Teleology Biology

Teleological Explanations in Evolutionary Biology
Author(s): Francisco J. Ayala

Source: Philosophy of Science, Vol. 37, No. 1 (Mar., 1970), pp. 1-15
Published by: The University of Chicago Press on behalf of the Philosophy of Science Association
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Philosophy of Science
March,
1970
TELEOLOGICAL EXPLANATIONS IN EVOLUTIONARY BIOLOGY*
FRANCISCO J. AYALAt
The Rockef eller University
The ultimate source of explanation in biology is the principle of natural selection.
Natural selection means dif f erential reproduction of genes and gene combinations. It
is a mechanistic process which accounts f or the existence in living organisms of end-
directed structures and processes. It is argued that teleological explanations in biology
are not only acceptable but indeed indispensable. There are at least three categories of
biological phenomena where teleological explanations are appropriate.
Early in the nineteenth century, William Paley in his Natural Theology [6] pointed
out the obvious f unctional design of the human eye. For Paley, it was absurd to
suppose that the human eye, by mere chance, "should have consisted, f irst, of a
series of transparent lenses (very dif f erent, by and by, even in their substance f rom
the opaque materials of which the rest of the body is, in general at least, composed;
and with which the whole of its surf ace, this single portion of it excepted, is covered)
secondly of a black cloth or canvas (the only membrane of the body which is black)
spread out behind these lenses so as to receive the image f ormed by pencils of light
transmitted through them; and placed at the precise geometrical distance at which,
and at which alone, a distinct image could be f ormed, namely at the concourse of
the ref racted rays: thirdly, of a large nerve communicating between this membrane
and the brain."
The adaptive character of the structures, organs, and behavior of plants and
animals is an incontrovertible f act. The vertebrate eye, with its complicated
anatomy of highly specialized tissues, is obviously adapted f or vision; the hand of
man is made f or grasping, and the bird's wing f or f lying. Organisms show them-
selves to be adapted to live where they live and the way they live. To explain the
phenomenon of the adaptation of lif e is one of the main objectives of natural
science and of natural philosophy.
Bef ore 1859, the year Darwin published The Origin of the Species, the adaptation
of organisms was either accepted as a f act without any explanation of its origin, or
more f requently, it was attributed to the omniscient design
of the Creator. God
*
Received September 1967.
t
I wish to express my appreciation to Prof essors Th. Dobzhansky, E. Mayr, and E. Nagel,
who read an earlier version of this paper and made
many
valuable
suggestions.
I
2 FRANCISCO J. AYALA
had given wings to birds so that they might f ly, and had provided man with kidneys
to regulate the composition of his blood. For Paley, living nature is a manif estation
of the existence and wisdom of the Creator.
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 explanation of evolutionary
processes-the theory of natural selection. The principle of natural selection, as
Darwin saw it, makes it possible to give a natural explanation of the adaptation of
organisms to their environment. With The Origin of the Species the study of
adaptation, the problem of design in nature came f ully into the domain of natural
science.
Darwin recognized, and accepted without reservation, that organisms are
adapted to their environments, and that their parts are adapted to the f unctions
they serve. Fish are adapted to live in water, the hand of man is made f or grasping,
and the eye is made to see. Darwin accepted the f acts of adaptation, and then
provided a natural explanation f or the f acts. One of his greatest accomplishments
was to bring the teleological aspects of nature into the realm of science. He sub-
stituted a scientif ic teleology f or a theological one. The teleology of nature could
now be explained, at least in principle, as the result of natural laws manif ested in
natural processes, without recourse to an external Creator or to spiritual or non-
material f orces. At that point biology came into maturity as a science.
1. Hereditary Variability. About the time Darwin published The Origin of the
Species, Gregor Mendel was perf orming in his Augustinian monastery in Briinn
(Austria) experiments with peas. The results of such experiments, published in
1866, provided the f undamental principles of heredity. The Mendelian principles
remained generally unknown until 1900, when they were independently and nearly
simultaneously rediscovered by three biologists. The principles of heredity were
extended during the nineteen hundreds to a considerable number of species of
plants and animals. A whole body of knowledge concerning heredity blossomed.
The biological or synthetic theory of evolution as we know it today is a synthesis
of Darwin's principle of natural selection and genetic knowledge. It is in essence a
two-f actor theory. Mutation is the ultimate source of hereditary variability;
natural selection is the directional f actor that results in organized complexes of
hereditary material and in adaptation.
Heredity is the transmission, f rom parent to of f spring, of the inf ormation that
directs the development of the f ertilized egg to its adult stage and controls the
living activity of the organism. The hereditary inf ormation is carried in a chemical
substance known as deoxyribonucleic acid (DNA). Molecules of DNA exist in
discrete but complexly interacting units called genes. The genes are organized in
chromosomes, which exist in sets. One or more sets of chromosomes-most
f requently two in higher organisms-exist in the f ertilized egg cell (zygote) f rom
which the adult individual develops. In sexually reproducing organisms, one of the
two sets of chromosomes is inherited f rom each parent via the sex cells.
The genes of a population are shuf f led and combined in dif f erent ways every
TELEOLOGICAL EXPLANATIONS IN EVOLUTIONARY BIOLOGY 3
generation. In the process of genetic recombination during the f ormation of the
sex cells (gametes), the two sets of hereditary material received by each individual
f rom its parents are combined in dif f erent ways. The sex cells carry a single set of
genes each, representing combinations in dif f erent proportions of the two sets
possessed by the individual. Fertilization brings together two sex cells in the zygote
f rom which the mature individual develops. Gametic recombination and f ertiliza-
tion create new combinations of genes and chromosomes every generation. These
new sets of inf ormation are tested against the environment where the individual
lives. Thus, genetic experimentation, so to speak, occurs in all natural populations
every generation.
The sum total of genetic inf ormation in a population of sexually interbreeding
individuals can be thought of as the "gene pool" of the population. The gene pool
of a population is characterized by the totality of genes in the population, their
combinations, and the relative f requencies of both among the individuals of the
population. Evolution consists in changes in the gene pool of a population.
Recombination produces new combinations of genes but by itself it does not change
the gene pool. There are f our known processes which can do so-mutation, random
f luctuation of genetic f requencies known as "sampling errors," migration of
individuals in and out of the population, and natural selection. The f irst three of
these processes are essentially random. Although the relative importance f or evolu-
tion of random genetic sampling has been questioned, it must have played a role
in certain instances-in particular, when a new environment is colonized by a small
number of individuals and when populations are reduced to f ew individuals in their
usual environments by drastic environmental stresses. For our present purpose we
need consider neither random sampling nor migration.
Genes are f airly stable entities but not completely so. Occasionally, mutations
occur. The f requencies of mutations vary f or dif f erent genes and f or dif f erent
organisms. It is probably f air to estimate the f requency of a majority of mutations
in higher organisms between one in ten thousand, and one in a million per gene per
generation. Mutations in a broad sense include not only changes in the hereditary
inf ormation of single genes, but also changes in the arrangement and distribution of
genes in chromosomes, and in the number of chromosomes and sets of chromo-
somes. Mutations have sometimes been described as "errors" in the replication of
the hereditary material. Such a description may be misleading, since the alleged
"errors", are the ultimate source of evolutionary change. Mutation provides the raw
materials of evolution, i.e., mutation is the ultimate source of genetic variability.
Mutations are random changes of the hereditary material. They are random in
the sense that they occur independently of the needs of the organism in which they
happen. Most new mutations are in f act harmf ul to the organism. If mutation were
the only f actor promoting genetic change in a population, it would result in an
array of f reaks and f inally in total disorder. The genetic inf ormation stored in the
DNA of the population would ultimately disintegrate. However, there is a directive
process that counteracts mutation and results in order and adaptation-natural
selection. Natural selection is able to produce and to preserve the stored inf orma-
tion transmitted by the hereditary process.
2. Natural Selection. Natural selection was Darwin's major contribution to the
explanation of the evolution of lif e. For Darwin, natural selection was primarily
dif f erential survival. The modern understanding of the principle of natural selection
derives f rom Darwin's concept, although it is f ormulated in a somewhat dif f erent
way. Natural selection is understood today in genetic and statistical terms as
dif f erential reproduction. Dif f erential reproduction is a compound process, the
elements of which are dif f erential survival, dif f erential mating success, and dif f er-
ential f ecundity. Natural selection implies that some genes and genetic combina-
tions are transmitted to the f ollowing generation on the average more f requently
than their alternates. Such genetic units will become more common in every
subsequent generation and their alternates less common. Natural selection is a
statistical bias in the relative rate of reproduction of alternative genetic units.
Genes and gene combinations are the entities subject to natural selection. Genes
do not exist by themselves but in organisms. Genes increase or decrease in relative
f requency depending on their average ef f ects in the organisms which carry them.
The process of natural selection can be also predicated of individual organisms-
and in a less precise sense, of populations of organisms as well-in the sense that
some organisms leave more progeny than others. Individual organisms are not last-
ing, however. Genes persist in the progenies of the organisms which carry them.
Natural selection is a process determined by the environment. The selective
advantage of certain genetic variants must be understood in relation to the environ-
ment where the population lives. A genetic unit which is f avorably selected in one
environment may be selected again in a dif f erent one. A trivial example is that
wings-and theref ore the
genes responsible
f or the
development
of
wings-may
increase the reproductive success of a bird, but will probably be of no advantage,
and presumably will be disadvantageous, to a deep sea f ish. To speak of the environ-
ment of a population is, however, an oversimplif ication. The environment is highly
heterogeneous both in the dimension of time and in the dimension of space. The
environment of a population includes all the physical and biotic elements af f ecting
the individuals of the population in the whole range of their geographic distribution.
Small or large dif f erences in climate, f ood resources, competitors, etc. exist within
the spatial distribution of any population. Moreover, no environment remains
constant in time. It changes f rom morning to night, f rom one season to another,
f rom one year to the next. The reproductive f itness of a genetic variant is then the
average result of the ef f ects of that genetic unit in all the environments where the
population lives. It may change f rom one to another generation as the biotic and
physical environments of the population change. Environmental diversity and
environmental change are responsible f or the continuous evolution of natural popu-
lations. If lif e existed in only a single unif orm and constant environment, evolu-
tion might conceivably have produced a genotype optimally f itted to that
environment with no f urther change. An absolutely unif orm and constant environ-
ment is an abstraction; it does not exist in nature.
Genes act in concert with other genes. The average ef f ect of a gene in a population
may vary depending on the other genes and genetic combinations existing in the
population. The reproductive f itness of a genetic unit must be understood as the
average ef f ect it has on all the individuals carrying it. That average ef f ect is likely
to change as the genetic composition of the population changes f rom generation
to generation.
The numbers of alternative genetic variants existing in a natural population is a
debated question, but they vary widely f or dif f erent kinds of organisms. If two
variants, A1 and A2, of a gene exist in a population, there are in diploid organisms
three possible dif f erent genotypes with respect to that gene, namely A1A1, A1A2,
and A2A2. If the number of genes existing in two alternative f orms is n, 3n dif f erent
genotypes are possible. That number becomes very large as n increases. For instance,
if n equals 10, the number of possible dif f erent genotypes is nearly one hundred
thousand; if n equals 20, there are more than one billion potential genotypes; and
if n equals 30, there are nearly one million billion possible dif f erent genotypes. The
number of possible genetic combinations in a population of diploid organisms,
even in those organisms carrying relatively f ew alternative genetic units,
is enormous.
Most of them will never occur because the number of individuals in the population
is much less than the number of possible dif f erent genetic combinations. Natural
selection operates exclusively on the genetic combinations actually realized in the
population.
If a gene or genetic combination increases on the average the reproductive
success of the individuals carrying it, its f requency in the population will increase
gradually. It has been shown both theoretically and experimentally that a newly
arisen genetic unit will swamp the population in relatively f ew generations, even
if the advantage over its alternative f orms is moderately small.
Natural selection has been compared to a sieve which retains the rarely arising
usef ul and lets go the more f requently arising harmf ul mutants. Natural selection
acts in that way, but it is much more than a purely negative process, f or it is able to
generate novelty by increasing the probability of otherwise extremely improbable
genetic combinations. Natural selection is creative in a way. It does not "create"
the genetic entities upon which it operates, but it produces adaptive genetic combina-
tions which would not have existed otherwise. The creative role of natural selection
must not be understood in the sense of the "absolute" creation that traditional
Christian theology predicates of the Divine act by which the universe was brought
into being ex nihilo. Natural selection may be compared rather to a painter which
creates a picture by mixing and distributing pigments in various ways over the
canvas. The canvas and the pigments are not created by the artist but the painting
is. It is conceivable that a random combination of the pigments might result in the
orderly whole which is the f inal work of art. Some modern paintings look very much
like a random association of materials, to be sure. But the probability of , say,
Leonardo's Mona Lisa resulting f rom a random combination of pigments
is
nearly inf initely small. In the same way, the combination of genetic units which
carries the hereditary inf ormation responsible f or the f ormation of the vertebrate
eye could have never been produced by a random process like mutation. Not even
if we allow f or the three billion years plus during which lif e has existed on earth.
The complicated anatomy of the eye like the exact f unctioning of the kidney are the
result of a nonrandom process-natural selection.
How natural selection, a purely material process, can generate novelty in the
f orm of accumulated hereditary inf ormation may be illustrated by the f ollowing
example. Some strains of the colon bacterium, Escherichia coli, to be able to
reproduce in a culture medium, require that a certain substance, the amino acid
histidine, be provided in the medium. When a f ew such bacteria are added to a
cubic centimeter of liquid culture medium, they multiply rapidly and produce
between two and three billion bacteria in a f ew hours. Spontaneous mutations to
streptomycin resistance occur in normal, i.e., sensitive, bacteria at rates of the order
of one in one hundred million (1 x 10-8) cells. In our bacterial culture we expect
between twenty and thirty bacteria to be resistant to streptomycin due to spontan-
eous mutation. If a proper concentration of the antibiotic is added to the culture,
only the resistant cells survive. The twenty plus surviving bacteria will start re-
producing, however, and allowing a f ew hours f or the necessary number of cell
divisions, several billion bacteria are produced, all resistant to streptomycin.
Among cells requiring histidine as a growth f actor, spontaneous mutants able to
reproduce in the absence of histidine arise at rates of about f our in one hundred
million (4 x 10-8) bacteria. The streptomycin resistant cells may now be transf erred
to an agar-medium plate with streptomycin but with no histidine. Most of them
will not be able to reproduce, but about a hundred will start dividing and f orm
colonies until the available medium is saturated. Natural selection has produced
in two steps bacterial cells resistant to streptomycin and not requiring histidine f or
growth. The probability of the two mutational events happening in the same
bacterium is of about f our in ten million billion (1 x 10-8 x 4 x 10-8
-
4 x 10-16) cells. An event of such low probability is unlikely to occur even in a
large laboratory culture of bacterial cells. With natural selection cells having both
properties are the common result.
Natural selection produces highly improbable combinations of genes by pro-
ceeding step-wise. The human eye did not appear suddenly in all its present perf ec-
tion. It requires the appropriate integration of many genetic units, and thus it
could not have resulted f rom a random process. Our ancestors have had f or at least
the last half billion years some kind of organs sensitive to light. Perception of light,
and later vision, were important f or their survival and reproductive success.
Natural selection accordingly f avored genes and gene combinations increasing the
f unctional ef f iciency of the eye. Such genetic units gradually accumulated eventually
leading to the highly complex and ef f icient human eye.
Natural selection can account f or the rise and spread of genetic constitutions,
and theref ore of types of organisms, that would never have existed under the
uncontrolled action of random mutation and recombination of the hereditary
materials. In this sense, although it does not create the raw materials, that is, the
genes, selection is def initely creative.
3. Natural Selection and Adaptation. Evolutionary changes in the gene pool of a
population f requently occur in the direction of increased adaptation. The organisms
likely to leave more descendants are those whose variations are most advantageous
as adaptations
to the environment. Natural selection, however, occurs in ref erence
to the environment where the population presently lives. Evolutionary adaptations
are not anticipatory of the f uture. The environmental challenges that a population
may meet in the f uture cannot af f ect in any way the reproductive f itness of the
organisms in the present environment. If the population is unable to react adaptively
to a new environmental challenge, the result may be extinction. The f ossil record
bears witness that a majority of the species living in the past became eventually
extinct without issue.
The evolutionary course of a population is conditioned by the past history of the
population. The genetic conf iguration of a population is determined by the environ-
ments where the population has lived in the past. Those genes and genetic combina-
tions were f avorably selected which increased the reproductive f itness of their
carriers in the environments where the population lived. The present conf iguration
of its gene pool sets limits to the evolutionary potentialities of a population. The
only genes that may be f avored by natural selection are those actually present in the
population. An obvious example is the colonization of the land by organisms. The
colonization of the land by plants occurred during the Silurian geological period,
and by animals during the Devonian period. New and diversif ied environments
were open to the evolution of lif e. New f orms of plants evolved, but the basic
adaptations to plant lif e remained in all of them. These adaptations had occurred
in the past and set limits to the evolutionary potentialities of their descendants.
The considerable diversif ication of anatomic and physiological characteristics
that occurred in animals were not open to plants and vice versa.
Natural selection is thoroughly opportunistic. A new environmental challenge
is responded to by appropriate adaptations in the population or results in its
extinction. Adaptation to the same environment may occur in a variety of dif f erent
ways. An example may be taken f rom the adaptations of plant lif e to desert climate
(Dobzhansky [1]). The f undamental adaptation is to the condition of dryness which
carries the danger of desiccation. During a major part of the year, sometimes f or
several years in succession, there is no rain. Plants have accomplished the urgent
necessity of saving water in dif f erent ways. Cacti have transf ormed their leaves into
spines, having made their stems into barrels containing a reserve of water. Photo-
synthesis is perf ormed in the surf ace of the stem instead of in the leaves. Other
plants have no leaves during the dry season, but af ter it rains they burst into leaves
and f lowers and produce seeds. A third type of adaptation exists. Ephemeral plants
germinate f rom seeds, grow, f lower, and produce seeds-all within the space of the
f ew weeks while water is available. The rest of the year the seeds lie quiescent in the
soil.
Natural selection can explain the f acts of the adaptation of living organisms to
their environments and to their ways of lif e. The account of natural selection given
here is also consistent with the history of lif e as obtained f rom the f ossil record and
with the diversity of plants and animals existing today (Simpson [101). The f ossil
record shows that the evolution of lif e occurred in a haphazard f ashion. The
phenomena of radiation, expansion, relaying of one f orm by another, diversif ica-
tion, occasional trends and extinction shown by the f ossil record, are best explained
by the synthetic theory of evolution. They are not compatible with a preordained
plan whether imprinted f rom without by an omniscient Creator, or the result of the
orthogenetic activity of any immanent nonmaterial f orce, be it called "elan vital,"
"radial energy" or "vital f orce."
4. Teleological Explanations in Biology. Nagel
([5],
p. 24) has written that "the
notion of teleology is neither hopelessly archaic nor necessarily a mark of super-
stition." The concept of teleology is in general disrepute in modern science. The
main reason f or this discredit is that the notion of teleology is equated with the
belief that f uture events are active agents in their own realization. Such belief ,
however, is not necessarily implied in the concept of teleology. Teleological explana-
tions are appropriate in certain areas of natural science. In particular, I shall
attempt to show that teleological explanations are appropriate and indispensable
in biology, and that they are f ully compatible with causal accounts, although they
cannot be reduced to nonteleological explanations without loss of explanatory
content.
The notion of teleology arose most probably as a result of man's ref lection 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 towards
which he directs his activity. Human actions can be said to be purposef ul when they
are intentionally directed towards the obtention of a goal.
The plan or purpose of the human agent may f requently be inf erred f rom the
actions he perf orms. That is, his actions can be seen to be purposef ully or teleo-
logically ordained towards the obtention of the goal. In this sense the concept of
teleology can be extended, and has been extended, to describe actions, objects or
processes which exhibit an orientation towards a certain goal or end-state. No
requirement is necessarily implied that the objects or processes tend consciously
towards their specif ic goals, nor that there is any external agent directing the process
or the object towards its end-state or goal. In this generic sense, teleological explana-
tions are those explanations where the presence of an object or a process in a system
is explained by exhibiting its connection with a specif ic 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 exist-
ence of a certain state or property of the system. Moreover, they imply that such
contribution is the explanatory reason f or the presence of the process or object in
the system. It is appropriate to give a teleological explanation of the operation of
the kidney in regulating the concentration of salt in the blood, or of the structure
of the hand obviously adapted f or grasping. But it makes no sense to explain
teleologically the f alling of a stone, or a chemical reaction.
There are at least three categories of biological phenomena where teleological
explanations are appropriate, although the distinction between the categories need
not always be clearly def ined. These three classes of teleological phenomena are
established according to the mode of relationship between the object or process
and the end-state or property that accounts f or its presence. Other classif ications of
teleological phenomena are possible according to other principles of distinction. A
second classif ication will be suggested below.
(1) When the end-state or goal is consciously anticipated by the agent. This is
purposef ul activity and it occurs in man and probably in other animals. I am
acting teleologically when I pick up a pencil and paper in order to express in
writing my ideas about teleology. A deer running away f rom a mountain lion, or a
bird building its nest, has at least the appearance of purposef ul behavior.
(2) In connection with self -regulating or teleonomic systems, when there exists a
mechanism that enables the system to reach or to maintain a specif ic property in
spite of environmental f luctuations. The regulation of body temperature in mammals
is of this kind. In general the homeostatic reactions of organisms belong to this
category of teleological phenomena. Two types of homeostasis are usually distin-
guished by biologists-physiological and developmental homeostasis, although
intermediate situations may exist. Physiological homeostatic reactions 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 a structure like muscle due to strenuous use, are examples of this
type of homeostasis. Developmental homeostasis ref ers to the regulation of the
dif f erent paths that an organism may f ollow in its progression f rom zygote to
adult.
Self -regulating systems or servo-mechanisms built by man are teleological in
this second sense. The simplest example of such servo-mechanisms is a thermostat
unit that maintains a specif ied room temperature by turning on and of f the source
of heat. Self -regulating mechanisms of this kind, living or man-made, are controlled
by a f eed-back system of inf ormation.
(3) In ref erence to structures anatomically and physiologically designed to
perf orm a certain f unction. The hand of man is made f or grasping, and his eye f or
vision. Tools and certain types of machines made by man are teleological in this
sense. A watch f or instance, is made to tell time, and a f aucet to draw water. The
distinction between this and the previous category of teleological systems is some-
times blurred. Thus the human eye is able to regulate itself within a certain range
to the conditions of brightness and distance so as to perf orm its f unction more
ef f ectively.
Teleological mechanisms in living organisms are biological adaptations. They
have arisen as a result of the process of natural selection. The adaptations of
organisms-whether organs, homeostatic mechanisms, or patterns of behavior-
are explained teleologically in that their existence is accounted f or in terms of their
contribution to the reproductive f itness of the population. As explained above, a
f eature of an organism that increases its reproductive f itness will be selectively
f avored. Given enough time it will extend to all the members of the population.
Patterns of behavior, such as the nesting habits of birds or the web-spinning of
spiders, have developed because they f avored the reproductive success of their
possessors in the environments where the population lived. Similarly, natural
selection can account f or the presence of homeostatic mechanisms. Some processes
can be operative only within a certain range of conditions. If the conditions are
af f ected by the environment, natural selection will f avor self -regulating mechanisms
that maintain the system within the f unction range. In man death results if the body
temperature is allowed to rise or f all by more than a f ew degrees above or below
normal. Body temperature is regulated by dissipating heat in warm environments
through perspiration and dilatation 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 f unction is
explained teleologically in that its presence is accounted f or in terms of the con-
tribution it makes to reproductive success in the population. The vertebrate eye
arose because genetic mutations responsible f or its development arose and increased
the reproductive f itness of their possessors.
There are two levels of teleology in organisms. There usually exists a specif ic and
proximate end f or every f eature of an animal or plant. The existence of the f eature
is explained in terms of the f unction or end-state it serves. But there is also an ulti-
mate goal to which all f eatures contribute or have contributed in the past-
reproductive success. The ultimate end to which all other f unctions and ends
contribute is increased reproductive ef f iciency. 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 two ways. Firstly,
natural selection is a mechanistic end-directed process which results in increased
reproductive ef f iciency. Reproductive f itness can, then, be said to be the end result
or goal of natural selection. Secondly, natural selection is teleological in the sense
that it produces and maintains end-directed organs and processes, when the
f unction or end-state served by the organ or process contributes to the reproductive
f itness of the organisms.
However, the process of natural selection is not at all teleological in a dif f erent
sense. Natural selection does not tend in any way towards the production of
specif ic kinds of organisms or towards organisms having certain specif ic properties.
The over-all process of evolution cannot be said to be teleological in the sense of
proceeding towards certain specif ied goals, preconceived or not. The only non-
random process in evolution is natural selection understood as dif f erential re-
production. Natural selection is a purely mechanistic process and it is opportunistic
in the sense discussed above. The f inal result of natural selection f or any species
may be extinction, as shown by the f ossil record, if the species f ails to cope with
environmental change.
The presence of organs, processes and patterns of behavior can be explained
teleologically by exhibiting their contribution to the reproductive f itness of the
organisms in which they occur. This need not imply that reproductive f itness is a
consciously intended goal. Such intent must in f act be denied, except in the case of
the voluntary behavior of man and perhaps of some animals. In teleological
explanation the end-state is not to be understood as the ef f icient cause of the object
or process that it explains. The end-state is causally-and in general temporally
also-posterior.
Mayr (cf . [3], p. 42) has pointed out that the term "teleology" has been applied
to two dif f erent sets of phenomena. "On one hand is the production and perf ection
throughout the history of the animal and plant kingdoms of ever new and ever
improved DNA programs of inf ormation. On the other hand is the testing of these
programs and their decoding throughout the lif etime of each individual." The
behavioral activities or developmental processes of an individual are controlled by
the program of inf ormation encoded in the DNA inherited by the organism f rom
its parents. The decoding of the DNA programs of inf ormation can properly be
said to be a teleological-or as Mayr pref ers to call it, teleonomic-process.
Teleology has also been applied to the evolution of organisms, that is, to the
production and perf ection of DNA codes of inf ormation. The overall process of
evolution cannot be said to be teleological in the sense of directed towards the pro-
duction of specif ied DNA codes of inf ormation, i.e., organisms. But it is my con-
tention that it can be said to be teleological in the sense of being directed towards
the production of DNA codes of inf ormation which improve the reproductive
f itness of a population in the environments where it lives. The process of evolution
can also be said to be teleological in that it has the potentiality of producing end-
directed DNA codes of inf ormation, and has in f act resulted in teleologically
oriented structures, patterns of behavior, and self -regulating mechanisms.
Three categories of teleological systems have been distinguished above, according
to the nature of the relationship existing between the object or process and its
end-state or goal. Another classif ication of teleology may be suggested in ref erence
to the agency giving origin to the teleological mechanism. The end-directedness of
living organisms and their f eatures 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 ref er to these two kinds of teleology as "natural"
and "artif icial," but the other two terms, "internal" and "external," have already
been used (cf . [2], p. 193). Internal teleological systems are accounted f or by
natural selection which is a strictly mechanistic process. External teleological
mechanism are products of the human mind, or more generally, are the result of
purposef ul activity consciously intending specif ied ends.
Living organisms, then, exhibit internal teleology, but do not in general possess
external teleology. The overall process of evolution is not teleological in the external
sense. Evolution can be explained without recourse to a Creator or planning agent
external to the organisms themselves. There is no evidence either of any vital f orce
or immanent energy directing the process towards production of specif ied kinds
of organisms. The evidence of the f ossil record is against any necessitating f orce,
external or immanent, leading the process towards specif ied goals.
5. Teleology and Causality. Nagel ([5], p. 24, 25) has convincingly argued that
"teleological explanations are f ully compatible with causal accounts.... Indeed, a
teleological explanation can always be transf ormed into a causal one." Teleological
explanations can be ref ormulated, without loss of explicit content, to take the f orm
of nonteleological ones. A typical teleological statement in biology is the f ollowing,
"The f unction of gills in f ishes is respiration, that is the exchange of oxygen and
carbon dioxide between the blood and the external water." Statements of this kind
account f or the presence of a certain f eature A
(gills)
in
every member of a class of
systems S (f ish) which possess a certain organization C (the characteristic anatomy
and physiology of f ishes). It does so by declaring that when S is placed in a certain
environment E (water with dissolved oxygen) it will perf orm a f unction F (respira-
tion) only if S (f ish) has A (gills). The teleological statement, says Nagel, is a
telescoped argument the content of which can be unravelled approximately as
f ollows: When supplied with water containing dissolved oxygen, f ish respire; if
f ish have no gills, they do not respire even if supplied with water containing
dissolved oxygen; theref ore f ish have gills. More generally, a statement of the
f orm "The f unction of A in a system S with organization C is to enable S in
environment E to engage in process F" can be f ormulated more explicitly; "Every
system S with organization C and in environment E engage in f unction F; if S with
organization C and in environment E does not have A, then S cannot engage in F;
hence, S must have A." The dif f erence between a teleological explanation and a
nonteleological one is, then, one of emphasis rather than of asserted content. A
teleological explanation directs our attention to "the consequences f or a given
system of a constituent part or process." The equivalent nonteleological f ormula-
tion f ocuses attention on "some of the conditions ... under which the system per-
sists in its characteristic organization and activities" ([4], p. 405).
Although a teleological explanation can be ref ormulated in a nonteleological
one, the teleological explanation connotes something more than the equivalent
nonteleological one. A teleological explanation imples that the system under
consideration is directively organized. For that reason, teleological explanations
are appropriate in biology and in the domain of cybernetics but make no sense
when used in the physical sciences to describe phenomena like the f all of a stone.
Moreover, and most importantly, teleological explanations imply that the end result
is the explanatory reason f or the existence of the object or process which serves or
leads to it. A teleological account of the gills of f ish implies that gills came to
existence precisely because they serve f or respiration.
If the above reasoning is correct, the use of teleological explanations in biology
is not only acceptable but indeed indispensable. Biological organisms are systems
directively organized towards reproductive f itness. Parts of organisms are direct-
ively organized towards specif ic ends that, generally, contribute to the ultimate
goal of reproductive survival. One question biologists ask about organic structures
and activities is "What f or ?" That is, "What is the f unction or role of such structure
or such process ?" The answer to this question must be f ormulated in teleological
language. Only teleological explanations connote the important f act that plants and
animals are directively organized systems. That such connotation-or, in Nagel's
expression, "surplus meaning"-can always be expressed in nonteleological
language is beside the point. As Nagel ([4], p. 423) has written questions about the
value of an explanation "can be answered only by examining the ef f ective role an
explanation plays in inquiry and in the communication of ideas."
It has been noted by some authors that the distinction between systems that are
goal-directed and those which are not is highly vague. The classif ication of certain
systems as end-directed is allegedly rather arbitrary. A chemical buf f er, and elastic
solid or a pendulum at rest are examples of physical-systems that appear to be
goal-directed. I suggest the use of the criterion 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. A f eature of a system will be teleological in the
sense of internal teleology if the f eature has utility f or the system in which it exists
and if such utility explains the presence of the f eature in the systems. Utility in
living organisms is def ined in ref erence to survival or reproduction. A structure or
process of an organism is teleological if it contributes to the reproductive ef f iciency
of the organism itself , and if such contribution accounts f or the existence of the
structure or process. Man-made tools or mechanisms are teleological with external
teleology if they have utility, i.e., if they have been designed to serve a specif ied
purpose, which theref ore explains their existence and properties. If the criterion of
utility cannot be applied, a system is not teleological. Chemical buf f ers, elastic solids
and a pendulum at rest are not teleological systems.
The utility of f eatures of organisms is with respect to the individual or the
species in which they exist at any given time. It does not include usef ulness to any
other organisms. The elaborate plumage and display of the peacock serves the
peacock in its attempt to f ind a mate. The beautif ul display is not teleologically
directed towards pleasing man's aesthetic sense. That it pleases the human eye is
accidental, because it does not contribute to the reproductive f itness of the peacock
(except, of course, in the case of artif icial selection by man). The criterion of
utility introduces needed objectivity in the determination 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 development of corn seeds into corn plants is sometimes
said to be natural, while their transf ormation into the f lesh of birds or men is
asserted to be merely accidental" ([4], p. 424). The adaptation of corn seeds have
developed to serve the f unction of corn reproduction, not to become a palatable
f ood f or birds or man. The role of wild corn as f ood is accidental, and cannot be
considered a biological f unction of the corn seed in the teleological sense.
Some f eatures of organisms are not adaptive nor usef ul by themselves. They
have arisen because they are concommitant of other f eatures that are adaptive or
usef ul. Features of organisms may also be present because they were usef ul to the
organisms in the past although they are no longer adaptive. Vestigial organs like
the vermif orm appendix of man are f eatures of this kind. If they are neutral
to reproductive f itness they may remain in the population indef initely.
6. Teleology, Teleonomy, and Aristotle. I want to take up, very brief ly, two more
issues; the f irst is a semantic question, the second a historical one. Pittendrigh [7],
Simpson [10], Mayr [3], Williams [11], and others, have proposed to use the term
"teleonomic" to describe end-directed processes which do not imply that f uture
events are active agents in their own realization, nor that things or activities are
conscious agents or the product of such agents. They argue that the term "teleo-
logy" has sometimes been used to explain the animal and plant kingdoms as the
result of a preordained plan necessarily leading to the existing kinds of organisms.
To avoid such connotation, the authors argue, the term teleonomy should be used
to explain adaptation in nature as the result of natural selection.
Although the notion of teleology has been used, and it is still being used, in the
alleged sense, it is also true that other authors, like Nagel [4], [5], Goudge [2], etc.,
employ the term "teleology" without implying a preordained relationship of means
to an end. Thus, it might originate more conf usion than clarity to repudiate the
notion of teleology on the grounds that it connotes an intentional relationship of
means to an end. The point is that what is needed is to clarif y the notion of teleo-
logy by explaining the various meanings the term may have. One may then explicitly
express in which sense the term is used in a particular context.
Should the term "teleology" eventually be discarded f rom the scientif ic vocabul-
ary, or restricted in its meaning to preordained end-directed processes, I shall
welcome such event. But the substitution of a term by another does not necessarily
clarif y the issues at stake. It would still be necessary to explicate whatever term is
used instead of teleology, whether teleonomy or any other. It may f urther be noted
that the term "teleonomic" is commonly employed in the restricted sense of self -
regulating mechanisms. There are phenomena in biology that are end-directed
without being self -regulating mechanisms in the usual sense. The hand of man, f or
example.
Pittendrigh ([7], p. 394) has written that "It seems unf ortunate that the term
'teleology' should be resurrected.... The biologists' long-standing conf usion would
be more f ully removed if all end-directed systems were described by some other
term, like 'teleonomic,' in order to emphasize that the recognition and description
of end-directedness does not carry a commitment to Aristotelian teleology as an
ef f icient causal principle." The Aristotelian concept of teleology allegedly implies
that f uture events are active agents in their own realization. According to other
authors, Aristotelian teleology connotes that there exists an overall design in the
world attributable to a Deity, or at least that nature exists only f or and in relation
to man, considered as the ultimate purpose of creation (cf . Simpson [10], Mayr
[3]).
Science, f or
Aristotle,
is a knowledge of the "whys," the "reasons f or" true
statements. Of a thing we can ask f our dif f erent kinds of questions: "What is it ?",
"Out of what is it made?", "By what agent?", "What f or?" The f our kinds of
answers that can be elicited to these questions are his f our causes-f ormal, material,
ef f icient, and f inal. Only the third type of answer is causal in the modern scientif ic
sense. Aition, the Greek term that Cicero translated "cause" (causa, in Latin)
means literally ground of explanation, i.e., what can be answered to a question.
It does not necessarily mean causality in the sense of ef f icient agency.
According to Aristotle, to f ully understand an object we need to f ind out,
among other things, its end; what f unction does it serve or what results it produces.
An egg can be understood f ully only if we consider it as a possible chicken. The
structures and organs of animals have f unctions, are organized towards certain
ends. Living processes proceed towards certain goals. Final causes, f or Aristotle,
are principles of intelligibility; they are not in any sense active agents in their own
realization. For Aristotle, ends "never do anything. Ends do not act or operate,
they are never ef f icient causes." (cf . Randall, [8], p. 128).
According to Aristotle there is no intelligent maker of the world. The ends of
things are not consciously intended. Nature, man excepted, has no purposes. The
teleology of nature is objective, and empirically observable. It does not require the
inf erence of unobservable causes. (cf . Ross [9], Randall [8]) There is no God
designer of nature. According to Aristotle, if there is a God, He cannot have
purposes (Randall [8], p. 125).
Finally, f or Aristotle, the teleology of nature is wholly "immanent." The end
served by any structure or process is the good or survival of that kind of thing in
which they exist. Animals, plants, or their parts do not exist f or the benef it of any
other thing but themselves. Aristotle makes it clear that nutritious as acorns may
be f or a squirrel, they do not exist to serve as a squirrel's meal. The natural end of
an acorn is to become an oak tree. Anything else that may happen to the acorn is
accidental and may not be explained teleologically.
Aristotle's main concern was the study of organisms, and their processes and
structures. He observed the f acts of adaptation and explained them with consider-
able insight considering that he did not know about biological evolution. His error
was not that he used teleological explanations in biology, but that he extended the
concept of teleology to the nonliving world.
REFERENCES
(11
Dobzhansky, Th., "Determinism and Indeterminism in Biological Evolution," in Philo-
sophical Problems in Biology (ed. E. Smith), St. John's University Press, New York, 1966,
pp. 55-66.
(2] Goudge, T. A., The Ascent of Lif e, University of Toronto Press, Toronto, 1961.
(3] Mayr, E., "Cause and Ef f ect in Biology," in Cause and Ef f ect (ed. D. Lerner), Free Press,
New York, 1965, pp. 33-50.
(4] Nagel, E., The Structure of Science, Harcourt, Brace and World, New York, 1961.
(5] Nagel, E., "Types of Causal Explanation in Science," in Cause and Ef f ect, (ed. D. Lerner),
Free Press, New York, 1965, pp. 11-32.
[6] Paley, W., Natural Theology, Charles Knight, London, 1836.
[7] Pittendrigh, C. S., "Adaptation, Natural Selection and Behavior," in Behavior and
Evolution (eds. A. Roe and G. G. Simpson), Yale University Press, New Haven, 1958,
pp. 390-416.
[8] Randall, J. H., Aristotle, Columbia University Press, New York, 1960.
[9] Ross, D., Aristotle, 5th edit., Barnes and Noble, New York, 1949.
(10] Simpson, G. G., This View of Lif e, Harcourt, Brace and World, New York, 1964.
(11] Williams, G. C., Adaptation and Natural Selection, Princeton University Press, Princeton,
N.J., 1966.
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Teleological Explanations in Evolutionary Biology

Author(s): Francisco J. Ayala

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