Karl Popper and Lamarckism

Elena Aronova Abstract

The Salk Institute for Biological Studies The article discusses Karl Popper’s account of Lamarckism.
San Diego, CA, USA In this article I use Popper’s published and unpublished state-
& ments regarding Lamarckism as well as his correspondence
Department of History/Science Studies Program with the Australian immunologist Edward Steele and other
University of California at San Diego, CA, USA
biologists to examine why Popper was interested in Lamar-
earonova@ucsd.edu
ckism, how his account of Lamarckism can be understood in
the context of his philosophy, and what, if any, new context
Popper provided for the discussion of this abandoned doc-
trine. I begin by discussing Popper’s frame of reference re-
garding Lamarckism, examining how it changed following his
involvement with different “transitory groups” of biologists
concerned with redefining the boundaries between traditional
disciplines. I then discuss Popper’s account of Lamarckism in
positioning it in relation to three aspects of his philosophy:
his distinction between the principles of induction/instruction
and deduction/selection, his distinction between scientific and
teleological concepts, and his concept of progress in science.
Then, I follow the debate concerning Steele’s Lamarckian ex-
periments with a closer look at Popper’s role and his position
in this controversy. I conclude that Popper’s writings provided
a new context for the discussion of Lamarckism in the 1970s
and 1980s, thus making it “scientifically acceptable” again. I
preface this discussion by a brief outline of Steele’s Lamar-
ckian hypothesis and Popper’s comments on Steele’s work.

Keywords
evolutionary biology, molecular biology, molecular Lamarck-
ism, philosophy of biology, Karl R. Popper, unity of science

February 2, 2007; accepted February 7, 2007

Biological Theory 2(1) 2007, 37–51. 
c 2007 Konrad Lorenz Institute for Evolution and Cognition Research 37
Karl Popper and Lamarckism
1. Introduction
In early 1979, Karl Popper received a letter from the unknown
Australian immunologist Edward J. (Ted) Steele, who at that
time was working at the Ontario Cancer Institute in Toronto.
Steele requested that the influential philosopher read and com-
ment on his manuscript Somatic Selection and Adaptive Evo-
lution: On the Inheritance of Acquired Characteristics.1 Steele
explained that the book, although intended for a scientific au-
dience, was influenced greatly by Popper’s philosophy.2
Steele expressed his dissatisfaction with neo-Darwinian
theory, arguing that a variety of biological “enigmas” resist
plausible explanations under the neo-Darwinian paradigm.
Steele insisted that they could be explained if one combined
Darwinian and Lamarckian principles as applied to adaptation
processes on the cellular and molecular levels. “The break-
through,” he explained to Popper, “came when I saw that a
fusion between two theories was possible.”3
Steele was an immunologist and used the immune re-
sponse as a model for evolutionary adaptation. The “clonal se-
lection theory” of antibody formation proposed by Niles Jerne
and Frank Burnet in the mid-1950s stated that an antigen is rec-
ognized by a preexisting complementary structure (the selec-
tive principle), and that it stimulates selectively the appropriate
cell’s proliferation (the clonal principle). Steele’s intention was
not to infringe on the clonal selection theory, which became
the “central dogma of postwar immunology” (see Söderqvist
1994), but rather to cast doubt on the credibility of the common
view concerning where the Darwinian process took place. His
hypothesis suggested that this locus was not so much the genes
in the germ cells but rather the genes at large, and constantly
changing, population of differentiated cells of the soma. Ac-
cording to Steele, by the Darwinian selection of one or more
mutations within the “ecological network” of the whole or-
ganism, the new pattern of somatic genes was established. By
itself, this new somatic gene ordering was insufficient to influ-
ence the germ line. Moreover, such an influence would violate
the doctrine of Weismann’s Barrier—the central presumption
of neo-Darwinian theory, which holds that the germ cells are
protected from changes in the somatic cells. However, Steele
(1981: 5) argued that “the current knowledge in immunology
and molecular biology . . . allows us to approach the belief that
somatic gene mutations can be genetically inherited.”
Steele’s hypothesis suggested that adaptation at the cellu-
lar level involved three stages. The first stage comprised a Dar-
winian process of somatic mutations and subsequent elective
proliferation of the mutant cells under favorable environmental
conditions. In the normal state (in the absence of environmen-
tal stimulation and selection), the frequency of the mutant
gene in both compartments, soma and germline, would be
low. However, under environmental pressure the mutant cells
could be clonally selected and come to dominate the somatic
38
cell compartment. In this case, there was a good chance that
this clonally selected gene in the somatic cells could be “cap-
tured” by endogeneous RNA viruses, cross the tissue barrier,
enter the ova or sperm, and by a process involving reverse tran-
scription integrate the selected gene into the germline DNA.
Thus, transposable genetic elements ensured a Lamarckian
“inheritance of acquired characters,” which operated during
Steele’s second phase of cellular level adaptation. The third
and final step involved classical Darwinian selection operating
on members of the progeny generation. Steele noted that the
Lamarckian “phase” could accelerate the rate of evolutionary
changes, by the hereditary fixation of somatic genetic changes,
and that this mechanism could be extended to any system of
a higher organism where clonal growth was operating (i.e., all
but the nervous system; see Steele 1979).
The suggested mechanism, Steele argued, violated Weis-
mann’s Barrier, a “Central Dogma of Neo-Darwinism,”
whereas Francis Crick’s “Central Dogma of Molecular Bi-
ology” was not violated. The “Central Dogma” describes the
key assumption of molecular biology, namely, the unidirec-
tional flow of information from nucleic acids to proteins (Crick
1958, 1970). It thus forbids a “Lamarckian feedback loop” that
would require some mechanism by which the proteins of the
phenotype could alter the DNA of the genotype. Although the
scenario Steele suggested was hypothetical, he pointed out that
there are “no direct observations which unequivocally prove
that genetic information (DNA, RNA) cannot pass, in princi-
ple, from differentiated body cells to ova or sperm” (Steele
1981: 47).4
The peculiarity of Steele’s book, besides his heretical
claims about the existence of “Lamarckian heredity,” a no-
tion that was by then firmly rejected by biologists, was his
strong appeal to Popper’s philosophy of science. Two sec-
tions of his book were devoted entirely to Popper’s theory
of knowledge. A chapter that outlined the “somatic selection
hypothesis” opened with a reference to Objective Knowledge
and the properties Popper held to be necessary for scientific
theories: “In our journey, we must be conscious of those fac-
tors which speed up or slow down the natural attrition rate
of scientific hypotheses. At least four criteria can be identi-
fied: (a) coherence, (b) simplicity and number of assumptions,
(c) explanatory and predictive power, and, in addition, (d) the
hypothesis must possess a clear plan for its refutation (Popper
1972)” (Steele 1981: 43).
Steele claimed that the “somatic selection hypothesis” met
all the prerequisites of a viable scientific theory, according to
Popper’s scientific methodology. His hypothesis was coherent
in the sense that it bridged two theories that enjoyed con-
siderable experimental support: the clonal selection theory of
antibody formation, and Howard Temin’s protovirus and pro-
virus hypotheses, which speculated on the significance of the
transposable genetic elements in evolution. Moreover, Steele’s
Biological Theory 2(1) 2007

hypothetical mechanism provided an explanation of a good
number of “biological enigmas.” Finally, Steele emphasized
that his hypothesis was “falsifiable” and suggested an exper-
imental design that would “provide—all other factors being
equal—a fairly decisive answer as to whether “Lamarckian”
modes of inheritance, as anticipated by the hypothesis, occur
in nature” (Steele 1981: 53). Indeed, according to Popper’s
methodology, a single example falsifying Weismann’s dogma
would outweigh countless observations apparently supporting
Weismann’s doctrine. Steele emphasized this quality of his
speculation: “[My] hypothesis . . . is not ad hoc and . . . entails
a rational plan for direct experimental testing.”5
Popper replied quickly, although reservedly, that, being
“overworked and overloaded with all kinds of duties, among
them the reading of many manuscripts” he cannot promise to
read Steele’s manuscript. However, since the subject “looks
both extremely important and extremely interesting,” he as-
sured that he would try to find time for the book, which
he tellingly misspelled as “Semantic Selection and Adaptive
Evolution.”6 Shortly after, Popper wrote to Steele with a great
deal of enthusiasm: “My impression is that your book is one
of the most important I have ever read. It is a real break-
through . . . Whether the particular form of your hypothesis is
correct, is, of course, more than doubtful. However, it is the
first serious attempt (as far as I know) to break through Weis-
mann’s barrier. I think that it is very important that you publish
the work as soon as possible.”7
Popper’s encouragement concerning the publication of
Steele’s book was timely. For Steele, the experiments he men-
tioned in his book were much more important than their theo-
retical substantiation. In the preface, Steele (1981: vii) wrote
about himself as an “experimental immunologist with an in-
terest in the evolution of multicellar organisms.” The book
was just the first step in the bold experimental program Steele
envisioned. He explained to Popper:
Your comments on the manuscript have been the first indication that
my efforts might be considered (i.e. read) beyond my small circle. . . . I
was relieved to see that your judgment coincides with mine—that
the book should be published as soon as possible. . . . My haste in
publishing the book (via . . . [a] somewhat unorthodox route) is bound
up not only with my desire to establish authorship, but also in . . . my
desire that the important experimental results do not appear in a
complete intellectual vacuum (at least at the experimental level of
biology).8
With regard to Steele’s discussion of Popper’s philosophy
in the Epilogue, Popper suggested that Steele read his other
works, in addition to the cited Objective Knowledge: an au-
tobiographical account in Unended Quest, Logic of Scientific
Discovery, and Conjectures and Refutations.9 Steele was keen
to expand the discussion of Popper’s work in his book. “I am
convinced that an expanded commentary on your philosophy
Biological Theory 2(1) 2007
Elena Aronova
is required. Initial thoughts include: (1) compare and contrast
the metaphysical research programme of Neo-Darwinism with
the testable research programme of Neo-Lamarckism, (2) state
more clearly your approximations to Lamarckism. . . . ”10 He
also asked Popper to write a preface or even a special chapter
where Popper would summarize his views on Lamarckism.
Popper declined politely, referring to the problems of his own
and his wife’s health, and the burden of other commitments.
Apparently Popper wasn’t quite happy with Steele presenting
his views on the distinction between Darwinism and Lamar-
ckism as an opposition of a “metaphysical research program
of Neo-Darwinism” to a “testable research program of Lamar-
ckism,” since in this form it never appeared in the book. Nev-
ertheless, once Steele’s book was published, Popper wrote a
very favorable review repeating his high appraisal: “I found
Dr. Steele’s book the most exciting scientific book which I
had read for a very long time. . . . [The Lamarckian] hypothe-
sis was killed and buried a hundred years ago by Weismann,
Steele now shows that, in the light of molecular virology, both
Darwin and Lamarck may have been right” (Popper 1979: 5).
Why, one may ask, did this eccentric but still rather techni-
cal book of a young immunologist draw so much attention from
the famous critical rationalist? Mark Parascandola (1995), in
his elegant essay about the controversy over the replication of
Steele’s experiments, which followed the publication of the
book, discusses Steele’s appeal to Popper’s philosophy and
his public support of a heretical scientist. In addressing Pop-
per’s methodological role in Steele’s controversy, Parascan-
dola (1995: 482) notes that Popper’s interest in Steele’s work
is understandable in light of Popper’s methodology, although
“it is not immediately apparent why Popper took an interest
in Steele’s work.” Scientists’ appeal to Popper’s philosophy
was not uncommon in the 1970s and 1980s. For example,
David Hull describes how both the proponents and opponents
of cladistic taxonomy used Popper’s philosophy to argue that
their classifications were genuinely scientific.11 While scien-
tists effectively used his philosophy as a rhetorical device and
capitalized on Popper’s authority to strengthen their ends in
their scientific battles, Popper’s own role in those controver-
sies appears ambiguous. He rarely stated his position in those
debates explicitly, let alone publicly, be it the debate over the
science of cladistics, or the controversy over Steele’s Lamarck-
ian claims. Popper, as Parascandola (1995: 482) puts it, “[did]
not appear to have been so much concerned with Steele’s
Lamarckian hypothesis as with the challenges it present[ed] to
the current theory.” Although Popper’s interest in furthering
the debate was certainly in line with his philosophy, my sug-
gestion is that Popper was also greatly interested in Steele’s
Lamarckian hypothesis as such. In what follows, I attempt to
analyze Popper’s views on Lamarckism and Popper’s role in
Lamarckian controversy in biology, by retrieving the historical
detail from Popper’s published and unpublished writings, as
39
Karl Popper and Lamarckism
well as his correspondence with biologists throughout his life,
thus extending Parascandola’s and Hull’s analyses.
2. Popper and Biology
Popper’s views on evolutionary theory and on the status of
evolutionary biology as a genuine science have been exam-
ined by many authors, primarily in connection with the for-
mation of Popper’s evolutionary epistemology (for example,
Watkins 1995; Stamos 1996; Settle 1996; Sadovsky 2000).
The common view is that before adopting his “evolutionary
epistemology” Popper made some critical remarks about evo-
lutionary theory, which he had to reevaluate later, and so had to
“eat some humble pie,” in Popper’s words (Hull 1999). As Hull
(1999) points out, the actual story is more complicated. Hull
argues that exactly why Popper thought he was eating hum-
ble pie on the score of his rejection of “evolutionary laws”
is puzzling, since he continued to reject the lawful character
of such statements throughout his life, as did, in fact, many
Darwinians at the time.
Reference group theory could provide a possible frame-
work for understanding why Popper changed his attitude to-
ward evolutionary theory over time, while remaining firm in
his opinion on “evolutionary laws” throughout his life.12 In
different periods of his life Popper joined, actually or symbol-
ically, different “biological” transitory groups that explored
the prospects of recasting the historical relations between the
sciences, and between the sciences and the humanities. I would
suggest that two of them, namely the “Bio-theoretical Club”
of the 1930s–1940s, and the cross-disciplinary gatherings or-
chestrated by the architects of the evolutionary synthesis in
the 1960s and 1970s, were the principal “biological” refer-
ence groups for Popper, which shaped his attitude and his
public strategy regarding the evolutionary theory and biology
in general.
Popper’s first books, including Logik der Forschung
(1934), never mention any biological theories as a form of
scientific knowledge. At the same time, biological metaphors
appeared quite often in Popper’s early writings,13 for example:
“Why do we accept one theory in preference to others? . . . We
choose the theory which best holds its own in competition
with other theories, the one which, by natural selection, proves
itself the fittest to survive . . . ” (Popper [1934] 1961: 108). Al-
though Popper was often criticized for his abuse of biological
metaphors and for “clamant anthropomorphism” (Sadovsky
2000: 27), his figurative comparisons fit well the narrative
genres of the time.14 Many intellectuals in the beginning of
the 20th century considered evolutionary theory as a bridge
between the natural and human sciences.15 In the 1920s, evolu-
tionary biology informed many social and economic theories,
including those of language, psychology and pedagogy, which
constituted the intellectual environment of early Popper, as
40
a schoolteacher and a social worker engaged in the Austrian
school reform in 1920s (Bartley 1974; Hacohen 2000). In
reconstructing his own path toward “evolutionary epistemol-
ogy,” Popper denied being influenced by various “biological
trends” of the time; however, he mentioned that before writing
his first book, he read with a great interest “not only Darwin,
but also Lloyd Morgan and Jennings” (Popper 1976: 67).16
When viewed against this background, Popper’s evolutionary
analogies in his early writings in philosophy of science appear
neither striking nor original. What is probably more astonish-
ing is that in his treatise on social science methodology, The
Poverty of Historicism, written in 1935–1940 and revised in
1944–45, Popper severely criticizes a general mode of evolu-
tionism, of which his own usage of the evolutionary analogies
serve as a good example.17
It was during his work on Poverty that Popper became
acquainted with a group, which would change considerably
his frame of reference on evolutionary theory and biology in
general. In 1935, Popper was invited (supposedly by J. H.
Woodger) to attend a meeting of the informal “Bio-theoretical
Club,” a group of philosophically and theoretically inclined
British leftist biologists, many of whom were already distin-
guished, concerned with the relationship between the physical
and biological sciences.18 The core group included Joseph
Woodger, John Desmond Bernal, Joseph Needham, William
F. Floyd, Conrad Waddington, Bertold Wiesner and mathe-
matician Dorothy Wrinch. The extended group included Pe-
ter Medawar, the polymath J. B. S Haldane and philosophers
Max Black and Popper. The group articulated an interdis-
ciplinary research program, which emerged as a variety of
early molecular biology (Abir-Am 1987). The program, named
“mathematico-physico-chemical morphology,” aimed at de-
veloping a joint perspective of the “exact sciences” on the
problem of biological organization.19 However, the scope of
the participants’ inquiry was much broader, as it retained an
isomorphism between organized entities in all disciplines. An
epistemological parity was seen as a new theme of scientific
unity, which would oust the classical, atomistic and reduction-
istic view discredited at that time by relativity and quantum
theories (Abir-Am 1987; see also Graham 1982).
With their positivist search for pluralistic lawfulness and
their aim of recasting the relationships between science and
philosophy, and between political ideology and social utopia,
the Bio-theorists held positions Popper called “historicist” and
that he criticized in his Poverty of Historicism. Malachi Haco-
hen in his intellectual biography of Popper’s early life notices,
however, that the first version of Poverty, written in 1938–
1940, doesn’t contain a systematic critique of evolutionary
thinking (Hacohen 2000: 316). Hacohen suggests that it was
Popper’s encounter in the mid-1940s at the London School of
Economics with another Austrian émigré, economist Friedrich
von Hayek, which shaped Popper’s attitude toward the unity of
Biological Theory 2(1) 2007

science as advocated by the Bio-theorists. Hayek’s essays on
“Scientism” (1942) and the “Counter-revolution of Science”
(1941) directly attacked the values of the group of leftist biolo-
gists (Bernal, Haldane, Hogben) for their positivism, Marxism,
and “historicism.” Waddington’s The Scientific Attitude (1941)
and Science and Ethics (1942), with their proclamation of a
sort of moral futurism based on evolutionist progressivism,
provided Popper with a model of “historicist ethics” and a tar-
get for criticism in his later version of Poverty of Historicism
(Hacohen 2000).
In 1946, when Popper moved to England to become a
reader in logic and scientific method at the London School of
Economics, he renewed his contacts with the Bio-theorists. He
wrote to Woodger announcing his appointment at LSE and ask-
ing for the address of another member of the “Bio-theoretical
Gatherings,” Berthold Wiesner.20 Popper made clear his in-
terest in biology: “I am just [a] little sad that I shall now,
professionally, have to go on with the methodology of the so-
cial sciences: I am much more attracted to that of the natural
sciences. I am very interested in your plan to write a Method-
ology of Biology: I have (quite) a few results bearing on the
problem of the methodological differences between Physics
and Biology” (“quite” crossed out by Popper–E.A.).21 After
his move to London, Popper became a permanent member
of the “Club of Bio-theorists.” With his appointment at LSE
in 1946, the meetings of the Theoretical Biology Club, which
were discontinued for six years during WWII, were resumed.22
The program of the post-war Bio-theoretical Gatherings,
which lasted usually two or three days, comprised a num-
ber of short papers intended mostly to raise discussion on
controversial or difficult issues in biology.23 Although meet-
ings sometimes included “after-dinner” discussions on topics
of broad interest, such as “Scientific Statements and Value
Judgments,” essentially their programs revolved around spe-
cific biological problems. For example, the program of the
July 1946 meeting lists the following topics for discussion:
“(1) Cytoskeletal hypothesis; (2) Relations between anatomy
and physiology; (3) Heterogonic Growth; (4) Morphogenetic
fields; (5) Differences between living and non-living systems;
(6) Directiveness of organic activities; (7) Michael Chance’s
contribution; (8) Environment; (9) Homology; (10) Proposals
for a reform of the language of genetics; (11) Principles of
Classification.”24 Popper was engaged not only in the discus-
sions of mathematical and methodological problems, but in the
biological discussions as well. In 1946, Woodger wrote to him
concerning the above mentioned list of topics: “I anticipate
that there will be a number of points arising here with which
you will be able to help us. Could you possibly undertake to
open the discussion on item (2)? Then I am prepared to follow
you.”25
Despite Popper’s interest in the methodology of biology
in the 1940s and 1950s, biology occupied only a scant place
Biological Theory 2(1) 2007
Elena Aronova
in his published writings until the 1960s. In the early 1960s
his attitude changed drastically; he became a prolific writer
on biological issues. Evolutionary theory became the object
of his critical analysis and the source for his speculations, and
biology began to occupy a place in his writings comparable to
that of physics in his early works. Popper’s shift to biology in
his published works in the 1960s and 1970s might be under-
stood in the context of the emergence of new transitory groups
articulating a new rationale for scientific unity, which became
apparent in the 1960s in both philosophy and biology.
The beginning of molecular biology in the 1950s intro-
duced a new scientific ideology into the biological sciences.26
The approaches and methods of quantum physics were applied
toward the investigation of living phenomena,27 resulting in the
emergence of molecular biology by the late 1960s as a central
and most fashionable science. The rise of molecular biology
entailed the reformulation of basic biological notions into the
terms of molecular biology. At the same time, the rise of the
“DNA bandwagon” in the 1960s was marked by the split be-
tween proponents of “organismic” and molecular biology, the
latter threatening the complete absorption of biology by the
physical sciences (see discussion in Beatty 1990). The archi-
tects of the evolutionary synthesis raised a sound alert about
the danger of the physicalist reduction that accompanied the
unification of sciences under the rubric of biochemistry and
molecular biology (see Smocovitis 1996). In addressing a wide
audience of scientists in 1961, Ernst Mayr argued in support
of the special status of evolutionary biology, which would lift
biology from complete reduction to the physical sciences and,
at the same time, would serve as a unifying element in biology
(Mayr 1961). In so arguing, Mayr, Theodosius Dobzhansky,
and other architects of the evolutionary synthesis were eager
to create a niche for a new philosophy of science, one based
on biology. A series of conferences were organized by them
in order to set up the problem space for the new philosophy
of biology. A distinct feature of these interdisciplinary confer-
ences was the attempt by organizers to involve philosophers,
physicists and mathematicians into discussions of biology and
evolutionary theory as an attractive object for philosophical
study (Smocovitis 1996).
Many authors have noticed that despite the rhetoric of
scientific unity, logical positivism neglected the analysis of
biology, focusing instead on physics, and, to a lesser extent,
the social sciences. Betty Smocovitis (1996: 105) suggests that
philosophy of biology came out of both the 1960s “antireduc-
tionist movements in biology and anti-positivist movements in
philosophy.” Werner Callebaut, Elliott Sober, and Alexander
Rosenberg also explicitly tie the emergence of philosophy of
biology to the decline of logical positivism in the 1960s and
1970s (Callebaut 1993: 73–74). Whether it was logical posi-
tivism that impeded the philosophers’ interest in biology as an
object of study or not is still open for discussion (Byron, in
41
Karl Popper and Lamarckism
press). In any event, a shift of interest to biology as an object
of study became apparent among philosophers in the 1960s.
Popper was engaged in both movements—in philosophy
and in biology. In the 1960s, he found himself again in the
focus of biologists’ attention, at that time the evolutionary
biologists—that is, the architects of the evolutionary synthe-
sis. Mayr wrote Popper in 1973: “I am delighted to see that you
are taking such an interest in philosophical problems raised by
biology. These were sadly neglected by the philosophers of
science until quite recently. I do not know how many volumes
I have on my shelves with the words “philosophy of science”
in the title which deal only with the physical sciences. . . . ”28
In the 1970s and 1980s Popper became an active participant of
various conferences in theoretical and evolutionary biology.29
His association with this transitory group in the 1960s and
1970s changed drastically his public strategy regarding bio-
logical issues. Although he considered himself “an extreme
layman” in biology for the rest of his life,30 since the 1960s
he published on various biological issues, including the prob-
lems of Darwinism and Lamarckism, which became the central
themes in his writing.
3. Popper’s Philosophy—A Possible Framework for
Lamarckism?
Popper presented his account of evolutionary theory publicly
in a series of lectures he delivered in the 1960s: the Herbert
Spencer Lecture (1961), the Compton Lecture (1965), the lec-
ture at Emory University (1969), and various talks given at di-
verse occasions in the 1960s (these lectures became drafts for
his most “biologically-oriented” book, Objective Knowledge:
An Evolutionary Approach, 1972). His intellectual autobiog-
raphy also contained a chapter devoted to the discussion of the
scientific status of evolutionary theory (Popper 1976).
Popper responded to the appeal for a new philosophy of
science based on biology. In examining evolutionary theory
as a distinctive form of scientific knowledge he came to the
conclusion that Darwinian theory cannot be falsified and was
thus a distinctive type of scientific theory—“a metaphysical
research programme—a possible framework for testable sci-
entific hypotheses” (Popper 1976: 134). As Popper explains,
“metaphysical research programmes” are “criticizable, though
not testable”, and can be to some extent “improved” (Popper
1976: 151).31
Popper (1972: 281) suggested his own “improvement”
of Darwinian theory: the concept of “genetic dualism” (or
“Spearhead Model”), which had to explain “orthogenetic
trends . . . without making any concession to the Lamarck-
ian doctrine of the inheritance of acquired characteristics”
(pp. 173–180).32 The “Spearhead Model” suggests that be-
havior is the major force of evolution. Only by altering its
behavior, argued Popper, can an organism solve various sur-
42
vival problems it faces: “This is, I think, a point the biologists
have missed. For an explanation of evolution, the main point
is really the behavioral spearheads. Everything else follows
this. . . . A behavioral tradition may also become the spearhead
of a genetic entrenchment—that is, of a genetic change in the
species of the race” (Popper 1994: 57–61). In order to explain
how the successful “behavioral patterns” can be fixed hered-
itarily, Popper suggested the existence of two types of genes:
genes that control the “anatomy” and those that control behav-
ior (Popper 1994: 138). The latter he subdivided into genes that
control “preferences or ‘aims’ ” and genes that control skills
(ibid). Popper speculated further that in addition to external
(or environmental) selection, there is an “internal selection
pressure” that exists within the organism itself (Popper 1994:
61). This latter type of selection, according to Popper, plays a
decisive role in determining the overall fitness of an organism.
“This is still Darwinism,” explained Popper: “The main
thing in my form of the theory is that mutations can suc-
ceed only if they fall in with an already established behavioral
pattern. That is to say, what comes before the mutation is a
behavioral change, and the mutation comes afterwards” (Pop-
per 1994: 59, 69). Another important feature of his model, as
Popper insisted, is that its different constituents are falsifiable:
“There is quite a lot in this theory that is testable. I don’t know
whether my theory is true but I think it is testable . . . and so
far a prediction in principle” (Popper 1994: 69–70).
Popper’s remark that his model “is still Darwinism” is
not incidental. It was not obvious for many of his listeners
and readers. After his lecture delivered at Emory University in
1969, where he discussed his Spearhead model, he was asked
directly:
How does that differ from what Lamarck said?” Popper replied: “The
difference . . . is very great. According to my theory and Darwin’s
theory, Lamarckism is wrong in so far as behavioral traits are not
inherited. They only stimulate a selection pattern, not the inheritance
as such. . . . A new behavior creates a new ecological niche, and then
the selection pressure operates so that this niche is filled. . . . It is only
a simulation of Lamarck and not a Lamarckian theory. That is really
the decisive point in the whole thing. You have to simulate Lamarck’s
theory, but it is a question of how you do it. (Popper 1994: 70)
There is no need to criticize Popper’s “Spearhead” hypoth-
esis since many authors have already pointed out the weak-
nesses, unoriginality, and amateurism of Popper’s biological
arguments, let alone the principal difficulty about testing it
(contrary to its design).33 What seems puzzling is Popper’s
firm persistence about his hypothesis, in spite of the rather cold
reception of his “improvements” by biologists,34 including his
biological friends from the old days of the “Bio-theoretical
Gatherings” in London. Peter Medawar, who not only ad-
mired Popper as the “greatest philosopher of science that has
ever been,”35 but also adopted his philosophical views in his
Biological Theory 2(1) 2007

own philosophizing on the methodology of biology was rather
skeptical about his friend’s biological concepts. He warned
Popper about his Spencer Lecture, which Popper had prepared
for publication in 1965: “. . . I think . . . you come adrift badly
over your exposition and criticisms of Neo Darwinism and
your conception of mutations generally.”36 Medawar wrote
a several page-long letter to his friend explaining what was
wrong with Popper’s hypothesis: he pointed out Popper’s con-
fusion about modern understandings of natural selection and
mutations; the weakness of his concept from the side of genet-
ics; and Popper’s apparent ignorance of Waddington’s work,
which provided a modern reading of the old concepts of Lloyd
Morgan and Baldwin that attributed behavioral changes a ma-
jor role in evolution. Medawar concluded: “I am not quite
sure that you are saying anything basically new enough to
justify publication in extenso.”37 A year later Medawar pa-
tiently repeated basically the same points in commenting on
Popper’s Compton lecture: “I have read the lecture several
times, and my trouble is that, on the one hand, I find almost
nothing to disagree with, and on the other hand I think the
entire formulation is rather out of date and inexpert (as well
it might be, since you are not a biologist.)”38 Popper thanked
his friend for his critique (assuring him that his silence during
four months was not because he disliked Medawar’s letter39 )
and published the lectures mentioning the fact that “an expert
discouraged me from publishing” (Popper 1972: 281). Pop-
per’s firmness concerning his “Spearhead hypothesis” may
seem puzzling, especially since his views on the status of
evolutionary biology change radically over time. Popper’s hy-
pothesis, however, never changed. His last book published in
his lifetime, Knowledge and the Body-Mind Problem, which
he put together in 1993, includes his widely criticized behav-
ioral “Spearhead hypothesis” without any revisions (Popper
1994).
Hull (1999: 491) noticed that the prominence of the
distinction between Lamarckian and Darwinian evolution in
Popper’s writings can be explained by his primary interest
in cultural, not biological, evolution—the context in which
Lamarckism was still alive, but astonishing for biologists to
whom such an “old chestnut” didn’t deserve so prominent a
place. However, the distinction itself may seem astonishing as
well. Whenever Popper comes to Lamarckism he contrasts it
to Darwinism. Some readers may see more similarity than dif-
ference between a Lamarckian’s attack on a non-Lamarckian
viewpoint and Popper’s continuously contrasting Darwinism
to Lamarckism. In his philosophy, however, dichotomies and
oppositions play an essential role. His account of logical asym-
metry between induction and deduction, instruction and selec-
tion, verification and falsification was at the heart of Popper’s
philosophy. Arguing for a logical continuity between Darwin-
ism and the theory of the growth of scientific knowledge, Pop-
per held that the distinction between Darwinism and Lamarck-
Biological Theory 2(1) 2007
Elena Aronova
ism is of the same nature as between deduction and induction,
selection and instruction, verification and falsification.
Since the 1960s the contraposition of Darwinism and
Lamarckism became one of the recurrent themes in Pop-
per’s writings. The discussion of Lamarckism in his writings
can be plugged into three more general contexts of Popper’s
philosophy: his distinction between the principles of induc-
tion/instruction and deduction/selection; Popper’s distinction
between scientific and teleological concepts, and his concept
of progress in science.
3.1. Instruction and Selection
The opposition of Lamarckism versus Darwinism in Popper’s
writings was engendered in his distinction between the prin-
ciples of induction and deduction in the concepts of teaching
and learning. “Instruction,” the method of teaching by means
of repetition, represents the inductive method of the acquisi-
tion of knowledge. Popper argued that inductivist models of
teaching and learning are methodologically unviable, holding
that human knowledge is conjectural and hypothetical.40 He
considered the principles of instruction and selection as uni-
versal principles, which characterize adaptations at different
levels: genetic, behavioral, and at the level of scientific dis-
covery (Popper 1985a: 78). At each level, the process of the
acquisition of novelties (i.e., new genetic information, new
behavioral patterns, or new knowledge) can be explained ra-
tionally by selection—Darwinian selection of genetic vari-
ations, or selection of the falsifiable hypotheses—however,
the results might imitate the instructive (or Lamarckian) pro-
cesses. Accordingly, Popper saw the solution of the problem of
Lamarckism in biology: seemingly Lamarckian effects (such
as inherited behavioral patterns or bacterial adaptation in a new
nutritional environment) are Darwinian processes of selection
that imitate Lamarckian evolution.
3.2. Science and Religion: The “Lamarckian
Daemon”
The principle of “instruction” in Popper’s writings was
bounded not only to the concepts of teaching and learning
but also to the teleological concept of intelligent design. In
referring to William Paley, the 18th-century philosopher of
religion remembered best for his watchmaker analogy (an ar-
gument for the existence of God in his 1802 book Natural
Theology), Popper makes a case for J.-B. Lamarck as the
one who transferred an “instruction”-type explanation from
the sphere of religion into the province of science. Concern-
ing Paley’s argument about the design of the watch, which
implies an intelligent designer, Popper (1985b: 244) writes:
“The theistic theory of Paley is instructionist. It is the Cre-
ator, who, by His design, moulds matter, and instructs it which
shape to take. . . .” If Lamarck presented a scientific version of
instructionist theory, Darwin gave an answer “to the problem
43
Karl Popper and Lamarckism
posed by Paley” (p. 245). A passing remark about the “Lamar-
ckian daemon” at the Altenberg meeting41 is pertinent in this
context. At this meeting a creative element of evolution was
called by one of the participants a “Lamarckian daemon”: often
in disguise, as a Bergsonian “l’esprit du vivant,” or Prigogine’s
“bifurcations,” the “Lamarckian daemon” impels biologists to
complement Darwinian evolution by a creative, directive and
accelerating element, to overcome the difficulties of Darwinian
theory (Popper and Lorenz 1988: 15). For Popper, the scientific
“daemon” is not the omniscient godhead but rather a super-
human intelligence, a sort of “super-scientist.” Akin to the
Laplacean daemon in classical physics, whose unlimited cal-
culative power allows him to make perfect predictions about
the future, the Lamarckian daemon opens the deepest meta-
physical, or theological, commitments to rational scrutiny and
rational debate.42
3.3. Dogmas in Science
Popper’s warning against dogmatism in science provided ad-
ditional ground for his interest in the Lamarckian tradition in
biology. Progress in science is realized through challenges to
the current dogmas. Warning against a “too dogmatic adher-
ence to Darwinism” (Popper 1985a: 85), he emphasized the
necessity to maintain a Lamarckian reasoning in biology as an
alternative to the dominant neo-Darwinian theory: the “Lamar-
ckian line . . . seems to have been mistaken. Yet it may be worth
while to speculate about possible limits to Darwinism; for we
should always be on the look-out for possible alternatives to
any dominant theory” (Popper 1985a: 85).
In sending Popper his manuscript, Steele hit the center
of Popper’s interests at the time. Popper would certainly be
pleased by Steele’s attempt to present a challenge to the exist-
ing theory. Popper’s reasoning regarding Lamarckism is strik-
ingly similar to that of Steele’s. Popper wrote in 1973: “The
argument against the genetic inheritance of acquired character-
istics . . . depends upon the existence of a genetic mechanism
in which there is a fairly sharp distinction between the gene
structure and the remaining part of the organism: the soma. But
this genetic mechanism must itself be a late product of evo-
lution . . . This point . . . is made merely as a warning against
too dogmatic adherence to Darwinism” (Popper 1985a: 85).
Compare this to Steele’s writing of 1979: “It is my view that
the central block to any consideration of the inheritance of the
acquired characteristics can be directly traced to Weismann’s
doctrine . . . which states that no acquired genetic character-
istic can penetrate the barrier protecting the germ cells from
the soma. . . . The scheme for transmission of somatic genes
to the germline is posed to help our thinking and to illuminate
a deficiency in the central axiom of neo-Darwinian theory. . . .
Weismann’s doctrine should no longer be a stumbling block to
potential (or teetotalling) “neo-Lamarckians” who may wish
to embark on animal breeding experiments designed to test
44
whether the inheritance of acquired characters occurs in na-
ture” (Steele 1981: 47–49).
The details of the molecular mechanism of Steele’s hy-
pothesis should have been of great interest to Popper as well.
Popper himself has been speculating himself about the possi-
ble molecular mechanism of Lamarckian inheritance. In the
draft of his paper “Lamarckism and DNA” (1973), which was
intended for publication in Nature, but apparently was rejected
and was never completely finished, Popper stated: “The Main
argument against any form of modified Lamarckism is not
only that we do not know of anything’s happening to an organ
having acted back upon DNA, but that we can hardly con-
ceive of such a mechanism.”43 Popper argued that, on the con-
trary, such a mechanism can be conceived and, like Steele, he
speculated about the possible role of an enzyme, reverse tran-
scriptase, and the transfer of genetic information from RNA to
DNA:
If this [reverse transcription—E.A.] is taken as established . . . this
leads to the following possibility. If there is a great demand for a
certain protein, there may be increased demand for the RNA which
controls its production, and it is possible that this may have a backfir-
ing effect on the DNA in the cells. . . . Increased demand for a certain
protein may lead to an enzyme controlled feedback from RNA to
DNA, for instance by having the DNA sequence which represents
the RNA (and thus the protein) retranslated and incorporated into
the DNA sequence. The result of this would be that the DNA has
an additional, identical sequence for the production of the protein.
This would mean that certain sequences within the genetic code, on
an external stimulus, may reappear on neighboring pieces of chro-
mosome. . . . This would constitute a mutation, one having, perhaps
other consequences than the increased production of the particular
protein.44
In such a way, Popper argued, a stimulus/response mechanism
can make a case for the inheritance of acquired characters. “It is
not being suggested that there is such a mechanism, merely that
the existence of such a mechanism . . . cannot be excluded.”45
He concluded: “This is not Lamarckism, but would in some
respect resemble it. It would constitute a break with current
dogma . . . ”46
Popper’s speculations about “molecular Lamarckism” re-
veal much in common with those of Steele’s. Both challenged
Weismann dogma in a way that would not violate Crick’s ‘Cen-
tral Dogma’ of molecular biology. Both suggested a hypothet-
ical mechanism that bears on the premise of the exceptional
evolutionary role of the enzyme reverse transcriptase (not to
mention that both were adherents of Popper’s philosophy).
They were not alone in such speculations. The central dogma
of molecular biology occupied an influential place in biology
in the 1960s and 1970s, serving as a distinctive icon for the
proponents of the new science of molecular biology of the late
1950s and 1960s. At the same time, formulated in negative
terms, the central dogma also provided a target for criticism
Biological Theory 2(1) 2007

for those who questioned the exclusive role of nucleic acids in
inheritance (Commoner 1961, 1964, 1968; Sarkar 1996).47
Although revealing much in common, Popper’s and
Steele’s positions differed in some principal aspects. For Pop-
per, the fusion of Darwinism and Lamarckism would make
as little sense as the fusion of the principles of induction and
deduction, or instruction and selection. The question for Pop-
per was still in the imitation of Lamarckian processes. What
distinguished them even more was that Popper’s interest was
exclusively in theory. He was interested in the development
of Steele’s hypothesis rather than in the results of its exper-
imental testing. His replies to Steele’s detailed reports about
his experiments were always laconic: “. . . whatever the out-
come of the tests of your hypothesis, you have certainly shown
that a kind of Lamarckism is compatible with modern genetic
findings. . . . ”48
Instead, Popper generated new ideas in developing
Steele’s scheme. When they met in November 1979 in Pop-
per’s home in Penn, Popper made the following note about
their conversation: “Main point: I told him about the diffi-
culties of embryology—the fact that, cells develop according
to position into . . . different organs. . . . All this, strange to
say, was quite new to him. I explained briefly the field the-
ory (which is just a restatement of the difficulty), and said
that the cell receives, obviously, information about its position
that is not in its DNA. This information may be explained by
Steele’s theory, or something like that. . . . ”49 Contrary to Pop-
per, Steele was not so much interested in developing his theory
as in strengthening his position as an experimental scientist.
He had a solid reason for that at the moment of his meeting
with Popper: at that time Steele was in the middle of a big
controversy over his experiments.
4. “Popperian Science” at Work: The Debate over
Ted Steele’s “Lamarckian Experiments”
The opinions on Steele’s work voiced by the scientific com-
munity were divergent. Although some of his colleagues were
enthusiastically supportive of his work and ideas, it met a
chilly reception at his Institute in Ontario. “As might be ex-
pected,” Steele confessed to Popper, “the initial reception of
my work here at the Institute ran onto some hostile resistance
. . . No doubt some resistance will be encountered when the
book is published.”50 The publication of his book also didn’t
go through straightforwardly. Steele’s colleague, Jeffrey W.
Pollard, who was “keenly interested” in Steele’s work, of-
fered the facilities of a small publishing house in Toronto he
co-owned, for the publication of Steele’s book.51 The book
was published in 1979 in a small run of 1000 copies, and
distributed by Steele and his friend’s company among the uni-
versities, college libraries, and research institutes throughout
Canada, North America, Europe and Australia.
Biological Theory 2(1) 2007
Elena Aronova
The publication of the book preceded the announcement
of the results of experiments Steele performed at the On-
tario Cancer Institute together with his colleague at the In-
stitute, Polish immunologist Reginald Gorczynski. Steele and
Gorczynski claimed that they had proven that induced im-
mune tolerance can be passed on to the next generation, thus
reviving an old controversy of Lamarckian heredity. Their
publication appeared first in a form of a modest abstract in
American Zoologist in 1979, and soon, supported by Howard
Temin,52 the extended articles were published in the lead-
ing scientific journals—Proceedings of the National Academy
of Sciences and Nature (Gorczynski and Steele 1979, 1980,
1981).
Their experiments consisted of inducing a state of immune
tolerance in new-born mice, and subsequently testing whether
the offspring showed an immune response to the same cells
of the foreign strain that were used to induce tolerance in the
parental mice. The authors reported that approximately half of
the first generation offspring showed an increased tolerance to
the spleen and bone marrow cells from hybrid mice strains used
to induce tolerance in the parents. Since only male mice were
used in the experiments, the likelihood of non-genetic transfer-
ence of tolerance was minimized (Gorczynski and Steele 1980,
1981). If the publication of the book went almost unnoticed,53
the articles published in established and widely read journals
led to a significant controversy within the scientific community
and outside of it. The Sunday Times proclaimed “an intellec-
tual revolution, a landmark in science history” (Parascandola
1995: 417).
Within the scientific community, Steele’s work came into
the area of expertise and interests of Peter Medawar, at that
time at the Clinical Research Center in Middlesex, and Leslie
Brent from St. Mary’s Hospital Medical School in London.
In 1960, Medawar and Frank Macfarlane Burnet received the
Nobel Prize for the discovery of acquired immunological tol-
erance, which laid the foundations for organ transplantations.
The techniques Medawar developed became widely used as a
classical way for inducing immunological tolerance. Using a
modified version of Medawar’s classical method for inducing
tolerance, Steele and Gorczynski focused their efforts on trac-
ing the after-effects of the induced tolerance in the offspring
generations. Medawar was so interested in this work that he in-
vited Steele to continue his experimental project in Medawar’s
laboratory “in a friendly but critical environment.”54
Medawar was a dedicated popularizer and defender of
neo-Darwinism, with a strong distrust of anything Lamar-
ckian (see Parascandola 1995). Steele’s case should have
reminded him of his other encounter with a Lamarckian
experimentalist—immunologist. Earlier in his career, in 1952–
1953, when Medawar was engaged in his most fruitful re-
search on immunological tolerance, inducing the Czechoslo-
vakian biologist Milan Hašek was conducting experiments on
45
Karl Popper and Lamarckism
“acquired immunity” in chickens. Hašek developed an ele-
gant technique, which consisted in joining two hen embryos
together by removing their shell membranes and by placing
the blastoderm of another embryo between the opening. In
this way Hašek connected the blood circulations of both eggs,
which resulted in an exchange of red blood corpuscles in the
two chicken embryos as if they had developed naturally as
twins. The result of this unusual technique was that both chick-
ens became tolerant of each other’s blood elements. Although
Hašek interpreted his results in terms of Michurinist biology,
as an example of “vegetative hybridization in chicken” (Hašek
1953a,b,c, 1954), his experimental technique exceeded all ap-
proaches at that time and presented a significant interest for
Medawar, since the mature spleen cells Medawar used for
inducing immune tolerance caused high mortality of the inoc-
ulated newborn mice, because they reacted with the host tis-
sue producing a classical graft versus host reaction. Medawar
successfully reproduced and confirmed Hašek’s results, and
this work was one of the important steps in the discovery of
immunological tolerance (Medawar 1986: 132–137). Hašek
finally adopted Medawar’s terminology and rejected his ini-
tial Michurinist interpretation of his results (Svoboda et al.
2005).
Besides his professional interest in Steele’s work,
Medawar’s interest stemmed from his own “Popperianism.” In
numerous essays and lectures (assembled in Medawar 1982),
he described the development of the hypothetico-deductive
scheme from the 19th century, arguing that Popper’s theory of
science provided the most useful and true account of scientific
growth among those currently available. Medawar interpreted
Hašek’s work from the perspective of Popper’s methodology
of science:
Simultaneous discoveries are not uncommon in science, but what was
remarkable was that Hašek’s motivation was entirely different from
our own. . . . With his experiments he hoped to repudiate Mendelian
genetics. There could not be a better example of how false premises,
metaphysical fences even, may lead to empirically sound conclu-
sions, a circumstance that reinforces Popper’s warning against the
vulgar error of dismissing all metaphysical speculation as so much
gobbledegook. (Medawar 1986: 133).
Similar considerations should have fed Medawar’s strong in-
terest in Steele’s case. Medawar was keenly interested in
replicating Steele’s experiments. The direct replication of an-
other’s work rarely happens in modern laboratory science.
The ‘weight’ of scientific publication dealing with the repeti-
tion of another’s work hardly justifies the investment of time,
labor and personnel it requires (on the unreliability of replica-
tion in the laboratory science see Collins 1992; Mulkay and
Gilbert 1986). However, in the case of Steele’s experiments,
Medawar was keenly interested in having his work repeated.
If confirmed, it would lead to significant discoveries in im-
46
munology, as in the case of Hašek’s work. If refuted, it would
falsify Steele’s Lamarckian doctrine.
Steele was enthusiastic about Medawar’s offer. It was right
in line with Popper’s ideal of science. He described his move to
London to Popper: “As you know independent experiments are
underway here . . . I am not privy to the details of these trials at
present i.e. it is totally independent of my involvement. . . . It
is obvious that confidence and morale is high. Later in the year
there will be an open discussion of these results.”55
The public discussion exploded very soon. Medawar,
Brent and his co-workers reported that the results they obtained
were not consistent with Steele’s conclusions, although, they
claimed that they had reproduced Steele’s technique (Brent
et al. 1981a,b). However, observers pointed out that it was
not easy to reproduce it precisely, and that Medawar changed
it significantly. Although using the classical technique of in-
ducing tolerance by inoculating newborn mice with mature
spleen cells, Steele and Gorczynski modified it by repeatedly
inoculating foreign cells into parental mice from birth until
the end of the breeding period. Jonathan C. Howard from the
Institute of Animal Physiology in Cambridge, UK, pointed out
the significant discrepancy of the techniques used: “Gorczyn-
ski and Steele’s tolerance—inducing protocol was eccentric in
the extreme: enormous doses of cells were injected repeatedly
into the father from birth until the end of the breeding period.
Heroically, Brent and his co-workers repeated this anoma-
lous procedure. . . . They made one mistake. The . . . injection
of cells . . . consisted of a total of 108 . . . of bone marrow
and spleen cells in Gorczynski and Steele’s experiments, and
5×107 . . . in Brent and colleagues’ experiments” (Howard
1981: 442).
Few other attempts were made to reproduce Steele’s work
(see Parascandola 1995 for a detailed account of attempts by
various groups to reproduce Steele’s experiments). Although
the results were contradictory—some of the groups reported
even the reverse effect: a heightened immune response in the
offspring instead of tolerance (McLaren et al. 1981)—none
of them firmly confirmed Steele’s results. Numerous expla-
nations for the divergent results were proposed, including the
use of different techniques (Howard 1981), the impact of ex-
ternal factors such as endogenous viruses (Hašek et al. 1981),
and others (Robertson 1981). Steele’s professional compe-
tence was not questioned: “Steele, beyond any question, is an
excellent practical scientist” (Tudge 1981: 485). All the par-
ticipants agreed that the conditions in the testing experiments
differed considerably from those in Steele’s original work,
however, they disagreed upon the significance of those dif-
ferences. Some observers even questioned whether the testing
group had actually reproduced Steele’s experiment (Howard
1981). The discussion ended with no consensus reached.
Popper’s philosophy of science was used as a reference
point by both sides of the controversy. All of the major
Biological Theory 2(1) 2007

participants, explicitly or implicitly, appealed to Popper’s phi-
losophy. The inability to reach a conclusion in the debate was
beneficial for Steele since it meant that his hypothesis was not
falsified. According to Popper, the better scientific theory is
one that stands the test of time and resists falsification. Thus,
due to the experiment’s resistance to a clear falsification, as
well as the involvement of established scientists like Medawar
and Brent, Steele gained some support. As one of the partici-
pants summarized, “although the results [of the testing experi-
ments] clearly cannot confirm those of Gorczynski and Steele,
they do not falsify their theory that a virus, which includes in
its genome some DNA coding for the tolerated transplanta-
tion antigens, could infect spermatogenic germinal epithelium
. . . [and thus] render progeny tolerant” (Smith 1981: 768).
Although no verdict upon Steele’s experiments was
reached, Steele might have felt himself in a “pit of despair.”
He bewailed to Popper: “I am doing my best to remain tolerant
during what has been a difficult time. . . . If the opposition I
have met was only scientific, I would not have had much cause
for distress. Unfortunately, however, the “opposition” . . . has
been almost exclusively non-scientific. . . . Sir Peter Medawar
and colleagues informed me . . . that I should change my field
of scientific interest and publish no more on the soma →
germline topic. I refused to comply with this and . . . I was told
that my grant would not be renewed. . . . I was most distressed
by all this during this period—not the least because the de-
cisions were made before any of the experimental evidence
from various workers here was gathered. . . . I was distressed
because I was put in a position where defending myself scien-
tifically was made most difficult. . . . I look forward very much
to my move to Australia. . . . ”56 Steele never changed the
topic, though. Indeed, he moved to Australia, and continued to
publish on possible soma-to-germline feedback mechanisms
in the immune system throughout his life.57
Popper, though he didn’t take sides in the debate, remained
supportive of Steele and his ideas. It was not without Popper’s
help that Steele received a job at the John Curtin School of
Medical Research at the Australian National University. Pop-
per wrote the most favorable letters of recommendation for
Steele. In 1984, when the continuation of Steele’s contract
with the ANU was under question, Popper wrote in reply to
the Deputy Vice-Chancellor of the ANU, I. G. Ross:
Indeed I was consulted by Dr. Steele several years ago in connection
with his book Somatic Selection and Adaptive Evolution, and . . . his
original manuscript was modified in response to my comments. I am
well aware that Dr. Steele has been criticized by some biologists, in
fact even by some of my biological friends whom I had told about
him: he has been criticized because of some alleged Lamarckist ten-
dencies. And as you know, the allegation of Lamarckism is regarded
by biologists as a very severe one . . . I do not know how far Dr.
Steele’s theories will ultimately be found correct, or in need of mod-
ification. However, there is no doubt in my mind that Dr. Steele was
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Elena Aronova
one of the first to see the great importance of the role of viruses in ge-
netics. In fact, he can be described in that respect as pathbreaking. . . .
I am worried that Dr. Steele may not get the recognition which in my
opinion he deserves.58
Ross responded: “I am grateful for the tolerance with which
you contemplate the possibility that Steele might be wrong
and yet that his work may still be seen as commendable. Most
of the biologists we have contacted do not have such shades
of grey in their palettes.”59
5. Conclusion
Lamarckism has a long tradition in biology, being associated
with scientific programs based on a mechanistic ontology, the
thoroughgoing acceptance of environmental action, the inher-
itance of acquired characteristics, and epigenetic conceptions
of heredity. Although the revival of Lamarckian interpreta-
tions in molecular biology and the emergence of a kind of
“molecular Lamarckism” in the late 20th century can hardly
be conceived of as a school of thought, it constitutes one of the
areas of non-selectionist biology, which stood (and stands) in
opposition to Darwinian evolution and Mendelian genetics.
Over the course of the 20th century, Lamarckism dra-
matically changed its status several times. Being considered a
perfectly scientific doctrine during the 1920s, it became scien-
tifically unacceptable by the 1950s because some of its essen-
tial theoretical statements were taken over by “Lysenkoists.”
This antipathy to Lamarckism as an identity and the fear of
seemingly Lamarckian explanations lasted throughout the sec-
ond half of the 20th century owing to the generation of fighters
who took part in the defeat of Lysenkoists and who insistently
tried to expunge any traces of what they considered Lamar-
ckian reasoning from biology. The reaction of the scientists
outside of Russia to the “Lysenko affair” was sometimes even
more dramatic than in Russia at the time of Lysenko’s decline
(see, e.g., Krementsov 1996; Harman 2003).
A new generation of scientists, mainly those who entered
science in the 1960s and later, made Lamarckism “scientif-
ically acceptable” again. Surprisingly, this unexpected, and
uneasily welcomed, support came out of the camp of molec-
ular biologists working in the fields of molecular genetics
and immunology. As Steele’s case shows, the new genera-
tion of Lamarckians can capitalize on their unorthodox views
and lead rather successful scientific careers. Although modern
Lamarckians are still marginal, they have already accumulated
some symbolic capital.
One of the unexpected sources of support for “mod-
ern” Lamarckism was Popper’s philosophy. Popper’s account
of Lamarckism provided a new context (“a possible frame-
work”) for the discussion of Lamarckism, if we paraphrase
his own definition of Darwinism as “a metaphysical re-
search program—a possible framework for testable scientific
47
Karl Popper and Lamarckism
theories.” As I have attempted to show, Popper’s interest in
Lamarckism was rooted in his philosophy, which shaped his
attitude toward Lamarckian reasoning, while his participation
in the two distinct “transitory groups” of biologists, the Bio-
theoretical Club of the 1930s–1940s, and the cross-disciplinary
gatherings orchestrated by the architects of the evolutionary
synthesis in the 1960s and 1970s, affected his public strategy
regarding Lamarckism. Although he had been keenly inter-
ested in the methodology of biology since the 1930s, he did
not express his “extreme layman” opinion publicly until the
1960s. The efforts made by the architects of the evolutionary
synthesis to create space for a new philosophy of science based
on biology, and the professionalization and institutionalization
of the philosophy of biology that followed thereafter, was one
of the reasons why Popper started to publish prolifically on
various biological issues, including Lamarckism.
Was Popper a “hidden” Lamarckian himself? For someone
of a consistently Darwinian persuasion, he seems inordinately
preoccupied with Lamarckism. Since the 1960s, the Lamarck-
ian motif was omnipresent in many of his works. At the same
time, he explicitly proclaimed himself a true Darwinian, both
in public and in his private correspondence. He explained his
position to Ernst Mayr in 1977:
I should have thought that I can call myself a Darwinist. I cannot
say that you are wrong when you say that I am holding out for
an alternative to Darwinism, for I think that every theory can be
surpassed. But is this what you mean? (At any rate I am a more
orthodox Darwinist than Darwin . . . , who believed in the occasional
inheritance of acquired characters.)60
A “Popperian” methodology of science, as the case of
Steele’s “Lamarckian” experiments shows, didn’t help sci-
entists determine whether there was anything in Lamarck-
ian reasoning that was compatible with modern genetics and
molecular biology. Rather, this case demonstrates how “non-
Popperian” real science is. As Steele’s case, and other cases
of the “use and abuse” of Popper’s philosophy show, scientists
often conceptualize retrospectively their intellectual positions
in terms of Popper’s methodology, but their scientific practices
remain rooted in the classical ideal of science, irrespective of
the rhetorical devices they use.
Popper’s discussion of Lamarckism illustrates the role and
place of philosophy in biology in the 1960s–1980s. Although
his account of Lamarckism met a chilly reception from leading
evolutionary biologists, the epistemic position to which Popper
subscribed made Lamarckism scientifically acceptable, thus
keeping discussions of Lamarckism alive.
Acknowledgments
This essay is based on research in Karl Popper Papers (the Popper Archive)
kept in the Klagenfurt University Library in Austria and the Hoover Insti-
tution Archives at Stanford University. I am extremely grateful to the staff
48
of both these institutions, in particular to Ronald Bulatoff from the Hoover
Institution Archives and to Manfred Lube from the Universität Klagenfurt,
for their substantial help with archival research. Exceptionally helpful feed-
back from Aleksey Kouprianov on earlier drafts, as well as comments and
editorial suggestions by Marta Halina, Matthew Shindell, and two anonymous
reviewers are all gratefully acknowledged. I am very grateful to Melvin Cohn
for his reading this article at final stages of preparation and for his insightful
comments and lovely discussion, which has also helped to correct some fac-
tual mistakes. Any remaining errors are of course only my own responsibility,
as well as views expressed here. Research was supported by the Research
Support Scheme of the Open Society Institute (grant no. RSS 148/1999).
Notes
1. Self-published in 1979 (Steele 1979). The second edition was published by
the University of Chicago Press in 1981.
2. Steele to Popper, January 8, 1979, Popper Archive (“PA” henceforth), b.
352, f. 17.
3. Steele to Popper, u/d (February 1979), PA, b. 352, f. 17.
4. “There is also no good evidence that it can,” Steele added (ibid).
5. See note 2.
6. Popper to Steele, January 17, 1979 and February 8, 1979, PA, b. 352, f. 17.
7. Popper to Steele, February 9, 1979, PA, b. 352, f. 17 (emphasis by Popper—
EA).
8. Steele to Popper, February 23, 1979, PA, b. 352, f. 17 (emphasis by E.
Steele—EA).
9. Popper to Steele, February 8, 1979, PA, b. 352, f. 17.
10. Steele to Popper, February 23, 1979, PA, b. 352, f. 17 (emphasis by
Steele—EA).
11. See Hull (1988) and also Hull (1999) for an analysis of the influence of
Popper’s philosophy on the reception of cladistic analysis, and Mulkay and
Gilbert (1981) for the sociological analysis of Popper’s influence on scientists.
12. The general idea of reference group theory is that people shape their at-
titudes by reference to groups other than their own, and that they evaluate
themselves by selecting particular points of comparison. Since Merton’s clas-
sical studies, concepts of reference group theory have been used in studies
involving science and scientists (Merton 1968; Merton and Rossi 1968).
13. Popper’s analogy with Darwinian natural selection drew on Herbert
Spencer’s formula of “survival of the fittest,” which Spencer used to rein-
force his parallels between economic theories and Darwin’s biological ones
and which became shorthand for the topics related or analogous to evolution
and natural selection in the early 20th century. (On “social” analogies to Dar-
winian evolution at the end of the 19th and the beginning of the 20th century
see Bowler 1984, 2004; Todes 1989, Rosenberg 2000.)
14. On the influence of Darwinism on literary narratives see Morton (1984);
Zirkle (1959) discusses the influence of neo-Lamarckism on literature of the
19th and 20th centuries.
15. See Smith (1997) for a “big picture” analysis of the impact of evolutionary
theory in the human sciences.
16. The books of psychologists C. Lloyd Morgan and James Mark Baldwin,
who independently stated that the evolutionary process can be guided by a
learning process, were widely read outside their field. Herbert S. Jennings
in his Prometheus, or Biology and the Advancement of Man (1925), The
Biological Basis of Human Nature (1930), The Universe and Life (1933), as
well as in innumerable shorter articles, philosophized on the significance of
various aspects of the rapidly accumulating work in evolutionary biology for
the understanding of human nature and society (see Kevles 1999).
17. Popper (1960: 3) defines “historicism” as “an approach to the social
sciences which assumes that historical prediction is their principal aim, and
Biological Theory 2(1) 2007

which assumes this aim is attainable by discovering the ‘rhythms’ or the
‘patterns’, the ‘laws’ or the ‘trends’ that underlie the evolution of history.”
He distinguishes “historicist” doctrines, which claim that the methods of the
natural sciences (which were poorly understood, in Popper’s view) can be
applied to the social sciences, from “anti-naturalist” doctrines, which do not
make this claim. (On Popper’s notion of “historicism” and its distinction from
both German Historismus and Hegelian historicism see Hacohen 2000: 355.)
In both cases, however, according to Popper, these doctrines stem from an
uncritical belief in the evolutionary laws of nature. Popper (1960: 106-107),
on the contrary, disclaimed not only the existence of historical laws, but also
the existence of evolutionary laws and thus came to question the scientific
status of evolutionism.
18. On the history of the “Bio-theoretical Gathering” in the 1930s see Abir-
Am (1987).
19. On the Bio-theorists’ vision of molecular biology as a genuine empirical
science although holding its own logic distinguishable from the logic of phys-
ical sciences, see Woodger (1929); Haldane (1931); overview in Abir-Am
(1987).
20. Popper to Woodger, August 28, 1945. PA, b. 363, f. 7.
21. Popper to Woodger, u/d (Fall 1945), PA, b. 363, f. 7 (emphasis by Popper—
EA).
22. It is unclear from the survived correspondence in the PA whether the
Bio-Theoretical Club was resumed simultaneously with Popper’s moving to
London, or his relocation and interest in the gatherings was one of the stimuli
for the bio-theorists to resume their meetings.
23. W. F. Floyd’s memo on Theoretical Biology Club, June 1948. PA, b. 363,
f. 7.
24. Program for the Bio-theoretical Conference, Oxford July 26–29, 1946.
PA, b. 363, f. 7.
25. Woodger to Popper, July 16 1946. PA, b. 363, f. 7.
26. For an overview of the history of molecular biology see: Kay (1993);
Chadarevian (2002); Morange (1994); Olby (1994).
27. On the role of physicists in the molecular revolution in biology see Abir-
Am (1980); Kay (1985, 1992); Beese (1991).
28. Mayr to Popper, September 24, 1973. PA, b. 551, f. 4.
29. Popper was one of the organizers of two philosophical-biological sympo-
siums (see published proceedings in Krebs and Shelley [1975]; and Medawar
and Shelley [1980]), was a participant of the conference on reductionism in
biology, organized by Francisco Ayala and Theodosius Dobzhansky (Popper
1974), and was invited to contribute in the collected volume of papers devoted
to the problems of evolutionary theory (Popper 1984).
30. In 1991 he wrote to Ernst Mayr, explaining how he had become interested
in the biochemical aspects of Darwinism: “I am an extreme layman in this
field—the extreme opposition of a specialist. I am, simply, very curious about
evolution, and I love to think about it. And I love theories” (Popper to Mayr,
January 1, 1991. PA, b. 551, f. 4).
31. The weakness and vagueness of Popper’s arguments were criticized many
times, especially his statement about “tautological character” of evolutionary
theory which, as Hull (1999) noted, was based on the formulations accepted
in the 19th century but widely rejected by his Neo-Darwinian contemporaries
(see also Stamos 1996).
32. Popper discussed his “Spearhead model” (or “genetic dualism” model)
for the first time in his Spencer lecture (1961), in the Compton lecture (1965),
in lectures delivered at Emory University (1969), and in his major books
discussing evolutionary theory (Popper 1972, 1976, 1994).
33. See Hull (1999) and Stamos (1996).
34. See Hull (1999) for the general reception of Popper’s suggestions by
biologists.
Biological Theory 2(1) 2007
Elena Aronova
35. Cit. in Magee (1974).
36. Medawar to Popper, March 29, 1965. PA, b. 325, f. 26.
37. See note 37.
38. Medawar to Popper, August 16, 1966. PA, b. 325, f. 26.
39. “I hear from Watkins that you thought I might not have answered . . . your
letter of August 16 . . . because I didn’t like your letter. On the contrary, I was
deeply grateful for your criticism . . . ” (Popper to Medawar, December 19,
1966. PA, b. 325, f. 26).
40. Bartley (1974) argues that Popper’s views originated in the debates over
the Austrian school system, in particular over the competing concepts of
“method of instruction” and method of Selbsttätigkeit (self-activity”), and
their implications for philosophy and psychology.
41. At the “Altenberger Gesprach” Popper discussed the problem of the di-
rectedness of evolution with Konrad Lorenz (Popper and Lorenz 1988).
42. For Popper’s account of “Laplacean daemon” see Popper 1982: 30-31.
43. “Lamarckism and DNA” [1973]. PA, b. 128, f. 4.
44. “Lamarckism and DNA” [1973]. PA, b. 128, f. 4.
45. See note 45.
46. See note 45.
47. In immunology and bacteriology, the selective (“Darwinian”) models of
antibody production and enzyme formation received strong support from
molecular biologists. However, various “instructive” theories, which implied
that the external stimulus could direct the formation of the molecular struc-
ture of proteins, didn’t disappear with the rise of molecular biology, but were
reformulated in molecular biological terms (on the discussion of “adaptive
enzymes” in bacteriology see Gaudillière (1991, 1993); for the controversy
on “selective” and “instructive” models of antibody formation in immunology
see Silverstein (1989) and Moulin (1991).
48. Popper to Steele, October 12, 1979. PA, b. 352, f. 17.
49. “Memo on a conversation with Ted Steele,” November 11, 1979. PA, b.
352, f. 17.
50. Steele to Popper, February 23, 1979. PA, b. 352, f. 17.
51. See note 51.
52. Temin was awarded the Nobel Prize in 1975 for his work on reverse
transcriptase. His speculations on the significance of RNA-directed DNA
synthesis for normal development and for carcinogenesis played a significant
role in Steele’s hypothesis and in Popper’s speculations about Lamarckism and
DNA. Temin submitted Steele’s and Gorczynski’s article to the U.S. National
Academy of Sciences for publication. Once published in PNAS their article
was accepted for publication in Nature.
53. Besides the aforementioned favorable review by Popper (1979), the book
was reviewed neutrally by Maynard Smith (1980) and Young (1980), and
received a strictly negative response from immunologist John Klein (1980).
54. Steele to Popper, October 7, 1980. PA, b. 352, f. 17.
55. See note 55.
56. Steele to Popper, March 1, 1981. PA, b. 352, f. 17.
57. Steele’s recent publication on this subject includes a book Lamarck’s Sig-
nature: How Retrogenes are Changing Darwin’s Natural Selection Paradigm
(Steele et al. 1998). The book was written in co-authorship with Steele’s wife,
R. Lindley, and R. V. Blanden, an immunologist who participated in testing
Steele’s experiments in the 1980s and who supported his claims concerning
the results of his experiments (Mullbacher et al. 1983); on Steele’s subsequent
career see Parascandola (1995).
58. Popper to Ross, July 12, 1984. PA, b. 352, f. 17.
59. Ross to Popper, July 26, 1984. PA, b. 393, f. 7.
60. Popper to Mayr, April 18, 1977. PA, b. 551, f. 4.
49
Karl Popper and Lamarckism
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