Cause and Effect in Biology

Author(s): Ernst Mayr

Source: Science, New Series, Vol. 134, No. 3489 (Nov. 10, 1961), pp. 1501-1506
Published by: American Association for the Advancement of Science
Stable URL: http://www.jstor.org/stable/1707986
Accessed: 15-08-2017 20:13 UTC

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 L. Berthet, C. DeDuve, Bull. soc. chim. biol.
33, 21 (1951).
6. R. Wu and E. Racker, J. Biol. Chem. 234,
1029 (1959).
7. F. Lynen, in Proceedings International Sym-
posium on Enzyme Chemistry (Maruzen, To-
kyo, 1958), p. 25.
8. B. Chance and B. Hess, Science 129, 700
(1959); J. Biol. Chem. 234, 2421 (1959).
9. R. Balazs, Biochem. J. 72, 561 (1959).
10. L. G. Abood, E. Bunngraber, M. Taylor, J.
Biol. Chem. 239, 1307 (1959).
11. P. Hochstein, M. Woods, D. Burk, Federa-
tion Proc. 14, 227 (1955); P. Hochstein,
Science 125, 496 (1957).
12. H. G. DuBuy and M. L. Hesselbach, J. Natl.
Cancer Inst. 20, 403 (1958); J. Histochem. and
Cytochem. 4, 363 (1956).
13. A. C. Eisenberg, The Glycolysis and Respira-
tion of Tumors (Academic Press, New York,
1961).
14. N. O. Kaplan, in The Enzymes, P. D. Boyer,
H. Lardy, K. Myrbick, Eds. (Academic Press,
New York, 1960), vol. 3, pt. B, p. 105; C.
Carruthers and V. Suntzeff, Cancer Research
14, 29 (1954); S. Weinhouse, in Advances in
Cancer Research, J. P. Greenstein and A.
Haddow, Eds. (Academic Press, New York,
1955), vol. 3, p. 268.
15. A. L. Lehninger, J. Biol. Chem. 190, 345
(1951).
16. B. Chance and G. R. Williams, ibid. 217, 409
(1955); G. F. Maley, ibid. 224, 1029 (1957).
17 C. Cooper and A. L. Lelninger, ibid. 219,
489 (1956).
18. D. E. Green and F. L. Crane, in Proceedings
International Symposium on Enzyme Chemis-
try (Maruzen, Tokyo, 1958), p. 275.
19. W. W. Kielley and J. R. Bronk, J. Biol. Chem.
230, 521 (1958).
2(. B. Chance and B. Sacktor, Arch. Biochem.
Biophys. 76, 509 (1958).
21. T. M. Devlin and B. Bedell, in preparation.
22. E. C. Slater and K. W. Cleland, Biochem. J.
53, 557 (1953).
23. W. Chefurka, Enzymologia 17, 73 (1954);
R. W. Estabrook and B. Sactor, J. Biol. Chem.
233, 1014 (1958).
24. E. Zebe, A. Delbriick, T. Biicher, Biochem.
Z. 331, 254 (1959).
25. G. E. Boxer and C. E. Shonk, Cancer Re-
search 20, 85 (1960).
26. E. I. Ciaccio and D. L. Keller, Federation
Proc. 19, 34 (1960).
27. D. E. Green, D. M. Needham, J. G. Dewan,
Biochem J. 31, 2327 (1937).
28. R. L. Ringler and T. P. Singer, Biochim. et
Biophys. Acta 29, 661 (1958).
29. B. Sacktor, L. Packer, R. W. Estabrook, Arch.
Biochem. Biophys. 80, 68 (1959).
30. M. Klingenberg and W. Slenczka, Biochem.
Z. 331, 254 (1959).
31. E. I. Ciaccio, D. L. Keller, G. E. Boxer,
Biochim. et Biophys. Acta 37, 191 (1960).
32. E. Leva and S. Rapoport, J. Biol. Chem. 149,
47 (1943); T. Bicher and M. Klingenberg,
Angew. Chem. 70, 552 (1958).
33. P. Holzer, P. Glogner, G. Sedlmayr, Biochem.
Z. 330, 59 (1958).
34. A. Delbriick, H. Schimassek, K. Bartsch, T.
Bucher, ibid. 331, 297 (1959).
35. B. Sacktor and A. Dick, Cancer Research 20,
1408 (1960).
36. E. I. Ciaccio and J. B. Orange, Abstr. Am.
Chem. Soc. Meeting, Div. Biol. Chem., Sept.
1960, p. 27C.
37. H. P. Morris, H. Sidransky, B. P. Wagner,
H. M. Dyer, Cancer Research 20, 1252 (1960).
38. M. Woods, personal communication.
39. T. M. Devlin and B. Bedell, J. Biol. Chem.
235, 2134 (1960).
40. - , Federation Proc. 19, 34 (1960).
41. A. L. Lehninger, H. C. Sudduth, J. B. Wise,
J. Biol. Chem. 235, 2450 (1960).
42. H. A. Krebs, L. V. Eggleston, A. D'Alessan-
dro, Biochem. J. 79, 537 (1961).
43. T. M. Devlin and B. Bedell, in preparation;
T. M. Devlin and G. E. Boxer, Proc. Am.
Assoc. Cancer Research 3, 220 (1961).
44. A. Delbriick, E. Zebe, T. Biucher, Biochem. Z.
331, 273 (1959).
45. T. E. Conover and L. Ernster, Biochem.
Biophys. Research Communs. 2, 26 (1960).
46. N. O. Kaplan, S. P. Colowick, E. F. Neufeld,
J. Biol. Chem. 205, 1 (1953); N. O. Kaplan,
S. P. Colowick, L. J. Zatman, M. M. Ciotti,
ibid. 205, 31 (1953).
47. A. M. Stein, N. O. Kaplan, M. M. Ciotti,
ibid. 234, 979 (1959).
48. T. M. Devlin, ibid. 234, 962 (1959).
49. B. Reynafarje and V. R. Potter, Cancer Re-
search 17, 1112 (1957).
50. P. Talalay and H. G. Williams-Ashman, Proc.
Natl. Acad. Sci. U.S. 44, 15 (1958).
51. C. E. Wenner, M. A. Spirtes, S. Weinhouse,
Cancer Research 12, 44 (1952).
52. H. Busch, ibid. 15, 365 (1955).
53. , M. H. Goldberg, D. C. Anderson,
ibid. 16, 175 (1956).
54. H. Busch and P. V. Nair, J. Biol. Chem. 229,
377 (1957); J. Papaconstantinou and S. P.
Colowick, J. Biol. Chem. 236, 278, 285
(1961).
55. V. R. Potter, Federation Proc. 17, 691 (1958).

planation, prediction, and teleology)

must be the cardinal points in any dis-
cussion of causality and were quite
rightly singled out as such by Nagel (1).
Biology can make a significant contri-

Cause
Cause and
and
Effect
Effect
in Biology
in Biology bution to all three of them. But before I
can discuss this contribution in detail,
I must say a few words about biology
as a science.
Kinds
Kinds ofof
causes,
causes,
predictability,
predictability,
and teleology
and teleology
are
areviewed
viewedby aby
practicing
a practicing
biologist.biologist.
Biology

Ernst Mayr
The word biology suggests a uniform
and unified science. Yet recent develop-
ments have made it increasingly clear
that biology is a most complex area-
Being
Beinga apracticing
practicingbiologist
biologist
I feelI that
feel
tal),
tal), that
and
and Lecomte
Lecomtedu Noiiy,
du Noiiy,
among theamong
indeed,the
that the word biology is a label
II cannot
cannotattempt
attempt thethe
kindkind
of analysis
of analysis
of
more of
moreprominent
prominent authors
authors
of the of
recent
the recent
for two largely separate fields which
cause
causeandand effect
effect
in biological
in biological
phenom-
phenom-
past.
past.Though
Though these
these
authorsauthors
may differ
may differ
differ greatly in method, Fragestellung,
ena
ena that
thata alogician
logicianwould
would
undertake.
undertake.
I particulars,
in
in I
particulars, theythey
all agree
all in
agree
claiming
in claiming
and basic concepts. As soon as one goes
would instead like to concentrate on the that
thatliving
livingbeings
beings
and life
andprocesses
life processes
can- beyondcan-the level of purely descriptive
special difficulties presented by the clas- not
notbebe causally
causally
explained
explained
in terms inof terms of biology, one finds two very
structural
sical concept of causality in biology. physical
physical andand
chemical
chemical
phenomena.
phenomena.
It different
Itareas, which may be designated
From the first attempts to achieve ais our task to ask whether this assertion functional biology and evolutionary bi-
unitary concept of cause, the student of is justified, and if we answer this ques- ology. To be sure, the two fields have
causality has been bedeviled by these tion with "no," to determine the source many points of contact and overlap.
difficulties. Descartes's grossly mecha- of the misunderstanding. Any biologist working in one of these
nistic interpretation of life, and the log- Causality, no matter how it is defined fields must have a knowledge and ap-
ical extreme to which his ideas were in terms of logic, is believed to contain preciation of the other field if he wants
carried by Holbach and de la Mettrie, three elements: (i) an explanation of
The author is Alexander Agassiz professor of
inevitably provoked a reaction leading past events ("a posteriori causality"); zoology at Harvard University and director of the
to vitalistic theories which have been in (ii) prediction of future events; and Museum of Comparative Zoology, Harvard Col-
lege, Cambridge, Mass. This article is adapted
vogue, off and on, to the present day.(iii) interpretation of teleological-that from a lecture presented 1 February 1961 at
I have only to mention names like is, "goal-directed"-phenomena. Massachusetts Institute of Technology in the
1960-61 series of Hayden lectures on "Cause
Driesch (entelechy), Bergson (elan vi- The three aspects of causality (ex- and Effect."

10 NOVEMBER 1961
1501

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to avoid the label of a narrow-minded phenomena' in biology. Everything is the specifications of the code. The
specialist. Yet in his own research he time-bound and space-bound. The ani- codes, therefore, are in some respects
will be occupied with problems of either mal or plant or micro-organism he is highly specific; in other respects they
one or the other field. We cannot dis- working with is but a link in an evolu- merely specify "reaction norms" or gen-
cuss cause and effect in biology without tionary chain of changing forms, none eral capacities and potentialities.
first having characterized these two of which has any permanent validity." Let me illustrate this duality of codes
fields. There is hardly any structure or func- by the difference between two kinds of
Functional biology. The functional tion in an organism that can be fully birds with respect to "species recogni-
biologist is vitally concerned with the understood unless it is studied against tion." The young cowbird is raised by
operation and interaction of structural this historical background. To find the foster parents-let us say, in the nest
elements, from molecules up to organs causes for the existing characteristics, of a song sparrow or warbler. As soon
and whole individuals. His ever-repeated and particularly adaptations, of organ- as it becomes independent of its foster
question is "How?" How does some- isms is the main preoccupation of the parents it seeks the company of other
thing operate, how does it function? evolutionary biologist. He is impressedyoung cowbirds, even though it has
The functional anatomist who studies by the enormous diversity of the or- never seen a cowbird before! In con-
an articulation shares this method and ganic world. He wants to know the rea- trast, after hatching from the egg, a
approach with the molecular biologist sons for this diversity as well as the young goose will accept as its parent
who studies the function of a DNA pathway by which it has been achieved. the first moving (and preferably also
molecule in the transfer of genetic He studies
in- the forces that bring aboutcalling) object it can follow and become
formation. The functional biologist changesat-in faunas and floras (as in part "imprinted" to. What is programmed
tempts to isolate the particular compo- documented by paleontology), and he is, in one case, a definite "gestalt," in
nent he studies, and in any givenstudies studythe steps by which have evolved the other, merely the capacity to be-
the miraculous adaptations so charac- come imprinted to a "gestalt." Similar
he usually deals with a single individual,
a single organ, a single cell, or a teristic
single of every aspect of the organic differences in the specificity of the in-
part of a cell. He attempts to eliminate,world. herited program are universal through-
or control, all variables, and he repeats We can use the language of informa- out the organic world.
his experiments under constant or tion theory to attempt still another char- Let us now get back to our main
vary-
ing conditions until he believes he acterization
has of these two fields of biol- topic and ask: Is cause the same thing
clarified the function of the element he ogy. The functional biologist deals with in functional and evolutionary biology?
studies. The chief technique of the func-all aspects of the decoding of the pro-Max Delbriick, again, has reminded
tional biologist is the experiment, and grammed information contained in the us (2) that as recently as 1870 Helm-
his approach is essentially the same as DNA code of the fertilized zygote. holtz
The postulated "that the behavior of
that of the physicist and the chemist. evolutionary biologist, on the other living cells should be accountable in
Indeed, by isolating the studied phe-hand, is interested in the history of terms of motions of molecules acting
these codes of information and in the
nomenon sufficiently from the complex- under certain fixed force laws." Now,
ities of the organism, he may achievelaws that control the changes of these says Delbriick correctly, we cannot even
the ideal of a purely physical or chem- codes from generation to account
generation. In for the behavior of a single
ical experiment. In spite of certain other words, he is interested in the hydrogen atom. As he also says, "any
limitations of this method, one must causes of these changes. living cell carries with it the experiences
agree with the functional biologist that Many of the old arguments of biolog-of a billion years of experimentation by
such a simplified approach is an ab- ical philosophy can be stated far more its ancestors."
solute necessity for achieving his par- precisely in terms of these genetic codes. Let me illustrate the difficulties of the
ticular objectives. The spectacular suc- For instance, as Schmalhausen, in Rus- concept of causality in biology by an
cess of biochemical and biophysical sia, and I have pointed out independ- example. Let us ask: What is the cause
research justifies this direct, althoughently, the inheritance of acquired char- of bird migration? Or more specifically:
distinctly simplistic, approach. acteristics becomes quite unthinkable Why did the warbler on my summer
Evolutionary biology. The evolution- when applied to the model of the trans- place in New Hampshire start his south-
ary biologist differs in his method and fer of genetic information from a pe- ward migration on the night of the 25th
in the problems in which he is inter- ripheral phenotype to the DNA of the of August?
ested. His basic question is "Why?" germ cells. I can list four equally legitimate
When we say "why" we must always be But let us not have an erroneous causes for this migration.
aware of the ambiguity of this term. It concept of these codes. It is character- 1) An ecological cause. The warbler,
may mean "how come?," but it may istic of these genetic codes that beingthean insect eater, must migrate, be-
also mean the finalistic "what for?" It cause
programming is only in part rigid. it would starve to death if it
Such
is obvious that the evolutionist has inphenomena as learning, memory, should
non-try to winter in New Hampshire.
mind the historical "how come?" when genetic structural modification, and2) Are-genetic cause. The warbler has
he asks "why?" Every organism, wheth- generation show how "open" acquired
these pro- a genetic constitution in the
grams are. Yet, even here there course
er individual or species, is the product is greatof the evolutionary history of its
of a long history, a history which in- species to
specificity, for instance with respect which induces it to respond ap-
deed dates back more than 2000 mil- propriately
what can be "learned," at what stage in to the proper stimuli from
lion years. As Max Delbriick (2) has the environment.
the life cycle "learning" takes place, and On the other hand,
said, "a mature physicist, acquainting the screech owl, nesting right next to it,
how long a memory engram is retained.
himself for the first time with the prob- The program, then, may be in lacks this constitution and does not re-
part
lems of biology, is puzzled by the cir- quite unspecific, and yet the rangespond to ofthese stimuli. As a result, it is
cumstance that there are no 'absolute possible variation is itself included in
sedentary.
1502 SCIENCE, VOL. 134

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 3) An intrinsic physiological cause. case of adaptation to life on land. We many other phenomena that are not
The warbler flew south because its mi- know through Gudernatsch that the necessarily goal-seeking in nature.
gration is tied in with photoperiodicity. growth of the legs can be produced at Aristotelian scholars have rightly em-
It responds to the decrease in day length any time even in the youngest tadpole, phasized that Aristotle-by training and
and is ready to migrate as soon as the which is unable to live on land, by feed- interest-was first and foremost a bi-
number of hours of daylight have ing the animal with the thyroid gland." ologist, and that it was his preoccupa-
dropped below a certain level. Let us now get back to the definition tion with biological phenomena which
4) An extrinsic physiological cause. of "cause" in formal philosophy and dominated his ideas on causes and in-
Finally, the warbler migrated on the see how it fits with the usual explana- duced him to postulate final causes in
25th of August because a cold air mass, tory "cause" of functional and evolu- addition to the material, formal, and
with northerly winds, passed over ourtionary biology. We might, for instance,efficient causes. Thinkers from Aristotle
area on that day. The sudden drop in define cause as "a nonsufficient condi- to the present have been challenged by
temperature and the associated weather tion without which an event would not the apparent contradiction between a
conditions affected the bird, already in have happened," or as "a member of a mechanistic interpretation of natural
a general physiological readiness for set of jointly sufficient reasons without processes and the seemingly purposive
migration, so that it actually took offwhich the event would not happen" sequence of events in organic growth, in
on that particular day. [after Scriven (4)]. Definitions such as reproduction, and in animal behavior.
Now, if we look over the four causa- these describe causal relations quite ade- Such a rational thinker as Bernard (5)
tions of the migration of this bird oncequately in certain branches of biology, has stated the paradox in these words.
more we can readily see that there is anparticularly in those which deal with There is, so to speak, a preestablished de-
immediate set of causes of the migra- chemical and physical unit phenomena. sign of each being and of each organ of
tion, consisting of the physiological con-In a strictly formal sense they are also such a kind that each phenomenon by
dition of the bird interacting with applicable to more complex phenom- itself depends upon the general forces of
nature, but when taken in connection with
photoperiodicity and drop in tempera- ena, and yet they seem to have little
the others it seems directed by some in-
ture. We might call these the proximate operational value in those branches of visible guide on the road it follows and
causes of migration. The other two biology that deal with complex systems. led to the place it occupies.
causes, the lack of food during winter I doubt that there is a scientist who We admit that the life phenomena are
and the genetic disposition of the bird, would question the ultimate causality attached to physicochemical manifesta-
tions, but it is true that the essential is not
are the ultimate causes. These are of all biological phenomena-that is, explained thereby; for no fortuitous com-
causes that have a history and that that have
a causal explanation can be given ing together of physicochemical phenom-
been incorporated into the system for past biological events. Yet such an ena constructs each organism after a plan
through many thousands of generations explanation will often have to be so un- and a fixed design (which are foreseen in
of natural selection. It is evident that advance) and arouses the admirable sub-
specific and so purely formal that its
ordination and harmonious agreement of
the functional biologist would be con- explanatory value can certainly be chal- the acts of life. . . Determinism can
cerned with analysis of the proximate lenged. In dealing with a complex sys- never be [anything] but physicochemical
causes, while the evolutionary biologist tem, an explanation can hardly be con- determinism. The vital force and life be-
would be concerned with analysis of sidered very illuminating that states: long to the metaphysical world.
the ultimate causes. This is the case with "Phenomenon A is caused by a complex What is the x, this seemingly purpo-
almost any biological phenomenon we set of interacting factors, one of which sive agent, this "vital force," in organic
might want to study. There is always a is b." Yet often this is about all one phenomena? It is only in our lifetime
proximate set of causes and an ultimate can say. We will have to come back to that explanations have been advanced
set of causes; both have to be ex- this difficulty in connection with the which deal adequately with this paradox.
plained and interpreted for a complete problem of prediction. However, let us The many dualistic, finalistic, and vi-
understanding of the given phenomenon. first consider the problem of teleology. talistic philosophies of the past merely
Still another way to express these replaced the unknown x by a different
differences would be to say that proxi- unknown, y or z, for calling an un-
mate causes govern the responses of theTeleology known factor entelechia or elan vital
individual (and his organs) to immediate is not an explanation. I shall not waste
factors of the environment while ulti- No discussion of causality is complete time showing how wrong most of these
mate causes are responsible for the evo- which does not come to grips with the past attempts were. Even though some
lution of the particular DNA code of problem of teleology. This problem had of the underlying observations of these
information with which every individual its beginning with Aristotle's classifica- conceptual schemes are quite correct,
of every species is endowed. The logi- tion of causes, one of the categories the supernaturalistic conclusions drawn
cian will, presumably, be little con- being the "final" causes. This category from these observations are altogether
cerned with these distinctions. Yet, theis based on the observation of the or- misleading.
biologist knows that many heated argu-derly and purposive development of theWhere, then, is it legitimate to speak
ments about the "cause" of a certain individual from the egg to the "final" of purpose and purposiveness in nature,
biological phenomenon could have been stage of the adult, and of the develop- and where is it not? To this question
avoided if the two opponents had real- ment of the whole world from its be- we can now give a firm and unambigu-
ized that one of them was concerned ginnings (chaos?) to its present order. ous answer. An individual who-to use
with proximate and the other with ulti- Final cause has been defined as "the the language of the computer-has
mate causes. I might illustrate this bycause a responsible for the orderly reach- been "programmed" can act purpose-
quotation from Loeb (3): "The earlier ing of a preconceived ultimate goal." fully. Historical processes, however, can
writers explained the growth of the legs All goal-seeking behavior has been clas- not act purposefully. A bird that starts
in the tadpole of the frog or toad as a
sified as "teleological," but so have its migration, an insect that selects its
10 NOVEMBER 1961
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host plant, an animal that avoids a It is now evident that the terms We can summarize this discussion by
predator, a male that displays to a fe- teleology and teleological have been statingap-
that there is no conflict between
male-they all act purposefully because plied to two entirely different causality
sets andof teleonomy, but that scien-
they have been programmed to do so. phenomena. On one hand is the pro-
tific biology has not found any evidence
When I speak of the programmed "in- duction and perfecting throughout the support teleology in the sense
that would
dividual," I do so in a broad sense. A history of the animal and plant of various
king-vitalistic or finalistic theories
programmed computer itself is an "in- doms of ever-new programs and (9, of10). All the so-called teleological
ever-
improved DNA codes of information.
dividual" in this sense, but so is, during systems which Nagel discusses (11) are
reproduction, a pair of birds whose in-On the other hand there is the testing actually illustrations of teleonomy.
stinctive and learned actions and inter- of these programs and the decoding of
actions obey, so to speak, a single these codes throughout the lifetime of
program. each individual. There is a fundamental The Problem of Prediction
The completely individualistic and difference between, on the one hand,
yet also species-specific DNA code of end-directed behavioral activities or de- The third great problem of causality
every zygote (fertilized egg cell), which velopmental processes of an individual in biology is that of prediction. In the
controls the development of the central or system, which are controlled by a classical theory of causality the touch-
and peripheral nervous systems, of the program, and, on the other hand, the stone of the goodness of a causal ex-
sense organs, of the hormones, of physi-steady improvement of genetic codes. planation was its predictive value. This
ology and morphology, is the program This genetic improvement is evolution- view is still maintained in Bunge's mod-
for the behavior computer of this in-ary adaptation controlled by natural ern classic (12): "A theory can predict
dividual. selection. to the extent to which it can describe
Natural selection does its best to In order to avoid confusion between and explain." It is evident that Bunge
the two entirely different types of
favor the production of codes guaran- is a physicist; no biologist would have
end
direction,
teeing behavior that increases fitness. A Pittendrigh (6) has intro- made such a statement. The theory of
behavior program that guarantees ducedin-
the term teleonomic as a descrip- natural selection can describe and ex-

stantaneous correct reaction to a tive term for all end-directed systems

poten- plain phenomena with considerable pre-
"not committed to Aristotelian teleol-
tial food source, to a potential enemy, cision, but it cannot make reliable pre-
or to a potential mate will certainly dictions, except through such trivial and
ogy." Not only does this negative defini-
give greater fitness in the Darwinian tion place the entire burden on the word meaningless circular statements as, for
sense than a program that lacks these system, but it makes no clear distinc-instance: "the fitter individuals will on
properties. Again, a behavior program tion between the two teleologies of the average leave more offspring." Scri-
that allows for appropriate learning and Aristotle. It would seem useful to re- ven (13) has emphasized quite correctly
the improvement of behavior reactions strict the term teleonomic rigidly that
to one of the most important contri-
by various types of feedbacks gives systems operating on the basis of abutions
pro- to philosophy made by the
evolutionary theory is that it has
greater likelihood of survival than a gram, a code of information. Teleono-
program that lacks these properties. demonstrated the independence of ex-
my in biology designates "the apparent
The purposive action of an individ- purposefulness of organisms and their planation and prediction.
ual, insofar as it is based on the prop- characteristics," as Julian Huxley ex- Although prediction is not an In-
erties of its genetic code, therefore is pressed it (7). separable concomitant of causality, ev-
no more nor less purposive than the Such a clear-cut separation of tele- ery scientist is nevertheless happy if his
actions of a computer that has been onomy, which has an analyzable physi- causal explanations simultaneously have
programmed to respond appropriately cochemical basis, from teleology, which high predictive value. We, can dis-
to various inputs. It is, if I may say so, deals more broadly with the over-all tinguish many categories of prediction
a purely mechanistic purposiveness. harmony of the organic world, is most in biological explanation. Indeed, it is
We biologists have long felt that it is useful because these two entirely dif- even doubtful how to define "predic-
ambiguous to designate such pro- ferent phenomena have so often been tion" in biology. A competent zoogeog-
grammed, goal-directed behavior "tele- confused with each other. rapher can predict with high accuracy
ological," because the word teleological The development or behavior of anwhat animals will be found on a previ-
has also been used in a very differentindividual is purposive, natural selection ously unexplored mountain range or
sense, for the final stage in evolutionaryis definitely not. When MacLeod (8) island. A paleontologist likewise can
adaptive processes. When Aristotle stated, "What is most challenging about predict with high probability what kind
spoke of final causes he was particularly Darwin, however, is his re-introduction of fossils can be expected in a newly
concerned with the marvelous adapta- of purpose into the natural world," he accessible geological horizon. Is such
tions found throughout the plant and chose the wrong word. The word pur- correct guessing of the results of past
animal kingdom. He was concerned pose is singularly inapplicable to evolu-events genuine prediction? A similar
with what later authors have called de- tionary change, which is, after all, whatdoubt pertains to taxonomic predictions,
sign or plan in nature. He ascribed to Darwin was considering. If an organism as discussed in the next paragraph. The
final causes not only mimicry or sym- is well adapted, if it shows superiorterm prediction is, however, surely legit-
biosis but all the other adaptations of fitness, this is not due to any purposeimately used for future events. Let me
animals and plants to each other and to of its ancestors or of an outside agency,give you four examples to illustrate the
their physical environment. The Aristo- such as "Nature" or "God," who cre- range of predictability.
ated a superior design or plan. Darwin 1) Prediction in classification. If I
telians and their successors asked them-
"has swept out such finalistic teleology have identified a fruit fly as an individ-
selves what goal-directed process could
have produced such a well-ordered byde-the front door," as Simpson (9) has ual of Drosophila melanogaster on the
sign in nature. rightly said. basis of bristle pattern and the propor-
1504 SCIENCE, VOL. 134

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tions of face and eye, I can "predict" Table 1. Two species of Tribolium in differences
compe- between biology and the
tition [from Park (14)].
numerous structural and behavioral physical sciences. Physicists and chem-
characteristics which -I will find if I Condition Victorious species ists often have genuine difficulty in un-
____ _ ~Replicas (No. of trials)derstanding the biologist's stress of the
study other aspects of this individual. If
Temp. Humidity (No.) T. con- T. cast- unique, although such an understanding
I find a new species with the diagnostic (?C) (%) fusum aneum
key characters of the genus Drosophila, has been greatly facilitated by the de-
34 70 30 30 velopments in modern physics. If a
I can at once "predict" a whole set of 29 70 66 11 55
biological properties. 24 70 30 21 9 physicist says "ice floats on water," his
2) Prediction of most physicochem- 34,29 30 60 53 7 statement is true for any piece of ice
24 30 20 20
ical phenomena on the molecular level. and any body of water. The members
Predictions of very high accuracy can of a class usually lack the individuality
be made with respect to most biochem- that is so characteristic of the organic
ical unit processes in organisms, such asand again that world, where all individuals are unique;
independent
metabolic pathways, and with respectlines exposed allto stages in the life cycle
the same are unique; selec
to biophysical phenomena in simple sure will respond all populations areat unique; all species
different
systems, such as the action of light, with different and higher categories
correlated are unique; all effe
heat, and electricity in physiology. of them predictable. interindividual contacts are unique; all
In examples 1 and 2 the predictive As is true in natural
many associations of species are
other bra
value of causal statements is usually science, the unique; validity and all evolutionary of events are
pred
very high. Yet there are numerous otherbiological phenomena (except for a unique. Where these statements are ap-
generalizations or causal statements in few chemical or physical unit processes) plicable to man, their validity is self-
biology that have low predictive values.is nearly always statistical. We can pre- evident. However, they are equally valid
The following examples are of this kind.dict with high accuracy that slightly for all sexually reproducing animals
3) Prediction of the outcome of more than 500 of the next 1000 new- and plants. Uniqueness, of course, does
complex ecological interactions. The borns will be boys. We cannot predict not entirely preclude prediction. We
statement, "An abandoned pasture in the sex of a particular unborn child. can make many valid statements about
southern New England will be replaced the attributes and behavior of man, and
by a stand of grey birch (Betula populi- the same is true for other organisms.
folia) and white pine (Pinus strobus)" Reasons for Indeterminacy in Biology But most of these statements (except
is often correct. Even more often, how- for those pertaining to taxonomy) have
ever, the replacement may be an almost Without claiming to exhaust all the statistical validity. Uniqueness is
purely
solid stand of P. strobus, or P. strobus possible reasons for indeterminacy, particularly
I characteristic for evolu-
may be missing altogether and in its can list four classes. Although they tionary biology. It is quite impossible
stead will be cherry (Prunus), red cedar somewhat overlap each other, each de- to have for unique phenomena general
(Juniperus virginianus), maples, sumac, serves to be treated separately. laws like those that exist in classical
and several other species. 1) Randomness of an event with re- mechanics.
Another example also illustrates this spect to the significance of the event. 3) Extreme complexity. The physi-
unpredictability. When two species of Spontaneous mutation, caused by an cist Elsisser stated in a recent sympo-
flour beetles (Tribolium confusum and "error" in DNA replication, illustrates sium: "[an] outstanding feature of all
T. castaneum) are brought together in this cause for indeterminacy very well. organisms is their well-nigh unlimited
a uniform environment (sifted wheat The occurrence of a given mutation is structural and dynamical complexity."
flour), one of the two species will al- in no way related to the evolutionary This is true. Every organic system is so
ways displace the other. At high tem- needs of the particular organism or of rich in feedbacks, homeostatic devices,
peratures and humidities, T. castaneum the population to which it belongs. The and potential multiple pathways that a
will win out; at low temperatures and precise results of a given selection pres- complete description is quite impossible.
humidities, T. confusum will be the sure are unpredictable because muta- Furthermore, the analysis of such a sys-
victor. Under intermediate conditions tion, recombination, and developmental tem would require its destruction and
the outcome is indeterminate and hence homeostasis are making indeterminate would thus be futile.
unpredictable (Table 1) (14). contributions to the response to this 4) Emergence of new qualities at
4) Prediction of evolutionary events. pressure. All the steps in the determi- higher levels of integration. It would
Probably nothing in biology is less pre- nation of the genetic contents of a lead too far to discuss in this context
dictable than the future course of evo- zygote contain a large component the thorny problem of "emergence."
lution. Looking at the Permian reptiles, of this type of randomness. What we All I can do here is to state its principle
who would have predicted that most of have described for mutation is also dogmatically: "When two entities are
the more flourishing groups would be- true for crossing over, chromosomal combined at a higher level of integra-
come extinct (many rather rapidly), and segregation, gametic selection, mate tion, not all the properties of the new
that one of the most undistinguished selection, and early survival of the entity are necessarily a logical or pre-
branches would give rise to the mam- zygotes. Neither underlying molecular dictable consequence of the properties
mals? Which student of the Cambrian phenomena nor the mechanical mo- of the components." This difficulty is
fauna would have predicted the revolu- tions responsible for this randomness by no means confined to biology, but
tionary changes in the marine life areofrelated to their biological effects. it is certainly one of the major sources
the subsequent geological eras? Unpre- 2) Uniqueness of all entities at the of indeterminacy in biology. Let us re-
dictability also characterizes small-scale
higher levels of biological integration. member that indeterminacy does not
evolution. Breeders and students of In the uniqueness of biological entitiesmean lack of cause, but merely un-
natural selection have discoveredand phenomena lies one of the major predictability.
again
10 NOVEMBER 1961 1505

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 All
All four
fourcauses
causes
of indeterminacy,
of indeterminacy, in- in- 1) Causality
Causality in in biology
biologyisisa afar
farcry
cry Finally,
Finally, causality
causalityininbiology
biologyis is
not not
in in
dividually
dividually and
andcombined,
combined,reducereduce
the the from
from causality
causality in
in classical
classicalmechanics.
mechanics.real
real conflict
conflictwith
withthe
the causality
causalityof of
clas-
clas-
precision
precisionofofprediction.
prediction. 2) Explanations
Explanations of of all
allbut
butthe
thesim-
sim-
sical
sical mechanics.
mechanics.As Asmodern
modern physics
physics hashas
One
Onemaymayraise
raise
thethe
question
question
at this plest
plest biological
at this biological phenomena
phenomenausuallyusuallycon-
con-
also
also demonstrated,
demonstrated,the thecausality
causalityof of
clas-
clas-
point
pointwhether
whether predictability
predictability
in classical sist of
in classical of sets
sets of
of causes.
causes.This sical
sical mechanics
Thisisisparticularly
particularly mechanicsisisonly
only a very
a very simple,
simple,
mechanics
mechanics andand
unpredictability
unpredictability
in biol- true for those biological phenomena
in biol- special
special case
caseof
ofcausality.
causality.Predictability,
Predictability,
ogy
ogyare
aredue
dueto to
a difference
a difference
of degree
of degree
or that
or can be understood only if their for
for instance,
instance,isisnotnota anecessary
necessarycom-
com-
of
of kind.
kind.There
Thereis much
is much
to suggest
to suggest
that that
evolutionary history is also considered. ponent
ponent of ofcausality.
causality.The
The complexities
complexities
of of
the
thedifference
differenceis, is,
in considerable
in considerable Each set is like a pair of brackets which biological
part, part, biologicalcausality
causalitydodonotnot
justify
justifyem-em-
merely a matter of degree. Classical contains much that is unanalyzed and bracing
bracing nonscientific
nonscientificideologies,
ideologies,
such
such
as as
mechanics is, so to speak, at one end much that can presumably never be vitalism or finalism, but should en-
of a continuous spectrum, and biologyanalyzed completely. courage all those who have been trying
is at the other. Let us take the classical 3) In view of the high number of to give a broader basis to the concept
example of the gas laws. Essentially multiple pathways possible for most of causality.
biological processes (except for the
they are only statistically true, but the
References and Notes
population of molecules in a gas obey- purely physicochemical ones) and in
1. E. Nagel, lecture presented at the Massachu-
view of the randomness of many of the setts Institute of Technology in the 1960-61
ing the gas laws is so enormous that the
actions of individual molecules become biological processes, particularly on the Hayden Lectures series.
molecular level (as well as for other 2. M. Delbriick, Trans. Conn. Acad. Arts Sci.
integrated into a predictable-one might 38, 173 (1949).
say "absolute"-result. Samples of reasons),
five causality in biological sys- 3. J. Loeb, The Organism as a Whole (Putnam,
New York, 1916).
or 20 molecules would show definite
tems is not predictive, or at best is only 4. M. Scriven, unpublished manuscript.
statistically
individuality. The difference in the size predictive. 5. C. Bernard, Lemons sur les phenomenes de la
vie (1885), vol. 1.
of the studied "populations" certainly 4) The existence of complex codes of 6. C. S. Pittendrigh, in Behavior and Evolution,
contributes to the difference between information in the DNA of the germ A. Roe and G. G. Simpson, Eds. (Yale Univ.
Press, New Haven, Conn., 1958), p. 394.
the physical sciences and biology.plasm permits teleonomic purposive- 7. J. Huxley, Zool. Jahrb. Abt. Anat. u. Ontog.
ness. On the other hand, evolutionary Tiere 88, 9 (1960).
research has found no evidence what- 8. R. B. MacLeod, Science 125, 477 (1957).
9. G. G. Simpson, ibid. 131, 966 (1960).
Conclusions soever for a "goal-seeking" of evolu- 10. ?, Sci. Monthly 71, 262 (1950); L. F.
Koch, ibid. 85, 245 (1957).
tionary lines, as postulated in that kind
11. E. Nagel, The Structure of Science (Harcourt
Let us now return to our initial ques-of teleology which sees "plan and de- Brace and World, Inc., New York, 1961).
tion and try to summarize some of our sign" in nature. The harmony of 12. the M. Bunge, Causality (Harvard Univ. Press,
Cambridge, Mass. 1959), p. 307.
conclusions on the nature of the cause- living universe, so far as it exists, is
13. an
M. Scriven, Science 130, 477 (1959).
and-effect relations in biology. a posteriori product of natural selection.
14. T. Park, Physiol. Zool. 27, 177 (1954).

preliminary
preliminarycalculations
calculations
which
which
are of
are of
physiological
physiologicalinterest
interest
in in
terms
terms
of the
of the
hazard associated with laser beams and
their potential employment as biological
and clinical tools.

Physiological Implications
Properties
of Laser Beams From the point of view of physio-
logical interest there are two important
The very high radiation flux densities of optical properties of laser beams, the extremely
collimated character of the light and its
masers point to important biomedical applications. high degree of monochromaticity. The
collimation property implies the possi-
bility of obtaining large energy densities
Leonard R. Solon, Raphael Aronson, Gordon Gould
in narrow beams. The optimum diver-
gence angle of a laser beam, 4min, is
limited only by the wavelength of the
light emitted and the diameter of the
Development of molecular amplifiers light amplification by by stimulated
stimulated emis-
emis- laser source in accord with the Fraun-
in the visible and near-visible region sion of radiation. Such devices have hofer diffraction relationship:
(1) of the electromagnetic spectrum been successfully demonstrated at sev-
5min = 2.44 X/DL (1)
has been in progress at several labora- eral places (2), and several industrial
tories. Such amplifiers go under the organizations have made them available Here Here X is
Xthe
is wavelength
the wavelength
of the emitted
of the emitted
designation of "laser" or optical maser, commercially. It is almost certain that laser
laserlight
light
and Di
and
is the
Didiameter
is the of diameter of
the former term being an acronym for lasers will be incorporated into com- the
thebeam
beam emerging
emerging
from thefrom
laser source
the laser source
The authors are affiliated with TRG, Inc. munications and other technologies at or
orfrom
froma subsequent
a subsequent
lens system,
lens
if system, if
(Technical Research Group), Syosset, N.Y. a rapid rate. This article presents some one is used.
1506 SCIENCE, VOL. 134

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