Studies in History and Philosophy of Biological and Biomedical Sciences 59 (2016) 145e153

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Studies in History and Philosophy of Biological and

Biomedical Sciences
journal homepage: www.elsevier.com/locate/shpsc

Writing the history of virology in the twentieth century: Discovery,

disciplines, and conceptual change
Pierre-Olivier Méthot a, b, *
a
Faculté de philosophie, Université Laval, 2325 rue des Bibliothèques, Québec, Québec G1R 1V7, Canada
b
Centre interuniversitaire de recherche sur la science et la technologie, Université du Québec à Montréal, C.P. 8888, succ., Centre-ville, Montréal, Québec
H3C 3P8, Canada

a r t i c l e i n f o a b s t r a c t

Article history: Concerned with the study of viruses and the diseases they cause, virology is now a well-established
Received 21 January 2016 scientific discipline. Whereas aspects of its history from the late nineteenth to the mid-twentieth cen-
Accepted 27 February 2016 tury have often been recounted through a number of detailed case studies, few general discussions of the
Available online 29 March 2016
historiography of virology have been offered. Looking at the ways in which the history of virology has
been told, this article examines a number of debates among scientists and historians of biology and show
Keywords:
how they are based on a different understanding of notions such as “discipline”, of processes such as
Historiography
“scientific discovery” as well as on distinct views about what the history of science is and how it should
Virus concept
Experimental practices
be written (the opposition between “longue durée” and “micro-history” or between history of “concepts”
Discipline formation versus “experimental methods”). The analysis provided here also suggests that a richer historiography of
Tobacco mosaic virus virology will require looking at the variations over time of the relations between conceptual, techno-
Discovery and scientific change logical, and institutional factors that fostered its development at the intersection of several other sci-
entific fields in the life sciences.
Ó 2016 Elsevier Ltd. All rights reserved.

When citing this paper, please use the full journal title Studies in History and Philosophy of Biological and Biomedical Sciences

1. Introduction: from the history to the historiography of
virology
Virology is concerned with the study of viruses and the several
diseases they cause as well as with the prevention of disease
transmission within and among populations. Now a well-
established discipline, the historical trajectory of virology from
the late nineteenth century to the present has often been recounted
(Burnet, 1955; Chastel, 1992; Creager, 2001; Galperin, 1994; Grafe,
1991; Grmek, 1994; van Helvoort, 1991, 1992, 1993, 1994a, 1994b,
1996; Smith Hughes, 1977; Sankaran, 2006; Scholthof, 2014;
Summers, 2014; Waterson & Wilkinson, 1978). However, whereas
a number of detailed case studies bearing on aspects of the history
of virology have been written, very few general discussions of the
* Faculté de philosophie, Université Laval, 2325 rue des Bibliothèques, Québec,
Québec G1R 1V7, Canada.
E-mail address: p.olivier.methot@gmail.com.
historiography have been offered. As a first step in this direction, this
article is thus primarily concerned with historiographical questions.
Looking at the ways in which the history of virology has been
told, this article discusses a number of points of contention among
scientists and historians of science and biology that have been
neglected in the literature, on the one hand, and examines how
they are based on a different understanding of notions such as
“discipline”, of processes such as “scientific discovery” as well as on
distinct views about what the history of science is and how it
should be written (the opposition between “longue durée” and
“micro-history” or between history of “concepts” versus “experi-
mental methods”), on the other.
Examining the historiography of virology can shed light on the
evolution of methodological approaches among historians of sci-
ence, on the more general problems they face, and inform us about
past and present diverging interpretations they proposed con-
cerning the significance of key episodes in the early days of
virology. For instance, one shall see that historical accounts of the
development of virology as a field from the 1970s to the 1990s were

http://dx.doi.org/10.1016/j.shpsc.2016.02.011
1369-8486/Ó 2016 Elsevier Ltd. All rights reserved.
146 P.-O. Méthot / Studies in History and Philosophy of Biological and Biomedical Sciences 59 (2016) 145e153
often wide-ranging in scope and organized around the emergence
of the concept of virus, from Pasteur until the rise of molecular
biology; in contrast, more recent accounts have embraced the shift
toward ‘micro-history’ and stressed the role of experimental prac-
tices (Creager, 2001; Scholthof, 2014; Summers, 2014) and, to some
extent, institutions (Kevles & Geison, 1995) in shaping the field.
After introducing aspects of contemporary historiography of
virology (section 2) and the early works on tobacco mosaic disease
in order to outline the historiographical issues to be examined later
(section 3), this essay revisits three important points of discussions
in the history of virology, namely: the discovery of viruses in the
late nineteenth century, as debated by scientists (section 4), the
beginnings of virology as a discipline (section 5), and the nature of
conceptual change in virology during the past century, as mostly
discussed by historians of science and biology (section 6). In
conclusion, I suggest that one way to provide a richer historiogra-
phy of virology would be to look at the variations over time of the
relations between conceptual, technological, and institutional
factors.
2. Writing the history of virology
The history of virology has often been written as the coming into
being of a discipline, namely as a process going from the study of ill-
defined biological or chemical objects called ‘viruses’ in the late
nineteenth century to the experimental and professionalized
laboratory-based science known as virology some seventy years
later. The creation of scientific disciplines offers historians of sci-
ence a privileged way to access and explore the multiplicity of
factors (conceptual, technical, cultural, etc.) involved in the growth
of science, which has largely developed by means of a “continual
branching out into fields of investigation previously unexplored
and often totally unexpected” (Lemaine, Macleod, Mulkay, &
Weingart, 1976, 2). The organized study and control of biological
entities known as ‘viruses’ is often seen as one such example of a
domain that has ‘branched out’ from previously established fields
of inquiry e plant pathology, bacteriology, chemistry, etc. e before
achieving disciplinary status, a process once described as “a model
of the emergence and establishment of a new branch of biological
knowledge” (Waterson & Wilkinson, 1978, ix).
Virology’s emergence as a discipline, of course, does not merely
result from the discovery of new forms of biological entities,
standing between the living and inert matter; it is also the product
of new techniques and apparatuses being used to advance knowl-
edge, notably in the medical and biomedical realms, as well as a
consequence of scientific infrastructures and social practices,
including the foundation of scientific journals and the development
of research communities and institutions such as the Rockefeller
Institute for Medical Research in New York, the California Institute
of Technology, the Walter and Eliza Hall Institute of Medical
Research in Melbourne, the Pasteur Institute in Paris, and the
Friedrich Loeffler Institute in Greifswald.
Despite the risk of taking for granted established knowledge-
structures (e.g., disciplines) instead of historicizing them
(Jordanova, 1993), the ‘disciplinary framework’ remains useful for
following the development and organization of scientific knowl-
edge. This is the case notably because of the disciplinary frame-
work’s social role: disciplines are important ‘institutional
mechanisms’ that regulate the circulation and production of
knowledge while they also help to ground scientific expertise. On a
broader level, they are also essential to the functioning of what Tim
Lenoir called the global ‘political economy’ of modern science
(Lenoir, 1997, 72).
Coming now to the historiography of virology itself, Daniel
Kevles and Gerald Geison have observed twenty years ago that this
literature has “tended to emphasize conceptual developments”
with little attention to “features of the research environment” that
allowed fundamental advances to be made (1995, 109). Indeed even
a cursory look shows that most accounts of the history of virology
mentioned in section 1 closely follow the development of the
concept of virus across time and place and focus on its role in
shaping virology as a distinct field of research and practice. In her
study of Wendell Stanley’s work on the tobacco mosaic virus, his-
torian of science Angela Creager noted this methodological point:
“the origin of virology”, she remarked, “is most often narrated as
the history of a concept, that of the virus as a nonbacterial path-
ogen” (2001, 45; emphasis added).
Creager’s comment was directed to Sally Smith Hughes’ earlier
study, The Virus: a History of the Concept, in which the historical
emergence of virology was characterized in terms of a shift from
bacterial to nonbacterial concepts of virus; but several other his-
torical accounts of the long history of virology are equally concept-
oriented. In 1978, London virologist P. A. Waterson wrote an
Introduction to the History of Virology together with historian Lise
Wilkinson that, he acknowledged, was “essentially, and indeed
deliberately, conceptual” (1978, xii; emphasis in original), a book
later perceived to be both “comprehensive and authoritative” by
historians of virology (van Helvoort, 2014, 258). Although Waterson
recognized the close connections between concepts and techniques
as well as between institutions and persons, his account focused
explicitly on “the evolution of the present concept of a virus” (1978,
xii). Around the same time, in a series of articles appearing from the
mid- to late-1970s in Medical History, Wilkinson pursued her
studies of “the development of the virus concept as reflected in
corpora of studies on individual pathogens” (1974, 1975, 1976, 1977,
1979; emphasis added).
A decade later, scientist and historian of science Ton van Hel-
voort argued in a number of foundational papers (1991, 1992, 1993,
1994a, b, 1996) that virology as a field emerged out of the ‘decon-
struction’ of the concept of ‘filterable virus’ and the subsequent rise
of the ‘modern’ concept of virus from the 1950s onward (1994a,
186). While these works do follow the development of techniques
such as filtration, cell and tissue cultures, electron microscopy, etc.,
what explains broad, even paradigmatic changes in the field of
virology are usually changes in the concept of virus itself (e.g. bac-
terial to non-bacterial, filterable virus to the modern concept of a
virus, etc.) In other words, concepts e and not so much techniques,
institutions, research environments, or their interaction e are seen
as the main drivers of scientific change.
Historians of science often single-out the role of concepts in
establishing disciplinary status because they are the “most con-
spicuous ‘tools of cognition and communication’” (Lenoir, 1997,
cited in Suárez-Díaz, 2009, 44). For instance, in van Helvoort’s
words the ‘plasticity’ of the virus concept “offered opportunities for
communication between members of thought collectives of
competing approaches” (1994a, 221). Yet as William Summers
recently pointed out, “The refinement or precision of the virus
concept is a story of technologies and methods” (2014, 26; emphasis
added). Following the so-called ‘practice-turn’ in the history and
philosophy of science, fueled by ‘laboratory studies’ and ethno-
methodological approaches, the role of experimentation in estab-
lishing knowledge-claims has gained prominence among historians
who paid particular attention to specific episodes and instruments,
located within a given scientific institution such as the Rockefeller
Foundation, rather than to the history of virology as a whole (see
Gaudillière, 2000; Rheinberger, 2010). Beginning in the mid-1990s,
this trend in ‘microstudies’ is perhaps best reflected in Creager
(2001) account of Wendell Stanley’s laboratory and and his works
on the purification and crystallization of tobacco mosaic virus
(TMV). There, the author construed viruses such as tobacco mosaic
 P.-O. Méthot / Studies in History and Philosophy of Biological and Biomedical Sciences 59 (2016) 145e153
viruses and bacteriophages as ‘model systems’, and emphasized the
role of material culture and experimental practices “as the leading
edge of scientific change” (2001, 7).
The history of virology has been intensely researched by
scholars, as the many books and articles mentioned in the first
section underline. One could even say that the history of virology
enjoyed momentum in the early 1990s. This was due to several
factors, including the growing interest in the history of molecular
biology in which viruses (especially TMV) were a key model or-
ganism, but also to the social and political consequences of the AIDS
pandemic and the international spur of ‘emerging viruses’. Amer-
ican architects of the concept of emerging disease Stephen Morse
and Joshua Lederberg, for instance, organized a conference in
Washington in May 1989 on ‘Emerging Viruses: The Evolution of
Viruses and Viral Diseases’ that attracted much scientific, political,
and media attention (see Méthot & Fantini, 2014).
In order to take stock on the present and past situation, French-
Croatian medical historian Mirko Grmek, together with the Amer-
ican Nobel Prize laureate in physiology or medicine Baruch S.
Blumberg, organized the bi-annual Ischia summer school in June
1990, one year after the Washington meeting, on the ‘History of
Virology: Biology of Viruses and Epidemiology and Clinical Viral
Diseases’. A few years later, in an extensive ‘essay review’ of ten
books on viruses, Grmek (1994) delineated five stages in the for-
mation of virology as a field e a longue durée history of the disci-
pline. Reaching back to antiquity, he first situated the ‘prehistory’ of
virology at a time when its object was ill defined and had yet to
become a part of ‘scientific’ knowledge. This ‘proto-historical
period’, in turn, led to the attribution of the word ‘virus’ to any
putative pathogenic germ e a period that ended only in the late
nineteenth century. Following that, Grmek went on to define what
he called the ‘heroic’ period that covered the discovery of ‘filterable
agents’ (i.e. entities able to pass through porcelain filters) in the last
decade of the nineteenth century up until works on crystallography
and the use of electronic microscopes in the 1930s. The fourth
phase in the recent history of virology e labeled ‘constitutive’e is
shorter and spanned a decade at most. It was concerned with the
growing recognition that viruses form a ‘particular class of bio-
logical objects’ although their nature (chemical or living entities)
was still in dispute. The ‘actual’ stage in the history of virology,
according to Grmek, began in 1957 when French molecular biolo-
gist André Lwoff famously stated: ‘viruses are viruses’ (i.e., not
bacteria). Following the development of these five stages, Grmek
concluded, “virology becomes an autonomous scientific discipline,
located at the crossroad between genetics, cytology, bacteriology,
and oncology” (1994, 341).
Despite virology having achieved the status of an autonomous
science, historians and scientists have continued to debate many
issues related to the origins and evolution of this discipline,
namely: who ‘discovered’ viruses? When exactly did virology
become an autonomous discipline? And was the development of
virology during the twentieth century continuous or discontin-
uous? Before exploring those questions, I turn to earlier works on
viruses in the late nineteenth and early twentieth century that, in
large part, set the stage for the present historiographical debates.
3. Tobacco mosaic virus (TMV): from microbes to ‘filterable
viruses’
The word ‘virus’ in Latin means ‘poison’ (Smith Hughes, 1977). In
the context of the rise of medical bacteriology in Europe during the
1880s, however, the term virus acquired two distinct meanings, one
more specific and the other more general. The more specific defi-
nition of a virus is linked to vaccine. In Paris, Louis Pasteur spoke for
instance of a ‘vaccine virus’ (virus-vaccin) to describe some form of
147
immunizing procedure inspired by Jenner’s earlier practice
(Moulin, 1992). The more general meaning of the term, then used in
the context of the nascent ‘germ theory’ to designate infectious
agents (bacteria, viruses, parasites), emerged when Pasteur asser-
ted that: “every virus is a microbe” (Pasteur, 1890, 673, quoted in
Smith Hughes, 1977, 112; see also van Helvoort, 1994a).
In this latter sense, the concept of ‘virus’ was operationalized
thanks to a procedure consisting in a series of methodological steps
known as ‘Koch’s postulates’, which were considered applicable to
all microscopic agents. First spelled out by Robert Koch’s younger
colleague Friederich Loeffler in Berlin, those postulates rapidly
became the cornerstone of the germ theory of disease. Whereas
this method led to the identification of several pathogenic agents
such as the tubercle bacillus and anthrax, its application to other
disease entities also revealed a number of ‘anomalies’. Those were
first observed in the case of the tobacco mosaic disease, which
could not be cultivated in vitro, before being noticed in several other
(viral) diseases such as yellow fever and influenza (Gradmann,
2014).
The most popular example of a plant virus disease is the ‘mosaic’
pattern observed on the leaves of tulips, first described in 1576 by
Dutch physician and botanist Carolus Clusius (1526e1609). Un-
known for centuries, the cause of this disease began to be better
understood only in the late nineteenth century, thanks to the
German industrial chemist Adolph Mayer who had been working
on disease in tobacco plants. The director of an Experimental Sta-
tion in the region of Wageningen in the Netherlands, Mayer’s in-
terest in tobacco mosaic disease (TMV) (or ‘spot disease’) was
sparked by the impediments the disease was causing to the tobacco
industry, which in some places have “caused the cultivation of to-
bacco to be given up entirely” (cited in Zaitlin, 1998).
Using the sap obtained from crushing the leaves of diseased
plants, Mayer showed he could infect healthy ones and generate
the same disease pattern. Yet the nature of the infectious agent or
substance remained unclear. Mayer hypothesized it could be an
enzyme but settled for a bacterium in the end. Drawing on Koch’s
postulates, he attempted e and failed e to cultivate the infectious
substance of the mosaic disease from the extracted juice in the
laboratory. Experimenting with double filter paper, Mayer obtained
a clear filtrate of the sap of diseased plants that lost its infective
power, and thus concluded that he was “concerned with a bacterial
disease” (Mayer in Hahon, 1964, 44 [1886]).
At Mayer’s request, the Dutch scientist Martinus Beijerinck
attempted in turn to isolate the agent responsible for the tobacco
mosaic disease using standard bacteriological procedures, which
also proved unsuccessful. In 1895, after taking up a professorship in
bacteriology at the Polytechnical School in Delft, Beijerinck
resumed his work on tobacco mosaic disease in his new laboratory.
Using porcelain Chamberland filter-candles, an apparatus initially
developed to obtain clean water (Fig. 1) and to clarify chemical
solutions, he demonstrated that the juice of diseased plants
remained infectious after filtration. Performing a ‘diffusion exper-
iment’, he showed that a drop of the sap from infected plants would
diffuse slowly into a thin layer of agar gel, a finding that convinced
him of the soluble (and non corpuscular) nature of the infectious
agent. Observing that the material did not lose its infectivity over
time, Beijerinck concluded that “the spot disease is an infectious
one that is not caused by microbes” but by an active and soluble
substance: a ‘virus’ or as he called it, a contagium vivum fluidum
(Beijerinck in Hahon, 1964, 54 [1898]).
According to most historical accounts, “Beijerinck’s ideas” of a
soluble and contagious agent “met with strong opposition and were
not readily accepted” (van Kammen, 1999, 4; see also Waterson &
Wilkinson, 1978; Smith Hughes, 1977). Shortly after Beijerinck’s
paper was published in 1898, a priority dispute flared as Russian
148 P.-O. Méthot / Studies in History and Philosophy of Biological and Biomedical Sciences 59 (2016) 145e153

and theoretical debates about the beginnings of virology and the

discovery of the nature of viruses.1 It is against this historical
background that a number of historiographical issues have arisen.

4. Who ‘discovered’ viruses?

As the centenary commemoration of the discovery of filterable

viruses was approaching in the early 1990s, scientists began
assessing the beginnings of their own field, which meant reflecting
on founding figures as well as on significant historical episodes.
Often, inquiries regarding the origins of virology were framed in
terms of ‘who first discovered viruses?’ e an apparently simple
question that leads to the identification of different beginnings of
virology as a science, and thus raises issues in the historiography of
virology itself. For example, did virology start in 1886 e thus
commemorating Mayer’s publication on the transmission of mosaic
tobacco disease? Or in 1892 e thus taking the work of Ivanovsky on
Fig. 1. A Pasteur-Chamberland filter. First designed to produce clean water, it was later
used in the context of bacteriological research, notably by Ivanovsky and Beijerinck. the filterability of the tobacco mosaic virus as a starting point? Or in
Source: Courtesy of the National Library of Medicine. 1898 e thus celebrating the publication of the paper by Beijerinck
on the concept of a contagium vivum fluidum?
In the 1990s, two American scientists opted to consider Iva-
bacteriologist Dmitrii Ivanovsky claimed he had already discovered
novsky’s work as genuinely foundational whereas others consid-
that the tobacco mosaic disease could pass through the porcelain
ered that the beginnings of virology were to be found in the
filter and that its virulence was maintained through serial in-
introduction of Beijerinck’s novel concept of contagium vivum flu-
oculations, a point Beijerinck willingly acknowledged. As it turned
idum (itself based on Mayer’s previous research). Recalling the
out, Ivanovsky had read a paper at the Academy of Sciences in St-
words of Wendell Stanley, for whom “there is considerable justifi-
Petersburg in 1892 where he presented experimental results that
cation for regarding Ivanovksy as the father of the new science of
contradicted Mayer’s earlier observation of a gradual loss of infec-
virology”, virologists Alice Lustig and Levine (1992) at Princeton
tivity in the filtrate. Using porcelain Chamberland filter-candles,
have argued that Ivanovksy indeed “fathered the field of virology”
Ivanovsky demonstrated that the filtrate remains infective despite
e and determined the foundation of the field in 1892 (1992, 4631).
the filtration process, although his microscopic analyses did not
This claim prompted a letter of protest from the Dutch virologist
reveal any microorganism and he was equally unable to grow the
Lute Bos (1995a), who considered that the use of the term ‘virus’ by
causative agent in pure culture (Lechevalier, 1972).
Beijerinck to designate “a new class of pathogens” is what truly
To Ivanovsky’s mind, Beijerinck’s concept of a contagium vivum
“marks the beginning of a new era in biology” (1995b, 618). Bos
fluidum was definitely a bacterial concept, as “he had been able to
criticized and ultimately rejected Lustig and Levine’s statement
transmit the disease via bacterial colonies” (van Helvoort, 1991,
because, he claimed, Ivanovsky “did not grasp the scope of his
561). Beijerinck’s diffusion experiment, Ivanovsky thought, did not
observations” (1995b, 614). Beijerinck’s 1898 paper, for him,
prove the soluble nature of the agent but on the contrary, “indicates
“marked the beginning of a far-reaching change in thinking in
that we are dealing with a contagium fixum” (Ivanovski, 1901, cited
microbiology” (Bos, 2000, 82).
in Lechevalier, 1972, 141; italics in original). Ivanovsky, still, enter-
In a more conciliatory tone, Marian Horzinek, from the Veteri-
tained the possibility that the agent might be a soluble toxin, since
nary Faculty at the University of Utrecht in the Netherlands,
Émile Roux and Alexandre Yersin at the Pasteur Institute in Paris
considered that even if Beijerinck did not act alone in fostering the
had only recently demonstrated such a toxin to be the cause of
development of virology he is nevertheless the one who accom-
diphtheria (Lechevalier, 1972, 140). More fundamentally, Beijer-
plished the “conceptual quantum leap” in discovering a new
inck’s concept was perceived as problematic because it contra-
kingdom of infectious agents, and he thereby “gave birth to a new
dicted the Pastorian framework in which every virus is a living
discipline” (Horzinek, 1997, 15&18). In German-speaking journals,
microorganism, and should thus be particulate (i.e. ‘organized’), not
in contrast, Friedrich Loeffler, Koch’s colleague, is typically credited
fluid and soluble (van Helvoort, 1991, 560).
as the ‘father of virology’ for his work on foot-and-mouth disease
In the late nineteenth century, the filtration apparatus used by
conducted with Frosch (Schmiedebach, 1999, 9). Despite those
Beijerinck and Ivanovski became a ‘defining technological criterion’
disagreements, a number of scientists celebrated ‘100 years
for the identification of contagious diseases attributed to ‘filterable
of virology’ in 1998 (Calisher and Horzinek, 1999; see also Bos,
viruses’ such as foot-and-mouth disease and fowl plague, in
2000).
contrast to toxins that could pass through the candles but were not
Looking back on this debate, it appears that singling out one
infectious (Summers, 2014, 28). Although they had remained
particular moment in the discovery process is problematic for, as
virtually invisible to scientists using light microscopes and were
Ludwik Fleck and more recently Dominique Pestre have argued, a
impossible to grow in pure culture, ‘filterable viruses’ could be
discovery “never appears full-blown”, but is rather “put together
detected thanks to this technology. At the turn of the past century,
contradictorily and retrospectively by the actors” (Fleck, 1979
filterable viruses were thus operationally defined by three (nega-
[1935]; Pestre, 1999, 205). Not unlike the present case, Pestre
tive) features: they “were not retained by bacteriological filters;
observed further that scientists often become “trapped in endless
they could not been seen in the light microscope; and they could
confrontations” when attempting to arbitrate priority disputes and
not be grown on artificial media” (Waterson & Wilkinson, 1978, 78).
The debate over the nature of the contagium vivum fluidum,
nevertheless, set the groundwork for lasting divergent in- 1
On the issue of whether viruses are alive or not, see the contributions of
terpretations of viruses as being alive or as being mere chemical Forterre, Koonin and Starokadomskyy, van Regenmortel as well as Kostyrka in this
products and engaged scientists on productive paths of research issue.
 P.-O. Méthot / Studies in History and Philosophy of Biological and Biomedical Sciences 59 (2016) 145e153 149

“to establish for posterity what the most important discoveries

were and by whom they were made” (1999, 204e205).
The main problem with “founder narratives” of scientific disci-
plines, such as the one proposed by virologists in the 1990s, is not
simply “the complexity of the task of building disciplines”, as Pierre
Bourdieu once pointed out; “the problem is that disciplines do not
have a single originary source, but are more appropriately grasped
as interactive systems” (Bourdieu, 1988, 52, cited in Suárez-Díaz,
2009, 51). The only way to characterize the early days of virology
in terms of founding narratives would be to consider the network of
concepts, techniques, and institutions within which Mayer, Iva-
novsky, and Beijerinck interacted e that is, to look at their contri-
butions together, not in isolation. Scientific discoveries are rarely (if
ever) the result of a single brilliant individual acting on his own e a
claim Grmek denounced as “the myth of the unity of the discov-
erer” (1981, 19). Rather, as Fleck, Kuhn, and others argued, they are
generally due to several scientists working over different periods of
time and across different places (Fleck, 1979 [1935]; Kuhn, 1962).
In the present case, deciding who discovered viruses is even
more difficult because none of the protagonists involved in the
early days of virology research (Mayer, Ivanovsky, Beijerinck) had a Fig. 2. An image of Wendell Stanley’s crystallization of the tobacco mosaic virus (TMV)
using the electron microscope. Source: Work in the public domain.
complete understanding of the nature of viruses. In fact, as Lwoff
(1957) pointed out, the status of viruses remained unclear until
the 1950s as they were considered to be either microorganisms or viruses but for life itself, as its simplest incarnation” (Creager, 2001,
chemical agents. And although Mayer, Ivanovsky, and Beijerinck 2). Standing on the border of life and non-life, viruses e long
suspected that ‘filterable viruses’ differed in some ways from the regarded as “naked genes” (see Sapp, 1994, 151) e were progres-
agents causing fungal diseases e then the most common etiology sively turned into more general models to study the properties of
for plant disease e this fell short of accepting the possibility of a living systems.
new kind of disease agents called ‘viruses’. Indeed, it should be
remembered that ‘viruses’ could not be seen, either by the naked
5. When did virology become a ‘discipline’?
eye or with a light microscope. The passing of tobacco mosaic vi-
ruses through the porcelain filters did not in and of itself establish
Another debated aspect of the modern history of virology con-
the claim for the existence of a new kingdom of life forms; viruses
cerns its historical beginnings as a scientific discipline in the
might have just been small bacteria that could be retained in
twentieth century, not just from a conceptual but also from an
principle by filters with more minute pores. “Their small size
institutional point of view. As Ton van Helvoort (1996) has recalled,
alone”, as US virologist Thomas Rivers argued, “should not make
Nobel laureate immunologist and virologist Frank Macfarlane
them insusceptible to cultivation” (quoted in van Helvoort, 1994a,
Burnet believed that prior to the late 1950s virology did not exist as
b, 197).
an independent field of research. In fact, Burnet recalled that when
As mentioned above, the claim that viruses are distinct from
he began his career in the 1920s, virology was not yet an inde-
bacteria began to gain a foothold when several animal, plant, and
pendent science (see Burnet 1953a,b). “Virology - there was no such
human diseases failed to conform to Koch’s postulates. Frederic
thing. There were filtrable viruses as an appendix to end of a bac-
Twort and Félix d’Hérelle’s work brought additional support to this
terial textbook”, he said to Sexton, his biographer, in a 1985 inter-
view when they discovered curious entities called ‘bacteriophages’
view (quoted in Sankaran 2006, p. 52). Burnet also did not use the
able to prey on bacteria, but whose identity was still unclear
word ‘virology’ e and much less called himself a ‘virologist’ e in his
(Summers, 2005; on phage research see Billiau ‘online first’;
influential book Virus as Organism (1946). However, the use of the
Galperin, 1987; Sankaran, 2006). “The idea that they [viruses] made
term grew rapidly from the beginning of the 1950s to the late
up a class of microbes in their own right”, Christoph Gradmann
1960s, just a few years after Burnet’s book was published.2 Ac-
pointed out, “evolved only slowly over the first three decades of the
cording to the Australian immunologist, the field turned into a
twentieth century” (2014, 4). Up until then, the concept of a virus as
cohesive discipline when scientists started to create virology jour-
Beijerinck defined it, remained virtually “unintelligible to the then
nals (e.g. Virology, Journal of General Virology), to write scientific
current paradigms in microbiology” (van Loon, 2002, 108; van
textbooks on the biology of viruses e efforts to which Burnet
Helvoort, 1994a).
himself contributed e and when they proposed a unifying
New knowledge about viruses was also made possible thanks to
conception of virus understood as genetic and protein-based en-
a number of technical advances in the first third of the twentieth
tities unable to grow by binary fission.3 Key textbooks of general
century such as the development of the electron microscope that
virology, and not only on plant virus diseases, including Burnet’s
allowed scientists to visualize the rod-shaped form of the tobacco
mosaic virus, a technology which had a broader transformative
effect on biology (Rasmussen, 1997). The construction of ultracen- 2
A research in Google Ngram Viewer illustrates a steep rise in the use of the term
trifuges and the development of X-Ray crystallography also ‘virologist’ from the 1950s up to the late 1960s. The same is true for the French
fostered the acquisition of more detailed knowledge of the minute word ‘virologue’. I thank Yves Gingras for this observation. Click here for the
structure of the viral particles. Here, the tobacco mosaic virus graphic curve: https://books.google.com/ngrams/graph?content¼virologist&year_
start¼1800&year_end¼2000&corpus¼15&smoothing¼3&share¼&direct_url¼t1%
played a crucial role as a model, for it was the organism Wendell
3B%2Cvirologist%3B%2Cc0.
Stanley selected to crystallize those viral particles (Fig. 2), which he 3
Lwoff (1957) introduced this molecular definition of the virus in a paper of
then showed to remain infectious when placed in soluble solution, historical significance that remains frequently cited in today’s textbooks (Morange,
and thus contributed to make TMV a model “not only for other 2005).
150 P.-O. Méthot / Studies in History and Philosophy of Biological and Biomedical Sciences 59 (2016) 145e153
own Principles of Animal Virology (1955) and his comprehensive
three volumes edited together with Wendell Stanley on The Viruses
shifted the focus away from viruses as mere “agents of disease” and
encouraged scientists to look at them as “biological entities whose
properties can be studied in the laboratory” (Burnet & Stanley, 1959,
1).
In The Virus: a History of the Concept, Sally Smith Hughes also
suggested that virology emerged as an independent science
“around the midpoint of the twentieth century” because: (1) vi-
ruses were now distinguished from other kinds of microorganisms;
(2) distinct techniques for studying them had been developed; (3)
researchers working on viruses began calling themselves ‘virolo-
gists’ instead of parasitologists or microbiologists; (4) the nature of
viruses, and not only the diseases they cause, became an object of
study; (5) the first journals of virology in English language were
launched around that time; and (6) specific institutions were
created to further research on viruses (1977, 104e105).
In the first three decades of the twentieth century however, a
similar ‘burst’ of scholarly activities emerged, starting with the 1928
volume edited by Rockefeller researcher Thomas Rivers, Filterable
Viruses as well as with the foundation of the Archives of Virology in
Vienna in 1939 (van Helvoort, 1996, 142). Furthermore, Rivers also
offered a new definition of viruses as “obligate parasites in the sense
that their reproduction is dependent on living cells” (Rivers, 1927,
228). It was even “Rivers’ stubborn insistence on this fundamental
point of difference between viruses and bacteria”, his biographer
argued, that “did much to establish virology as a separate discipline”
(Horsfall,1965, 270e271). Did virology exist or not prior to the 1950s?
This discrepancy between the interpretation of the pre- and
post-1950s areas of virology research can serve to illustrate that
tracing the beginnings of a scientific discipline often depends on
the criteria used to ascertain what ‘disciplinarity’ means. “Whether
something is, or is not a discipline is in every epoch”, Frederick L.
Holmes emphasized, indeed “partly a question of fact, and partly an
object of controversy” (2004, 167). That is, whether a scientific field
is seen as a discipline (or not) requires setting boundaries between
what disciplines are taken to be (in contrast to other areas of
knowledge such as ‘schools’, ‘paradigms’, ‘research traditions’,
‘styles of reasoning’, or ‘ways of knowing’).
It is usually agreed upon that to be recognized as a full-fledged
discipline a scientific sector must fulfill at least two requirements,
one cognitive and another institutional: first there must be a
consensus on the relevant problems to be explored, with what
methods and according to what norms of validation for knowledge
claims (see Burnet and Stanley who defined the ‘problems of
virology’, 1959, 1); second, for a sector of research to exist as a
discipline, there must be institutional structures that allow its
recognition as well as its persistence through time. Education
programs, university degrees, and scientific journals exemplify this
second criterion (Gayon, 2004, 153).
Looking at Sally Smith Hughes’ criteria, one sees that post-1950s
virology research satisfied both the cognitive and the institutional
requirements. On this account, the pre-1950s period, in contrast,
was not part of virology because criteria 1, 3, 4, and 5 were not met,
that is, viruses remained within the category of bacteria; virology
had not yet ‘branched off’ from the bacteriological paradigm; the
study of viruses was pursued in medical research, veterinary
medicine, and botany but viruses were not studied for themselves;
and finally, no international journal devoted to the understanding
of viruses as biological entities existed.
Yet one might question the choice of those criteria. Indeed by
standards such as academic journals (and not exclusively Anglo-
phone ones), concepts, and techniques, some strands of research
conducted in the first half of the twentieth century (especially
d’Hérelle and Rivers’ works) could very well fall within the fold of
what is today known as ‘virology’. The upshot here is that the
coming into being of a scientific discipline strongly depends on
criteria chosen to define what a discipline is. However, there is no
general agreement concerning what these criteria should be
(except, perhaps, that there should be an institutional and a
cognitive component). A more interesting question, therefore, is
perhaps not whether ‘virology’ existed as a whole or not before and
after the 1950s taken as a benchmark, but which aspects of its
progressive ‘disciplinarisation’ were achieved prior to this moment.
According to sociologist and historian of science Yves Gingras,
three ‘steps’ can be identified in the formation of scientific disci-
plines: 1) the emergence of a new practice; 2) its institutionaliza-
tion, allowing for its reproduction and its systematic diffusion
within the scientific community; and 3) the shaping of a social
identity, which can take on many distinct forms, including the
professionalization of the field (Gingras, 1991, 43e44). This tri-
partite framework helps to clarify whether virology ‘existed’ e
and under what form e before the 1950s and to resolve some of the
aforementioned tensions. For instance, whereas the practice of
virology (step 1) certainly existed in the first third of the twentieth
century (i.e., the works of Rivers on filterable viruses, of Twort and
d’Hérelle on bacteriophage, on Stanley on tobacco mosaic disease),
and this practice was to some extent institutionalized (step 2), i.e., it
was embedded into university and non-university based research
centers in the United States as well as in Europe, virologists had yet
to acquire a distinct social identity that would set them apart from
bacteriologists, immunologists, and epidemiologists (step 3). As
mentioned previously, Burnet did not refer to himself as a virologist
in the mid- to late-1940s. The latter phase, the acquisition of a
definite social identity, like for most nascent disciplines, certainly
benefited from the recurrent use of a name e virology e that
testified of a well-defined community of research, for ‘naming’ is a
powerful tool in constructing a new field (see Powell, O’Malley,
Müller-Wille, Calvert & Dupré, 2007).
Historians have yet to trace more precisely the emergence of the
term ‘virologist’ and its growing use among scientists that coincides
with the organization of several conferences on the nature of vi-
ruses and virulence, notably by the influential CIBA Foundation in
London during the 1950s and 1960s (e.g. Wolstenholme & Millar.,
1957; Wolstenholme & O’Connor, 1960) but also with the organi-
zation, in the mid-1950s, of large-scale, global eradication cam-
paigns targeting viral diseases, particularly in tropical countries
(see Listios, 1997). In the next section I turn to the question of
whether such a process of ‘disciplinarisation’ occurred in a pro-
gressive or rather revolutionary manner in the history of virology.
6. The development of virology: continuity or revolution?
Whether science is characterized by the progressive growth of
knowledge or whether it is punctuated by sudden and sharp con-
ceptual breaks has polarized historians, philosophers, and sociol-
ogists of science for well over a century now (see Kragh, 1987).
Historians of biology often characterized changes in the history of
virology during the twentieth century in terms of continuity and
discontinuity e even though the application of Kuhn’s concepts of
‘paradigm’ and ‘normal science’ to the life sciences has been
questioned (Wilkins, 1996). This schema is especially present in Ton
van Helvoort, who used it to promote a discontinuous picture of the
history of virology in place of what he perceived to be a shared
continuist narrative among both scientific practitioners and histo-
rians of science. “The historiography of virus research in the
twentieth century”, as he put it, “is usually presented as a process of
continuity and as a smooth path of progress” (1994a, 187). A “rev-
olutionary upheaval”, he noted, “is not claimed to exist” (van
Helvoort, 1994a).
 P.-O. Méthot / Studies in History and Philosophy of Biological and Biomedical Sciences 59 (2016) 145e153
van Helvoort’s historiographical position, defended in several
papers, takes issue with a rather bold claim put forward by
Waterson, according to whom the modern history of virology is
“the story of the progressive unveiling of the nature of the virus
particle” (Waterson & Wilkinson, 1978, xii). Instead of this realist
reading, on which viruses form a ‘natural category’, for van Hel-
voort “the description of nature that is given in the natural sciences
is the result of the formation of consensus” (1994b, 95; emphasis in
original).
In several articles, van Helvoort argued against the character-
ization of virus studies provided by Waterson and rejected the
‘mistaken metaphor’ of “a smooth process of growing knowledge of
the virus particle” (van Helvoort, 1994a, 187; emphasis in original).
For, despite the regular appearance of the concept of ‘filterable vi-
ruses’ in textbooks and journals during the 1930s and 1940s, he
claimed, this concept “lacked clarity and certainty” (1996) and,
furthermore, the links between bacteriology and virology pre-
vented a neat separation between bacteria and viruses. For van
Helvoort “the 1950s marked the maturing of the virological para-
digm with which the study of viruses truly emancipated from
bacteriology” (1993, 21).
Waterson’s approach to the history of virology is also prob-
lematic, according to van Helvoort, because it “ignores the deep
controversies in virus research during the first half of the twentieth
century” that led to the formation of consensus and later to the
deconstruction of the filterable virus concept.4 “If we consider the
definition of viruses as filterable agents and the modern concept of
viruses as agents with an ‘eclipse’ phase”, van Helvoort argued,
“one can speak of two paradigms, to use Thomas Kuhn’s termi-
nology” (1996, 142; see also 1994a). In sum, for him “an abrupt
conceptual shift or scientific revolution took place in the 1950s”
(1996, 142).
Waterson’s remark about the ‘unveiling’ of the virus particle was
made only in passing, in the preface of his book and was not sup-
ported by an historical argument. A stronger contender for a con-
tinuist position is Yale historian of biology William Summers
(2014), who recently sketched a different account of the develop-
ment of virology. According to Summers, scientists tend to think of
viruses as a ‘natural kind’, that is, as an objective category of the
biological world scientists go about discovering empirically. By-
passing the philosophical debates on natural kinds, what he seeks
to get at is “how did the category ‘virus’ come to be recognized, and
what are its essential, defining qualities” (2014, 26). For this scholar,
while viruses are ‘natural objects’ our beliefs about them have
profoundly changed over time (2014, 26). “It can be said”, Summers
continues, “that virologists invent (and continually reinvent) the
concept of a virus as part of the normal progress of their science”
(Summers, 2014, 26). The history of virology Summers proposes is
not
a story of the triumphant accumulation of knowledge by the he-
roic scientists of the past so much as it is an examination of the
continual struggle to understand and organize observations that
challenged and made obsolete the comfortable certainties of the
often recent past (2014, 26; emphasis added).
While this part of Summers’ account finds an echo in van
Helvoort’s focus on scientific controversies, he also rejects Kuhn’s
epistemology and goes on to argue that the history of virology is
“a story of technologies and methods more than a story of paradigm
4
Those controversies include questions such as: is a virus an exogenous agent or
the endogenous product of host cells? Are bacteriophages viruses? Are viruses alive
and can they evolve by natural selection? (van Helvoort, 1992, 1994a, 1994b, 1996).
151
shifts, eureka moments, and discoveries” (Summers, 2014; emphasis
added). Departing from van Helvoort’s perspective, Summers’ ac-
count, rooted in the practice-turn in the history and philosophy of
science, focuses on how our understanding of the properties of
viruses has changed over time and space, “not in the sense of
replacement of former knowledge with current knowledge, but
rather in the sense of enlargement of our knowledge and increased
clarity in place of earlier innocence” (Summers, 2014.; emphasis
added). For him, therefore, there will be no ‘revolutionary
moment’ to single out around the mid-twentieth century that
purportedly turned virology into a full-fledged discipline.
Despite Summers’ claim to the contrary, it would be hard to
deny that virology developed its own paradigm in the mid-
twentieth century by breaking with bacteriology, along the lines
van Helvoort suggested. It was also around this time that the name
‘virology’ began to gain prominence and helped establish the social
identity of this research community, demarcating it from bacteri-
ologists or immunologists, among other fields. Burnet was an
exception as he did pioneering work in bacteriology, immunology,
and virology throughout his career. Back then, moreover, many of
the controversies van Helvoort has studied in detail were stabilized
and put to rest by the 1940s and 1950s, which speaks in favor of a
period of ‘normal science’ that followed a period of ‘revolution’.
Lastly, a number of research institutions and research groups
dedicated to the study of viruses flourished in Europe and the
United States (Creager, 2001; Rheinberger, 2010), not least the
Phage-group led by Max Delbrück, and other techniques such as
cell and tissue cultures were also developed, all of which helped
“making a virus visible” (Scholthof, 2014).
But can this process of ‘disciplinarisation’ really be called a
‘revolution’ e a paradigm-shifting moment e in the Kuhnian
sense? There was no former paradigm to overthrow in favor of a
new one. van Helvoort convincingly argues for the existence of a
‘bacteriological paradigm’ in virology research but the emergence
of virology in the 1950s did not displace it; rather, it com-
plemented it with both fields now occupying a space within the
larger realm of microbiology. What happened in the 1950s was
perhaps more the coming into being of a new disciplinary matrix
that consolidated and, as Summers argued, enlarged previously
existing lines of research (pursued elsewhere, notably in bacteri-
ology but also in chemistry, etc.) that had remained apart than a
paradigm-shift as such. At any rate, the bringing together of plant,
animal, and bacterial virology into a unified field contributed to
shape virology as a whole around a common and well-defined
object of biological research worthy of study for its own sake,
not just as agent of disease, and opened up a space to rethink
many questions in general biology, including issues in the classi-
fication and the ecology of viruses (see O’Malley, 2016). There are
thus at once elements of continuity and rupture in the recent his-
tory of virology, as probably in the development of most life sci-
ences: while breaking from existing fields of research, virology
emerged as a separate and also fundamental discipline by making
visible sub-microscopic entities that became, in turn, central to
understanding the properties of life.
7. Conclusion
This essay has been concerned with historiographical issues in
the history of virology, from the late nineteenth century until the
end of the past century. Two elements come into view at the end of
this inquiry. First, whereas histories of virology published between
the 1970s and the mid-1990s were mostly written from a concep-
tual point of view (Waterson and Wilkinson, Smith Hughes, van
Helvoort), more recent accounts (Creager, Summers) have stressed
152 P.-O. Méthot / Studies in History and Philosophy of Biological and Biomedical Sciences 59 (2016) 145e153
the role of scientific practice, methods, and instruments in gener-
ating knowledge and the formation of concepts.5
A second difference concerns the scale of this historical project.
Prior to the ‘practice-turn’, histories of virology often attempted to
provide a broad overview of the whole history of virology, going
from the late nineteenth century until the present. In contrast,
contemporary scholars have significantly departed from this ‘big
picture’ approach and have concentrated on detailed case studies
that provide access to otherwise invisible dynamics in the pro-
duction of scientific knowledge. In this recent history of virology,
Creager’s study of Wendell Stanley’s laboratory is a case in point.
van Helvoort’s work, however, sits between the two poles as his
account, scattered across several articles, extensively covers the
‘long history’ of virology but it does so by attending to specific cases
as well.
A more complete image of the coming into being of virology as a
discipline, nevertheless, will require greater attention to what
Kevles and Geison have called ‘goals, patronage, and institutions’
(1995) that would complement the attention to both concepts and
practices. Research centers, philanthropists, and research chairs are
crucial institutional factors to be considered in the development,
change, and stabilization of any scientific field. Combining the
depth of ‘micro-history’ and the breadth of ‘longue durée’ ap-
proaches in considering the variations over time of the relation-
ships between conceptual, methodological, and institutional
factors would provide us with a richer and more comprehensive
history of virology in the twentieth century (on the relation be-
tween ‘micro-history’ and ‘big picture’, see de Chadarevian, 2009).
The writing of the institutional history of virology was possibly
delayed because in contrast to molecular biology, which was often
taken to be a genuine turning point that led to the rapid flourishing
of many molecular biology institutes, about which an institutional
history could be told, it still was to be determined more precisely at
the time whether or not virology was a separate discipline (van
Helvoort, personal communication). At any rate, now that the tra-
jectories of several twentieth-century sciences are being explored
e bacteriology, immunology, molecular biology, etc. e we may be
better placed to recuperate the institutional dimension of the sci-
ence of virology and to situate it within biology.
This article also considered three related questions that gener-
ated debates among historians as well as among scientists, namely:
who discovered viruses? When did virology become a scientific
discipline? And what was the nature of the conceptual change
before and after the 1950s? As we have seen, the discovery of vi-
ruses could hardly be credited to a single individual, for all of the
three main protagonists, who worked in different places, used
similar bacteriological procedures but different concepts, shed
some light on the nature of the virus without being able to convince
the wider scientific community of the existence of a distinct bio-
logical category. The establishment of virology as a field, in turn, is a
question that cannot solely be answered empirically, for ascer-
taining ‘disciplinarity’ involves not only the widespread use of a
given term but also a decision about what counts as a discipline at a
given time. Thanks to the fast-changing period in virology research
in the post-1950s era, historians have proposed various ways of
characterizing this change, either in terms of continuity or in terms
of rupture. While continuity narratives have largely dominated the
writing of the history of virology in the 1970s, Ton van Helvoort has
argued that there should be room for the claim of a ‘revolutionary
upheaval’ in virology’s past, a claim implicitly contested by
5
For recent contributions to the history of virology, see the special section of
Studies in History and Philosophy of the Biological and Biomedical Sciences (48,
2014) entitled 100 Years of Cancer and Viruses, edited by Robin Wolfe Scheffler.
Summers’ more recent article. This point surely deserves more
historical attention than what was provided here and should be
connected to other recent ‘continuists’ accounts of key episodes in
the historiography of biology (e.g. the ‘Darwinian revolution’
(Hodge, 2005) and the ‘bacteriological revolution’ (Worboys,
2007)).
Advances in virology during the post-1950s have mostly been
studied from the perspective of molecular biology (see Morange,
1998), and issues raised in virology itself have more rarely been
given close attention during those following years, perhaps because
we are still too close to them. At the moment, the history of
virology, still based on the ‘modern’ concept of virus as described by
van Helvoort, continues to grow and includes a number of new
entities such as prions and viroids. Perhaps paradoxically, whereas
viruses are now studied as models to understand living phenomena
in general, their precise classification in the ‘tree of life’ next to the
evolutionary history of other biological entities remains a matter of
controversy and debate (O’Malley, 2014; see also Morgan, 2016). By
intersecting with issues in causation, classification, and definitions
of life viruses are prototypical candidates for future historical and
epistemological reflections on the life sciences and their
historiography.
Acknowledgments
I would like to thank Ann-Sophie Barwich, Yves Gingras, Gladys
Kostyrka, Michel Morange, Alex Powell, Thomas Pradeu, and
Neeraja Sankaran for their very detailed and helpful critical com-
ments on both the structure and contents of this paper. I am
especially grateful to Ton van Helvoort for his comments on the
penultimate draft, to Alexandra Bacopoulos-Viau for her help in
editing the paper in its final stage as well as to the guest editors of
this special issue for inviting me to contribute. Funding from the
Fonds Recherche du Québec Société et Culture is warmly
acknowledged.
References
Beijerinck, M. W. (1964). Concerning a contagium vivum fluidum as cause of the
spot disease of tobacco leaves. In N. Hahon (Ed.), Selected papers on virology (pp.
52e63). Englewood Cliffs: Prentice-Hall [1898]
Billiau, A. (1881e1957). (‘online first’) At the centennial of the bacteriophage:
reviving the overlooked contribution of a forgotten pioneer, Richard Bruynoghe.
Journal of the History of Biology. http://dx.doi.org/10.1007/s10739-015-9429-0.
Bos, L. (1995a). One hundred years of virology? ASM News, 61, 53e54.
Bos, L. (1995b). The embryonic beginning of virology: Unbiased thinking and
dogmatic stagnation. Archives of Virology, 140, 613e619.
Bos, L. (2000). 100 years of virology: From vitalism via molecular biology to genetic
engineering. Trends in Microbiology, 8, 82e87.
Bourdieu, P. (1988). Homo academicus. Oxford: Polity Press.
Burnet, F. M. (1946). Virus as organism. Evolutionary and ecological aspects of some
human viruses. London: Oxford University Press.
Burnet, F. M. (1953a). Virology as an independent science: lecture I: the foundations
of virology. (Mathison Memorial Lecture series) Medical Journal of Australia, 22,
809e813.
Burnet, F. M. (1953b). Virology as an independent science: lecture II: the substance
of virology. (Mathison Memorial Lecture series) Medical Journal of Australia, 23,
841e845.
Burnet, F. M. (1955). An historical introduction to animal viruses. In Principles of
animal virology. New York: Academic Press.
Burnet, F. M., & Stanley, W. (Eds.). (1959). The viruses, general virology (Vol. 1)New
York, London: Academic Press.
Calisher, C. H., & Horzinek, M. C. (Eds.). (1999). 100 Years of virology. The birth and
growth of a discipline. Vienna, New York: Springer.
Chastel, C. (1992). Histoire des virus. De la variole au sida. Paris: Société nouvelle des
éditions Boubée.
Creager, A. N. (2001). The life of a virus. Tobacco mosaic virus as an experimental
model, 1930e1965. Chicago: University of Chicago Press.
de Chadarevian, S. (2009). Microstudies versus big picture accounts? Studies in
History and Philosophy of the Biological and Biomedical Sciences, 40, 13e19.
Fleck, L. (1979). Genesis and development of a scientific fact. Chicago: Chicago Uni-
versity Press [1935].
 P.-O. Méthot / Studies in History and Philosophy of Biological and Biomedical Sciences 59 (2016) 145e153
Forterre, P. (2016). To be or not to be alive: how recent discoveries challenge the
traditional definitions of viruses and life. Studies in History and Philosophy of
Biological and Biomedical Sciences, 59, 100e108.
Galperin, C. (1987). Le bactériophage, la lysogénie et son déterminisme. History and
Philosophy of the Life Sciences, 9(2), 175e227.
Galperin, C. (1994). Virus, provirus et cancer. Revue d’histoire des sciences, 47(1), 7e
56.
Gaudillière, J.-P. (2000). Rockefeller strategies for scientific medicine: molecular
machines, viruses and vaccines. Studies in History and Philosophy of the Bio-
logical and Biomedical Sciences, 31(3), 491e509.
Gayon, J. (2004). Introduction to a special issue on “Dynamiques des disciplines
scientifiques: classification, embryologie, génétique”. Bulletin d’histoire et
d’épistémologie des sciences de la vie, 11, 151e155.
Gingras, Y. (1991). L’institutionnalisation de la recherche en milieu universitaire.
Sociologie et sociétés, 23, 41e54.
Gradmann, C. (2014). A spirit of scientific rigour: Koch’s postulates in twentieth-
century medicine. Microbes and Infection, 16, 885e892.
Grafe, A. (1991). A history of experimental virology. Berlin: Springer-Verlag.
Grmek, M. D. (1981). A plea for freeing the history of science from myth. In
M. D. Grmek, R. S. Cohen, & G. Cimino (Eds.), On scientific discovery, the Erice
Lecture 1977, Boston studies in the philosophy of science (pp. 9e42).
Grmek, M. D. (1994). Histoire de la virologie, des viroses et des virologues. History
and Philosophy of the Life Sciences, 16, 339e354.
Hodge, J. (2005). Against ‘revolution’ and ‘evolution’. Journal of the History of Biology,
38(1), 101e121.
Holmes, F. L. (2004). The dynamics of disciplines in the history of science. Bulletin
d’histoire et d’épistémologie des sciences de la vie, 11, 157e174.
Horsfall, F. L., Jr. (1965). Thomas Milton Rivers, 1888-1962. A biographical memoir.
Washington: National Academy of Sciences.
Horzinek, M. C. (1997). The birth of virology. Antonie van Leeuwenhoek, 71, 15e20.
Hughes, S. S. (1977). The virus: A history of the concept. London: Heinemann
Educational Books.
Jordanova, L. (1993). Gender and the historiography of science. British Journal for the
History of Science, 26, 469e483.
Kevles, D. J., & Geison, G. L. (1995). The experimental life sciences in the twentieth
century. Osiris, 10, 97e121.
Koonin, E. V., & Starokadomskyy, P. (2016). Are viruses alive? The replicator para-
digm sheds decisive light on an old but misguided question. Studies in History
and Philosophy of the Biological and Biomedical Sciences, 59, 125e134.
Kostyrka, G. (2016). What roles for viruses in origin of life scenarios? Studies in
History and Philosophy of the Biological and Biomedical Sciences, 59, 135e144.
Kragh, H. (1987). An introduction to the historiography of science. Cambridge: Cam-
bridge University Press.
Kuhn, T. (1962). The structure of scientific revolutions. Chicago: University of Chicago
Press.
Lechevalier, H. (1972). Dmitri Iosifovich Ivanovski (1864-1920). Bacteriological Re-
views, 36, 135e145.
Lemaine, G., Macleod, R., Mulkay, M., & Weingart, P. (1976). Problems in the
emergence of new disciplines. In G. Lemaine, R. Macleod, M. Mulkay, &
P. Weingart (Eds.), Perspectives on the emergence of scientific disciplines (pp. 1e
23). Paris: Maison des Sciences de l’Homme.
Lenoir, T. (1997). Instituting science. The cultural production of scientific disciplines.
Stanford: Stanford University Press.
Listios, S. (1997). René J. Dubos and Fred L. Soper: Their contrasting views on vector
and disease eradication. Perspectives in Biology and Medicine, 41, 138e149.
van Loon, J. (2002). A contagious living fluid. Objectification and assemblage in the
history of virology. Theory, Culture & Society, 19, 107e124.
Lustig, A., & Levine, A. J. (1992). One hundred years of virology. Journal of Virology,
66, 4629e4631.
Lwoff, A. (1957). The concept of virus. Journal of General Microbiology, 17, 239e253.
Mayer, A. (1964). Concerning the mosaic disease of tobacco. In N. Hahon (Ed.),
Selected papers on virology (pp. 37e45). Englewood Cliffs: Prentice-Hall [1886]
Méthot, P.-O., & Fantini, B. (2014). Medicine and ecology: historical and critical
perspectives on the concept of ‘emerging disease’. Archives Internationales
d’histoire des sciences, 64(172e173), 214e230.
Morange, M. (1998). A history of molecular biology. Cambridge: Harvard University
Press [1993]
Morange, M. (2005). André Lwoff: from protozoology to molecular definitions of
viruses. Journal of Biosciences, 30, 591e594.
Morgan, G. J. (2016). What is a virus species? Radical pluralism in viral taxonomy. Studies
in History and Philosophy of the Biological and Biomedical Sciences, 59, 64e70.
Moulin, A.-M. (1992). La métaphore vaccine. De l’inoculation à la vaccinologie.
History and Philosophy of the Life Sciences, 14, 271e297.
O’Malley, M. A. (2014). Philosophy of microbiology. Cambridge: Cambridge University
Press.
Pasteur, L. (1890). La rage. In P. Vallery-Radot (Ed.), Oeuvres de Pasteur (Vol. 6). Paris:
Masson, 1922e1939, pp. 672e688.
Pestre, D. (1999). Commemorative practices at CERN: Between physicists’ memories
and historians’ narratives. Osiris, 14, 203e216.
Powell, A., O’Malley, M. A., Müller-Wille, S., Calvert, J., & Dupré, J. (2007). Disci-
plinary baptisms: a comparison of the naming stories of genetics, molecular
153
biology, genomics, and systems biology. History and Philosophy of the Life Sci-
ences, 29, 5e32.
Rasmussen, N. (1997). Picture control. The electron microscope and the transformation
of biology in America, 1940-1960. Stanford: Stanford University Press.
Rheinberger, H.-J. (2010). Virus research at the Kaiser Wilhelm institutes for
biochemistry and biology, 1937-45. In An epistemology of the Concrete.
Twentieth-Century histories of life (pp. 128e149). Durham & London: Duke
University Press.
Rivers, T. (1927). Filterable viruses. A critical review. Journal of Bacteriology, 14, 217e
258.
Sankaran, N. (2006). Frank Macfarlane Burnet and the nature of the bacteriophage,
1924e1937. Ph.D. thesis. Yale University.
Sapp, J. (1994). Evolution by association. A history of symbiosis. Oxford: Oxford
University Press.
Schmiedebach, H.-P. (1999). The Prussian state and microbiological research e
Friedrich Loeffler and his approach to the “invisible virus”. In C. H. Calisher, &
M. C. Horzinek (Eds.), 100 years of virology. The birth and growth of a discipline
(pp. 9e23). Vienna, New York: Springer.
Scholthof, K.-B. G. (2014). Making a virus visible: Francis O. Holmes and a biological
assay for tobacco mosaic virus. Journal of the History of Biology, 47, 107e145.
Suárez-Díaz, E. (2009). Molecular evolution: concepts and the origin of disciplines.
Studies in History and Philosophy of the Biological and Biomedical Sciences, 40,
43e53.
Summers, W. (2005). Bacteriophage research: early history. In E. Kutter, &
Al Sulakvelidze (Eds.), Bacteriophages: Biology and applications (pp. 5e27). Boca
Raton: CRC Press.
Summers, W. (2014). Inventing viruses. Annual Review of Virology, 1, 25e35.
van Helvoort, T. (1991). What is a virus? The case of tobacco mosaic disease. Studies
in History and Philosophy of Science, 22(4), 577e588.
van Helvoort, T. (1992). Bacteriological and physiological research styles in the early
controversy on the nature of the bacteriophage phenenomenon. Medical His-
tory, 36, 243e270.
van Helvoort, T. (1993). A bacteriological paradigm in influenza research in the first
half of the twentieth century. History and Philosophy of the Life Sciences, 15, 3e
21.
van Helvoort, T. (1994a). History of virus research in the twentieth century: the
problem of conceptual continuity. History of Science, 32, 185e235.
van Helvoort, T. (1994b). The construction of bacteriophage as bacterial virus:
linking endogenous and exogenous thought styles. Journal of the History of
Biology, 27, 91e139.
van Helvoort, T. (2014). Virus and cancer studies e still fascinating after all these
years. Studies in History and Philosophy of the Biological and Biomedical Sciences,
48, 258e259.
van Kammen, A. (1999). Beijerinck’s contributions to the virus concept e an
introduction. In C. H. Calisher, & M. C. Horzinek (Eds.), 100 Years of virology. The
birth and growth of a discipline (pp. 1e8). Vienna, New York: Springer.
van Helvoort, T. (1996). When did virology start? ASM News, 62, 142e145.
van Regenmortel. (2016). The metaphor that viruses are living is alive and well, but
it is no more than a metaphor. Studies in History and Philosophy of the Biological
and Biomedical Sciences, 59, 117e124.
Waterson, P. A., & Wilkinson, L. (1978). An introduction to the history of virology.
Cambridge: Cambridge University Press.
Wilkins, A. S. (1996). Are there ‘Kuhnian’ revolutions in biology? BioEssays, 18, 695e
696.
Wilkinson, L. (1974). The development of the virus concept as reflected in corpora
of studies on individual pathogens. Beginnings at the turn of the century.
Medical History, 18, 211e221.
Wilkinson, L. (1975). The development of the virus concept as reflected in corpora
of studies on individual pathogens. The agent of fowl plague e a model virus?
Medical History, 19, 52e72.
Wilkinson, L. (1976). The development of the virus concept as reflected in corpora
of studies on individual pathogens. Lessons of the plant viruses e tobacco
mosaic virus. Medical History, 20(2), 111e134.
Wilkinson, L. (1977). The development of the virus concept as reflected in corpora
of studies on individual pathogens. Rabies etwo millennia of ideas and
conjecture on the aetiology of a virus disease. Medical History, 21, 15e31.
Wilkinson, L. (1979). The development of the virus concept as reflected in corpora
of studies on individual pathogens. Smallpox and the evolution of ideas on
acute (viral) infections. Medical History, 23, 1e28.
Wolstenholme, G. E. W., & O’Connor, C. M. (1960). CIBA foundation symposium on
virus, virulence and pathogenicity. London: J. & A. Churchill.
Wolstenholme, G. E. W., & Millar, E. C. P. (Eds.). (1957). CIBA foundation symposium
on the nature of viruses. London: J. & A. Churchill.
Worboys, M. (2007). Was there a bacteriological revolution in the late nineteenth-
century medicine? Studies in History and Philosophy of the Biological and
Biomedical Sciences, 38(1), 20e42.
Zaitlin, M. (1998). The discovery of the causal agent of the tobacco mosaic disease.
In S. D. Kung, & S. F. Yang (Eds.), Discoveries in plant biology (pp. 105e110). Hong
Kong: World Publishing.