BOOK REVIEW

doi:10.1111/j.1558-5646.2009.00754.x

WILLIAM BATESON: VARIATION, HEREDITY,AND SPECIATIONMichael J. Wade

Department of Biology, 1001 East 3rd Street, Bloomington, Indiana 47405

E-mail: mjwade@indiana.edu

Received June 4, 2009

Accepted June 4, 2009

Review of: Treasure Your Exceptions: the Science and Life of

William Bateson. Alan G. Cock and Donald R. Forsdyke. 2008.

Springer Science and Business Media: LLC, New York, NY. xxvi,

745 pp. 50 illustrations, HC $59.95, ISBN 978-0-387-75687-5.

The term controversial is conveniently used, by those who arewrong, to apply to persons who correct them.

W. Bateson (Feb. 1907).

This is a densely detailed biography of William Bateson, an

important and controversial figure in the origins of evolutionary

genetics, whose influence as the leading exponent of Mendelism

helped to shape the modern synthesis and whose disagreements

with T. H. Morgan over the chromosome theory of heredity may

well have impeded its acceptance. The book is based on the ev-

idence from a voluminous correspondence, his papers and ad-

dresses, as well as other historical records and tends to follow a

temporal trajectory, breaking Bateson’s intellectual and personal

life into five parts: Genesis of a Geneticist, Mendelism, The Innes

Years, Politics, and Eclipse.

This scholarly work has an unusual style and flavor owing

to its origins as a comingled rather than a coauthored book. This

character is best revealed in the three-part Preface, one by Fors-

dyke, one by Cock, followed by a more typical description of the

content of the coming chapters again by Forsdyke. It combines

a book-length manuscript nearly completed in the mid-1980s by

Cock that lay unpublished for a variety of reasons fused with

extensive revisions and considerable additional research and in-

terpretive perspective added by Forsdyke. Cock, who died in 2004,

began his scientific career at the Poultry Genetics Unit at Cam-

bridge, using the same record-keeping notations and methods as

those of Bateson and Punnett. Forsdyke is a Professor of Bio-

chemistry at Queen’s University whose career spans the pre- and

postgenomics and bio-informatics periods of Genetics, much like

Bateson’s spanned the era before and after the rediscovery of

Mendel.

The temporal trajectory works well within some sections

but not in others. In the period after the rediscovery of Mendel,

where new and synthetic scientific observations and inferences

from a wide variety of sources came quite rapidly, the authors’

approach gives a feeling of immediacy and excitement. It is most

interesting to read about the evolution of Bateson ideas and his

conceptual synthesis of the empirical observations from scientists

and breeders on which they rested, especially in the context of

a blow-by-blow description of the controversy with the biome-

tricians. Bateson was a fierce protagonist in the battle between

the Mendelians and the biometricians before the modern syn-

thesis established the compatibility between discrete Mendelian

genes and the continuous variation of traits like height and weight.

However, the time-line approach and the minutia of his correspon-

dence made the going fairly tedious in the early chapters of the

first section. The many extensive quotations from his correspon-

dence give the reader a strong sense for Bateson, the man, and

perhaps this criticism reflects my own bias and delight in seek-

ing out Bateson, the early geneticist. Much evidence is presented

that, throughout his career, Bateson was relentless but generally

ineffective in his pursuit of funding for experimental studies of

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BOOK REVIEW

heredity and variation—a feature that will resonate with anyone

conducting research in evolutionary genetics today.

William Bateson, 1861–1926, was a central figure in a

number of landmark events and discoveries at the birth of the

field of Genetics and in the study of speciation. He translated

Mendel’s singular work into English, and initiated the rediscov-

ery of Mendel at the turn of the century, remarking in a paper

in May, 1900, “we are in the presence of a new principle of the

highest importance” (p. 202). Bateson is best known in modern

textbooks for introducing the terms “genetics” and “epistasis” and

for his codiscovery with Punnett of epistasis and linkage (which

they called “coupling of characters”) between pairs of Mendelian

factors. They also cofounded the Journal of Genetics in 1910.

The book covers all of this material in exquisite detail. There

is truly a wealth of information here, including how Bateson’s

hostile reception of Morgan’s chromosome theory of heredity led

Morgan to found the competing journal, Genetics, in 1916. And it

shows Bateson’s central role in the evolution–creationism debate

in the United States (where he earned the distinction of being

denounced by William Jennings Bryan) as well as in the Kam-

merer episode over the inheritance of acquired characters. Also

included, are excerpts from several of Bateson’s obituaries written

by both affectionate colleagues and by his opponents in contro-

versy! (Chapter 24 is entirely devoted to “Bateson Bashing,” in

which contemporary authors are found guilty of lumping Bateson

erroneously with mutationists like de Vries.)

In his earliest research, Bateson engaged in the systematic

study of natural variation and its causes, motivated by his con-

viction that an understanding of variation was essential for an

understanding of evolution by natural selection. Notably, he sepa-

rated the causes of variation into those attributable to the gametic

state of the parents and those attributable to other, nonheritable

sources of variation: “There is definiteness of variation and no

one has suggested that this is due to adaptation” (p. 138). He un-

derstood that natural selection could not presume the phenotypic

variation on which it acted and required a firm foundation in the

science of transmissible variations: “Variation leads; the breeder

follows” (p. 234). He labored in his empirical work to sort traits

into those that appeared to be a “blend” of the parental types and

those that tended to be discrete, resembling one parent and not

the other. His early career and the conceptual issues that drove

him read very much like the historical introductions to Darwinism

found in introductory college texts, only Bateson did not know

how these conceptual issues would be resolved.

For his seminal work on variation and heredity, Bateson was

awarded the Darwin Medal of the Royal Society in 1904 and was

named Professor of Genetics at St John’s College, Cambridge, in

1908. There he “fought for the rights of women to have access

to study and research facilities, to be issued degrees, to take part

in university business, and to be eligible for academic positions”

(p. 177). The “end of the Batesonian era” began in 1911 when

Bateson’s “reduplication hypothesis” for the mechanistic basis for

coupling was superseded by T. H. Morgan’s chromosomal theory

of association among Mendelian factors. Unfortunately, in my

opinion, lost in the controversy over mechanism was discussion

of the importance of gene interaction in adaptive evolution.

I found it most interesting to read Bateson’s views on the

reasons for the decades’ long neglect of Mendel’s foundational

work (p. 204). The authors’ detailed discussion of science and

correspondence from this period helped me better understand that

the rediscovery of Mendel was not the smooth, inevitable, and

uncontested process of enlightenment which it sometimes seems

from textbook accounts. Weldon in his 1902 analysis of Mendel’s

experiments in Biometrika remarked that the “numbers are not

large enough to get really smooth results,” and that there was

a “grave discrepancy” between his results and those of other

“equally competent and trustworthy breeders” (p. 206). In a later

paper, Weldon noted that the variation within Mendel’s categories

might have been accounted for by ancestry and his neglect of this

was “a fundamental mistake which vitiates all work based upon

Mendel’s method” (p. 205). Bateson (1902) soundly refuted Wel-

don’s criticisms in his Cambridge University Press monograph,

Mendel’s Principles of Heredity: a Defense. In this same book,

Bateson pointed out that even so few as four or five pairs of

Mendelian elements would give rise to a nearly continuous curve

of variation, clearly stating his belief that discrete and continuous

characters shared the same underlying mechanism of inheritance.

This is 16 years before the publication of R. A. Fisher’s famous

1918 paper on the correlations among relatives, which proved his

point and much more.

An important finding, not typically associated with Bateson’s

legacy, is his understanding of sex in terms of Mendelian factors.

In 1908, Bateson and Punnett published a Science paper entitled,

The Heredity of Sex, concluding “On general rounds it seems to

us probably that one and not both sexes of the same organism will

be shown to be heterozygous for sex and that the approximately

equal output of the two sexes in ordinary cases is a consequence of

this.” He recognized that the male was the heterozygote in some

organisms but not in others; however, his quest for unity of the

underlying genetics led him to associate this evidence with his

flawed “reduplication hypothesis.”

In recent literature, the genetic basis of speciation has been

referred to as the Bateson–Dobzhansky–Muller (BDM) Model.

The “BDM meme” arose in 2000 and arises from an interpretation

of Bateson that is mistaken according to Forsdyke (p. 637). The

Dobzhansky–Muller (D & M) model for the genetic origins of

reproductive isolation rests on two observations. The first is that

gene combinations that work well within each of a pair of species

are incompatible and cause sterility or inviability when brought

together in hybrids. Thus, it is a genetic model of speciation based

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BOOK REVIEW

explicitly on epistasis between genes. The second observation is

that such incompatible gene combinations are in theory unstable

within populations and will be removed from them by natural

selection. The insight of D & M is that the incompatible gene

combinations that separate species need never co-occur within a

population, because they arise by mutation and spread in allopatry

as a result of positive natural selection. The negative epistatic

interactions between derived alleles at different loci are manifest

only in the hybrids between the species. What was Bateson’s

conception of the genetics of speciation? Should he be added to

or given priority over D & M when referring to this iconic model

of the genetic basis of reproductive isolation, which guides so

much current research?

Certainly, Bateson recognized that understanding the origin

of species was in large part a genetic problem. In his Silliman

Lecture in 1907, he said, “Nowhere does our new knowledge of

heredity and variation apply more directly than to the problem

what is a species and what is a variety?” (p. 399). His experi-

mental prescription sounds much like modern speciation genetic

research: “ . . . the first step is to discover the nature of the factors

which by their complementary action . . . cause the sterility of the

hybrid” (p. 400). To my reading, Bateson in his Centenary Essay

of 1909 was also clear in recognizing that the infertility manifest

by the viable hybrids between species had a genetic basis and

that this basis was founded on incompatible gene interactions:

“When two species, both perfectly fertile severally, produce in

crossing a sterile progeny, there is a presumption that the sterility

is due to the development in the hybrid of some substance which

can only be formed by the meeting of two complementary fac-

tors” (p. 331). He was equally clear that “the phenomenon could

only be produced among the divergent [sterile] offspring of one

species by the acquisition of at least two new factors; for if the

acquisition of a single factor caused sterility the line would then

end. Moreover each factor must be separately acquired by distinct

individuals, for if both were present together, the possessor would

by hypothesis be sterile. . . . Next, if the factors responsible for

sterility were acquired, they would in all probability be peculiar

to certain individuals and would not readily be distributed to the

whole breed. Any member of the breed also into which both the

factors were introduced would drop out of the pedigree by virtue

of its sterility” (Bateson’s emphasis; p. 331–332). Like Dobzhan-

sky and Muller, Bateson clearly understood the need for epistasis

as the cause of hybrid sterility as well as the problem of main-

taining the coexistence of incompatible gene combinations within

the same population. Importantly, Bateson also assisted J. B. S.

Haldane in the formulation and publication of his well-known

1922 paper on what is now called Haldane’s Rule. However, in

his invited Nature paper in 1925, celebrating the centenary of

T. Huxley’s birth, Bateson affirmed that “. . . we come to suspect

that no amount of selection or accumulation of such variations as

we commonly see contemporaneously occurring can ever culmi-

nate in the production of that ‘complete physiological divergence’

to which the term species is critically applied” (p. 510). He ap-

pears here to reject a role for natural selection in fixing differences,

which is the last key component of the Dobzhansky and Muller

Model. And, in notes shortly before his death in 1926, Bateson’s

“last word on evolution” contains this remark on the problem of

the genetic basis of hybrid sterility: “the more genetical experi-

ence extends, the more serious does this hiatus in the evidence

become” (p. 516). On balance, I would conclude that Bateson

throughout his career recognized the centrality of the problem of

the origin of species defined by the criterion of intrinsic barriers to

interbreeding. He also clearly formulated the necessity for epista-

sis and the experimental path toward understanding such barriers

in terms of hereditary factors. I find it remarkable that he did so

while Genetics was in its infancy and that his ideas were similar

in so many respects to those of Dobzhansky and Muller, members

of the second wave of great geneticists, who were in their early

teens during what is called the Bateson Era.

I do have some other complaints about the book. I wish that it

had included more of the original diagrams from the various sci-

entists involved, instead of schematics constructed by the authors.

The earliest Punnett squares were laid out somewhat differently

from those of modern texts and they often included pictures as

opposed to cartoons of the animals and plants used for illustration.

It is not clear, despite the dense and freely cited correspondence,

what role such diagrams played in the give and take over the

Mendelian interpretation of data or when they first began appear-

ing in texts. The pictorial representation of key insights and its

role in the rise of genetics is not found here. The authors’ voice

can also be intrusive, especially when interpretively paraphrasing

Bateson’s extensively quoted words; I found this tendency added

a certain amount of conceptual redundancy to the prose. In addi-

tion, in assembling my comments on speciation genetics above, it

was necessary to search back and forth among the major sections

of the book. Nevertheless, these are somewhat minor quibbles

that stem from the nature of this jointly authored work and the

commitments of both authors.

Overall, I strongly recommend this book for its information

on an important central figure and for its bringing to life the several

controversies at the origins of Genetics. It greatly illuminates the

conceptual foundations of evolutionary genetics.

Book Review Editor: J. Thompson

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