History of Science SCIENCE IN THE TWENTIETH CENTURY

Jon Agar, Science in the Twentieth Century and Beyond AND BEYOND
David Knight, The Making or Modern Science

JON AGAR

polity
 NHW SCIItNCHS (lI' 1.llIft

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in diffcrcnt location
I" IHlllccd diffcrcnt li vinil. COI'I11S,
NEW SCIENCES OF L1FE
Heredity and Mendelism

'"11 win's theory was less satisfactory as an explanation of tempo-

IltI vnriation. He proposed that changes between generations were
I "d,,::II - smooth, and continuous. But what was tbe mechanism
111111 curried or caused tbese changes? In later writings, Darwin had
\,1 uposcd a mechanism for accounting for similarity and difference
IlI'OSSthe time-gap of generations. He called it pangenesis, a theory
"I hercdity that proposed that every part of the body - and by the late
ulurrcenth century the unit part was taken to be the cell- shed tiny
But what of the living world? Across space, organisms cxhihlll'll
patterns of similarity and difference. Horses were similar to ~~,I" ,I
1'111'[icles, 'gemmules', which collected in the reproductive organs and
II'pr'csented features of the parent. If the nineteenth century was sliced
Zebras differed from buffalo or crocodiles or baobab trees. 1I1til
Iwny from the twentieth century and we asked what ma de the living
eighteenth and nineteenth centuries the spatial diversity of thc liV1I1
tlI'ld of the old century similar and different to the new, Darwin
world had been classified, measured and mapped. The results, bio
would say it was the gemmules leaping the gap that carried any simi-
ography, were the achievement of scientists such as Alexandor VIIII
lurity or difference.
Humboldt, whose travels through South America were inspiratioll,d
Notice that such a mechanism for heredity made this later Darwin
to the host of explorers who followed in his footsteps. Humboldt IHld
nuhcr like Lamarck: organisms could acquire characteristics that
not only recorded the spatial variations of the living world, but 111111
nuld be passed on. Nevertheless, Lamarckism was vehemently
mapped out such variation alongside other measurable quantities 111
upposed in some quarters. The German zoologist August Weismann,
the natural world: height, temperature, rainfall, and so on. (And, Iri
who worked for much of his life at the University of Freiburg, argued,
course, in order to obtain measurements he had had to take standm ti
lrorn investigations of hydrozoa (such as the Hydra of ponds and
instruments and measures. Comparability across space had requircd
the movement of similar values.) iquaria), that there must exist some hereditary substance - he called
It 'gerrn plasm' - that was passed on between generations. Germ
One such Humboldtian follower had been the young ChaJ'1
plnsrn was not affected by changes in the body, so acquired character-
Darwin, who, as a companion to the captain of HMS Beagl»,
isrics could not be inherited. This 'hard heredity', the absolute inde-
had witnessed the global variety of the living world as the vessel
pcndence of germ plasm from the rest of the body, was demonstrated,
crossed the Atlantic via Tenerife (pausing for Darwin to climl
irgued Weismann, by experiments in which the tails of mice were cut
Mount Teide in emulation of his hero, Humboldt), travcl
ror many generations: since the tails of the last-born mice were just
ling along the South American coastline and stopping at the i
18 long as the first, there was no evidence for the acquisition of the
Iated Galapagos isles, before returning home to London via thc
.haracteristic of 'short-tailedness'. Germ plasm threading across the
Pacifico Darwin's theory of natural selection, finally published in
nineteenth to the twentieth century explained why twentieth-century
1859, offered an explanation of the variety found in the natural
world. organisms were like nineteenth-century organisms. Different combi-
nations of germ plasm in a new generation explained why individuals
Darwin's theory accounted for spatial variation, among, fOI'
differed.
example, the finches of the Galapagos, by proposing that variations in
Many scientists in the late nineteenth century found Darwin's or
organisms, passed on between generations, were selected for, if they
Weismann's accounts implausible. Perhaps the greatest difficulty
were to survive, by the environment, living and non-living, in whicb
they had was with the role of gradual continuous change between
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sei HNCI' li II'I'HII .1tJI
gcncrations. Tnkc, for cxurnplc, 111l OI'gl1lliNlll Ihul WllS CVCI' so slighcly
better cquippcd to survivc rhc rigoUI's o/' lhe natural world. [t rnay
even be able to pass on this charactcristic to its offspring. However
to reproduce, it would have to mate with an organism with lesscr
talents, and any advantage would be reduced. Repeat this process a
few generations and it was clear, at least to many, that any advantag
would be irretrievably diluted: 'swamped' was a term used. Another
dissenting argument carne from the towering figure of William
Thomson, Lord Kelvin, who had pronounced with all the author-
ity of a physicist and a peer of the realm that the relatively rapid
cooling of the earth had not left time enough for evolution by natural
selection.
The success of such arguments left Darwin a relatively marginal
figure by 1900. While the fact of evolution was largely accepted,
many scientists - including the late r Darwin - were non-Darwinian
in two senses. First, they may be Lamarckian, at least in the sense
of accepting some kind of acquired characteristics as an impor-
tant route by which features could be passed to later generations.
Most scientists around the turn of the nineteenth into the twentieth
century accepted evolution and preferred Lamarckian mechanisms to
Darwinian natural selection.' Second, Darwin's theory of the natural
selection of continuously changing characteristics was rejected in
favour of selection over discontinuously changing characteristics.?
A 'sport', a member of a new generation that was radically differ-
ent in size or shape or performance from the parent generation - an
observed and familiar phenomenon in artificial breeding - was seen as
the source of evolutionary steps. These jumps were also called muta-
tions or 'saltations'.
On the other hand, a new group of scientists and mathemati-
cians, who looked and measured different organisms, clung to
the Darwinian picture of continuous variation. If the heights of a
hundred men are measured and plotted on a graph, numbers against
height, then a peak is observed. Few men are very short, and few men
are very tal1. More men are of heights in between. Take enough meas-
urernents and the graph begins to look quite smooth. lndeed the more
measurements taken, the smoother the graph. Every range of height
between the shortest and the tallest is eventually taken, and the peak
reveals an average. A similar pattern can be found when many natural
variables are measured. As an image of the natural world, such
graphs powerfully support the idea of continuous gradual change
across a population. The scientists and mathematicians who made
such measurements, 'biometricians' ('measurers of life') such as Karl
46
NHW ~CIHNt;nS () I' 1,11'1
bomnists such os W, lo', R. Wcldon in London,
backed Darwin and continuous variation.
The potential for a fierce dispute bctwccn the biometricians and the
altationists turns out to be critical to how the work of an obscure
mid-nineteenth-century monk, Gregor Mendel, was received. One
outcorne of the debate would be a new science, genetics, which pro-
vided the twentieth century with its dominant language for talking
about heredity. However, the reworking of Mendel happened in
two phases or contexts.:' The first concerns the so-called rediscovery
of Mendel around 1900, and the second is the debate between the
biometricians and the saltationists. We will examine both in turn.
But, first, let's rewind to say something about who Mendel was and
what he did.
Johann Mendel had been born into a family of peasant farmers in
1822.4 When novices enter a monastery they can change their name,
and, on admission to an Augustinian monastery at Brünn in Moravia
(now Brno in the Czech Republic), Johann Mendel became Gregor
Mendel. Rather unusually, the novice was allowed to study at univer-
sity, attending in the early 1850s the University of Vienna, where he
was lectured by Franz Unger in cytology (the science of cells), evolu-
tion and even hybridization experiments, and in physics by Christian
Doppler. After returning to the monastery, Mendel grew plants in the
gardens and investigated hybrids. In his famous experiment, he noted
seven pairs of character differences in Pisum (pea) plants - some seeds
were yellow and others were green, or some seeds were wrinkled and
others were smooth - and he cross-bred plants with opposing char-
acteristics. ln results presented to the Brno Natural History Society
in 1865, and published in their journal the following year, Mendel
reported that, while a character might entirely disappear for one gen-
eration, it would reappear in a fixed 3:1 ratio in the second genera-
tion. Characters seemed to be carried by discrete 'elements' passed to
the next generation.
The later mythologizers of Mendel would hold that the monk's
work was ignored by the scientific world, only to be triumphantly
vindicated thirty-five years later. Loren Eiseley, in Darwin's Century
(1958), for example, dwells on the moment of Mendel's presenta-
tion: 'Stolidly the audience had listened ... Not one had ventured a
question, not a single heartbeat had quickened ... Not a solitary soul
had understood him." Had Mendel been neglected, it would have
been for good reasons: the paper contained mathematics that would
have been forbidding to an audience interested in hybridization, and
the paper being presented was by a scientist of little status and was
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 N I'.W 1\t:lltNCI!~ \)1' 1.1\11
sei HNCI' 1111'1'1(11
IUI
HlllhCl'ing alurm. 110, LOO, hnd cxp<.:rjmenl\\1 cvidcnce for" 3: I rutio,
publishcd in un OhSCLll'C jourunl. RCI'I'()sp~'~livtdy, Mcnd
Iht' product of ycars of pClinSLl1king lnbour. Whac should he cio?
been accused (for cxarnplc, by Ronnld Aylmcr Fishcr]
hould he concede dcfcat graccfu Ily, as scicntists are su pposed to
clear-cut, the nurnbers almost unbclievablc." ln face Mcndcl's worl
lo, ncknowlcdging that De Vries had pipped hirn tO the post, ar
drew him into correspondence with some of the bcst botanists of rh
.ould he find a compromise that recorcled his contribution? Correns
day, men such as Carl Nãgeli. So Mendel was not ignored. A ralhei
.hosc to publish his results, but, unlike De Vries, he cited the work
different factor in the fate of his science was Mendel's own changinu
of the obscure monk, Gregor Mendel. What Correns was saying was
research interests: he increasingly left peas for another organism '
rhis: Correns may be second to De Vries, but De Vries and Correns
Hieracium (hawkweed), a small dandelion-like plant common acr
1 ogcther were following an older ancestor. As Augustine Brannigan, a
Europe but possessing (we now know) an immensely complex proccss
historian of scientific discovery, explains, Correns framed
of reproduction, involving self-pollination and hosts of rnicro-speci
Furthermore, Mendel was elected abbot of the monastery in 18 his announcement so as to indicate that though he had lost priority to
which filled his time. de Vries, both had lost out to an earlier researcher, even though the
Mendel's work was known in the scientific world, but it was not initíal intent of that research ... [was] somewhat less than identical. ln
obvious that it was of great importance. Only late r, retrospectively other words he neutralises his loss ... This is accomplished decisively
would its relevance seem clear. As historian Peter Bowler has sum- by labelling the discovery 'Mendel's law'. This is perhaps the most
marized, the 'traditional image of Mendel is a myth created by th important fact in the reification of Mendel as founder of genetics.f
early geneticists to reinforce the belief that the laws of inheritance ar The reason why Mendel is famous now has much to with the way
obvious to anyone who looks closely enough at the problem.'? The
that this altercation between Hugo de Vries and Carl Correns was
scientific world needed a positive reason to return to Mendel. settled. Had Correns chosen a different strategy it is quite likely that
ln 1900, a Dutch botanist, Hugo de Vries, published a paper in the 'Mendel's Law' would be known as 'De Vries's Law', ar 'Correns's
Comptes Rendus, France's premier scientific journal - far more pres- Law', ar even 'von Tschermak-Seysenegg's law', after an Austrian
tigious than a provincial natural history newsletter. The Netherlands,
botanist who also interpreted Mendel's ideas around the same time as
of course, was a world centre of horticulture, especially the breeding De Vries and Correns but in terrns of largely pre-Mendelian concepts
of flower-bearing plants. Since 1866, De Vries had been growing and
comparing a plant called the evening primrose, in particular a species of heredity.
Following the 'rediscovery' of Mendel's ideas, Mendel's original
called Oenothera lamarckiana. A tall plant with large yellow flowers, paper was republished and probably read by many for the first time.
Oenothera, in fact, was rather like Hieracium, in appearing in many If the initial resurrection of Mendel had been brought about by a pri-
different forms, ar micro-species. Also, it was the kind of organ- ority dispute between botanists in continental Europe, his fame was
ism that frequently threw up sports, and De Vries was a convinced firmly secured by appeals to his name in a fierce British controversy,
believer in evolution by discontinuous variation. Indeed, in Oenothera
between the biometricians and the saltationists.
he thought he saw mutations in the process of becoming new species Recall that the biometricians saw patterns of smooth varia-
in front of his eyes. In 1899, he had written up some of his views tion within organisms of the same type. They saw themselves as
in Intracellular Pangenesis, which had divided Darwin's pangenesis Darwinian. On the other hand, the saltationists argued that evolu-
concept into, first, a concept of material particles responsible for each tion happened by the selection of qualitatively different 'sports',
character of an organism (which he accepted) and, second, the trans- dramatically different-shaped individuals thrown up once in a while.
portation hypothesis, the idea that these particles moved around the William Bateson, a Cambridge zoologist, thought such discontinu-
body, which he rejected. As De Vries's plant-breeding experiments ity was typical in populations. He rejected Darwinism, and wrote
expanded, he began to observe a 3:1 ratio in the second generation. books such as Materials for the Study of Variation (1894), saying 50.
The 1900 paper set out these results in support of his notion of the Around 1900 he too read the now publicized work of Mendel, and
segregation of characters. But he did not cite Mendel. was convinced not only of Mendel's evidence with regard to peas, but
Meanwhile, Carl Correns, a German botanist, received an advanced that Mendel's laws were universally valid, and that in the 3:1 ratio
version of Hugo de Vries's first article, and read it with interest and
49
48
 .~CIHN(::1l 1\1"11111 IIJtj()
thcre was cvidcnc
rnarshalled ali the evidence hc could lny his hunds on anel ar
general case in Mendel's Prmciples
'Mendel' had now come.
Why genetics? Breeding and eugenics
But Bateson added something else of significance: a new name.
Frustrated that the science of heredity, defined along saltationist
lines, was attracting little support or patronage, he put elear blue
water between his kind of science and that of the biometricians by
naming his work 'genetics'. lronically, then, 'genetics' was named and
promoted as a weapon against Darwinism. Several other core terms
of genetics were invented during these exchanges. The Danish bota-
nist Wilhelm Johanssen, for example, took 'pangene', a term of De
Vries's, and shortened it to 'gene'. (Yet what Johanssen thought the
term referred to was an energy state within the organism, probably
unobservable, and quite different from what 'genes' might mean later
in the century.) Johanssen also insisted that the 'genotype' (another
of his new words) was the correct subject for the study of heredity,
as opposed to the 'phenotype', or the body. Genetics can be seen
emerging as part of explicit attempts to rule out other experts, such
as biometricians or embryologists, who did know how to measure
and study 'phenotypes', from being able to speak authoritatively on
heredity.
But why was it worth fighting over?
There are two, intimately connected answers. First, in the working
world of agriculture, the possession of skills, or even merely the repu-
tation, backed with the prestige of science, to be able to breed better
plants or animals was lucrative. The possessors of a science of hered-
ity would expect to be encouraged and be able to attract patronage
from the working world of animal and plant breeding. Second, the
idea that these methods of 'good breeding' could be applied to human
reproduction, the project of eugenics, greatly appealed to many
groups and individuais in the late nineteenth and early twentieth cen-
turies. Let us consider each of these interests in turno
The organized application of science for agricultural and horticul-
tural improvement dates at least from the seventeenth century. In the
mid-nineteenth century Liebig's research school at Giessen had sys-
tematically moved between new chemistry and methods of increased
production of foodstuffs. The spirit of the eighteenth-century
50
NHW SClltNCH~ Olr t.l11J
improving gcntlcman-furrner had rnorlvnrcd ehc cstablishmcnt of thc
Rothamsrcd Experimental Station in Hcrtfordshire in 1843, where
john Bennet Lawes's long-terrn experirncnts on fertilizers and crop
yields began; they have run continuously ever since.? A trust fund set
up on Lawes's sale of his fertilizer business in 1889 provided funds
to keep Rothamsted an independent concern, collaborating but not
controlled by government or farmer interests. In the United States,
however, agricultural research was an early and highly significant
recipient of federal and state funding.!? In 1862, the Morrill Act had
provided federalland, which in turn could be sold to fund the expan-
sion or creation of 'land-grant colleges'. In the mid- to late nineteenth
century, the United States was an industrializing nation, which in turn
created a demand for increased agricultural productivity. In 1887, the
Hatch Act repeated the Morrill Act ploy, granting federal land for
sale for states to fund agricultural experiment stations. Many of these
stations were attached to land-grant colleges. The result was a large
number of institutions, each with a research and teaching staff.
In 1903, members of the American Association of Agricultural
Colleges and Experiment Stations established the American Breeders'
Association, attracting as members a broad audience of 'commercial
breeders, professors at agricultura I colleges and experiment stations,
and researchers at the US Department of Agriculture', with the latter
two providing the 'great majoriry'c-! Its leader was Willet B. Hays, of
Minnesota experimental station and, from 1904, assistant secretary
of agriculture in Theodore Roosevelt's administration. Hays had
a strong preference for broad cooperative alliances, ineluding ones
between researchers, plant breeders and animal breeders; the 'pro-
ducers of new values through breeding', Hays summarized in a 1905
speech to the association, 'are here brought together as an apprecia-
tive constituency of their servants, the scientists'i '?
The breeders had a primary and overwhelming interest in improv-
ing stock, to which they brought two established methods, the selec-
tion and breeding of best individuals and hybridization to breed in
desired characteristics. This combination, for example, was behind
the formidable successes in plant breeding at the Vilmorin company in
France.P However, unlike the French, the American breeders rapidly
embraced Mendel's theory. Between 1903 and 1909 the proportion
of papers presented at meetings of the American Breeders' Association
mentioning Mendel increased from one-ninth to one-fifth, and 'virtu-
ally all such citations corroborated the theory'; the 'breeders par-
ticularly appreciated the predictive value of the Mendelian pattern
of inheritance of various traits', which chimed with their practical
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 i>C/JlNCI' AII'I'llIt I N 1'\\1 !I(~IHNelti> (')11 LI fll

intcI'CSCS,I4 'lhc prncticul nnd PI'OfCNHiol1nl lnrcrese in WItH 'cnuse I'hc useful clllSSCS in lhe community
lO to contributc 11101""
dcsirable cha ractcrs cncournged thc breedcrs to em brace ot her 11 ,111111thcir proportion to the ncxr gcncrarion'."
methodoJogies, for instance rho biomctry of Pca rson, wh ich "rolld~1'I1 Eugcnics was dccisivcly shapcd by lhe devcloping sciences of
predictabiliry and control. In contrast, the brccders were far lcss "('l'll hvrcdiry in thc late ninctccnth and early twcnrieth centuries. Civen
on the emphasis on unpredictable sports in Hugo de Vries's l11utnllllll '''Ilt Weismann had undcrrnined Larnarckian evolution, and had
theory. The professional breeders were among the first, for CXQIIlpl usisrcd that there was a hard separation between the germ plasma
when compared to elite academic scientists, to distinguish shnrp] (111(, vchicle of heredity) and the somatic cells, then how could a race
between the two theories; the working world of the breeders thcrefo: hl' improved? Any improvement to the cells of the body during the
played a substantive role in the shaping of genetic science. lrlc of the organism would not be passed on to the next generation.
The second answer to the question as to why the science of hored I'hcrcfore, the only source of improvement was the germ plasm - the
ity was worth fighting for in 1900 can be uncovered by followhu l'I1CS, in the new jargon.
another saltationist, Darwin's cousin Francis Calton. Something (lj And many thought there was an urgent need for improvement.
a child prodigy, very much a sport, Calton had read mathernatics III 'l'he sprawling nineteenth-century metropolises such as London,
Trinity CoUege, Cambridge, had a nervous breakdown and travclled l'nris or New York, with their mobile and sometimes violent popula-
in Africa before reading his relative's great work, On the Crigin (lI tions, were theatres of fear. Social mobility threatened those on topo
Species, in 1859. Calton was possessed by the project to quantif l'ca rs of decline or degeneration were often nationalistically inflected.
Darwin's theory, to place the theory of evolution by natural selection I\ugenic campaigns, therefore, were often national campaigns to pre-
on a firm mathematical basis.P He began a life's work collecting datn serve the national race. Overall, it was a powerful strategic move to
on human variation, developing methods of measurement and rnath .lairn expertise in good breeding, which is why it was worth fighting
ematics to make sense of it. Calton published some of his quantitn- or the right to speak authoritatively on heredity. Of course, someone
tive arguments, such as the claim that the children of eminent fathcrs - of some social group - would have to choose who were and were
tended to inherit superior qualities, in Hereditary Genius in 1869. not good breeders. This fact that the eugenic official - 'a compound',
'Calton's achievement in founding statistical biometry should not bc noted biologist J. B. S. Haldane, 'of the policeman, the priest, and the
minimised', writes historian Theodore Porter. 'He did so by mergin procurer' (or pimp) - would be socially self-interested was eugenics'
evolutionary theory with "statistics" in the nineteenth-century sen political advantage and flaw.l?
of the term - that is, social numbers.vf In his English Men af Science: Eugenic movements were active in Britain, the United States,
Their Nature and Nurture of 1874, Calton also contributed, via ermany, Scandinavia and elsewhere. Members were mostly white
subtitle, an oppositional framework for debates on heredity that live and middle class, with large numbers of professionals, scientists (par-
on still.
ticularly, later, geneticists) in addition to non-scientists.ê? Eugenics
Calton reflected in these works on the pairing habits of the English rnovements provided 'a legitimation of the social position of the pro-
upper classes, a topic that encouraged attention to notions of good fessional middle class, and an argument for its enhancement'. 21 Their
breeding. He was convinced that society must ensure the more active aims were to prevent social 'degeneration'. Problems of industrial
reproduction of its fitter members if it was not to be swamped by ociety - such as crime, slums, prostitution - were attributed to bio-
the 'unfit'. He proposed, for example, that members of families of logical causes, first poor 'blood' and later poor 'genes'. In all eugenic
high rank should be encouraged to marry early. ln 1883, he coined a programmes there was a special focus on those identified as 'feeble-
term 'eugenics' - or 'good in birth' - to capture this programme: the minded', as well as certain medical conditions (such as epilepsy and
improvement of the human race along the lines of the plant or animal diabetes) and perceived social burdens (alcoholism, prostitution,
breeding. Discouragement ar prevention of human reproduction criminality, poverty). In particular, eugenicists distinguished and
judged undesirable is called 'negative' eugenics, while programmes described underclasses for special attention and action. 'Defective
such as the Galtori's, to encourage the higher ranks to breed early and delinquency' theory, for example, equated 'high-grade feeble-mind-
more often, 'to multiply reproduction judged desirable, is termed 'pos- edness' with criminality.
itive' eugenics.l? Overall, Calton wrote in 1904, 'the aim of Eugenics' In Britain, Calton encouraged the foundation of a Eugenics
52 53

sei HN(~H A 1I'1'lllt II
Records Officc l1C Univcrsity ColI~I-\U l.ondor: til 1904, followcd by
a Eugenics Laboratory in 1907, directcd hy Kl1l'1 Pcarscn. 111 ]91 J,
Galton's bequest of f45,000 creatcd a chair of cugcnics, held by
Pearson, followed by R, A. Fisher. A Eugenics Education Society -
later shortened to the Eugenics Society - was formed in 1907, and
journals were produced: The Eugenícs Revíew and Biometrika (from
1901), The political achievements of the British eugenics movement
included family allowance policies, a positive eugenic measure that
rewarded the right kinds of families, and the Mental Deficiency Act
of 1913, which 'allowed for eugenically motivated segregation' and
remained in force until1959.22
In the United States, the most active centre of eugenic research was
the Eugenics Record Office, at Cold Spring Harbor, Long Island, New
York. Its director, Charles B. Davenport, was a keen Mendelian.êê
Davenport was also, with Willet Hays, one of the leaders of the
American Breeders' Association, a strong indication that ideas about
'good' human, animal and plant breeding were considered of a piece
in the early twentieth century. The association set up a Committee on
Eugenics in 1906 and pushed eugenic programmes in its Amerícan
Breeders' Magazine; Hays in 1912 addressed the National Farmers'
Congress on the subject of 'The farm, the home of the race', outlin-
ing a vision in which corn-fed 'genetically efficient people' would be
encouraged to breed, all justified by appeal to Mendel.ê" Meanwhile,
Davenport's Heredity ín Relatíon to Eugenics of 1911 attributed to
single genes many mental and behavioural characteristics, including
insanity, epilepsy, alcoholism, 'pauperism', criminality, 'shiftlessness'
and even 'thalassophilia', or a love of the sea, found in families of
seafarers. According to research conducted at the Eugenics Research
Office by Henry H. Goddard, feeblemindedness was a 'condition
of mind or brain' which was 'transmitted as regularly and surely as
color of hair or eyes' .25The office also sponsored in-depth studies of
families, tracing the appearance of undesirable traits across whole
genealogies.ê"
The political impact of eugenic research in the United States was
extensive and lasting. Starting with Indiana in 1907, sterilization
laws, negative eugenic methods that brutally prevented undesirable
breeding, were introduced in twenty-eight states by the 1920s; 30,000
people had been 'sterilized on eugenic grounds' by 1939, nearly half
of them in California.ê? Such laws were challenged but declared
constitutional in a Supreme Court decision (Buck vs. Bell, 1927), in
which Justice Oliver Wendell Holmes declared that 'three generations
of imbeciles is enough'. The United States led the way with eugenic
54
NItW SCIItN<:HII ()II I.IIIH
laws. Many Amcrican sentes WCI'C OII'Ol\\.ly 1'lr\'\!.:t1sillg seorilizaelon
before they werc joincd by Buropcan stctcs: the Swiss canton of
Waadt (1928), Denmark (1929), Gcrmany (1933), Norway (1934),
Sweden and Finland (both 1935), arnong orhers.ê"
Until the 1930s, eugenics programmes and policies in Germany
were broadly comparable with those in Britain, the United States
and Scandinavia. Indeed, German eugenicists looked to states such
as California, which sterilized more people than any other, for inspi-
ration and example. The active centres included the Kaiser Wilhelm
Institute for Research in Psychiatry, established 1918 in Munich, and
the Kaiser Wilhelm Institute for Anthropology, Human Heredity and
Eugenics, founded in Berlin in 1927. German sterilization laws were
in place before the Nazis took power in 1933.
The technocratic appeal of eugenics - the opportunity to claim the
rights to steer social policy through an appeal to technical expertise -
and the credit, status and resources that might follow partly explain
the intense interest in sciences of heredity in the late nineteenth and
early twentieth century. It was not a prize that could be shared, and
the competition to secure the right to speak authoritatively was the
context in which key terms - not least 'genes' and 'genetics' - were
used. Nor was it only a say in human heredity that was at stake:
authority over plant and animal breeding would als o pay dividends.
Nevertheless, despite the pressures to compete and challenge, in
the 1910s the controversy between biometrical and Mendelian sei-
ences of heredity was significantly, if not completely, resolved. The
triumph of techniques of two kinds - statistical and biological - was
the heart of the resolution.
Galton's protégé was Karl Pearson, a fierce supporter of Darwinian
gradualism, who developed the mathematical techniques to under-
stand correlation between variables, techniques that became standard
statistical methods. His x-squared test, for example, was proposed in
1900 as a replacement for the established normal curve. Sociologist
Donald MacKenzie considers Pearson's statistical methods to have
been shaped by his eugenic interests, while historian Eileen Magnello
denies any influence between the two central projects of Pearson's
life.29 Nevertheless, writes historian Ted Porter, it is in Pearson's
projects that the ambition and reach of biometrics can be clearly
witnessed:
His Grammar af Science (1892) was, after all, a philosophical tract
designed to extend the reach of science into every domain. He idealized
statistics, a field long nourished by similar ambitions, as the true form
for all scientific knowledge. Statistics was descriptive and positivistic,
55
_.~--------------
 NCIIi,NCH Alrl'l'lI 1\ NI(W ~t:II!NCIt~ 01 1,IIIi'
'

to be cnergy sinres, yet [)\: Vrics thought them

IIHlw'inl, nnd in the hnnds of thc mnth\:I"uticians they were a manipu-
luhlc vnriablc with unkriown rcfcrcncc. The identification of genes
wlth a material substance - the 'chromosomes' that microscopists had
Iound in cell nuclei - is usually considered the achievement of Thomas
l lunt Morgan, although, as we shall see, it was the creation of an
'xrraordinarily rich and productive material culture that enabled him
10 sustain the claim. ln fact the identification of chromosomes as the
hereditary substance was another aspect of the emergence of genetics
It was precisely because 'genes' could not pass the positivist criteriou where the working world of plant and animal breeders at least pre-
that Pearson thought them metaphysical, and to be rejected. TI1\' pn red the ground.
debate between Mendelians and biometricians was therefore simul The idea that pairs of chromosomes, observable in cytological
taneously about the proper method of science - in particular what studies, might be associated with pairs of Mendelian factors occurred
could be measured and how such measures should be compared - and to many scientists after 1900. Evidence was put forward in 1902
about what scientific objects existed. by two scientists in particular; the venerable German embryologist
Pearson, the first Galton Professor of Eugenics at University 'I'heodor Boveri, working with sea urchin eggs at Naples, argued
College London, was an ideologue, highly unlikely to concedi rhat normal development required full sets of chromosomes, while
ground to his enemies, and while he held power the controversy wa in Columbia, a young postgraduate, Walter S. Sutton, drew similar
not going to end. However, his successor in the UCL chair, Ronald conclusions based on studies of an agricultural pest, the grasshopper,
Aylmer Fisher, was more flexible. ln 1918, Fisher published a paper in Kansas. Nevertheless, the theory remained controversial. However,
'The correlation between relatives on the supposition of Mendelian papers on breeding, genes and chromosomes appeared frequently in
inheritance', which gave a sharp mathematical sense to the concep the Proceedings of the American Breeders' Association. The United
of variance. Now defined as the square of the standard deviation, dif- States Department of Agriculture scientist W. J. Spillman argued for
ferent variables could be identified and measured in convenient and a connection in 1907; the Illinois swine breeders Q. L Simpson and
clear ways.ê! Deploying variance, Fisher argued that the gradually J. P. Simpson enthusiastically attributed Mendelian factors to chro-
changing data found by the biometricians could be produced by the mosomes; while W. Lawrence Balls, of the Khedivial Agricultural
action of combining large numbers of discontinuous variables such as Society, argued in 1909 from cotton research for the same attribu-
Mendelian genes. ln other words, Mendelian genetics could produce tion.32 The 'readiness to entertain' such theories was encouraged by
Darwinian results. the breeders' practical concerns and illustrate how the working world
This mathematical argument was not enough, in itself, to resolve was ready to receive, and in turn maintain, Morgan's chromosomal
the controversy. Just because Mendelian genes could produce smooth theory.
changes in the characteristics of large populations did not mean that Morgan had benefited from training at the American university that
Mendelian genes existed, except as a useful mathematical fiction. most closely followed the German model, Johns Hopkins University.
lnstead, the emergence of a scientific consensus of the material reality He had won his PhD in 1890. ln 1904 Morgan was appointed profes-
of genes required a solution that was part material technique, part sor of experimental zoology at Columbia University. At Columbia, he
social innovation. chose an organism that seemed ideal for biometric studies and, most
relevantly in a university following the German model, for teaching
by research.ê ' The fruit f1y, Drosophila melanogaster, was common,
Making and circulating Drosophila genetics was easily sustained - a population would happily breed in a glass jar
with some bananas as food - and multiplied rapidly. Large popula-
Recall that there was considerable disagreement in what scientists tions of flies provided plenty of material for measurements. lndeed,
took 'genes' to be, even if they admitted their existence. Johanssen Drosophila's cheapness made the fly suitable student project material.

56 57
 SClltN\;1tAII'I'HU I'
It was, howcver, [111unlikely olW1llism 011 which to sruke n pl'llI
sional ca reei' as a gcncricist: it hnd 1)()I1C of lhe stn rus or Ii eeruu 11li 01
experimental marnmal organisms, nor was it dorncsticatcd, ut\d !lUI
did it seem to show many visible characters or mutations,
However, the choice was fortuna te. The ceUs of thc Sl1IiV.\1
glands of Drosophila have four chromosomes of imrnense phYIlil:ul
size, some of the largest among all organisms." They were morphn
logically distinct, and their size made them easier to dissect OLl( 1\11I1
examine. Furthermore, as the biometrical studies were sca led u p, 111
populations under study became so large that mutations were mOi
likely to appear. From 1908, Morgan increasingly used Drosoplul«
as a mo del organism in Mendelian research. (lndeed, he had startcd
the investigation thoroughly sceptical of Mendelism.) His method
encouraged the building of a research schoo1. Students could h..
trained and fellow researchers could work together, coUaboratin
around the jars of Drosophila. The identification of new phenotypcs,
and the breeding of separate populations that displayed interest
ing traits, was doable but productive science - 'normal science', in
Kuhn's terms. Furthermore, Morgan's method could travel: sma11jar
of breeding fruit flies were exchanged with like-minded colleagu
around the United States, and eventually across oceans.P
Many eyes looking down microscopes are better than one. ln 1910,
Morgan's school noticed rare phenotypic variations and mutations,
including male flies that had white eyes as opposed to the typical
red. This seemed odd: why should a trait such as the colour of eye
be confined to one sex? Furthermore, some sex-linked traits wer
not inherited together, while others were. This prompted Morgan to
speculate that genes that were closer together were more likely to be
inherited together - in doing so he drew on the studies of the skilful
Belgian cytologist Frans Alfons Janssens, who had observed through
the microscope chromosomes approaching, breaking and recombin-
ing. From this insight, Alfred Henry Sturtevant, who had been trained
in Morgan's fly room and shared the work of identifying and isolating
the white eye mutants, drew up 'chromosome maps', which showed
which genes were dose and which were distant. The results were pub-
lished in 1913. Notice what was happening here: the articulation and
spread of material techniques that gave Morgan and his school the
ability to isolate and quantify traits in populations were accompanied
by the identification of the genes with a specific location in a material
substance - the. chromosome - and, indeed, a measurable position
within it.'
In 1915, Morgan, with students Sturtevant, Calvin Blackman
58
NIIW SCIHNCH~ (lI' 1,llil
Ill'idl-lCHnnd Ilel'I1W1111Mullcr, publishcd thc methods and rcsults
11'/,/)(1 Mecl)(;lIfisl/1. o( Melldelioll l/orcclity. This work of synthesis
dl.!ineJ lhe cornponcnts of thc classlcal theory of genes, induding the
lnlms that: 'chrornosomes are che bearers of hereditary material',
"'!-Icnes" (not "unir characters") are the fundamental units of hered-
t y', 'genes are arrayed linearly on chromosomes', 'the number of
llnkngc groups of genes ... equals the number of chromosomes', 'each
lisrinct gene, though it may have many alieles, remains unchanged
-xccpt by mutation', 'environmental factors (e.g. temperature and
uutrition) can influence the effects of some genes', 'some genes can
modify the effects of other genes, sometimes quite specifically', 'genes
t hcmsel ves are not altered when their effects are changed by modifier
!-Ienes', 'genes must cooperate in large numbers to yield observable
truits', and 'many mutations have large effects, but many more have
mall effects'; altogether, 'even though the pathways from genes to
.haracters are wholly unknown, Mendel's principles, interpreted via
gcne theory, provide "a scientific explanation of heredity [that] fufills
J Il the requirements of any causal explanation'''. 36
At a growing pace throughout the twentieth century, to be a certain
kind of geneticist was to be part of the network that sustained and
.xchanged bottles of fruit flies and interbred, dissected, observed and
correlated the features of their model organismo Similar people, in
the sense of trained students, passed through Morgan's fly room and
research school, and carried similar techniques and organisms with
them. With the network, and similar flies, people and techniques, in
place, a statement or measurement about the chromosorne was more
likely to be comparable, repeatable and stable - or, where necessary,
indeed, disputed in agreed, contained and productive ways. The mul-
tiplication and export of similar entities were the secret to Morgan's
success. In application to animal and plant breeding, too, new genetic
knowledge would contribute to other circulations of similar things.
Furthermore, the heredity that genetics was mastering was increas-
ingly an aspect of an artificial world. The mutants were chosen and
favoured in reproduction. The fruit fly became a standard in order
that mutations might be identified and isolated, and so that rneasure-
ments could be made and compared in different places and at different
times. The fruit fly was not really Drosopbila melanogaster any more,
at least in the sense that the population of the fly room diverged from
the population in the 'wild', but might be called, perhaps, Drosophila
laboratoriensis, a creature of a particular artificial contexto J ust as
in physics, the remarkable new natural sciences of the turn of the
nineteenth into the twentieth century were the result of the study of
59
 1-I(:lItNI:It AP'I'IlH 1\
an artificial world, And lhe
ished. Morgan, at Colurnbia and from 192H \H lhe Cnlifornia Institue
of Technology, was a world lcadcr in thc scicncc of genctics, whiJ
Drosophila rernained the leading mcdel organism for genetics int
the twenty-first century. William Bateson, the Englishman who had
coined the word 'genetics' in 1905, paid homage to the shift in power
at the gathering of scientists at the American Association for the
Advancement of Science in December 1921: 'I come at this Christmas
Season to lay my respectful homage before the stars that have arisen
in the W est. >37
Another consequence of the sheer success of Morgan's Drosophila
work was a narrowing of Mendelism as a scientific programme.
'Since the development and evolutionary consequences of Mendelism
were not readily tested', note historians Richard Burian and Doris
Zallen, 'those issues were gradually dismissed as speculative and set
aside.t " The roles of genetic change in evolution would not return
as research questions until the period of the so-called Evolutionary
Synthesis beginning in the 1930s. The separare, difficult, if profound,
questions about the relationship between development and genetic
make-up would return as highly active research topics towards the
end of the century.
Working worlds of the life sciences
We have seen that agricultura 1experimental stations and agricultural
colleges provided institutional homes for the new genetics. The first
American genetics departments - at Berkeley, CorneU and Wisconsin
- were located in the agricultural faculties.ê? University-trained
plant physiologists, employed at the United States Department of
Agriculture's Bureau of Plant Industry, began the development of
hybrid corn deploying Mendelian theory."? The Agricultural CoUege
in Berlin housed genetics in Germany - although it was unique in
Germany as a place where genetics researchers established strong
links with breeders."! Many early Mendelians found employment
at agricultural institutions - such as Hermann Nilsson-Ehle at the
Swedish plant-breeding station at Svalõf, Erich von Tschermak-
Seysenegg at the Agricultural College in Vienna, William Bateson
at the John Innes Horticultural Institution in Britain, and Raymond
Pearl at the Maine Agricultural Station in the United States.P This
working world context for genetics would be continued during the
Evolutionary Synthesis era, when, for example, R. A. Fisher worked
60
NItW SCIHNCI\1-I ()lI 1.1111'
at Rothamsrcd and Scwall WI'ight wns cmplnycd ne thc Madison agri-
cultural station.
But genetics was not the only life scicnce that was encouraged
within this working world. 'Biochemistry', notes historian of biology
Jonathan Harwood, 'also took root in agricultural soi1.'43 Eduard
Buchner had been appointed professor of general chemistry at the
Agricultural College in Berlin in 1898, after passing through lecture-
ships at Munich, Kiel and Tübingen. His research showed that the
liquid obtained from ground and pressed yeast cells would ferrnent
sugars imo a\cohol. Buchner would be awarded the Nobel Prize for
chemistry in 1907 for the discovery of this ceU-free fermentation,
and he is remembered as a father of a new discipline, biochemistry.
Biochemistry would flourish in the twentieth century, although less so
in the United States, where it was located in medica 1 schools in which
it was encouraged - and constrained - to develop analytical methods
useful in the clinic and in medical science; at Cambridge, under F.
Gowland Hopkins, and in Berlin, under Otto Warburg, biochemistry
was located away from medical institutions and targeted basic bio-
logical problems with success."
As well as at the Agricultural College, fermentation research
was carried out in Berlin at the Institute for Fermentation
(Garungsgewerbe), where Max Delbrück, uncle of the famous mid-
rwentieth-century physicist turned biologist, was also exploring
yeast. Delbrück saw the organism in engineering terms: at a confer-
ence in 1884 he had begun a papel' with the simple staternent 'yeast
is a machine'r'r Outside Berlin, two other cities where fermentation
research was being conducted in close contact with industry were
Copenhagen and Chicago. Denmark, indeed, would carve out a
reputation as home to the most scientifically shaped agriculture in
the world. The Copenhagen Polytechnic School employed Sigurd
Orla Jensen, who had spent several years as director of the Central
Institute of Cheese, perfecting the production of holes, as professor
of the physiology of fermentation and agricultural chemistry. At
Copenhagen, the Carlsberg Brewery established an institute under
E. Hansen in 1876, while, at Chicago, John Ewald Siebel built up a
brewing school, the Zymotechnical College.:" This zymotechnology
would eventually be renamed 'biotechnology'.? Indeed, Orla Jensen's
chair was renamed professor of biotechnical chemistry in 1913.48 The
working world context here was the manufacture of beer and cheese,
and the science was the first 'biotechnology'.
Finally, interconnections between the working world of agricul-
ture and the German universities had provided the context for the
61
 SCIENCE AFTER 1900

development of a new botany - laboratory-based plant physiology.

Two major research schools had grown up under Julius Sachs at
the University of Würzberg and Anton de Bary at the university at
Strasbourg. The intersection of practical agricultural research and
academic botany lay in interest in the 'nutritional requirements of
crop plants [which had] provided a strong impetus to the devel-
opment of plant physiology'r'? Sachs, for example, had taught in
Germany's agricultural colleges before setting up his Würzberg
laboratory. De Bary's students, the Russian Sergei Winogradsky and
the Dutchman Martinus Willem Beijerinck, who had als o taught
at an agricultural school, explored the wild occurrences of rnicro-
organisms, identifying new types of microbes in soils and plants,
respectively. In 1898, Beijerinck named the tobacco mosaic disease-
causing substance which passed through the fine filters that removed
known rnicro-organisrns a 'virus'. Winogradsky's techniques, on the
other hand, illustrate how scientists related to the working world of
agriculture: by isolating microbial components within the soil, or
working with simplified surrogates of the natural world (the common
pedagogical tool for illustrating bacterial ecology, the Winogradsky
column, for example), he was able to refine and describe processes of
interest to academic microbiology (chemoautotrophy, for example)
while generating findings, for instance the identification of iron and
sulphur bacteria, of direct interest to, and taken up by, the working
world of agriculture.t? The new life sciences, with new laboratories,
entities ('virus', 'gene') and theories, were intimately connected to
working worlds.