FRANCISCO J.

A Y ALA

BIOLOGY AS AN AUTONOMOUS SCIENCE

The goal of science is the systematic organization of knowledge about the

universe on the basis of explanatory principles that are genuinely testable.
The starting point of science is the formulation of statements about ob-
jectively observable phenomena. Common sense knowledge also provides
information about the world. The distinction between science and com-
mon sense knowledge is based upon the joint presence in science of at
least three distinctive characteristics. First, science seeks to organize
knowledge in a systematic way by exhibiting patterns of relations among
statements concerning facts which may not appear obviously as mutually
related.
The information obtained in the course of ordinary experience about
the universe is frequently accurate, but it seldom provides any explana-
tion of why the facts are as alleged. It is the second distinctive charac-
teristic of science that it strives to provide explanations of why the ob-
served events do in fact occur. Science attempts to discover and to formu-
late the conditions under which the observed facts and their mutual re-
lationships exist.
Thirdly, the explanatory hypotheses provided by science must be genu-
inely testable, and therefore subject to the possibility of rejection. It is
sometimes asserted that scientific explanatory hypotheses should allow
one to formulate predictions about their subject matter which can be
verified by further observation and experiment. However, in certain fields
of scientific knowledge, as in those fields concerned with historical ques-
tions, prediction is considerably restricted by the nature of the subject
matter itself. The criterion of testability can then be satisfied by requiring
that scientific explanations have precise logical consequences which can
be verified or falsified by observation and experiment. The word 'precise'
is essential in the previous sentence. To provide genuine verification, the
logical consequences of the proposed explanatory hypotheses must not
be compatible with alternate hypotheses.
It is the concern of science to formulate theories, that is, to discover

M. Grene and E. Mendelsohn (eds.) , Topics in the Philosophy 0/ Biology, 312-29.

This article Copyright © 1968 American Scientist, Sigma Xi.
M. Grene et al., Topics in the Philosophy of Biology
© D. Reidel Publishing Company, Dordrecht, Holland 1976
 BIOLOGY AS AN AUTONOMOUS SCIENCE 313

patterns of relations among vast kinds of phenomena in such a way that

a small number of principles can explain a large number of propositions
concerning these phenomena. In fact, science develops by discovering new
relationships which show that observational statements and theories that
had hitherto appeared as independent are in fact connected and can be
integrated into a more comprehensive theory. Thus, the Mendelian prin-
ciples of inheritance can explain, about many different kinds of organisms,
observations which appear as prima facie unrelated, like the proportions
in which characters are transmitted from parents to offspring, the dis-
continuous nature of many traits of organisms, and why in outbreeding
sexual organisms no two individuals are likely to be genetically identical
even when the number of individuals in the species is very large. Knowl-
edge about the formation of the sex cells and about the behavior of
chromosomes was eventually shown to be connected with the Mendelian
principles, and contributed to the explanation of additional facts, for ex-
ample, why certain traits are inherited independently from each other
while other traits are transmitted together more frequently than not. Fur-
ther discoveries have contributed to the formulation of a unified theory
of inheritance which explains many other diverse observations, including
the discreteness of natural species, the adaptive nature of organisms and
their features, and paleontological observations concerning the evolution
of organisms.
The connection among theories has sometimes been established by
showing that the principles of a certain theory or branch of science can
be explained by the principles of another theory or science shown to have
greater generality. The less general branch of science, called the secondary
science, is said to have been reduced to the more general or primary sci-
ence. A typical example is the reduction of Thermodynamics to Statistical
Mechanics. l The reduction of one branch of science to another simplifies
and unifies science.
Reduction of one theory or branch of science to another has repeatedly
occurred in the history of science. During the last hundred years, several
branches of Physics and Astronomy have been to a considerable extent
unified by their reduction to a few theories of great generality like Quan-
tum Mechanics and Relativity. A large sector of Chemistry has been re-
duced to Physics after it was discovered that the valence of an element
bears a simple relation to the number of electrons in the outer orbit of
314 FRANCISCO J. AYALA

the atom. The impressive success of these and other reductions has led
in certain circles to the conviction that the ideal of science is to reduce
all natural sciences, including biology, to a comprehensive theory that
will provide a common set of principles of maximum generality capable
of explaining all our observations about the material universe.
To evaluate the validity of such claims, I will briefly examine some of
the necessary conditions for the reduction of one theory to another. I
will, then, attempt to show that, at the present stage of development of
the two sciences, the reduction of biology to physics cannot be effected.
I will further claim, in the second part of this paper, that there are pat-
terns of explanation which are indispensable in biology while they do not
occur in the physical sciences. These are teleological explanations which
apply to organisms and only to them in the natural world, and that can-
not be reformulated in non-teleological form without loss of explanatory
content.
I. CONDITIONS FOR REDUCTION

In general, reduction can be defined in the present context as "the ex-

planation of a theory or a set of experimental laws established in an area
of inquiry, by a theory usually though not invariably formulated for some
other domain." 2 Nagel has stated the two formal conditions that must be
satisfied to effect the reduction of one science to another. First, all the
experimental laws and theories of the secondary science must be shown
to be logical consequences of the theoretical constructs of the primary
science. This has been called by Nagel the condition of derivabi/ity.
Generally, the experimental laws formulated in a certain branch of sci-
ence will contain terms which are specific to that area of inquiry. If the
laws of the secondary science contain some terms that do not occur in the
primary science, logical derivation of its laws from the primary science
will not be prima facie possible. No term can appear in the conclusion of
a formal demonstration unless the term appears also in the premises. To
make reduction possible it is then necessary to establish suitable connec-
tions between the terms of the secondary science and those used in the
primary science. This may be called the condition of connectability. It can
be satisfied by a redefinition of the terms of the secondary science using
terms of the primary science. For example, to effect the reduction of
genetics to physical science such concepts as gene, chromosome, etc., must
 BIOLOGY AS AN AUTONOMOUS SCIENCE 315

be redefined in physicochemical terms such as atom, molecule, electrical

charge, hydrogen bond, deoxyribonucleic acid, etc.
The problem of reduction is sometimes formulated as whether the prop-
erties of a certain kind of objects, for instance organisms, can be explained
as a function of the properties of another such group of objects, like the
organism's physical components organized in certain ways. This formula-
tion of the question is spurious and cannot lead to a satisfactory answer.
Indeed it is not clear what is meant by the 'properties' of a certain object
which enters as a part or component of some other object. If all the prop-
erties are included, it appears that reduction could always be accom-
plished, and it is in fact a trivial issue. Among the properties of a certain
object one will list the properties which it has when it is a component of
the larger whole. To use a simple example, one may list among the proper-
ties of hydrogen that of combining in a certain way with oxygen to form
water, a substance which possesses certain specified properties. The prop-
erties of water will then be included among the properties of oxygen and
hydrogen.
The reduction of one science to another is not a matter of deriving the
properties of a kind of objects from the properties of some other group of
objects. It is rather a matter of deriving a set of propositions from an-
other such set. It is a question about the possibility of deriving the ex-
perimental laws of the secondary science as the logical consequences of
the theoretical laws of the primary science. Scientific laws and theories
consist of propositions about the world, and the question of reduction can
only be settled by the concrete investigation of the logical consequences
of such propositions, and not by discussion of the properties or the nature
of things.
From the previous observation it follows that the question of reduction
can only be solved by a specific reference to the actual stage of develop-
ment of the two disciplines involved. Certain parts of chemistry were re-
duced to physics after the modern theories of atomic structure were devel-
oped some fifty years ago, but the reduction could not have been accom-
plished before such development. Ifthe reduction of one science to another
is not possible at the present stage of development of the two disciplines,
it is empirically meaningless to ask whether reduction will be possible at
some further time, since the question can only be answered dogmatically
or in terms of metaphysical preconceptions.
316 FRANCISCO J. AYALA

Simpson has suggested that the unification of the various natural sci-
ences be sought not "through principles that apply to all phenomena but
through phenomena to which all principles apply." Science, according to
Simpson, can truly become unified in biology, since the principles of all
natural sciences can be applied to the phenomena of life. 3 To be sure, the
theoretical laws of physics and chemistry apply to the physicochemical
phenomena occurring in organisms. Besides, there are biological theories
that explain observations concerning the living world but have no applica-
tion to non-living matter. To conclude therefore that biology stands at the
center of all science is true as far as it goes, but it is trivial and constitutes
no progress in scientific understanding that I can discern.
The goal of the reductionistic program is not, as Simpson seems to
believe, to establish a "body of theory that might ultimately be completely
general in the sense of applying to all material phenomena," nor a "search
for a least common denominator in science." It is rather a quest for a
comprehensive theory that would explain all phenomena - the living as
well as the inanimate world - with an economy of stated laws and a cor-
responding increase in our understanding of the world. Whether such an
ideal can be accomplished is a different issue that I shall consider presently.

II. THE REDUCTION OF BIOLOGY TO PHYSICAL SCIENCE

The question of the reducibility of biology to physicochemistry has been

raised again in the last decade particularly in connection with the spec-
tacular successes accomplished in certain areas of biology. In genetics,
research at the molecular level has contributed to establish the chemical
structure of the hereditary material, to decipher the genetic code, and to
provide some understanding of the mechanisms of gene action. Brilliant
achievements have also been obtained in neurophysiology and other fields
of biology. Some authors have claimed that the understanding of all bio-
logical phenomena in physicochemical terms is not only possible but the
task of the immediate future. It is thus proclaimed that the only worthy
and truly 'scientific' biological research is what is called in recent jargon
'molecular biology', that is, the attempt to explain biological phenomena
in terms of the underlying physicochemical components and processes.
It is easy to dispose of two extreme positions which seem equally un-
profitable. On one end of the spectrum there are substantive vitalists
 BIOLOGY AS AN AUTONOMOUS SCIENCE 317

who defend the irreducibility of biology to physical science because living

phenomena are the effect of a non-material principle which is variously
called vital force, entelechy, elan vital, radial energy, or the like. A non-
material principle cannot be subject to scientific observation nor lead to
genuinely testable scientific hypotheses. 4
At the other end of the spectrum stand those who claim that reduction
of biology to physicochemistry is in fact possible at present. In the current
stage of scientific development, a majority of biological concepts, such
as cell, organ, species, ecosystem, etc., cannot be formulated in physi-
cochemical terms. Nor is there at present any class of statements belong-
ing to physics and chemistry from which every biological law could be
logically derived. In other words, neither the condition of connectability
nor the condition of derivability - two necessary formal conditions of
reduction - are satisfied at the present stage of development of physical
and biological knowledge.
Two intermediate positions have also appeared in the recent literature.
First, a reductionist position maintains that although the reduction of
biology to physics cannot be effected at present, it is possible in principle.
The factual reduction is made contingent upon further progress in the
biological or in the physical sciences, or in both. Secondly, certain anti-
reductionist authors claim that reduction is not possible in principle be-
cause organisms are not merely assemblages of atoms and molecules, nor
even of organs and tissues standing in merely external relation to one
another. Organisms are alleged to be 'wholes' that must be studied as
wholes and not as the 'sum' of isolable parts. 5
Although biological laws are not in general derivable from any avail-
able theory of physics and chemistry, the reductionists claim that such
accomplishment will be possible in the future. Such assertions are fre-
quently based on metaphysical preconceptions about the nature of the
material world. In any case it cannot be convincingly argued empirically,
since it is only a statement of faith about the possibility of some future
event. It must be noted, however, that advances in various areas ofmolec-
ular biology are continuously extending the, as yet, exiguous realm of
biological phenomena that can be explained in terms of physicochemical
concepts and laws.
As for the anti-reductionist position that maintains that organisms and
their properties cannot be understood as mere 'sums' of their parts, I
318 FRANCISCO J. AYALA

have already stated that it rests on an unsatisfactory formulation of the

problem. The question of reduction is whether propositions concerning
organisms can be logically derived from physicochemical laws, and not
whether the properties of organisms can be explained as the result of the
properties of their physical components. It should perhaps be added that
the phenomenon of so-called 'emergent' properties occurs also in the non-
living world. Water is formed by the union of two atoms of hydrogen with
one atom of oxygen, but water exhibits properties which are not the im-
mediately apparent consequence of the properties of the two gases, hydro-
gen and oxygen. Another simple example can be taken from the field of
thermodynamics. A gas has a temperature although the individual mole-
cules of the gas cannot be said to possess a temperature. 6
The reduction of biology to physicochemistry admittedly cannot be ef-
fected at the present stage of scientific knowledge. Whether the reduction
will be possible in the future is an empirically meaningless question. A
majority of biological problems cannot be as yet approached at the mo-
lecular level. Biological research must then continue at the different levels
of integration of the living world, according to the laws and theories
developed for each order of complexity. The study of the molecular struc-
ture of organisms must proceed hand in hand with research at the levels
of the cell, the organ, the individual, the population, the species, the com-
munity, and the ecosystem. These levels of integration are not isolated
from each other. Laws formulated at one level of complexity illuminate
the other levels, both lower and higher, and suggest additional research
strategies. 7 It is perhaps worth pointing out that in fact biological laws
discovered at a higher level of organization have more frequently contri-
buted to guide research at the lower level than vice versa. To mention but
one example, the Mendelian theory of inheritance preceded the identifica-
tion of the chemical composition and structure of the genetic material,
made possible these discoveries, and has not become superogatory be-
cause of the latter.

III. THE NOTION OF TELEOLOGY

I will now proceed to discuss the role of teleological explanations in

biology. I shall attempt to show that teleological explanations constitute
patterns of explanation that apply to organisms while they do not apply
 BIOLOGY AS AN AUTONOMOUS SCIENCE 319

to any other kind of objects in the natural world. I shall further claim
that, although teleological explanations are compatible with causal ac-
counts, they cannot be reformulated in non-teleological form without loss
of explanatory content. Consequently, I shall conclude that teleological
explanations cannot be dispensed with in biology, and are therefore dis-
tinctive of biology as a natural science.
The concept of teleology is in general disrepute in modern science.
More frequently than not it is considered to be a mark of superstition,
or at least a vestige of the non-empirical, a prioristic approach to natural
phenomena characteristic of the prescientific era. The main reason for
this discredit is that the notion of teleology is equated with the belief that
future events - the goals or end-products of processes - are active agents
in their own realization. In evolutionary biology, teleological explana-
tions are understood to imply the belief that there is a planning agent
external to the world, or a force immanent to the organisms, directing
the evolutionary process toward the production of specified kinds of or-
ganisms. The nature and diversity of organisms are, then, explained tele-
ologically in such a view as the goals or ends-in-view intended from the
beginning by the Creator, or implicit in the nature of the first organisms.
Biological evolution can however be explained without recourse to a
Creator or a planning agent external to the organisms themselves. There
is no evidence either of any vital force or immanent energy directing the
evolutionary process toward the production of specified kinds of orga-
nisms. The evidence of the fossil record is against any directing force,
external or immanent, leading the evolutionary process toward specified
goals. Teleology in the stated sense is, then, appropriately rejected in
biology as a category of explanation.
In The Origin of the Species Darwin accumulated an impressive number
of observations supporting the evolutionary origin of living organisms.
Moreover, and perhaps most importantly, he provided a causal explana-
tion of evolutionary processes - the theory of natural selection. The prin-
ciple of natural selection makes it possible to give a natural explanation
of the adaptation of organisms to their environments. Darwin recognized,
and accepted without reservation, that organisms are adapted to their
environments, and that their parts are adapted to the functions they serve.
Penguins ale adapted to live in the cold, the wings of birds are made to fly,
and the eye is made to see. Darwin accepted the facts of adaptation, and
320 FRANCISCO J. AYALA

then provided a natural explanation for the facts. One of his greatest
accomplishments was to bring the teleological aspects of nature into the
realm of science. He substituted a scientific teleology for a theological one.
The teleology of nature could now be explained, at least in principle, as
the result of natural laws manifested in natural processes, without re-
course to an external Creator or to spiritual or non-material forces. At
that point biology came into maturity as a science.
The concept of teleology can be defined without implying that future
events are active agents in their own realization nor that the end-results
of a process are consciously intended as goals. The notion of teleology
arose most probably as a result of man's reflection on the circumstances
connected with his own voluntary actions. The anticipated outcome of
his actions can be envisaged by man as the goal or purpose toward which
he directs his activity. Human actions can be said to be purposeful when
they are intentionally directed toward the fulfilment of a goal.
The plan or purpose of the human agent may frequently be inferred
from the actions he performs. That is, his actions can be seen to be pur-
posefully or teleologically ordained toward the fulfilment of a goal. In
this sense the concept ofteleology can be extended, and has been extended,
to describe actions, objects or processes which exhibit an orientation to-
ward a certain goal or end-state. No requirement is necessarily implied
that the objects or processes tend consciously toward their specified end-
states, nor that there is any external agent directing the process or the
object toward its end-state or goal. In this generic sense, teleological ex-
planations are those explanations where the presence of an object or a
process in a system is explained by exhibiting its connection with a specific
state or property of the system to whose existence or maintenance the
object or process contributes. Teleological explanations require that the
object or process contribute to the existence of a certain state or property
of the system. Moreover, and this is the essential component of the con-
cept, teleological explanations imply that such contribution is the explan-
atory reason for the presence of the process or object in the system. Ac-
cordingly, it is appropriate to give a teleological explanation of the opera-
tion of the kidney in regulating the concentration of salt in the blood, or
of the structure of the hand of man obviously adapted for grasping. But
it makes no sense to explain teleologically the motions of a planet or a
chemical reaction. In general, as will be shown presently, teleological ex-
 BIOLOGY AS AN AUTONOMOUS SCIENCE 321

planations are appropriate to account for the existence of adaptations in

organisms while they are neither necessary nor appropriate in the realm
of non-living matter.
There are at least three categories of biological phenomena where tele-
ological explanations are appropriate, although the distinction between
the categories need not always be clearly defined. These three classes of
teleological phenomena are established according to the mode of relation-
ship between the structure or process and the property or end-state that
accounts for its presence. Other classifications of teleological phenomena
are possible according to other principles of distinction. A second classifi-
cation will be suggested later.
(1) When the end-state or goal is consciously anticipated by the agent.
This is purposeful activity and it occurs in man and probably, although
in a lesser degree, in other animals. I am acting teleologically when I buy
an airplane ticket to fly to Mexico City. A cheetah hunting a zebra has
at least the appearance of purposeful behavior. However, as I have said
above, there is no need to explain the existence of organisms and their
adaptations as the result of the consciously intended activity of a Cre-
ator. There is a purposeful activity in the living world, at least in man;
but the existence of the living world, including man, need not be ex-
plained as the result of purposeful behavior. When some critics reject the
notion of teleology from the natural sciences, they have in mind exclu-
sively this category of teleology.
(2) Self-regulating or teleonomic systems, when there exists a mecha-
nism that enables the system to reach or to maintain a specific property
in spite of environmental fluctuations. The regulation of body tempera-
ture in mammals is a teleological mechanism of this kind. In general, the
homeostatic reactions of organisms belong to this category of teleological
phenomena. Two types of homeostasis are usually distinguished by biolo-
gists - physiological and developmental homeostasis, although inter-
mediate and additional types do exist. 8 Physiological homeostatic reac-
tions enable the organism to maintain certain physiological steady states
in spite of environmental shocks. The regulation of the composition of the
blood by the kidneys, or the hypertrophy of muscle in case of strenuous
use, are examples of this type of homeostasis.
Developmental homeostasis refers to the regulation of the different
paths that an organism may follow in its progression from zygote to adult.
322 FRANCISCO J. AYALA

The development of a chicken from an egg is a typical example of de-

velopmental homeostasis. The process can be influenced by the environ-
ment in various ways, but the characteristics of the adult individual, at
least within a certain range, are largely predetermined in the fertilized
egg. Aristotle, Saint Augustine, and other ancient and medieval philoso-
phers, took developmental homeostasis as the paradigm of all teleological
mechanisms. According to Saint Augustine, God did not create directly all
living species of organisms, but these were implicit in the primeval forms
created by Him. The existing species arose by a natural 'unfolding' of
the potentialities implicit in the primeval forms or 'seeds' created by God.
Self-regulating systems or servo-mechanisms built by man belong in
this second category of teleological phenomena. A simple example of such
servo-mechanisms is a thermostat unit that maintains a specified room
temperature by turning on and off the source of heat. Self-regulating
mechanisms of this kind, living or man-made, are controlled by a feed-
back system of information.
(3) Structures anatomically and physiologically constituted to perform
a certain function. The hand of man is made for grasping, and his eye for
vision. Tools and certain types of machines made by man are teleological
in this third sense. A watch, for instance, is made to tell time, and a faucet
to draw water. The distinction between the (3) and (2) categories of tele-
ological systems is sometimes blurred. Thus, the human eye is able to
regulate itself within a certain range to the conditions of brightness and
distance so as to perfoIm its function more effectively.

IV. TELEOLOGY AND ADAPTATION

Teleological mechanisms and structures in organisms are biological adap-

tations. They have arisen as a result of the process of natural selection.
Natural selection is a mechanistic process defined in genetic and statistical
terms as differential reproduction. Some genes and genetic combinations
are transmitted to the following generations on the average more fre-
quently than their alternates. Such genetic units will become more com-
mon, and their alternates less common, in every subsequent generation.
The genetic variants arise by the random processes of genetic mutation
and recombination. Genetic variants increase in frequency and may even-
tually become fixed in the population if they happen to be advantangeous
 BIOLOGY AS AN AUTONOMOUS SCIENCE 323

as adaptations in the organisms which carry them, since such organisms

are likely to leave more descendants than those lacking such variants. If
a genetic variant is harmful or less adaptive than its alternates, it will be
eliminated from the population. The biological adaptations of the orga-
nisms to their environments are, then, the result of natural selection,
which is nevertheless a mechanistic and impersonal process.
The adaptations of organisms - whether organs, homeostatic mecha-
nisms, or patterns of behavior - are explained teleologically in that their
existence is ultimately accounted for in terms of their contribution to the
reproductive fitness of the species. A feature of an organism that increases
its reproductive fitness will be selectively favored. Given enough genera-
tions it will extend to all the members of the population.
Patterns of behavior, such as the migratory habits of certain birds or
the web-spinning of spiders, have developed because they favored the re-
productive success of their possessors in the environments where the popu-
lation lived. Similarly, natural selection can account for the existence of
homeostatic mechanisms. Some living processes can be operative only
within a certain range of conditions. If the environmental conditions oscil-
late frequently beyond the functional range of the process, natural selec-
tion will favor self-regulating mechanisms that maintain the system within
the functional range. In man, death results if the body temperature is
allowed to rise or fall by more than a few degrees above or below normal.
Body temperature is regulated by dissipating heat in warm environments
through perspiration and dilation of the blood vessels in the skin. In cool
weather the loss of heat is minimized, and additional heat is produced by
increased activity and shivering. Finally, the adaptation of an organ or
structure to its function is also explained teleologically in that its presence
is accounted for in terms of the contribution it makes to reproductive
success in the population. The vertebrate eye arose because genetic muta-
tions responsible for its development occurred, and were gradually com-
bined in progressively more efficient patterns, the successive changes in-
creasing the reproductive fitness of their possessors in the environments
in which they lived.
There are in all organisms two leveb of teleology that may be labeled
specific and generic. There usually exists a specific and proximate end for
every feature of an animal or plant. The existence of the feature is ex-
plained in terms of the function or property that it serves. This function
324 FRANCISCO J. AYALA

or property can be said to be the specific or proximate end of the feature.

There is also an ultimate goal to which all features contribute or have
contributed in the past - reproductive success. The generic or ultimate
end to which all features and their functions contribute is increased re-
productive efficiency. The presence of the functions themselves - and
therefore of the features which serve them - is ultimately explained by
their contribution to the reproductive fitness of the organisms in which
they exist. In this sense the ultimate source of explanation in biology is
the principle of natural selection.
Natural selection can be said to be a teleological process in a causal
sense. Natural selection is not an entity but a purely mechanistic process.
But natural selection can be said to be teleological in the sense that it
produces and maintains end-directed organs, when the functions served
by them contribute to the reproductive efficiency of the organism.
The process of natural selection is not at all teleological in a different
sense. Natural selection is not in any way directing toward the production
of specific kinds of organisms or toward organisms having certain specific
properties. The over-all process of evolution cannot be said to be teleo-
logical in the sense of proceeding toward certain specified goals, precon-
ceived or not. The only nonrandom process in evolution is natural selec-
tion understood as differential reproduction. Natural selection is a purely
mechanistic process and it is opportunistic. 9 The final result of natural
selection for any species may be extinction, as shown by the fossil record,
if the species fails to cope with environmental change.
The presence of organs, processes, and patterns of behavior can be ex-
plained teleologically by exhibiting their contribution to the reproductive
fitness of the organisms in which they occur. This need not imply that
reproductive fitness is a consciously intended goal. Such intent must in
fact be denied, except in the case of the voluntary behavior of man. In
teleological explanations the end-state or goal is not to be understood as
the efficient cause of the object or process that it explains. The end-state
is causally - and in general temporally also - posterior.

V. INTERNAL AND EXTERNAL TELEOLOGY

Three categories ofteleological phenomena have been distinguished above,

according to the nature of the relationship existing between the object or
 BIOLOGY AS AN AUTONOMOUS SCIENCE 325

mechanism and the function or property that it serves. Another classifica-

tion of teleology may be suggested attending to the process or agency
giving origin to the teleological system. The end-directedness of living
organisms and their features may be said to be internal teleology, while
that of man-made tools and servo-mechanisms may be called external
teleology. It might also be appropriate to refer to these two kinds of
teleology as natural and artificial, but the other two terms, 'internal' and
'external', have already been used.1°
Internal teleological systems are accounted for by natural selection
which is a strictly mechanistic process. External teleological systems are
the products of the human mind, or more generally, are the result of pur-
poseful activity consciously intending specified ends. An automobile, a
wrench, and a thermostat are teleological systems in the external sense;
their parts and mechanisms have been produced to serve certain func-
tions intended by man. Organisms and their parts are teleological systems
in the internal sense; their end-directedness is the result of the mechanistic
process of natural selection. Organisms are the only kind of systems
exhibiting internal teleology. In fact they are the only class of natural
systems that exhibit teleology. Among the natural sciences, then, only
biology, which is the study of organisms, requires teleology as a category
of explanation.
Organisms do not in general possess external teleology. As I have said
above, the existing kinds of organisms and their properties can be ex-
plained without recourse to a Creator or planning agent directing the
evolutionary process toward the production of such organisms. The evi-
dence from paleontology, genetics, and other evolutionary sciences is also
against the existence of any immanent force or vital principle directing
evolution toward the production of specified kinds of organisms.

VI. TELEOLOGICAL EXPLANATIONS IN BIOLOGY

Teleological explanations are fully compatible with causal accounts. "In-

deed, a teleological explanation can always be transformed into a causal
one." 11 Consider a typical teleological statement in biology, "The func-
tion of gills in fishes is respiration." This statement is a telescoped argu-
ment the content of which can be unraveled approximately as follows:
Fish respire; if fish have no gills, they do not respire; therefore fish have
326 FRANCISCO J. AYALA

gills. According to Nagel, the difference between a teleological explana-

tion and a non-teleological one is, then, one of emphasis rather than of
asserted content. A teleological explanation directs our attention to "the
consequences for a given system of a constituent part or process." The
equivalent non-teleological formulation focuses attention on "some of the
conditions ... under which the system persists in its characteristic organiza-
tion and activities." 12
Although a teleological explanation can be reformulated in a non-teleo-
logical one, the teleological explanation connotes something more than
the equivalent non-teleological one. In the first place, a teleological ex-
planation implies that the system under consideration is directively orga-
nized. For that reason teleological explanations are appropriate in biol-
ogy and in the domain of cybernetics but make no sense when used in
the physical sciences to describe phenomena like the fall of a stone.
Teleological explanations imply, while non-teleological ones do not, that
there exists a means-to-end relationship in the systems under description.
Besides connoting that the system under consideration is directively
organized, teleological explanations also account for the existence of spe-
cific functions in the system and more generally for the existence of the
directive organization itself. The teleological explanation accounts for the
presence in an organism of a certain feature, say the gills, because it con-
tributes to the performance or maintenance of a certain function, respira-
tion. In addition it implies that the function exists because it contributes
to the reproductive fitness of the organism. In the non-teleological transla-
tion given above, the major premise states that 'fish respire'. Such formu-
lation assumes the presence of a specified function, respiration, but it
does not account for its existence. The teleological explanation does in
fact account for the presence of the function itself by implying or stating
explicitly that the function in question contributes to the reproductive
fitness of the organism in which it exists. Finally, the teleological explana-
tion gives the reason why the system is directively organized. The apparent
purposefulness of the ends-to-means relationship existing in organisms is
a result of the process of natural selection which favors the development
of any organization that increases the reproductive fitness of the orga-
nisms.
If the above reasoning is correct, the use of teleological explanations in
biology is not only acceptable but indeed indispensable. Organisms are
 BIOLOGY AS AN AUTONOMOUS SCIENCE 327

systems directively organized. Parts of organisms serve specific functions

that, generally, contribute to the ultimate end of reproductive survival.
One question biologists ask about organic structures and activities is
'What for?, That is, 'What is the function or role of such a structure or
such a process?' The answer to this question must be formulated in teleo-
logical language. Only teleological explanations connote the important
fact that plants and animals are directively organized systems.
It has been argued by some authors that the distinction between systems
that are goal directed and those which are not is highly vague. The classifi-
cation of certain systems as teleological is allegedly rather arbitrary. A
chemical buffer, an elastic solid or a pendulum at rest are examples of
physical systems that appear to be goal directed. I suggest using the crite-
rion of utility to determine whether an entity is teleological or not. The
criterion of utility can be applied to both internal and external teleological
systems. Utility in an organism is defined in reference to the survival and
reproduction of the organism itself. A feature of a system will be teleo-
logical in the sense of internal teleology if the feature has utility for the
system in which it exists and if such utility explains the presence of the
feature in the system. Operationally, then, a structure or process of an
organism is teleological if it can be shown to contribute to the reproduc-
tive efficiency of the organism itself, and if such contribution accounts for
the existence of the structure or process.
In external teleology utility is defined in reference to the author of the
system. Man-made tools or mechanisms are teleological with external
teleology if they have been designed to serve a specified purpose, which
therefore explains their existence and properties. If the criterion of utility
cannot be applied, a system is not teleological. Chemical buffers, elastic
solids, and a pendulum at rest are not teleological systems.
The utility of features of organisms is with respect to the individual or
the species in which they exist at any given time. It does not include use-
fulness to any other organisms. The elaborate plumage and display is a
teleological feature of the peacock because it serves the peacock in its
attempt to find a mate. The beautiful display is not teleologically directed
toward pleasing man's aesthetic sense. That it pleases the human eye is
accidental, because it does not contribute to the reproductive fitness of the
peacock (except, of course, in the case of artificial selection by man).
The criterion of utility introduces needed objectivity in the determina-
328 FRANCISCO J. AYALA

tion of what biological mechanisms are end-directed. Provincial human

interests should be avoided when using teleological explanations, as Nagel
says. But he selects the wrong example when he observes that "the devel-
opment of corn seeds into corn plants is sometimes said to be natural,
while their transformation into the flesh of birds or men is asserted to be
merely accidental." 13 The adaptations of corn seeds have developed to
serve the function of corn reproduction, not to become a palatable food
for birds or men. The role of wild corn as food is accidental, and cannot
be considered a biological function of the corn seed in the teleological
sense.
Some features of organisms are not useful by themselves. They have
arisen as concomitant or incidental consequences of other features that
are adaptive or useful. In some cases, features which are not adaptive in
origin may become useful at a later time. For example, the sound pro-
duced by the beating of the heart has become adaptive for modern man
since it helps the physician to diagnose the condition of health of the pa-
tient. The origin of such features is not explained teleologically, although
their preservation might be so explained in certain cases.
Features of organisms may be present because they were useful to the
organisms in the past, although they are no longer adaptive. Vestigial
organs, like the vermiform appendix of man, are features of this kind. If
they are neutral to reproductive fitness these features may remain in the
population indefinitely. The origin of such organs and features, although
not their preservation, is accounted for in teleological terms.
To conclude, I will summarize the second part of this paper. Teleo-
logical explanations are appropriate to describe, and account for the
existence of, teleological systems and the directively organized structures,
mechanisms, and patterns of behavior which these systems exhibit. Orga-
nisms are the only natural systems exhibiting teleology; in fact they are
the only class of systems possessing internal teleology. Teleological expla-
nations are not appropriate in the physical sciences, while they are ap-
propriate, and indeed indispensable, in biology which is the scientific study
of organisms. Teleological explanations, then, are distinctive of biology
among all the natural sciences. 14

University of California, Davis

 BIOLOGY AS AN AUTONOMOUS SCIENCE 329

NOTES

1 E. Nagel, The Structure of Science, New York: Harcourt, Brace and World, 1961,
pp. 338-345.
2 Ibid. p. 338; see also pp. 336-397.
3 G. G. Simpson, This View of Life, New York: Harcourt, Brace and World, 1964,
p. 107. According to J. G. Kemeny (A Philosopher Looks at Science, Van Nostrand,
1959, pp. 215-216) the most likely solution of the question of the reduction of biology
to physics is that a new theory will be found, covering both fields, in new terms. Inan-
imate nature will appear as the simplest extreme case of this theory. In that case, one
would say that physics was reduced to biology and not biology to physics.
4 Except for the general conclusion that biological phenomena will never be satis-
factorily explained by mechanistic principles.
5 E. S. Russell, The Interpretation of Development and Heredity, Oxford, 1930; see also
E. Mayr, 'Cause and Effect in Biology', Cause and Effect, D. Lerner (ed.) New York:
Free Press, 1965, pp. 33-50.
6 The temperature of the gas is identical by definition with the mean kinetic energy of
the molecules.
7 Th. Dobzhansky, 'Biology, Molecular and Organismic', The Graduate Journal
Vll(I), 1965, pp. 11-25.
8 For instance, the maintenance of a genetic polymorphism in a population due to
heterosis can be considered a homeostatic mechanism acting at the population level.
9 F. J. Ayala, 'Teleological Explanations in Evolutionary Biology', Philosophy of Sci-
ence, 37 (1970), 1-15.
10 T. A. Goudge, The Ascent of Life, Toronto: Univ. of Toronto Press, 1961, p. 193.
11 E. Nagel, 'Types of Causal Explanation in Science', Cause and Effect, D. Lerner
(ed.), New York: Free Press, 1965, p. 25.
12 E. Nagel, The Structure of Science, p. 405; see also F. J. Ayala, ref. in note 9 above.
13 E. Nagel, The Structure of Science, p. 424.
14 The author is a recipient of a PHS Research Career Development Award from the
National Institute of General Medical Sciences.