Opposition before 1926
Walter Frank Raphael
Weldon and the Biometricians
Research and Teaching at Cambridge
Edward
B. Poulton
Opposition after 1926
R. A. Fisher and L. Darwin
Richard Goldschmidt
Ernst Mayr
H. Ledyard Stebbins, and Richard Dawkins
H. Allen Orr
Robin Marantz Henig
William Coleman
Speciation as a Non-Problem-
Dawkins
Steven Jay Gould
While opposition to Bateson
occurred during his lifetime, a
not unusual phenomenon in human affairs, it is of note that the opposition continued long
after his death in 1926. Indeed, opposition seemed to increase as the 20th century
progressed, reaching an apparent apogee in 1996 when an article entitled "Dobzhansky,
Bateson and the Genetics of Speciation" by H. Allen Orr was published in the
"Perspectives" section of Genetics, the journal of the Genetics Society of
America (144,
1331-1335).
|
|
|
Reginald
Punnett and William Bateson 1907 |
Opposition before 1926
The following is as much
of interest for what it does not contain, as for what it contains:
The Dictionary of
National Biography (Oxford University Press) had this
entry for
Walter Frank Raphael
Weldon
W.F.R.
Weldon (1860-1906) was born in London on 15 March 1860. In 1867 he matriculated at
University College, London, with the intention of qualifying for a medical career. After a
year he was transferred to King's College, London. In 1878 he entered St John's College,
Cambridge. At Cambridge, Weldon came under the influence of Francis Balfour and abandoned
medical studies for zoology.
In 1881 he obtained a first class degree in the natural sciences. Subsequently he spent a
year's research work at the zoological station in Naples. Returning to Cambridge in 1882
he became demonstrator in zoology, fellow of St John's College, and university lecturer in
invertebrate morphology. After his marriage in 1883 he and his wife spent their vacation
at such resorts as offered the best opportunities for the study of marine zoology. The
most important of their expeditions was to the Bahamas in the autumn of 1886. At Plymouth,
Weldon worked in the laboratory of the Marine Biological Association. Until 1888 he was
engaged on the morphological and embryological studies which seemed to contemporary
zoologists to afford the best hope of elucidating the problems of animal evolution.
In 1889, when Galton's recently published book on natural inheritance came into his hands,
he perceived that the statistical methods explained in that book might be extended to the
study of animals. He soon undertook a statistical study of the variation of the common
shrimp. This study was published in the Proceedings of the Royal
Society. In a second
paper, he calculated numerical measures of the degree of inter-relation between two organs
or characters in the same individual and tabled them for four local races of shrimps.
These two papers were the foundation of that branch of zoological study afterwards known
by the name of 'biometrics'.
Weldon was elected a Fellow of the Royal Society in 1890. In this year he succeeded
Professor Sir E. Ray Lankester as Jodrell professor of zoology at University College,
London. His tenure of the Jodrell chair (1891-99) was a period of intense activity. A
brilliant lecturer and endowed with the power of exciting enthusiasm, Weldon soon
attracted a large class, and his association with Professor Karl Pearson led to increased
energy in the special line of research which he had initiated. In 1894 Weldon became the
secretary of a committee of the Royal Society for conducting statistical inquiries into
the measurable characteristics of plants and animals. In 1894, Weldon stated that "the questions raised by the Darwinian hypothesis
are purely statistical, and the statistical method is the only one at present obvious by
which that hypothesis can be experimentally checked."
In
1899 Weldon was elected to the Linacre professorship of comparative anatomy at Oxford.
Though the Oxford life afforded opportunities for greater intellectual leisure, Weldon
disdained to make use of them. He had on hand numerous exacting projects and he tried to
deal with them all at once. His leisure hours at Oxford were spent in long bicycle rides,
during which he studied the fauna of the neighbourhood.
His vacations were spent in
journeys to various parts of the continent, where he worked at his statistical
calculations and collected material for fresh lines of research. In the Easter vacation of
1906, while he was staying with his wife at an inn at Woolstone, he was attacked by
influenza, which on his return to London on 11 April, developed into acute pneumonia. He
died in a nursing home on 13 April 1906. He was buried at Holywell, Oxford.
A Weldon Memorial Prize for the most noteworthy contribution to biometric science was
founded at Oxford in 1907. |
While the DNB tells us precisely where Weldon fell
terminally ill and where we may seek his tomb, there is no mention of his profound
opposition, together with his fellow "biometricians"
such as Karl Pearson, to Mendel's work.
"Karl Pearson is a biometrician,
And this, I think, is his position,
Bateson and co.,
Hope they may go
To monosyllabic perdition."
N. Mitchison in Haldane and Modern Biology (1969) ed by K. R.
Dronamraju |
This story is related in:
-
Beatrice Bateson's "William Bateson, Naturalist,"
(CUP 1928),
-
Reginald C. Punnett's "Early Days of
Genetics" (1950; Heredity 4, 1-10),
-
William
Provine's The Origins of Theoretical Population Genetics
(1971),
-
Marsha L. Richmond's "Women
in the early history of genetics" (2001; Isis
92, 55-90),
-
My brief biography of William Bateson in the
Nature Encyclopaedia of Life Sciences
(2001) (Click Here)
The
above record from the DNB was obtained from the internet
several years ago. The 2004 DNB internet biography has two
brief allusions to Bateson, with some indication of the
intensity of Weldon's opposition to Mendel:
"Following a
much heated debate initiated by William Bateson on one of
Pearson's papers on inheritance at the Royal Society on 15
November 1900, Weldon suggested to Pearson that they start their
own journal for papers dealing with matters of statistical
biology."
"Weldon's
statistical interpretation of Mendel's data led Bateson to publish
his fiercely polemical 100-page chapter to 'defend Mendel from
Professor Weldon' (W. Bateson, Mendel's
Principles of Heredity. A Defence, 1902, viii). Bateson's
criticisms seem actively to have discouraged Weldon from adopting
virtually any aspect of Bateson's work."
DRF
2007 |
Research
and Teaching at Cambridge
Bateson
founded the immensely successful Cambridge School of Genetics against much
local opposition and on a shoe-string budget. He was helped by his wife, Caroline Beatrice
Durham,
and a host of undergraduate and graduate helpers, many of whom were women.
 |
| Beatrice Bateson circa 1925 |
David Lipset in Gregory
Bateson: The Legacy of a Scientist (1980) cites one of
Bateson's helpers, Nora Barlow (a
grand-daughter of Charles Darwin),
as evidence that initially Bateson had difficulty even in getting formally
to teach Genetics at Cambridge:
"My first introduction
to the whole subject ... was when William Bateson was giving what
we called his bible class, in a remote lecture room, in the back
of one of the colleges. It was outside the ordinary curriculum. It
was a five or six o'clock lecture. And there he introduced a small
set of people into the elements of the new Genetics. Mendelism was
just coming in. ... He was a brilliant lecturer and, of course, he
had an entirely new view of ordinary heredity. ... It was very
inspiring indeed." |
Bateson was the opening
speaker at the International Conference on Plant Breeding and Hybridization
held in New York in 1902. It was here that he stated:
| "We have reason to believe
that the chromosomes of the father plant and mother plant, side by
side, represent blocks of parental characters." |
Many speakers paid tribute to Bateson, as did
cytologists such as Guyer, Cannon and Sutton (Click Here).
In 1910 he became the first director of the John Innes Institute, then based
at Merton in Surrey. This became a centre for studies of the genetics of
plants and animals, and received visits from geneticists of many countries.
The following photograph shows Bateson with Wilhelm Johannsen from Denmark, who
introduced the term "gene" as an "accounting unit" (Rechnungseinheiten).
He was Bateson's closest scientific 'soul-mate,' and many of the incorrect
criticisms launched at Bateson would also have been launched at Johannsen,
had the launchers read Danish or been aware of his English texts. For
details see:
Roll-Hansen, N. (2014) The holist tradition in twentieth century genetics.
Wilhelm Johannsen's genotype concept.
J Physiol 592, 2431-2438.
Roll-Hansen, N. (2014) Johannsen's criticism of the chromosome theory.
Perspectives in Biology and Medicine
57, 40-56.
Roll-Hansen, N (2014) Wilhelm Johannsen and the
problem of heredity at the turn of the 19th century.
International Journal of
Epidemiology, 2014, 1007-1013.
Both J. B. S. Haldane (of
"Haldane's rule" fame) and
A. E. Garrod (of "inborn errors
of metabolism" fame) later acknowledged
inspiration from Bateson. In a letter to Penrose (1934) Garrod wrote:
| "Hogben in his paper on
alkaptanuria has, I think, given me more credit than I am entitled
to, seeing that it was Bateson who saw the daylight."
(See A. G. Bearn's paper on "Archibald E. Garrod. The Reluctant
Geneticist." Genetics
May 1994). |
Bearn reports that what appears to be the
first letter from Garrod to Bateson (11th January 1902) begins:
| "It was a great pleasure
to receive your letter and to learn that you are interested in the
family occurrence of alkaptonuria." |
|
Poulton versus Bateson and
Punnett
The Hope Professor of Zoology at Oxford
was also not supportive. Click Here
. Much of the 36 page Introduction to his text Essays
on Evolution (1908) was devoted to an attack on Bateson and
his fellow Mendelians.
Opposition after 1926
R. A. Fisher and L. Darwin
The great pioneer in
statistics, biomathematician Ronald Fisher, was guided in his understanding
of the biology of evolution by one of Charles Darwin's sons, Major Leonard Darwin.
In the course of a long correspondence Darwin wrote (19 Jan. 1929):
Darwin:
"As to Bateson, if I had to write, I
should write something like the following. But I am not well up
in what he did do, and may well blunder ...
'In the future the great merit of Mendelism will be seen to
rest on the proof that the ingredients of germ plasm on which
heredity depends are located in pairs in each organism, one of
each pair selected by chance disappearing at each sexual union.
... The merit for this discovery must mainly rest with Mendel,
whilst among our countrymen, Bateson played the leading part in
its rediscovery. Unfortunately he was unable to grasp the
mathematical and statistical aspects of biology, and from this
and other causes, he was not only incapable of framing an
evolutionary theory himself, but entirely failed to see how
Mendelism supplied the missing parts of the structure first
erected by Darwin. Nothing but harm can come from following
Bateson in regard to evolution theory, though his name will come
to be honoured for his pioneer work in Mendelism when what he
failed to do as regards theory has been accomplished.' |
Having written it, I daresay I
should tear it up, and advise you to do ditto. ..."
Fisher: "Many
thanks for the note on Bateson; it puts the point admirably, and
though I have already altered the wording somewhat, it seems to
me just what was wanted. The only thing to do is to commend
Bateson's enthusiasm for genetics, without saying, which would
rather comfort my conscience, 'while
greatly retarding its progress in his own country.' ... I
have just been reading Samuel Butler's Luck
or Cunning; what a
malignant knave he must have been, yet Bateson borrowed his
sneers and quoted his opinions."
(See Natural
Selection, Heredity and Eugenics.
Edited by J.H. Bennett. 1983. Oxford University Press. pp. 95-96) |
At that time Fisher was
composing his major work, The Genetical Theory of
Natural Selection (1930,
1958). Disregarding Darwin's
caveat ("But I am not well up
in what he did do, and may well blunder") he
did not "tear
it up." Indeed, he closely
followed Darwin's wording.
While
noting that Bateson "played the leading part"
in the "early advocacy of Mendelism,"
Fisher wrote (p. ix):
"Unfortunately he was
unprepared to recognize the mathematical and statistical aspects
of biology, and from this and other causes he was not only
incapable of framing an evolutionary theory himself, but
entirely failed to see how Mendelism supplied the missing parts
of the structure first erected by Darwin. His interpretation of
Mendelian facts was from the first too exclusively coloured by
his earlier belief in the discontinuous origin of specific
forms. Though his influence upon
evolutionary theory was thus chiefly retrogressive,
the mighty body of Mendelian researches throughout the world has
evidently outgrown the fallacies with which it was first
fostered. As a pioneer of genetics he has done more than enough
to expiate the rash polemics of his earlier writings." |
Thus, Fisher, who shortly thereafter
was, through statistical analysis, to cast doubt on the soundness of the
data of Mendel himself, seemed to have based his attack on Bateson on the
words of someone who admitted to not having thoroughly read Bateson's
work. To compound the matter, Bateson's first book, Materials
for the Study of Variation (1894),
was described by Fisher (p. 138)
as:
|
"A work which owed its
influence to the acuteness less of its reasoning that of its
sarcasm."
|
Defending
Bateson, Alan G. Cock
(1973;
J. Hist. Biol. 6, 1-36)
concluded that "Fisher's criticisms are ...
unfair and in large part based on a misunderstanding of Bateson's
views." However, even Cock subscribed to the view that
"theoretical innovation was not where
Bateson's strength lay," basing his case on
-
(i) Bateson's
incorrect "reduplication" hypothesis for why certain characters were co-inherited
or "coupled" (in a
linkage group, - later equated with a particular chromosome), and
-
(ii) a
misunderstanding of what Bateson meant by his "presence
and absence" hypothesis to explain genetic dominance.
In his
Presidential Address to the 13th International Congress of Genetics (1974;
Genetics 78, 21-33),
Curt Stern noted that while the reduplication hypothesis "could
not beat the [later]
competition," nevertheless "it
was an ingenious suggestion" to propose that "Mendelian
segregation ... occurs during a somatic cell division followed by
differential multiplication of the different genotypes."
Indeed, some genetic phenomena can be explained along
these lines (see Stern's biography of Wilhelm Weinberg (1962;
Genetics 47, 1-5).
Richard Goldschmidt
Goldschmidt (1878-1958)
thought along similar lines as Bateson, yet he consistently
misrepresented Bateson's "presence and absence hypothesis"
(concerning genetic dominance). Furthermore, in his last major
scientific work, Goldschmidt wrote:
"The continuous
variation with which Darwin worked ... became a matter of
phenotype, while genetic variation was discontinuous. Such a
system makes it rather difficult to lead evolution beyond the
reshuffling of mutated genes, that is, limited permutation or
variations on the same theme. The early Mendelians felt this
rather keenly and assumed a skeptical attitude towards
evolutionary speculations, which found its extremest expression
in Bateson's Australian address (1914) with the embarrassing
idea of evolution by loss of inhibitors."
Theoretical
Genetics (1955) University
of California Press. p. 478.
|
You can read the address on these
web-pages (Click Here),
where you will see that Bateson was merely stating that Mendelian
effects can involve the loss of inhibitors, not that evolution as a
phenomenon occurs in this fashion.
Ernst
Mayr
Ernst
Mayr (1904-2005) was perhaps the most influential, and certainly the most
relentless of the disparagers and misrepresenters of William Bateson. There
was much celebration of Mayr's 100th birthday, which, at
least in one quarter, was considered to overstate his achievements.
"I
do not wish to belittle the work of Mayr and the geneticist
Theodosius Dobzhansky - but our impression that they solved the
species problem is illusory. They were merely the ones who
translated it from the technical literature, enunciating much
more clearly than before what had become the prevailing view of
species among those who had thought about the problem."
James Mallet, Nature
430, 503
|
To
give Mayr due credit, when considering linear and branching
evolution (which he called transformation and diversification,
respectively) he acknowledged his debt:
|
|
"Although
Darwin was aware of this difference ... unfortunately he
subsequently did not sufficiently stress the far-reaching
independence of these two components of evolution, and this has
been the cause of several of the post-Darwinian controversies. Two
post-Darwinians, however, made a clear distinction between the two
modes. Gulick (1888) used the term monotypic evolution
for transformation and polytypic evolution for
diversification. Romanes (1897), who adopted Gulick's terminology,
also referred to transformation as 'transformation in time' and
diversification as 'transformation in space.' Both Gulick and
particularly Romanes appreciated that these were two very
different components of evolution, an insight that was largely
forgotten again after 1897, until Mayr (1942) and others revived
it during the evolutionary synthesis."
Mayr
1982. The Growth of Biological
Thought |
Here is a
sampling from a 1973 paper of Mayr (J.
Hist. Biol. 6, 125-154):
- "Certain aspects of evolution
... were consistently misunderstood or ignored by
experimental scientists like Bateson and De Vries." (p.
127)
- "The early hybridizers were
completely right and the early Mendelians (Bateson, De Vries
and so on) who ignored them and believed the opposite, were
quite wrong." (p. 136)
- "Bateson was pig-headed,
intemperate, and intolerant ... uncompromising ... and ...
quite incapable of understanding the nature of natural
populations ... ." (p. 146)
- "Bateson was still totally
unable to grasp the nature of speciation in his Toronto
address of 1922." (P. 147)
- "Bateson's stubborn resistance
to the chromosome theory resulted in much effort by members
of the Morgan school that could have been devoted to new
frontiers in genetics." (P. 147)
- "Attempts to refute the power
of selection were truly a comedy of errors. Bateson tried to
do so ... . None of those who denied the efficiency of
selection so insistently bothered to visit the animal
breeders ... ." (P. 150)
|
For Bateson, Mayr and the Thrasybulus
principle (Click Here).
No less strident opposition came from other
major figures in evolutionary biology, including H. Ledyard Stebbins, and Richard Dawkins. In The Blind Watchmaker
and elsewhere
(Click Here),
Dawkins, shaking his head in bewilderment, found it "extremely
hard for the modern mind to respond ... with anything but mirth"
to the quaint ideas of the "mutationists"
Bateson and De Vries.
The
above "modern" minds may have
mischaracterized and dismissed Bateson, but at least they did not
claim he was really on their side. At the turn of the 20th century came the "Bateson,
Dobzhansky, Muller model of
speciation."
It is to Dobzhansky's credit that he
resurrected the Romanes-Gulick view that reproductive isolation was of paramount
importance for speciation, but he held that this reproductive isolation was
usually achieved initially by differences in gene products ("genic"
isolation), a view also held by H. J. Muller.
In "Dobzhansky,
Bateson, and
the Genetics of Speciation" (1996) H. Allen Orr asserted that:
"William
Bateson offered the "Dobzhansky- Muller" model in 1909, just nine years
after the rediscovery of Mendelism and a good quarter of a century before
Dobzhansky or Muller. And when I say that Bateson offered the model, I do not
mean he obliquely alluded to it. Rather, Bateson spells it out, step by step,
presenting it as the likely 'secret of interracial sterility'...
Those who
differ on larger issues, as Dobzhansky and Bateson surely did, can nevertheless
arrive at the same conclusion. Recent work on speciation renders this
coincidence all the happier: for Bateson and Dobzhansky not only arrived at the
same conclusion, but at the right conclusion." |
Of course, Dobzhansky and Muller are likely to have read enough of Bateson's
writings to know, even if they did not understand him, that he most certainly did
not support a genic model for speciation. Thus, they did not cite him in this
respect. Orr suggests, however, that the reason they did not cite Bateson was
that "neither Dobzhansky nor Muller knew of Bateson's model."
Orr believes that "Bateson apparently never repeated his
argument." For
numerous quotations from Bateson on his non-genic view of speciation. Click Here
Sadly, Orr's
misinterpretation was further relayed by Stewart Berlocher (1998) in chapter 1
of the multiauthor text Endless Forms. Species and
Speciation, and by Jerry Coyne (with Orr) in 1998 in the Philosophical
Transactions of the Royal Society of London (353,
287-305). In 2000 the misinterpretation was even dignified with an acronym
"BDM," standing for the "Bateson, Dobzhansky, Muller model of
speciation" (Lynch and Force. American
Naturalist 156, 590-605).
By 2001 "BDM" had
reached the textbooks. See:
|
The Evolution of
Developmental Pathways by Adam Wilkins, Sinauer Associates 2001;
chapter 12, pp. 459-461.
Fitness Landscapes and
The Origin of Species, Princeton Univ. Press 2004, by S.
Gavrilets; Part 2, chapters 5, 6 and 7).
Speciation
by J. A. Coyne & H. A. Orr, Sinauer Associates 2004; chapter 7, pp. 269-272.
For more:
(Click Here) |
The BDM "meme" lingered on. When
resurrected in Heredity in 2009 by
Norman A. Johnson, I responded with a Letter to the Editor that, while published
online in February 2010, did not get formal publication until 2011 Click Here.
In March 2011 Genome Biology and Evolution
accepted a paper by Masatoshi Nei and Masafumi Nozawa, who seemed to have
independently arrived at the conclusion that it should be "DM" not "BDM":
"Orr
(1996) argued that the first person who proposed the DM model is neither
Dobzhansky nor Muller but Bateson (1909) and
Bateson's model was identical with that of Dobzhansky and Muller. In our view,
this argument is disputable. It is certainly true that Bateson considered
the two-locus model of complementary genes in order to explain hybrid
sterility, but he never considered how such a system can evolve. By
contrast, Dobzhansky and Muller spelled out the evolutionary process of
hybrid sterility genes, albeit very crudely. In evolutionary biology, it
is important to understand the process of evolution. For this reason, we
will refer to the model as the DM model in this paper."
|
The paper concludes that "the currently
popular Dobzhansky-Muller model of evolution of reproductive isolation
is only one of many possible mechanisms."
|
Nei N, Nozawa M (2011) Roles of
mutation and selection in speciation: From Hugo de Vries to the
modern genomic era.
Genome Biology and Evolution
3, 812-829. |
Robin Marantz Henig
When scientists get the story wrong, then historians may be misled, and
popularizers and journalists who depend on the scientists and historians for
their books and stories may compound the error. In a letter to T. H. Huxley's
grandson, Julian, with whom he was collaborating on a book, H. G. Wells wrote in
1928:
"I am against any
further alterations of that Bateson paragraph. I know the man.
My last talk with him was with Morgan in N. Y. and he has a
schoolboy pleasure in making trouble and a Samuel Butler-like
hatred for Darwin. Any fool can play the negative game and no
doubt some of the young fools will go on with it."
Memories. J.
Huxley, p. 161.Harper & Row, N.Y. 1970 Here
is "that Bateson paragraph:"
"There remains one
other temperamental type which has found expression in
these discussions, and that is the brilliant sceptic as
typified by the late Professor William Bateson. He
accepted the facts of Evolution, if only on the
palaeontological evidence, but, as the outcome of a life
spent largely in the study of variation and especially of
Mendelism, he developed an increasing inability to satisfy
himself how any progressive variation could ever occur. He
crowned his scientific career by various lectures and
addresses in which he reiterated his imaginative failure.
This type of agnosticism was probably the negative aspect
of a passionate and unquestioning faith in the implacable
unteachableness and integrity of certain Mendelian units
of heredity we shall presently describe and discuss. Later
work has removed much of the point of his
criticisms."
The
Science of Life (1931) Wells,
Huxley & Wells. |
|
So what are we to make of The Monk in the Garden. The Lost and Found Genius of Gregor Mendel,
the Father of Genetics by Robin Marantz Henig (2000; Houghton & Mifflin;
Click
Here)?
This is how the book begins:
|
