Biology, Molecular and Organismic

Author(s): Theodosius Dobzhansky

Source: American Zoologist, Vol. 4, No. 4, (Nov., 1964), pp. 443-452
Published by: Oxford University Press
Stable URL: http://www.jstor.org/stable/3881145
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 BIOLOGY, MOLEGULAR AND ORGANISMIG

Theodosius Dobzhansky

The Rockefeller Institute, New York City

It is customary for presidents of scientific magnitude, although probably not in the

societies to address those who saw fit to elect depth of the philosophical repercussions,
them. The wise course is to choose a safe to that which occurred roughly a century
and noncontroversial topic; for a postpran- ago under the stimulus of Darwin's theory
dial address, a mildly humorous topic is of evolution. Every biologist feels gratified
to enable the to
preferable speaker display by this advance, and hopes that further
his wit, if he has any. I am following nei- research in molecular biology will be pur-
ther the custom nor the council of wisdom. sued with all possible What is de-
vigor.
This address has been postponed long batable is the situation of the organismic
enough for me to achieve the obscurity of
biology vis-a-vis its molecular sibling.
an ex-president, and my topic is controver-
Nothing succeeds like success. In molecu?
sial and not particularly amusing. How? lar biology, one spectacular discovery has
ever, the relationships of molecular and
followed closely on the heels of another.
organismic biology, zoology and botany, are Molecular has become a glamor
biology
most certainly of current interest; I am
field. It has attracted many able young
probably not far wrong to surmise that this students as well as older investigators.
topic is the most often discussed one among Glamor and brilliance generate enthusiasm
zoologists. I expect that what I am going and optimism; they may also dazzle and
to say may seem provocative to some of you; blindfold. The notion has gained some
I do not expect it to seem boring. that
currency the only worthwhile biology
Indisputably, molecular biology has is molecular biology. All else is "bird
achieved in our day advances of signal im- or "butterfly
watching" collecting." Bird
portance. Suffice it to mention the elucida-
watching and butterfly collecting are occu-
tion of the chemistry of heredity, the break-
pations manifestly unworthy of serious sci-
ing of the genetic code, studies on protein entists! I have heard a man whose official
structure and synthesis, and the unraveling title happens to be Professor of Zoology
of the sequences of chemical reactions in declare to an assembly of his colleagues
metabolic processes. All this has brought that "a good man cannot teach zoology."
biology considerably closer than it was a A good man can teach, of course, only mo?
generation ago to understanding the phe- lecular biology.
nomena of life. Although it is equally in-
Such pronunciamentos can be dismissed
disputable that this understanding is no-
as merely ridiculous. They are, however,
where near complete as yet, it is fair to say
caricatures of opinions entertained by some
that we are the living witnesses of a great
intelligent and reasonable people, whose
efflorescence of biological sciences. The
views deserve an honest and careful con-
modern advance is perhaps comparable in
sideration and analysis. Science must cope
It is a pleasure to acknowledge here the kindness with new problems that arise and devise
of my colleagues Messrs. W. Anderson, A. G. Bearn, new approaches to old problems. Some
E. Bosiger, R. J. Dubos, H. G. Frankfurt, A. E.
lines of research become less profitable and
Mirsky, and B. R. Voeller, who have read this
address in manuscript and have contributed valu- less exciting and others more so. The prog-
able suggestions and corrections. The responsi- ress in a given field of study may slacken
bility for the ideas expressed, as well as for the
errors of commission, or omission, is nevertheless because the approaches used have already
entirely my own. yielded most of what they are capable of
[This address was presented at the summer meet? scien-
ing of the American Society of Zoologists, Boulder, yielding. Probably every thoughtful
Colorado, August 27, 1964.?Ed.] tist can give examples of research efforts

44
444 Theodosius Dobzhansky

which have bogged down, and of types of organelles. Chromosomes and genes have
inquiry which seem to have run into at that extraordinary chemical substance, the
least temporarily impassable obstacles. If DNA, as the key constituent. But the DNA
such researches and inquiries are not aban- in the chromosome is something more than
doned altogether they usually drift into the DNA in a test tube. A chromosome is
more and more narrow specializations and an organized body, and its organization is
uninspired repetitiousness. On the other as essential as is its composition.
hand, an apparently depleted field may The supra-individual forms of integra-
burst into renewed fertility when a new tion seem less tangible in a spatio-temporal
idea or a new technique is invented. sense than the infra-individual ones, but
Is it, then, possible that biology other just as interesting and significant. Mankind
than molecular biology has entered upon is less clearly perceived by our sense organs
a period of doldrums? It is good for any than an individual man, but it is neverthe-
scientist from time to time to re-examine less as meaningful a biological entity as it
and to re-think his aims, purposes, and ap- is a cultural entity. The sexual mode of
proaches. Intellectual laziness has been the reproduction connects individuals into re?
undoing of many a capable scientist, who productive communities, Mendelian popu?
rested on the comforting assumption that lations. Mendelian populations are united
what was good a generation ago is good by reproductive bonds into inclusive repro?
enough today. A line of research is not ductive systems?biological species. An iso?
necessarily good because it is traditional, lated individual, especially an individual
and it is not necessarily worth pursuing of a sexual species, is at least as clearly an
because it has become an ingrown habit. anomaly as a cell isolated from a multi-
But neither is newness and fashionableness cellular body. With asexual modes of re?
a valid enough reason to choose one's line production, the bonds which integrate
of work. Let us face the problem squarely individuals in Mendelian populations and
and honestly. biological species are absent. Other bonds
Biology is structured rather differently operate, however, in sexual as well as in
from other natural sciences. Since this is asexual organisms. Individuals and species
equally true of zoology and botany, I prefer belong to ecological communities and eco-
to use here the inclusive word, biology, systems. An individual taken out of the
covering both. A biologist, more than, for system in which it normally occurs is in?
example, a physicist or a geologist, is faced complete and it may be inviable.
with several hierarchically superimposed The hierarchy of levels of biological in-
levels of integration in the objects which tegration may be represented schematically
he studies. Life presents itself to our view as the following sequence: molecule, cellu?
almost always in the form of discrete quanta lar organelle, cell, tissue, organ, individual,
?individuals. But unlike the atoms of Mendelian population, species, community,
classical physics, individuals are conspicu- ecosystem. This sequence is, to be sure, not
ously divisible, and, unlike the atoms of everywhere rigorously adhered to. There
modern physics, divisible into great num? are unicellular (or acellular) as well as
bers and a great variety of component ele- multicellular organisms; the sexual and
ments, cells. Cells are, in turn, complexly the asexual modes of reproduction impose,
structured and well-integrated entities. as indicated above, different modes of in-
They contain chemical substances of nu- tegration. Even the level of an individual
merous, probably thousands, molecular spe? is not always unambiguously distinct. Con-
cies. It is, however, a gross error to think sider a colonial form, such as a siphono-
of a cell simply as a mixture of chemicals, phore; an individual of the higher order
like a mixture that can be made in a test (colony) is composed of several individuals
tube. The chemical components are ar- of the lower order which are incapable of
ranged in cells in series of intricately built independent existence. Among social in-
 Biology, Molecular and Organismic 445

sects, the colony becomes an entity for other principle in Physics than in Geome-
which the designation "supraorganism" has try or abstract Mathematics, because all the
been suggested. phenomena of nature may be explained by
Biologists have studied the manifesta- their means, and sure demonstration can
tions of life at all levels of integration. It be given of them." Descartes also wrote
would therefore be logically possible to dis- "that the body of a living man differs from
tinguish molecular biology, cell biology, that of a dead man just as does a watch
individual biology, population biology, or other automation (i.e., a machine that
community biology, etc. This is neither moves of itself), when it is wound up and
necessary nor convenient in practice. It is, contains in itself the corporeal principle
however, desirable to have a simple di- of those movements for which it is designed
chotomy of molecular and organismic biol? along with all that is requisite for its ac?
ogy, the latter name subsuming studies on tion, from the same watch or other ma?
all levels above the molecular one. chine when it is broken and when the
The designation "organismic" is an ap- principle of its movement ceases to act."
propriate one, notwithstanding the fact that Most present-day biologists accept the
this adjective was utilized by the so-called Cartesian view of the nature of living bod?
"holists" for some of their special, and ies. Three centuries of research in biology
now almost completely forgotten, notions. have yielded abundant evidence in favor
This should not, I think, make the word of this view. Time and again, processes and
forever ineligible for use in a context which phenomena which appeared distinctive of
renders its different meaning unambigu- living matter were shown to be compounds
ously clear. of chemical and physical constituents.
Organismic biology, dealing with biologi- Driesch was probably the last outstanding
cal integration levels above the molecular biologist to espouse the classical vitalist
one, has in recent years been referred to, doctrine. He believed that a special force
sometimes pejoratively, as the classical or or energy, which he called by the Aristo-
traditional biology, or as natural history. telian name "entelechy," was active in
The opinion forcibly expressed by some living bodies. Vitalism is now not only
molecular biologists is that, to be "mod- very much a minority view but, and this is
ern," or even "scientific," organismic biol? characteristic, its present adherents are
ogy must be reduced to molecular biology. loath to admit that they are vitalists. For
All that this means in most cases is that example, Sinnott is convinced that the de?
many molecular biologists are so excited velopment of the organism is presided upon
about what they are doing that they are by a "psyche," but, if I understand him
unable to see why their organismic col- aright, this psyche neither substitutes for,
leagues can find excitement in something nor enters into any give-and-take with, ordi-
else. nary physical corporeal processes.
There are, however, also more rational The reason why mechanism has tri-
arguments with which the claims of a su- umphed in biology, and vitalism has faded
premacy of molecular biology are some? out of the picture, must be made unequivo-
times supported. One reason is simply the cally clear. Far from all life processes have
acceptance of the mechanistic hypothesis been, or for that matter are ever likely to be,
and rejection of vitalism. Biological phe- exhaustively described in chemical and
nomena are complex patterns of physico- physical terms. A universal negation is noto-
chemical ones; there is nothing in living riously hard to substantiate; there is no irre-
bodies, no special form of energy or any futable proof that some sort of an entelechy
other agency, that is not potentially ana- may not be lurking somewhere. The point
lyzable into physicochemical components. is rather that vitalism has turned out to be
More than three eenturies ago, Descartes unnecessary and unprofitable, while mecha?
wrote "That I do not accept or desire any nism has vindicated itself as a guide to
446 Theodosius Dobzhansky

discovery. For this and for no other reason, mental laws established in one area of in-
the contest of mechanism versus vitalism quiry, by a theory usually though not
has been a dead issue in biology for at least invariably formulated for some other do-
half a century. Not even the few surviving main." Reduction of organismic to molecu?
vitalists deny that physical and chemical lar biology, and of the latter to chemistry,
processes occur in living bodies, and more would be effected if biological laws and
examples would not impress them greatly. theories, such as for example Mendel's laws
To do research for the purpose of invali- of the theory of inheritance, were shown
dating vitalism is at a height of to be deducible as consequences of the laws
present
It is not unlike artil- and theories of chemistry, physics, or me-
futility. using heavy
chanics.
lery to kill mosquitoes.
Reductiori of the organismic biology to The reductionism is a more sophisticated
the molecular level may, however, be urged notion than the simple, and often a little
also on different grounds. This is the naive, wish to prove that biological phe?
proposition that chemistry and physics are nomena are not manifestations of some sort
sciences more "advanced," more exact, and of vital force or psyche. It must, however,
hence superior to biology. More than a be understood that, while under some con?
century ago (1830-1842), the positivist phi- ditions the reduction is useful and enlight-
losopher Auguste Comte set up an hier- ening, under others it merely detracts from
archy of sciences. In his opinion, the most the research effort better applied elsewhere.
basic science was mathematics; less basic This is a question of research strategy, not
were, in a descending order, mechanics, of some sacred and immutable law of scien-
astronomy, physics, chemistry, biology, and tific development. To be profitable, the
sociology. The progress of scientific in- reduction should open up new possibilities
quiry consists of reducing the description of using some powerful theories or concepts
of the phenomena studied by the less basic of a more advanced, or if you wish, more
sciences to the more basic ones. The aim basic, science. It must help making dis-
of biology is, then, to describe life in terms coveries in the field of science undergoing
first of chemistry, and eventually of physics reduction, and at that, discoveries which
and mechanics, and thus to dispense with could not otherwise be made or not made
biological concepts and ideas altogether. as easily. Such advantages have accrued,
The greatest conceivable success of biology for example, when thermodynamics was
would be to make itself obsolete and un- reduced to statistical mechanics. In biology,
necessary. at least some chapters of physiology are be?
The Comtian positivism had a powerful ing successfully reduced to biochemistry,
influence on the world view of nineteenth chemistry, biophysics, and physics. How?
century scientists, but its reputation has ever, here I must again quote Nagel, whose
not fared well among philosophers. Some philosophy is, let this be made clear, quite
of the greatest modern works on philosophy favorable to reductionism. According to
of science (e.g., Nagel, 1961) do not men- Nagel, "The question whether a given
tion Comte at all. At present not all scien? science is reducible to another cannot in
tists know his name either, and fewer still the abstract be usefully raised without ref-
have read any of his works. The belief in erence to some particular stage of devel?
the Comtian hierarchy of sciences, and the opment of the two disciplines. . . . The
faith in reduction as the intent of scientific possibility should not be ignored that little
inquiry, nevertheless persist and are sel- if any new knowledge or increased power
dom questioned among scientists, especially for significant research may actually be
among biologists. The matter is, however, gained from reducing one science to an?
far from simple; it deserves being consid- other at certain periods of their develop?
ered with care and caution. ment, however great may be the potential
Nagel (1961) defines reduction as "The advantages of such reduction at some later
explanation of a theory or a set of experi- time."
 Biology, Molecular and Organismic 447

Nagel's analysis has not been made espe? of coadapted constituents. Biotic commu?
cially with biology in view, but it describes nities or ecosystems are not miscellanies
splendidly the present situation of the of species which happen to live side by side
biological sciences. The progress of biology or in the same general region; they are
would not be furthered by frenetic efforts structured associations of more or less mu-
to reduce organismic biology to chemistry tually interdependent forms.
or physics. This is not because there is A follower of the philosophy of Francis
anything in living things that is inherently Bacon could perhaps hope that if one ac-
irreducible. It is rather because a different cumulates an abundance of accurate chemi?
research strategy is more expedient. Those cal and biochemical observations, then all
who urge an immediate absorption of the biological phenomena on all integration
organismic into molecular biology neglect levels could easily be deduced from these
the simple but basic fact that life has de? observations. Indeed, we have admitted
veloped several levels of organization. that what the organismic biology studies
These are levels of increasing complexity, are patterns, and patterns of patterns, of
and they are hierarchically superimposed. chemical and physical processes. In actual
The elementary phenomena and regulari- fact, the development of biology has fol?
ties on each succeeding level are organized lowed a quite different path, and really no
patterns of those on the preceding level. branch of science has trod the way which
Organismic biology can be said to be a Bacon, who was not himself a practicing
study of of molecular phenomena. scientist, imagined it would. In biology,
patterns
Such a definition of organismic biology is research was and is being carried on simul-
correct as far as it goes, but it does not go taneously, and discoveries are being made
It is a study not only of on all organismic and molecular levels.
quite far enough.
The discoveries in one branch of biology
the molecular patterns but also of patterns
of patterns. often suggest work to be done, and stimu-
Some examples should make the meaning late discovery in other biological disci-
of this clear. A gene, or at least its key plines. It is, however, not at all a general
constituent, is a double-stranded DNA rule that these discoveries are made by
molecule, or perhaps a part of such a mole? simple deduction. It was, for example, bio-
cule. A chromosome is, however, not a chemistry and spectroscopy that yielded the
of but a configuration of genes celebrated Watson-Crick model of the
heap genes,
arranged in a certain way which proved to structure of DNA. This represents a very
be adaptively in evolution. important advance in our understanding
advantageous
A cell is not a conglomeration of chromo- of what the genes are and how they work.
But the existence of genes was discovered
somes but a supremely orderly contrivance
not of chromosomes alone but with the aid of the methods of hybridiza-
consisting
also of many other organelles. An organ tion and of statistical analysis of hybrid
not of chemical methods. Even
and an individual body are, in turn, not progenies,
de- now, given the entire knowl-
simply piles of cells but beautifully present day
signed and often highly complex machines, edge of the chemistry of DNA, one could
in which the cellular are not hardly deduce from this knowledge that
components
diversified but often have lost their the genes exist and behave as Gregor Men-
only
identities. Mendelian del found them behaving in inheritance.
separate populations
and species of sexually reproducing A suggestion has also been made that
organ-
isms are not throngs or medleys of indi? biologists should exercise a kind of re-
viduals, they are reproductive communities straint, and leave the problems of organis?
of interdependent members. are mic biology in abeyance until the time
Species
of classification, but are not should come when these problems could
categories they
be reduced to the molecular level. This
only that. Evidence is rapidly accumulating
to show that the gene pool, the collective suggestion has at least the merit of a kind
of ruthless logic behind it, but like so many
genotype of a species, is an organized system
448 Theodosius Dobzhansky

other ruthlessly logical proposals, it is a instead of an address or an article. I hope,

practical impossibility. It is like the advice, however, that it is not out of place to ven-
also logically impeccable, that a morato- ture here a very general characterization of
rium ought to be declared on all scientific the subject matter of biology, both organ?
research, to give time for mankind to ab- ismic and molecular.
sorb the knowledge already available. Both The world of life can be studied from
proposals are futile, not to say silly, and two points of view?that of its unity and
for the same reason. Man's intellect will that of its diversity. All living things, from
not tolerate such shackles, not even were viruses to men, have basic similarities. And
it convinced that these shackles would be yet there is an apparently endless variety of
good for it. living beings. Knowledge and understand?
To make the situation ironic, some of the ing of both the unity and the diversity are
same people who would declare a morato- useful to man. I like, however, to stress
rium on organismic biology until such time here not the pragmatic aspect, not the ap-
when it can be reduced to molecular terms, plied biology, but the aesthetic appeal.
also argue that organismic biology is largely Both the unity and the diversity of life are
a finished business, worthy neither of much fascinating. Some biologists find the unity
attention nor support. Now, it is true that more inspiring, others are enthralled by
a method of investigation or a line of re? the diversity. This is evidently a matter of
search may be productive at one time, and personal taste, and a classical adage coun-
become like a squeezed-out lemon after- sels that tastes are not fit subjects for dis-
wards. However, he is a reckless, rather putation (although this is what most dis-
than a courageous man, who wraps himself putations are about). The consequence of
in a prophet's mantle. The history of the polymorphism of tastes is that there
science often deals roughly with fortune- always will be different kinds of biologists
tellers. Consider the discipline of human and different subdivisions of biology. Some
anatomy. It should have been dead four of the subdivisions may be offering more
centuries ago; after Vesalius not many new fleshpots than others, and hence will be
organs can be found in the human body. more popular, especially among those for
Yet we find anatomy prospering and for- whom the fleshpots are the prime considera-
ward-looking. Microscopes have opened tion. Other subdivisions will, however,
vistas which were not accessible to Vesalius. continue to attract some votaries.
Polymorphisms and variations, normal and The number of described species of ani?
pathological, individual and racial, have mals is estimated to be not less than one
acquired new meaning in the light of ge- million, and of plants about one-third as
netics and evolution. Finds of fossil human many. The total number of existing species
and prehuman remains create such excite- of organisms may only be guessed?from
ment that some discoveries are reported in two to four million. It is, however, not only
the daily press before they are buried in the great number of yet undescribed species
weighty monographs. that gives the lie to the oft-repeated con-
What is predictable about most discover? tention that the systematic and descripive
ies is that they are unpredictable. If they biology have already fulfilled their func-
were predictable, they would be made tions and may be relegated to amateurs and
sooner, but making them would be a less to museum drudges. Species identification
inspiring occupation than it is. We have and description is an indispensable prelimi-
been discussing the methodology and the nary, but only a preliminary, to other, and
strategy of modern biology. No synopsis perhaps more exciting and significant in-
of major or outstanding problems of either quiry. The ferreting-out of new species
the molecular or the organismic biology belongs to what Mayr has called the "ana-
will be attempted here. Even if I had the lytical stage," and what is sometimes re-
wisdom, or the brashness, needed for such ferred to also as the "alpha-taxonomy."
an undertaking, it would require a book, This is followed by the "synthetic stage,"
 Biology, Molecular and Organismic 449

and finally by the study of causes and regu- tesian and the organismic biology is basi-
larities of the evolutionary process. Birds cally Darwinian. I utter this generalization
are the group of animals which is attracting not without misgivings; it should not be
the greatest number of workers relative to misunderstood as creating a dichotomy, for
its size (i.e., to the number of species in such a dichotomy would be a false one.
the group); the species of birds are, how? Both the Cartesian and the Darwinian ap?
ever, so well explored that the chances of proaches are essential for understanding
finding a new one are probably smaller the unity and the diversity of life at all
than in any other group of animals of levels of integration. Nevertheless, at the
comparable size. Ornithologists are inter- lower levels of integration the type of ques?
ested surely not so much in finding new tion most frequently asked is "how things
species as in understanding the old ones. are," while at the higher levels an addi-
So are many, if not most, other systematists. tional question insistently obtrudes on the
Remarkably, even paradoxically, the mind of the investigator?"how things got
fundamental unity of all living things to be that way."
makes possible an understanding of their Perhaps the most significant and gratify-
prodigious diversity. Nutrition, respiration, ing trend during the last two decades or
irritability, and reproduction are found so has been the increasing unification of
everywhere. Some of the enzymes in my biology as a field of knowledge. Of course,
body are similar in function to the enzymes we are all specialists in some particular line
in the lowly yeast and bacterial cells. My or even technique of research. But now
genes are different sequences of the same more than ever before one can discern the
four "letters" of the "genetic alphabet" meaningful relationships between all these
which also compose the genes of a fish or of specialties and techniques. The spectacular
a corn plant. Genes reproduce themselves progress in molecular biology has surely
generally with an astonishing accuracy; the acted as a unifying agent. To treat molecu?
sequences of the four "letters," the nucleo? lar biology instead as a bludgeon with
tide bases, are usually identical in hundreds which to destroy, or to reduce to insignifi-
of billions of cells of the bodies of the cance, the organismic biology is to basically
parents and of their progeny. Occasionally, misunderstand the nature of life and the
there occur, however, changes, "misprints,,, requirement for its study.
mutations. Self-reproduction plus mutation I venture another, and perhaps equal-
make possible natural selection. Natural ly reckless, generalization?nothing makes
selection makes possible evolution. Evolu? sense in biology except in the light of evo?
tion is not always, and not necessarily, but lution, sub specie evolutionis. If the living
sometimes, progressive. world has not arisen from common ances-
The enterprise of biology rests chiefly on tors by means of an evolutionary process,
two patterns of explanation. One is the then the fundamental unity of living things
organism-the-machine theory, stated quite is a hoax and their diversity is a joke. The
clearly by Descartes. The other is the unity is understandable as a consequence
theory of evolution, creditable, despite of common descent and of universal neces-
some predecessors and anticipators, to sities imposed by common materials. The
Charles Darwin. Both mechanistic and diversity is intelligible as the outcome of
evolutionary explanations are pertinent to, adaptation of life to different environments,
and are made use of, in molecular as well or, if I may use this unfortunately ambigu-
as in organismic biology. These explana? ous and yet indispensable concept, to dif?
tions are not alternative or competing; they ferent ecological niches.
are complementary, without, however, be? If one could imagine a universe in which
ing either deducible from or reducible to the environment would be completely uni-
each other. It is nevertheless possible to form in space and in time, then in such a
say, as a broad generalization, that the universe a single kind of living inhabitant
molecular biology is preponderantly Car- could conceivably be all that an evolution-
450 Theodosius Dobzhansry

ary process might produce. The real uni- the wasp is conscious of the purpose of its
verse is certainly not uniform. The living activities. Reproduction is accomplished,
matter has responded to the diversity of and the offspring is fed, by a great many
physical environments by evolving a diver? methods other than those used by the mam-
sity of genotypes able to survive and to malian and by the wasp females.
reproduce in a variety of environments. The meaning of the above statements is
Organic diversity is necessary because no really simple and straightforward. When
single genotype can possess a superior certain hormones are produced in the body
adaptedness in all physical environments. of a mammalian female, and produced in
This is, if anything, even more true with a certain delicately balanced sequence, then
respect to the biotic environments. The and only then the chain of events takes
more different organisms inhabit a terri- place which eventuates in the birth of a
tory, the greater becomes the variety of viable infant. The wasp goes through a
ecological niches. In a sense, the growth series of complex actions, which result in
of the organic diversity is a self-accelerating her progeny's feeding and developing, in-
process. stead of starving to death or being poisoned
Although there is again no sharp dichot- by unsuitable food. A logical analysis of
omy here, the concepts of adaptedness and pseudo-teleological statements like the ones
adaptation occupy a more important posi? above has been made with a great discern-
tion in organismic than in molecular biol? ment by Nagel in his admirable book "The
ogy. The existence of several hierarchically Structure of Science." This pseudo-teleo?
superimposed levels of organic integration logical language can only be avoided by
is in itself understandable only as an adap? means of ponderous circumlocations, which
tation. Living beings survive and repro? are superfluous to a biologist acquainted
duce sometimes in apparently most hostile with the modern evolutionary thought.
environments. One can argue that all en? It is a striking and profoundly meaning-
vironments are hostile, and that death and ful fact that organisms are so constructed,
extinction are probable events, while sur- so function, and so behave that they survive
vival is improbable. Just how life has man- and perpetuate themselves in a certain
aged to overcome this improbability is a range of environments frequently enough
problem which many biologists find chal- for their species not to become extinct for
lenging and fascinating. In my opinion, long periods of time. Furthermore, the
this problem may well be used as the frame- ranges of the environments propitious for
work on which to build the teaching of survival and reproduction are widely differ?
biology. At least I found it so, both as a ent for different forms of life. A biologist
student and as a teacher. who chooses to ignore this widespread
I am, of course, not unaware of dissent- adaptedness overlooks a fundamental and
ing opinions about adaptation. It has been very nearly universal characteristic of all
argued that adaptation is either a tautology that can be meaningfully studied on every
(what can survive, survives), or a teleology level of biological integration, from the
(a belief that organisms are shaped by or strictly molecular to the highest organismic
for a purpose). Such opinions reveal a ?the ecosystem level. Even the exceptional
basic misconception. Darwin has, once and failures of the adaptedness, the phenomena
for all, taken the sting out of teleology. For of extinction, constitute an obviously mean-
example, the statement that the hormonal ingful and important subject of study.
mechanisms in a mammalian female serve Zoologists, and in fact all biologists,
the purpose of reproduction does not imply should never lose sight of this one highly
that these mechanisms were contrived by peculiar, and yet remarkably interesting,
some kind of entelechy which knew what animal species?Homo sapiens. The worth
it wanted to accomplish. Nor does the and utility of biology, and, indeed, of
statement that a wasp seeks a prey in order science and of intellectual endeavor as a
to provide food for its offspring mean that whole, will perhaps, in the fullness of time,
 Biology, Molecular and Organismic 451

be judged by the contribution they will highly complex and remarkable one. I sug?
have made to man's understanding of him- gest to you a single reason, but in itself
self and of his place in the universe. I do a sufficient reason, why organismic biology
not wish this statement to be misconstrued will always occupy a leading place in the
as urging that we jettison our zoology and enterprise of science. Man seeks to under-
all strive to become anthropologists or phi- stand himself. The pursuit of self-under-
losophers. By being good biologists, we standing is a never-ending quest. Darwin's
may make a real contribution to the Science work marked a turning point in the intel-
of Man, if not to anthropology in the strict lectual history of mankind because it
technical sense. It is a hoary fallacy to showed that mankind was a product of a
think that man is nothing but an animal; biological history. The evidence for this
however, man's nature is in part animal is now overwhelmingly convincing, except
nature, and man's not-so-remote ancestors to a few antievolutionists. But just how
were full-blown animals. Man's humanity and why man's bodily structures, physio-
and his animality are not independent or logical functions, and mental capacities
kept in isolated compartments; they are have developed as they did is by no means
interdependent and connected by reciprocal well understood. The working hypothesis
feedback relationships. now in vogue is that the process of adapta?
The parts played by the molecular and tion to the environment is the main propel-
the organismic biology come out with ex- lant of evolutionary change. Evidence is
traordinary clarity when viewed against the rapidly accumulating which, in my opinion,
background of the Science of Man. Like substantiates the hypothesis. It remains,
that of any other living body, the physio? however, not only to convince the doubters
logical machinery of the human body is but, what is more important, to discover
compounded of chemical and physical in- just how the challenges of the environment
gredients. Certain diseases, particularly are translated into evolutionary changes.
hereditary ones, are molecular diseases. Man is interested in his future no less
The elucidation of their etiologies makes than in his past. Evolution is not only a
some splendid pages in the story of modern history, it is also an actuality. Of course,
biology. Let me cite just one example? Homo sapiens evolves culturally more rap?
that of the sickle-cell anemia. This usually idly than it evolves biologically. Man must,
fatal disease is due to homozygosis for a however, face the problem of adapting his
single gene; the heterozygote for this gene culture to his genes, as well as adapting his
and its normal allele is healthy or only genes to his culture. Man is being forced
mildly anemic. The hemoglobin in the by his culture to take the management and
blood of homozygous individuals is chiefly direction of his evolution in his own hands.
the so-called S hemoglobin; the hetero? This is perhaps the greatest challenge
zygote has both S and the normal hemo? which mankind may ever have to face, and
globin A. Ingram and others found that this is far too large a problem to be more
hemoglobin S differs from A in the substi- than mentioned here. It is childish to think
tution of just a single amino acid, valine that it is solely a biological problem; the
in place of glutamic acid in the beta chain entire sum of human knowledge and of
of the hemoglobin molecule. The muta- human wisdom will be needed. Biology is,
tional change in the gene responsible for however, involved, and this necessarily
the synthesis of the beta chain must have means both the Cartesian and the Darwini?
involved the substitution of just a single an, the molecular and the organismic biol?
nucleotide, a single "letter" of the "genetic ogy. Fashions and fads come and go in
alphabet." At least 14 other abnormal he- science as they do in dress and in head gear.
moglobins, in addition to S, are known to The big question remains: What is Man?
have single amino acids substituted in cer? It remains not because it is hopelessly in-
tain definite positions in the molecule. soluble, but because every generation must
Man is, however, an organism, and a solve it in relation to the situation it faces.
452 Theodosius Dobzhansky

Biology is here relevant; a solution based REFERENCES

only on biology may well be wrong, but, Dobzhansky, Th. 1964. Heredity and the nature of
man. Hareourt, Brace, & World, New York.
surely, no solution ignoring either the or? Mayr, E. 1963. Animal species and evolution. Har?
or the molecular can be vard University Press, Cambridge.
ganismic biology
Nagel, E. 1961. The structure of science. Hareourt,
right and reasonable. Brace, & World, New York.