction-cycles; on the
balanced inter-connection of anabolism and catabolism the con-
tinuance of life depends. Thus in the elementary processes of
nutrition, each organism or each cell first breaks down its food
materials into simpler stable components, and then recombines
these into other compounds of complex and specific configurations
of which the chief are the proteins. The synthetic recombination
of elements isolated by analysis thus appears as the fundamental
physical feature of vital action. This synthesis, if it is to have
constant character, must be based on constancy in the properties
of the elementary units which are thus isolated, as well as on
constancy in their modes of activity and interaction (velocity con-
stants and other kinetic constants). On these characteristic
constants, static and kinetic, the constancy or stability of the sys-
tem as a whole depends. Accordingly constants of synthesis
must also be assumed (constants of emergence). Vital action, at
least on its routine side, is based on the constancy of these syn-
thetic processes. The question is whether the constants of syn-
thesis, whose existence we thus seem compelled to recognize, can
be referred in all their characteristics to constants of the kind

alreadyknownand formulatedin the physicalsciences,or whether
constants of a natureother than physical are also to be assumed.
Let us now turn to a brief considerationof the psychological
charactersof living organisms. Synthesisis also the predominant
feature of mental life, as normally experiencedin each human
being. This is perhapsbest seen in everyday visual experience,
where (e.g.) colored patches, or "sensa," presented in scattered
form in the visual field, are recombinedin consciousnessto form
a unifiedmental imageor "Gestalt."8a There is somethingmore
here than a mere analogy to the general physiologicalconditions
just reviewed. The psychologicalunificationexperiencedin the
Gestalt has its physiologicalcorrelatein the unificationof bodily
activity as observed in the response associated with the experi-
ence, e.g., the reaction of pursuit when the animal "sees" its
prey. In this case, the psychologicaland the physiological in-
tegrationscannot be separatedin reality; they must be regarded
as two aspects of one and the same biologicalprocessof response
or adjustment. In other words, the automatic action of the
psyche in unifying its data shouldbe considerednot as something
without physical analogy-peculiar to "mind" as such-but as
one form of the general biologicalactivity of integration. The
psychic unification represents the subjective or inner aspect of
the organismic unification. The synthetic action of mind ex-
hibits itself as one expressionof a unitary property or activity,
uniquely characteristicof living beings, which expresses itself
also in the integrative side of their physical (physiological)life.
We have already pointed out that the concept of integration in
biology is closely allied to the concept of individuation;thus in
embryonicdevelopment,the living individualwith its automati-
cally maintained unity of mental and physical life comes into
existence as a productof the orderlyassociationof materialsand
energies taken from the environment. Living organisms,
whether we consider them as physical or as psychical entities,
always exhibit this integrative character;they are synthesizing
agents, in which special physical and psychicalcharactersemerge
as the outcome of an orderly processof synthesis.
8 Similarlytemporalsuccessionsof auditory sensa are combinedto formwords,sen-
tences, rhythms,melodies,etc.

R. S. Lillie 325
As in the case of physical or physiological constants, the ex-
istence of a system of psychic constants is implied when the
organism is considered from the point of view of a scientific psy-
chology, i.e., as a psychical rather than a physical entity. Con-
stancy or stability in the constitutive elements of psychic life
must be assumed to underlie stability in the psychic life as a
whole. In general this principle is clearly recognized by psy-
chologists of the analytic school (Freud, Yung, and their suc-
cessors), with their assumption of "unconscious" mental elements
or factors. But the problem of how these psychic elements
are to be conceived, scientifically, presents great difficulties.
Doctrines of psychological atomism have not commanded the
same assent in psychology as atomic theories in physics. Ac-
tually, most modern psychologists, in their scientific search for
stable foundations, have tended to regard the underlying stable
determinants of psychic life as primarily physical in nature-
i.e., to picture them after physical rather than psychical models.
This tendency is reflected in their interest in the physiological
aspects of animal behavior-even though this behavior is usually
considered in its aspect as unified activity of the entire organism,
i.e., as a whole rather than partitively. Such preference for
physical types of explanation is understandable, since exact
scientific consideration, qua scientific, requires representation in
terms of constants,-i.e., of elements which are unambiguous
and whose stability can be relied upon. The conscious elements
of psychic life, as these are directly experienced (sensations,
images, volitions, qualities, etc.) are too transient or unstable to
serve as a permanent basis for synthesis. They disappear in
sleep, inattention, or anaesthesia; while physical life persists, and,
so far as we can observe, obeys all the physical rules of con-
servation, independently of the presence of consciousness.
Experimentally we find psychic life to be dependent on bodily life,
which has a firm basis in the physics of the organism. The
stability required by psychology, as natural science, would thus
seem to be found only in the physical substratum of the mental.
Physical or physiological stability, as we have seen, is based
ultimately on the stability of physical atomic units and their

modes of action and interaction. If conservation, permanency,
invariance are the criteria of scientific reality, the psychic ele-
ments, taken by themselves, would be unreal. But since they
have constant characters, they must be regarded as signs of some-
thing real underlying. Their recurrent character can be ac-
counted for only by assuming the presence of permanent factors
of determination, and it is of these that some scientific account is
required.
There would seem to be a theoretical impasse here. Little is
gained by assigning mental qualities to physical atoms or by as-
suming the existence of atomic psychic units (which have been
called "psychons") which combine to form units of higher men-
tality. Atomic hypotheses of mind are a concession to the in-
veterately logical character of scientific method, which takes
satisfaction in showing that complex and diversified wholes can
be built up by combinations of uniform elements; but in the
psychological field such hypotheses seem insufficient or mis-
applied. No doubt atomism must be regarded as an actuality
in the elementary structure of nature; the physical evidence is
overwhelming on this point. But when we consider impartially
all the characters of living organisms, it seems necessary to supple-
ment the physical atomic theories by theories of a different type,
which take into account the synthetic or individualizing side
shown not only by life, as a special development in nature, but
by other natural processes and in fact by the cosmic process in
its entirety. The scientific analysis of nature into atomic units
is an abstractional procedure, valid in its proper sphere, but not
inconsistent with the further assumption of special unifying or
integrative factors, effective on a larger scale than the atomic,
which act to bind together larger groups of units into coherent
wholes of various kinds. That such wholes are a reality in the
natural world needs no special proof. Heavenly bodies and
nebulae are examples of integration, although of a less highly
organized type than living beings. The significant fact of ob-
servation is that an immense diversity of natural composites
exist, all of which exhibit integration of definite types, exemplified
in numerous similar individuals, and ranging all the way from

R. S. Lillie 327
atoms to living beings. A scientific procedure which is ex-
clusively analytic, will never account for the propertiesof such
wholes, consideredas wholes. Each type has its special indi-
viduality; and the problem arises "why" certain types of in-
dividuationand not others have appearedin the courseof natural
evolution and now occupy a permanentplace in nature.
Here we are brought to the problem of creative evolution,
which has its metaphysical as well as scientific connotations.
Bergson'sdiscussion9is only in part scientific; his universe is a
flux rather than a mosaic, a view implying that the natural
constants are never entirely stable but shift, even though slowly,
in their values. In physical science many constants tend to be
regardedas final, and it may be that their stability is over-rated.
If physical characters which appear stable to observation are
identifiedwith the charactersof purely mathematicalconstants,
which are stable by definition, fallacies in the interpretationof
nature may arise; too rigid concepts of causation, especially of
biological causation, illustrate this danger. It must be remem-
bered that the intrinsic charactersof natural existence remain
what they are; physical description, however exact, can never
be more than partial and approximate;a scientific characteriza-
tion is not a reduplicationbut a simplificationand schematization
of nature. In the present discussion I am adhering to the
scientific point of view; and scientific conceptions of biological
integration must be based on the general verifiable facts of ex-
perience. So far as we can judge from our own experience, a
correlatingor unifying action which imposes a synthetic unity-
an orderly coherence,pattern, or organization-upon otherwise
unorganized data, seems always to have in it (initially at least)l?
somethingof the mental or psychic; the term mental is used here
to includethe volitionalor active as well as the passivelyreceptive
or perceptual qualities of mind. Such action differs from that
found in a purely external or physical world; some factor is
present over and above the physical (while including and con-
9 Henri
Bergson, Creative Evolution, New York, 1911.
10This reservation is made to allow for habit;
synthetic procedures, once discovered,
may become unconscious (or "mechanized") by repetition.

sistent with it), which we may call a synthetic (or creative) fac-
tor; this is directive" and individualizing in its action; apparently
it opposes or counteracts the general thermodynamic trend to-
ward uniformity. As to the general nature or method of this
factor, little can be said: description can be only by analogy.
If the analogy is with voluntary action, we should describe its
actuation as internal rather than external.12 To illustrate:
when an artist shapes a mass of modeling clay into a portrait,
the factors which make possible this outcome are both physical
and mental; i.e., both kinds are observable, at least in part.13
The clay has its constant physico-chemical characters upon which
the artist relies; so also has the physical organism of the artist
himself-that is, his personal activity, with its underlying physi-
cal and physiological constants, is presupposed. Nothing occurs
that is not consistent with the physical conditions, as these are
viewed externally; to an observing physicist (or physiologist)
the whole process would appear as physically "determined."
Nevertheless the precise form which is assumed by the mass of
clay is determined by factors not definable in purely physical
terms; i.e., by the image, ideal, or Gestalt of the artist. This is
the psychic or directive factor which, in this case, is indispensable
for ordered synthesis. But the Gestalt itself is a product of syn-
thesis, some of the factors in which can be observed, others not.
The transformation effected by the artist is accordingly to be
referred finally to causal conditions in which, in addition to a
variety of routine factors acting in accordance with fixed con-
1 For a fuller discussion of directive action cf. my recent paper, "Directive Action
and Life," Philosophy of Science, 1937, vol, 4, p. 202.
2 This would imply in physical analysis that the internal or intrinsic atomic energy
is in some way applied to determine the time, place, and direction in which quanta of
action are transferred.
13 I say "in part," because any natural system of observables is found, when analyzed
closely, to taper off into unobservables,-i.e., physically speaking, into systems consisting
of atomic units which become observable only when they transfer a portion of their own
(or reflected) energy to other complexes of atomic units, viz., those forming part of the
observer's organism. The preponderance of atomic energy remains (in this sense) un-
observable. To suppose that physical unobservables cannot act as determinants in
special events is like supposing that the externally unobservable factors in another
human being (his psychic life, volition, etc.) have no determining effect on his behavior
-which seems an unrealistic assumption!

R. S. Lillie 329
stants, novelty-producing or creative factors play an essential
part.
The analogy of natural creation with human creation has its
limits, and is suggestive rather than demonstrative; but it can
hardly be ignored by a science which aims at giving a comprehen-
sive and realistic account of living beings in all their essential
characters, both physical and psychical. Our illustration shows
that the decisive factors in many instances of biological causation
have an individual (monadic) character, which is additional to
the physical and mathematical factors which also play their
demonstrable part in the physiological organism. Just why
certain special types of individuation, each with its correlative
types of causation,14 have arisen in nature, and how they have
become perpetuated by a stable genetic routine, are problems
which cannot be considered theoretically to any purpose without
reference to the past history of nature. Now the essential
features of nature may be described as activity and process,
conditioned by certain underlying permanencies;15 historical
process, in so far as it is evolution and not merely unprogressive
repetition, is itself largely a process of individuation; and while
individual action necessarily has its foundation in general or
constant factors, its special character as individual cannot be
thus explained. No one would deny the dependence of indi-
vidual development and behavior on the orderly routine of
physical processes which are scientifically definable in large part;
the biological sciences of physiology and genetics are a standing
proof of this. But what must also be recognized is that the
stable, constant or routine elements and processes are not the
only factors to be considered in accounting theoretically for the
existing types of biological individuation and action. A further
reference to special whole-making or integrative factors is ap-
parently required; these express themselves largely by or through
physical modes of causation, but nevertheless have an inde-
pendent character of their own.
14
As will be indicated more fully below, each type of individuation has its own special
types of causation; to give a biological illustration, hormonal causation has reached a
high state of development in vertebrates, but apparently not in insects.
16A. N. Whitehead, "Process and Reality," passim.

According to the analogy just cited these factors, considered
in their individualizingor synthetic character,would appear to
be essentially of the nature of psychic or mental factors. The
special natural quality or characteristicwhich we refer to as
mentality (perhaps rather vaguely) would then be ascribed to
the whole-makingor unifying factor, consideredin its action as a
totality, rather than to the elementary components (such as
atoms) consideredsingly or as aggregates. There is always a
difficultyin conceivingmental life as a merelysummationalprod-
uct. The living organism-as-a-wholeis to be regarded as a
psycho-physical system with factors of both kinds, mental
and physical, neitherof which is completelydefinablein terms of
the other.16 Both kinds are essential, but the special integral
characteror maintainedvital unity would seem largely or mainly
attributable to factors which are not primarily physical but
representa distinguishingfeatureof mind as mind. The mental
unity is sui generis:a purely additive associationof non-mental
elements can never make a mind. Hypothetical psychic ele-
mentary units (e.g., "psychons")are conceivedafter the analogy
of physical units, whose action is additive. But in a psychic
unity, as immediately experienced,the single elements are held
organicallywithin the unity in their own peculiarway; and while
they may be analytically distinguishable,they are not (strictly
speaking) detachableor isolable without change of propertiesin
the same sense in which physicalunits are isolable. The psychic
constants can thus not be conceivedas merelyphysical constants;
they have a unique characterof their own, combining physical
stability with stability of another kind not physical,-which we
may label "psycho-physical"(if we will). As I need hardlypoint
out, the purely physical view of nature is now recognizedto be a
partialor abstractview; but so also is any other kind of scientific
view; and the problem of devising the proper type of abstractions,
which shall do justice to experimental and other fact, and at the
same time be scientifically serviceable (as in giving direction to
investigation), is an especially pressing one for biology. No one
16I.e. the concepts "purely mental" and "purely physical" are abstractional fictions.
Cf. H. Prinzhorn, "Psychotherapy," London, J. Cape, 1932, p. 97, for a fuller discussion.

R. S. Lillie 331
would deny that a large part of biological investigation can be
conducted on the basis of purely physical (physico-chemical)
conceptions and procedure, but this is not true of psychology;
and there are the theoretical and other demands of a general
biology to be met.
Scientific analysis resolves a complex phenomenon into simpler
components and factors which themselves exhibit constant char-
acters and constant types of interrelation, both actual and pos-
sible. Constant combinations or interactions give rise to con-
stant products or effects. To return to the physical point of
view: if we assume that all the materials and substances (chemi-
cal elements and compounds) entering into the composition of
the living organism have their constant properties, we may con-
sider analytically the vital organization as including basic struc-
tural features of two kinds: (I) static or purely spatial arrange-
ment of components (pure morphology), and (2) constant modes
of activity or inter-action of components; the latter may be called
temporal structure. The general concept of structure would
thus include all constant modes of spatio-temporal relationship
between components. In other words, the stable interconnec-
tions of components or events might be either spatial or temporal
(or both); the corresponding constants would be respectively
static and dynamic.17 In the latter case, sequences of activity,
having their characteristic constants, intersect and influence one
another in constant ways; and this type of temporal structure,
in which the component events (simultaneous or successive)
are definitely interdependent or interlocked, is equivalent to
causal structure. On causal structure of this kind, all biological
activity, as shown in physiological process or behavior, depends.
It is mainly because of this temporally interlocked character of
its separate processes that the living system, considered as a
whole, preserves its stability in nature. Failure or derangement
in any one member of the causally interlocked system may make
integration impossible; disintegration, or death, then follows.
17 The distinction between chemical "statics" and "dynamics" is familiar, referring
respectively to conditions of equilibrium and conditions of chemical change. (Cf. such
a work as Mellor's "Chemical Statics and Dynamics.")

How completely the living system is dependent on the stability
of its kineticconstantsmay be realizedif one imagineswhat would
happen if the relative chemical reaction-velocitiesin any cell
were suddenly to change. The disruptive effects of high tem-
peraturesor poisons (such as cyanide) are in fact generally at-
tributed by physiologists to derangementsof this type. It is
because of the stability of these constants that each living
organismshows constancy or reproducibilityin its physiological
or behavioristic as well as in its morphologicalcharacters.
In biology the ultimate physical basis of the organismic
stability is usually referredto the germinal constitution. Such
referenceaccounts logically for the fact that individuals of the
same species of animal or plant are similarin structureand type
of behavior, since they start their development from germs of
similar constitution and grow up under similar conditions. The
branch of biology called genetics is the scientific counterpartof
this conception of the living organism as not merely a static
system, complete at a given time, but as a process. Such a con-
ception does not imply lack of stable structure-quite the con-
trary; but this structure is conceived as a stable frameworkof
interrelationswhich has its temporal as well as its spatial side;
i.e., in the geometric sense it is four-dimensional. At the same
time, the system, consideredas a whole, exhibits a characteristic
unity and stability, in the sense of persistenceand reproducibility
of both active and static characters.18
Is it scientifically sufficient to assign this reproducibilityto
the existence of a static spatial structure or morphological
organization in the germ, as represented, e.g., in the specific
stereochemical configurationof certain proteins in the genes,
having a determinativerelation to development? There would
seem to be no doubt that a constant stereo-structureof the
nuclearproteinsis a necessaryfactor (somewhatin the sense of a
"governor"or stabilizer); but apparently this structure is not a
18 I referhere
particularlyto the fact that each cycle of development,as exhibitedin
different species of animal and plant, has its constant and specific characters. This
problemis discussedat some length in my recent address,"The Nature of Organizing
Action," Amer. Naturalist, 1938, vol. 72, p. 389.

R. S. Lillie 333
merely static one like the arrangement of parts in a machine, but
is actively maintained, by a metabolic balance between break-
down and synthesis. It represents a "steady state" rather than
a static equilibrium. All proteins taking an active part in proto-
plasmic processes are subject to continual breakdown, as is
shown by the general facts of nitrogen metabolism. They must,
therefore, be in continual process of replacement; i.e., the main-
tenance of molecular as well as of cellular and anatomical struc-
ture requires constant activity, with a corresponding constant
expenditure of energy. We come again to the essential fact (so
characteristic of living organisms) of a stability which is directly
dependent on activity and lapses when activity ceases. The
stream of activity must be kept above a certain level, otherwise
the system disintegrates or "dies." But so long as this level is
maintained, a correlative spatial structure, or distribution of
components, remains present, and this structure becomes itself a
constant factor determining the course of further activity in the
system.
In the last analysis the constancy of biological activity depends
on the constancy of the fundamental physical factors, both
static and kinetic. It has already been pointed out that the
maintenance of any activity at a constant level presupposes
constancy of the static conditions under which the activity pro-
ceeds. A corollary is that the kinetic constants which determine
the rates of elementary natural processes must have as their
background or stable condition a corresponding system of static
constants. The nature of these is fixed by the general cosmic
stability. Such general considerations are far from irrelevant
here, since they have an intimate bearing on the problem of evolu-
tion, an essentially biological problem. If processes are to be
exactly repeated, as is required if a progressively evolving rather
than a chaotic universe is to result, the conditions'under which
they occur must remain constant. Some definite relation of
interdependence must therefore exist between the fundamental
kinetic constants (like the velocity of light) and the fundamental
static constants (like the gravitational or inertial constants).
But just what this relation is has never been clearly defined by

physicists, so far as I am aware.l18 The existence of these con-
stants points to the further existence of some stable uniform
substratum or background to natural processes in general; if
such a stable physical medium does in "fact" exist, the several
properties of this matrix must be quantitatively interrelated;
and the fundamentalconstants of action, like the quantum con-
stant, must have a definiteplace in the same system of relations.19
The special problem of biological causation is a part of the
more general problem of "why" certain types of activity have
appeared in the course of natural evolution, and why, having
appeared,they persist. But the problemof origins is the prob-
lem of novelty or creativity. The appearanceof the genuinely
novel in the natural world means that something has occurred
which is not entirely referableto the establishedsystem of con-
stants. Science, which proceeds by rule, is at a loss when it
attempts to account for phenomena which involve departure
fromrule. Such departures,however,may give rise to new rules;
new constants of action, i.e. of causation, may appear in newly
1a Apparently Eddington believes this and similar relations are mathematically
deducible (cf. his recent Tamer Lectures, "The Philosophy of Physical Science,"
Cambridge University Press, 1939), but Einstein expresses himself with reservations
(Address before the Eighth American Scientific Congress, Washington, May I5, I940,
printed in Science, I940, vol. 91, p. 487).
19The hypothesis of an all-pervading uniform medium or ether has lately been dis-
credited as unnecessary by many physicists, under the influence of relativity theory;
but the question as to its physical existence or non-existence would seem after all to be
a matter of experimental evidence. As a naturalist, I do not quite see how we can
dispense with an ether, if by the term we mean some homogeneous neutral background,
matrix or container which provides the static conditions for the observed regularity of
physical action occurring within it. Such a concept corresponds closely with the New-
tonian concept of "empty" space. Of course the question of how its geometrical char-
acters are best represented is one for the mathematical physicists to decide. But the
fact that valid calculations can be made, without considering at the time the physical
nature of the conditions assumed as constant in the calculation, need not mean that these
conditions have in themselves nothing corresponding to physical existence. Facts like the
stable setting of the stars and nebulae, experiments like the Foucault pendulum and
others, were formerly thought to indicate the existence everywhere of a uniform medium
with fixed coordinates. Opinions differ as to the theoretical significance of the Michelson-
Morley experiment-and even (it seems) as to the facts. Recently the question of the
physical existence of an ether has been raised again, on purely physical grounds, by H. E.
Ives, in an address recently published in Science (1940, vol. 91, p. 79), "The Measurement
of Velocity with Atomic Clocks."

R. S. Lillie 335
evolved stable systems. The problem of causation is in fact
inseparable from the problem of evolution. Constancies 'in
modes of action have appeared in nature as stable conditions or
factors have come into existence. The problem of evolution has
been regarded as an essentially biological problem; biological
evolution, however, has cosmic evolution as its background and
condition;20 the two are inseparable. We cannot now con-
sider the problem of final or teleological causation which enters
at this point. The purely analytic study of nature discloses
organism within organism;21 it is sufficient to note that each or-
ganism must in some way achieve stability before it can beusedin
the building of higher organisms. Anything which happens
regularly in an organism (living or non-living) implies that a
station of stability, furnishing constancy of conditions, has been
reached. But activity under a given set of stable conditions
appears never to be completely predictable or stable. Some ten-
dency to spontaneous change seems always present; and this
tendency, if it asserts itself under favorable conditions, may be-
come "creative," i.e., may give origin to new and higher stations
of stability. The higher organisms, such as man, thus become
secondary centers of creative activity, but since their factors of
stability are routine processes, they may themselves, as units,
lapse into routine; or they may retrogress, since their stability
is precarious.
Obviously no scientific presumption of a complete and all-
pervasive stability which excludes spontaneous change can ac-
count for the historical fact of evolution, which involves deviation
from constancy. In living organisms deviation from the normal
routine of inheritance (mutation) is, according to modern bio-
logical theory, the prerequisite for evolutionary change. Species
are only relatively stable; the causal origin of variations, or of
mutations, appears to be partly environmental, partly internal.
But any attempt to assign definite causes to biological mutation
would be to raise again the problem of causation, in reference to a
20 This has been
emphasizedespeciallyby L. J. Henderson,"The Orderof Nature,"
Harvard University Press, I9I7.
a A. N. Whitehead:"Processand Reality,"passim.

3 36 Biological Causation
special type of change in living organisms. There would seem
to be no escape from the ascription of a certain variable element
of spontaneity, non-conformity to rule, or creativity to natural
processes in general. This condition reaches a maximum in the
more complex types of biological causation such as human
activity, although it is there limited in its possibilities by physical
and physiological constants already established, as well as by social
and other environmental conditions.
Iihe University of Chicago.


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velopments. Any single stable entity, say an atom, illustrates

"The word is etymologicallyequivalent to non-divisibility,or non-disintegrability,

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