Natural Histories of Infectious Disease: Ecological Vision in Twentieth-Century Biomedical Science Author(s): Warwick Anderson Source: Osiris, 2nd

Series, Vol. 19, Landscapes of Exposure: Knowledge and Illness in Modern Environments, (2004), pp. 39-61 Published by: The University of Chicago Press on behalf of The History of Science Society Stable URL: http://www.jstor.org/stable/3655231 Accessed: 04/08/2008 10:19
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Natural

Histories

of

Infectious

Disease:

Ecological Vision in Twentieth-Century Biomedical Science

Anderson* By Warwick
ABSTRACT During the twentieth century, disease ecology emerged as a distinct disciplinary network within infectious diseases research. The key figures were Theobald Smith, F. Macfarlane Bumet, Rene Dubos, and Frank Fenner. They all drew on Darwinian evolutionism to fashion an integrative (but rarely holistic) understanding of disease processes, distinguishing themselves from reductionist "chemists" and mere "microbe hunters."They sought a more complex, biologically informed epidemiology. Their emphasis on competition and mutualism in the animated environment differed from the physical determinism that prevailed in much medical geography and environmental health research. Disease ecology derived in part from studies of the interaction of organisms-micro and macro-in tropical medicine, veterinary pathology, and immunology. It developed in postcolonial settler societies. Once a minority interest, disease ecology has attracted more attention since the 1980s for its explanations of disease emergence, antibiotic resistance, bioterrorism, and the health impacts of climate change. INTRODUCTION

The end of the twentieth century found Joshua Lederberg reflecting on the history of infectious diseases research. "During the early acme of microbe hunting, from about 1880 to 1940," he wrote, "microbes were all but ignored by mainstream biologists." Moreover, "medical microbiology had a life of its own, but it was almost totally divorced from general biological studies." Bacteriologists "had scarcely heard of the conceptual revolutions in genetic and evolutionary theory."Although germs had long been "recognized as living entities . . . the realization that they must inexorably be evolving and changing" was slow to penetrate public health and medical practice.1Yet Lederberg could identify a few conceptual bridges linking bacteriology and general
*Department MedicalHistoryandBioethics,Universityof WisconsinMedicalSchool, 1440 Mediof cal Sciences Center,1300 UniversityAve., Madison,WI 53706-1532; whanderson@med.wisc.edu. I would like to thankGreggMitman,Michelle Murphy,andChrisSellers for askingme to writethis at Place, andHealth," the Universityof Wisconsinat Madiessay for the conferenceon "Environment, son. Susan Craddockprovidedhelpful comments at the meeting. BarbaraGutmannRosenkrantzand MarkVeitchhave fromthe startguidedmy workon disease ecology. I am also gratefulto DavidAbernathy,Nick King, LindaNash, CharlesRosenberg,andConeveryBoltonValenciusfor theiradvice on earlierversions of this essay. FrankFenner,JoshuaLederberg,Steve Boyden, and Tony McMichael cheerfullycorrectedsome misconceptions,thoughthey may not agree with all thatremains. JoshuaLederberg,"InfectiousHistory," Science 288 (2000): 287-93, on 288, 291. 2004 by The Historyof Science Society.All rightsreserved.0369-7827/04/1901-0003$10.00 OSIRIS 2004, 19: 039-061

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WARWICK ANDERSON

biology (most of them, admittedly, constructed after 1940). Once O. T. Avery, at the Rockefeller Institute, discovered that nucleic acid was the transmissible factor responsible for pneumococcal transformation, bacteria and bacterial viruses had "quickly supplanted fruit flies as the test-bed for many of the subsequent developments of molecular genetics and biotechnology."2 Indeed, in 1956 Lederberg himself had been awarded a Nobel Prize for his work in microbial genetics. He noted, too, that the synoptic texts of Macfarlane Bumet and Rene Dubos had described the nexus of microbiology and general biology; their pioneering integrative work characterized the relations of disease, environment, and evolutionary processes in newly fashionable ecological terms.3 Increasing confidence in antibiotic and vaccine development during the 1960s and 1970s, however, led to the neglect of such ecological interpretations of infectious disease. In the 1980s, nature struck back. Emergent diseases, such as AIDS, and problems of microbial resistance to antibiotics, prompted "widespread re-examination of our cohabitation with microbes." It was time, Lederberg wrote in the year 2000, for us to abandon the old metaphor of a war between germs and humans, replacing it with "a more ecologically informed metaphor, which includes the germ's-eye view of infection." Above all, he concluded, we need more "researchinto the microbial ecology of our own bodies."4 Historians generally have neglected ecological traditions in biomedical science. Like most of the scientists and physicians they study, historians have chosen instead to emphasize the development during the twentieth century of simplified laboratory models for complex pathophysiological mechanisms. Moreover, diagnosis and prevention are commonly framed in terms of "microbe hunting," and treatment in terms of "magic bullets." Sometimes the story will conclude with a monitory account of the pitfalls of progress: technically biomedicine is doing better than ever, but at the cost of its interpretive or exegetical power, its ability to define and represent our place in nature. Medical science, in these accounts, has concentrated on elucidating mechanisms of disease, abandoning the older efforts-frequently associated with the names of Hippocrates and Sydenham-to make sense of life forms and their relations to the environment. It is easy to find examples of such cautionary tales, with their typical mixture of satisfaction with contemporary achievement and nostalgia for a more integrative, or holistic, worldview. 5When Charles-Edward Amory Winslow came to write his history of the "conquest" of epidemic disease, he admitted that "the practical triumphs of bacteriology did indeed tend to over-simplify the problem and to cause medical men for nearly half a century to ignore the true manysidedness of disease."6 Following a similar line of reasoning, Mirko Grmek has
Ibid., 288. See Ren6 J. Dubos. TheProfessor,TheInstitute,and DNA (New York, 1976). 3 F. M. Buret, Biological Aspects ofInfectiousDisease (New York,1940); andRen6J. Dubos, Man Adapting(New Haven, Conn., 1965). See also JoshuaLederberg,"J.B. S. Haldane(1949) on Infectious Disease andEvolution,"Genetics 153 (1999): 1-3. 4 Lederberg,"InfectiousHistory"(cit. n. 1), 289, 290, 293. Even in 1988, Lederberg,citing Buret, had urgedus "to come to grips with the realitiesof our place in nature" ("MedicalScience, Infectious Journal of the AmericanMedical Association 260 [1988]: Disease, and the Unity of Humankind," 684-5, on 684). 5 For my own contributionto this literature,see WarwickAnderson,"Disease and Its Meanings," Lancet2000 354 (1999): SIV49. 6 C.-E. A. Winslow, The Conquestof EpidemicDisease: A Chapterin the History of Ideas (1943; Madison,Wis., 1980), 335. reprinted
2

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41

pointedout that since Pasteurand Koch, "muchmore importanceis attachedto investigationsof the biology of germs thanto knowledgeof the influenceof milieu."7 Recently,MichaelWorboyshas arguedthateven if germtheoriesdid not lead physicians simply to switch from holism to reductionism,interestin social and environmentalinfluencesgraduallydeclined, and increasingly"diseasewas constitutedin relationsbetweenbacteriaandindividualbodies."8 pace may be slower,but the The destinationis the same: medical science eventuallybecomes triumphantly, peryet reductionist. haps meretriciously, Anyone would think that modem biomedicineis just a matterof culturinggermsin the laboratory, identifyingtheirphysicochemical and trackingthem in the community-that is, little more than microbe properties, hunting. In this essay, however,I wantto review the historyof infectious diseases research in the twentieth century to recover various emerging forms of ecological understandingfrom what has sometimes seemed an arid waste of reductionism.In particular,I would like to sketchout personalconnectionsbetweenthe majoradvocates of the ecology-or "natural rehistory"-of disease, to describe an international searchnetwork,and to explore the variousinstitutionaland social niches these scientists occupied. Reconstructingthis social ecology of ecological knowledge in medical science is no easy task.9For one thing, each of the major pioneers of a broaderbiological conception of disease processes tended to representhimself as singular,as the sole authorof the idea, and rarelycited others,even those linked to him by educationand friendship.For many early proponents,this rhetoricof singularityand marginalitywas a crucial aspect of the argumentand partof theirown propheticself-fashioning.Yet the trainingand the careerpaths of key figures such as TheobaldSmith, F. MacfarlaneBurnet,and Rene Dubos structured intricate an networkof influence,counsel, and criticism.This is not to say thata uniform,welldefinedschool of disease ecology existed in the twentiethcentury.The differences between many of these scientists can at times be as greatas any similarity:Dubos, for example, was by the end of his careera significantoutlier from this cluster of scholars.But an emphasison singularity,and on variationin laboratory work, or in and appeal,often disguised a fundamentalsimilarityin approach,a famargument ily resemblancein theory, perspective, and career.These scientists had common points of referenceand a sharedrhetoricalrecourseto ecological andotherbroadly disease. That is, the intellectual interacintegrativeapproachesto understanding tions of these andotherscientistsforgeda recognizablesubdiscipline,a sharedconceptual frameworkand rhetoric,within infectious diseases research,even if their own local commitmentsand laboratorystyle sometimes differed. Itis important distinguish assertion an"environmental" to this of in perspective medical science from an earlierconcernwith medicalgeography, role of the physical the
7 Mirko medicale et histoiredes civilisations," Annales: Economies,Societes, Grmek,"G6ographie Civilisations 18 (1963): 1071-89, on 1085. For similar statements,see Fielding H. Garrison,"The Newer Epidemiology," MilitarySurgeon53 (1923): 1-14, 10; andErwinH. Ackerknecht, Historyand Diseases (New York, 1965), 1. Geographyof the Most Important 8 Michael Worboys, Spreading Germs: Disease Theories and Medical Practice in Britain, 1865-1900 (Cambridge,2000), 285. 9 CharlesE. Rosenberg,"Toward Ecology of Knowledge:On Discipline, Context,and History," an in The Organizationof Knowledgein ModernAmerica, 1860-1920, ed. AlexandraOleson and John Voss (Baltimore,1979), 440-55.

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WARWICK ANDERSON

was more disease.0Theworkof Burnet, particular, structured around in milieuin causing in a biologicallymediated environment derived, part,fromtheparasitological and tradito medicineandveterinary whichwas preadapted ecological tionin tropical pathology, of notionsof directenvironmental deexplanations thissort,notfromolderHippocratic in Unlikemostmedicalgeography-which persisted thetwentieth terminism. in century an medicalhistoryand in geographyitself-disease ecology postulated evolutionary and and time scale, modelsthatwere integrative interactive, a globalscope.Moreover, diseaseecology was less explicitlyracialin its arguments, thougha concernwithpopuof In general, spatial the lationquality imaginary diseaseecology undoubtedly persisted. and was moreabstract biologicallyanimated medicalgeography, theprocesses and than not or it described werevisibleonly to experts, readilydiscerned experienced by usually of was thegeneral interaction generally evless public.Thatis, thefinepattern microbial identthana changein seasonor a shiftin the winddirection. the ecologicalunderBut couldnonetheless comof theglobalas a siteof infectious diseaseemergence be standing it was, afterall, Duboswho coinedthe slogans"OnlyOne Earth," "Think and pelling: Act it the Furthermore,wouldbe diseaseecologythatprovided most Globally, Locally." of forthe emergence "new" diseasesin the 1980s. plausible explanation The disease ecologists whose careersI traceherewerenot, of course,the only ones in the seekingto reinterpret relationsof healthandenvironment the twentiethcentury. for Therewereresearchers, example,in toxicology andoccupationmedicine,who, as Sellers has pointed out, transformed those fields into "environmental Christopher health"afterWorldWarII, a metamorphosis most forcefullyexpressed,perhaps,in studiesof the etiology of cancerandrepresented popularlyin RachelCarson'sSilent 1 Althoughoften not explicitly "ecological"-the termis used only twice in Spring. Silent Spring-studies of the effect of the physical environmenton humanhealth could, at times, drawextensivelyon workin animalecology, andthe emphasison the balanceof nature the darkside of modernity and conveyeda broadlyecologicaltone.12 It is remarkable, how rarelythe proponents an ecology of infectiousdisof then,just eases andthe expertson the healthimpactof the physicalenvironment referto one another.They may have sharedintellectualinterests,even political concerns,but their social networks,careertrajectories, institutional and niches seem verydifferent.Such basic similarityalliedwith self-styleddifferenceatteststo boththe broadsalienceand the interpretive and flexibilityof notions of "environment" "ecology"in twentiethof centuryexplanations healthanddisease. of The relationship disease ecology to medicalholism, or constitutional medicine, andto theoriesof the pathogenicpotentialof civilizationis just as ambiguous.Both Burnetand Dubos severely criticizedwhat they saw as an exclusively reductionist
10See Grmek, and Mias"G6ographiemedicale" (cit. n. 7); Caroline Hannaway,"Environment in mata," Companion Encyclopediaof the Historyof Medicine,ed. W. F Bynum andRoy Porter(Lonin don, 1993), 292-308; ConeveryBolton Valencius,"Historiesof Medical Geography," Medical Geographyin HistoricalPerspective,ed. NicolaasA. Rupke(London,2000), 3-29; andFrankA. Barrett, Disease and Geography:TheHistory of an Idea (Toronto,2000). " Christopher Sellers, Hazardsof the Job: FromIndustrialDisease to Environmental C. Health Science (ChapelHill, N. C., 1997). 12 1 am gratefulto Gregg Mitmanand Chris Sellers for insisting on this. Carsonobserves that disease is "a problem of ecology, of interrelationships, interdependence" of (Silent Spring [1963; re1974], 169). But as Sellers suggests, "[E]venas [Carson's]own ecological printedHarmondsworth, habitsof mindprovedcrucialto her synthesis,not ecology itself butthe sciences of humanhealthsup(Hazardsof the Job [cit. n. 11], 2). plied the core of her argument"

NATURALHISTORIESOF INFECTIOUSDISEASE

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trendin modem biomedicine,but they did not displaymuchinterestin holism at the level of the humanorganism,andtheirecological modelsalwayscontainedsome key narrowand overly simreductionistelements.'3Rather,they distrustedunnaturally with physplified accountsof any sort, especially those revealinga preoccupation icochemical mechanism,promotinginsteada more inclusive and integratedunderand standingof natureandsociety.Theirannoyancewith reductionism dismayat the of were not unrelated.'4 character modernity CertainlyDubosbecamea faemerging mous critic of the dangersof industrialcapitalismand environmentally insensitive the end of his career,condemnation a destructivecivilization,and of By modernity. its associatedalienatedrationalism, came dominatedhis writing,and his arguments to assumea more traditionally holistic, humanistic,and even mystical,cast. Buret, towardmodernity, too, displayedambivalence pointingto the dangersof overpopulation, biological warfare,antibioticresistance,andenvironmental all, degradation, in his opinion,the fruitsof a narrowly reductionist-even obscurantist-worldview. As CharlesE. Rosenberg suggested,anxietiesaboutthe riskof modernways of has life were, duringthe courseof the twentiethcentury, "explained increasinglyin terms not of the city as a pathogenicenvironment, of evolutionary globalecological and but realities."15 is the purposeof this essay to explainhow one partof this complexexIt framework assembledandeventuallymadepopular. was planatory
THE NEWEREPIDEMIOLOGY AND HOST-PARASITE INTERACTIONS

and Writingin 1940,Iago Galdston,a New Yorkpsychiatrist historianof medicine, remarked the crudereductionism most "post-Pasteurian" on of medicine.16Epidemiand ology, the statisticalstudyof the character cause of disease,had once provideda multifactorial and concomitants, the richlytextured, accountingof the environmental social andmoralcircumstances, epidemicsthatravaged of nineteenth-century Europe andNorthAmerica.Since the demise of medicalgeographyand the generalacceptance of germtheories,however,an intricateand exactingcalculushad dwindledinto meremicrobehunting.This lamentation becomecommonplaceand,by this time, had was perhapsmorea necessaryritualthana realisticdescription. was certainlyconIt venient for reformersto representthe statusquo as a collection of narrow-minded
Moregenerally, holismin the twentieth on see "Holismin Twentiethcentury, CharlesE. Rosenberg, CenturyMedicine,"in Greaterthan the Parts: Holism in Biomedicine, 1920-1950, ed. Christopher LawrenceandGeorgeWeisz (New York, 1998), 335-55. 14 Nor are they unrelatedto the concernsthatpromptedsocial medicine, though the styles of analysis, and policy implications,were often quite different.Yet in his earlierwork, JohnA. Ryle could claim that"therearebetterinspirations thoughtfulmedicineto be foundin the Originof Species than to in a modem textbook of bacteriology"(The Natural History of Disease [1936; reprintedLondon, 1988], 382). See also Iago Galdston, The Meaning of Social Medicine (Cambridge,Mass., 1954); Bulletinof the History GeorgeRosen, "WhatIs Social Medicine?A GeneticAnalysis of the Concept," of Medicine21 (1947): 674-733; andDorothyPorter,"Changing Disciplines:JohnRyle andthe MakBritain," ing of Social Medicine in Twentieth-Century Historyof Science 30 (1992): 119-47. 15 Charles E. Rosenberg, "Pathologiesof Progress:The Idea of Civilization as Risk,"Bull. Hist. Med. 72 (1998): 714-30, on 723. Rosenbergdescribes"a more expansiveglobal perspectivein which inclusive and ecological styles of analysis have become increasinglypervasive"(726), citing Buret. For a helpful discussion of the epidemiologicaltropesof "contamination" "configuration" and (i.e., a more ecological understanding epidemics), see Rosenberg,"Explaining of Epidemics,"in Explaining Epidemics,and OtherStudiesin the History of Medicine (Cambridge,1992), 293-304. 16 Iago Galdston, Progress in Medicine:A Critical Review of the Last One HundredYears(New York, 1940).
13

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technicians, to represent the old guard as cloistered laboratory scientists dutifully cultivating their germs or as public health officials glibly documenting disease outbreaks, tracing germs through the community. The benefits of complexity and integrative analysis surely would stand out most clearly against such alleged routinism and artlessness. But those who studied tropical medicine and veterinary pathology-and, indeed, most field epidemiologists-understood that tracking the microbial cause of a disease outbreak rarely proved as easy as one might hope. Other biological factors had to be taken into account. The development of the notion of "carrier status" during the 1890s-the idea that susceptibility to germs might vary in a human population, allowing some individuals to spread a germ without succumbing to it-had already presented a challenge to more simplistic efforts to track microbes.17The full complexity of host-parasite relations, however, was perhaps better revealed in studies of tropical medicine and veterinary pathology, in which various life forms were commonly involved in disease processes. Just before World War I, Ronald Ross had tried to get his colleagues in tropical medicine to accept a sophisticated dynamic-equilibrium model of disease transmission, but by then few of them took his work seriously.'8 It was instead Theobald Smith, a comparative pathologist at Harvard, who became the major advocate of the study of disease as a general biological problem. Smith, the codiscoverer of the role of a tick in transmitting the parasite that causes Texas cattle fever, was fond of emphasizing the mutual dependence of host and microorganism, whether in health or disease. In an address at the 1904 St. Louis Universal Exposition, he had claimed that the "social and industrial movement of the human race is continually leading to disturbances of equilibrium in nature, one of whose direct or indirect manifestations is augmentation of disease."19Later, Smith would argue that the study of disease was no longer "in the hands of professional mystics" because "disease has become a biological problem.... [I]t is ranging itself among natural phenomena in our mind." He also positioned his comparative work in pathology and immunology, with its concern for dynamic biological processes, against the work of investigators trying to dig toward the "more fundamental concepts embodied in physics and chemistry."20Smith described health and disease as consequences of a struggle for existence between living things, predatory and parasitic. He reported on the life cycle of parasites, host-parasite conflict, symbiosis and mutualism, cell parasitism and phagocytosis, and variation and mutation among parasites. "Parasitism
17 On the see "healthycarrier," Winslow,Conquestof EpidemicDisease (cit. n. 6); andJudithWalzer Leavitt,Typhoid Mary: Captiveto the Public's Health (Boston, 1996). 18 RonaldRoss, "Some Quantitative Studies in Epidemiology," Nature 87 (1911): 466-7. See also J. Andrew Mendelsohn,"FromEradicationto Equilibrium:How Epidemics Became Complex after WorldWarI,"in LawrenceandWeisz, Greaterthan the Parts (cit. n. 13), 303-31. On the practiceof disease ecology in African tropical medicine during this period, see Helen Tilley, "Ecologies of AfricanTrypanosomiasis, the Science of Disease Controlin and Complexity:TropicalEnvironments, BritishColonialAfrica, 1900-1940" (this volume). 19 TheobaldSmith, "Some Problemsin the Life Science, History of PathogenicMircoorganisms," as n.s., 22 (1904): 817-32, on 817. See also his "Parasitism a Factorin Disease," Science, n.s., 54 (1921): 99-108. Smithbecame directorof the Division of Animal Pathologyof the RockefellerInstiGutmann Amer"Theobald Smith," tute,in Princeton,New Jersey,in 1914. See Barbara Rosenkrantz, ican NationalBiographyOnline,Feb. 2000, http://www.anb.org/articles/12/12-00861.html; Hans and Zinsser, "BiographicalMemoir of Theobald Smith, 1859-1934," Biographical Memoirs (National Academy of Sciences) 17 (1936): 261-303. 20 Theobald Smith,Parasitismand Disease (Princeton,1934), viii, x.

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may be regarded,"Smith wrote in 1934, "not as a pathological manifestation, but as a normal condition having its roots in the interdependence of all living organisms." The prevailing modus vivendi was always temporary, always evolving: the "face of nature and of civilization is steadily changing and thereby changing the host-parasite relations."21 Here was a naturalistic and evolutionary understanding of the interactions of organisms-micro and macro-in which human disease was decentered and environment, or milieu, became animated. Ultimately, "all that can be postulated is the universal struggle of living things to survive, and in this struggle the fundamental biological reactions gradually range themselves by natural selection under ... categories of offence and defence." The environment that mattered most was alive, and any effect of climate or topography would be mediated through the interactions of organisms. Smith reflected on the difference between his dynamic modeling and the work of earlier medical geographers: Sanitarians lookedto the variations atmospheric in moistureandtemperature, rise and the fall of the waterin subsoil,greatfluctuations temperatures, favoringcauses of epiin as demics.Todaywe areinclinedto narrow themdownto the humanandanimalworld,their the in intercourse, migrations, continualfluctuations habitsand modes of life, but espeof cially in the increasingsusceptibility populations duringthe disease-freeperiods.22 If "the human race is in a rather delicate, unstable relation to its environment," as Smith contended, then a more complex epidemiology was required. Hans Zinsser, in his obituary of Smith, saw his friend and colleague struggling against a tendency "for bacteriological investigation to segregate isolated fractions of a problem for analysis, with frequent neglect of the correlation of results with the problem as a whole."23 Complex, integrative statistical studies of disease were necessary. The relation of illness to population mobilization and host resistance, evident during World War I and in the subsequent influenza pandemic, had already attuned many epidemiologists to such appeals for a more intricate and naturalistic accounting of disease patterns. Major Greenwood, a statistician at the Ministry of Health in Britain, and later the first professor of epidemiology at the London School of Hygiene and Tropical Medicine, claimed that in the nineteenth century William Farr and John Simon had recognized that "the conditions of human life-both, as we now say, environmental and eugenic-were quite as important to the epidemiologist as the materia morbi."24Since then, however, epidemiology had become oversimplified, too close to a mere addendum on bacteriology. Greenwood wanted more complex and evolutionary studies of disease in populations, not mere microbe hunting. Epidemics, he believed, were the consequence of a disturbance in biological equilibrium and thus not reducible to invasion stories.25Simon Flexner, director of the Rockefeller Institute and an admirer of Smith's work, also observed a reaction against a facile bacteriological epidemiology during the 1920s. He noted that "each generation receives its
Ibid., 2, xi. Ibid., 162. 23 Zinsser,"TheobaldSmith"(cit. n. 19), 284. 24 "TheEpidemiological Pointof View," BritishMedicalJournal(hereafter MajorGreenwood, BMJ)2 see and (1919):405-7. Formoreon Greenwood, J. RosserMatthews,Quantification the QuestforMedical Certainty "From Eradication Equilibrium" n. 18). to (Princeton, 1995),chap.5; andMendelsohn, (cit. 25 to MajorGreenwood,Epidemicsand CrowdDiseases: An Introduction the Studyof Epidemiology (London, 1935).
22 21

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was particular impressionof epidemicdiseases"andthatthe postwargeneration making an effortto "defineepidemiologyin termswiderthanthose of the microbicincitantsof disease alone."26 Accordingto Flexner,this would mean developingan exinteractions Smithhad sketched perimental epidemiologyin which the host-parasite out mightbe exploredfurther. Studiesof the biological complexitiesof host-parasite interactionsoon reacheda morepopularreadership, which wantedto learnaboutthe latestscientifictheorieson man'splace in nature.In Rats,Lice, and History,HansZinsserrecountedthe "biography"of typhusfever,tracingthe impactof the disease on the rise and fall of civilizations.Zinsser,a professorof bacteriologyandimmunologyat Columbiaandlater at Harvard, respectedfor his investigations leucocytefunctionandthe precipwas of itin reaction,and he wrote extensivelyon allergyand the immuneresponse.Having trainedinitially as a biologist-with E. B. Wilson at Columbia-Zinsser knew that "physicochemical analysis will never give the final clue to life processes"and that TheobaldSmith'sstudyof "complexsystems"was required instead."Infectious disease is one of the greattragediesof living things," Zinsserwrote,"thestrugglefor existence betweendifferentformsof life."27 similarlytragicbiologicalview could be A seen in PercyAshbur's TheRanksof Death, a popularbook on the contribution of diseaseto the European of America.Influenced Zinsser,Ashburn evoked conquest by European migrationto the WesternHemisphereas "thegreatestmobilizationof disto ease, of its introduction new andsusceptiblepeoples, the most strikingexampleof the influenceof diseaseuponhistory,of which we can speakwith any certainty." The nativepeoples of theAmericashadlittle immunityto the diseasesEuropeans brought with them:the preexistingbiologicalequilibrium upset,with devastating was human who hadservedin the Philippines,worriedthatwhitesmight Ashburn, consequences. yet meet theirmatchif exposed to the "yellow man,"who was even more immunoin of logically competent,the resultof "thegeneraldearth sanitation Asia, andthe unrivaledopportunities ingestinga neighbour'sdung." for Ashbur's understanding of the natural of infectiousdiseaseled him to wonderif, in the comingcenturies, history it would be "machinery science or immunityto disease that will most influence or racialdominance?"28 Disease history,usually shornof its raciallinks, would continueto appealto bioat the logicallyinclinedhistorians, firstthrough Annalesschool andthen,in the 1970s, throughAmericanhistoriansseeking a broadconceptualframeworkfor ambitious surveys.In The Columbian Exchange,for example,AlfredW. CrosbyJr.expresseda desireto understand man above all as a "biologicalentity." Inspiredby Ashburnand inZinsser,andciting HenrySigeristandGaldston,Crosbydescribedhow European vadershad disrupted ecological stabilityof the New World,spreadingdisease to the vulnerable He the of populations. charted transfer plants,animals,andgermsbetween EuropeandtheAmericas,arguingthatthis "Columbian exchangehas left us with not
26 Simon Flexner, "Experimental Epidemiology,"Journal of ExperimentalMedicine 36 (1922): 9-14, on 9, 11. 27 Hans Zinsser,Rats, Lice, and History:A Studyin Biography(1934; reprintedBoston, 1963), 16, 7. DuringWorldWarI, Zinsserhad been a memberof the Red Cross SanitaryCommissionin Serbia, organized to control typhus by RichardP. Strong, later professor of tropical medicine at Harvard. to See Strongwas a minorcontributor the developmentof disease ecology at Harvard. Tilley, "Ecologies of Complexity"(cit. n. 18). 28 P. M. Ashburn,TheRanksof Death: A Medical History of the Conquestof America (New York, 1947), 5, 211, 212. Most of the book was finishedbefore 1937.

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A a richerbut a more impoverished genetic pool."29 few years later,WilliamH. McNeill also triedto applythetheoriesof SmithandZinsserto the historicalstudyof disease outbreaks. McNeill was concernedabouthumanpersistencein "tampering with Since WorldWarII, a new generationof disease ecological relationships." complex Macfarlane BurnetandRene Dubos,hadbeen warningof the ecologists, in particular of population and biologicaldangers growth,biologicalwarfare, environmental degradation.Like them, McNeill fearedthat "a sequenceof sharpalterationsand abrupt oscillationsin existing balancesbetween microparasitism macroparasitism and can be therefore expectedin the nearfutureas in the recentpast."30
MACFARLANE BURNETAND THE NATURAL OF HISTORY INFECTIOUS DISEASE

In Melbourne, Australia, Buret, a youngvirologist,was duringthe 1930s,Macfarlane the worksof Theobald SmithandHansZinsserwithgreatinterest. reading Growing up in rural Burnet takendelightin natural had a whichwouldremain source Victoria, history, for of pleasure therestof his life. In his youth,he becameanavidcollectorof beetlesand in Whilea medicalstuparticipated enthusiastically the local nature studymovement.31 dentat theUniversity Melbourne, continued beetlingandfurther of he his his developed underthe tutelageof Wilfred Agar,theprofessor zoology. E. of biologicalinclinations Agar,an experton the role of chromosomesin heredity,had become increasingly with wide-ranging preoccupied biological speculation. Inspired his readingof Alby fredNorthWhitehead, professorhadfashionedhimselfas an opponentof physicthe ochemicalreductionism an advocateof vitalismandholism. "Surelyit is not sciand he all entific," pleadedin 1926,"toinsistthatbiologistsmustinterpret theirphenomena in the same termsas those foundsufficientin physics andchemistry."32 Familiar with Gestaltpsychology and new work in ecology, Agar transmitted enthusiasmsto his Burnet. Agar,as forBurnet, organisms, For evenmiall manyof his students, including itself."33 crobes,were "linksin the causalprocesswhichis the courseof nature As a young doctor,beginninghis researchon animalviruses and bacteriophages, Burnet later by H. G. Wells, Huxley, and G. P. Wells.34"In one way or another,"
CrosbyJr.,The ColumbianExchange:Biological and CulturalConsequencesof 1492 (Westport,Conn., 1972), xiii, 219. 30William H. McNeill, Plagues and Peoples (New York, 1976), 254, 257. 31 TomGriffiths, Hunters Collectors: and inAustralia (Melbourne, 1996). TheAntiquarian Imagination 32 W. E. Agar, "SomeProblemsof Evolutionand Genetics,"Presidential Address, Section D.-Zoology,"in Reportof the 17thMeetingof theAustralasianAssociationfor theAdvancement Science, of Adelaide, 1924 (Adelaide, 1926), 347-58, on 358. Whitehead'sargumentsfor creativeforce, interrelations, and process in natureinfluenceda numberof ecologists in the 1930s: see his Process and Reality:An Essay in Cosmology(New York, 1929), 127-97. 33W. E. to Agar,A Contribution the Theoryof the LivingOrganism(Melbourne,1943).Agarrepeatedly cites RaymondPearl, D'A. W. Thompson,and C. M. Child.Agar's interestin populationpolicy and eugenics also infected Burnet.See Agar, "Some EugenicAspects of Australian PopulationProblems,"in ThePeoplingof Australia,ed. P. D. Phillips andG. L. Wood (Melbourne,1928), 128-44. 34 Julian Huxley, Essays of a Biologist (London, 1923); and H. G. Wells, JulianHuxley, and G. P. New York,1931). See F. M. Burnet,ChangingPatWells, TheScience of Life, 2 vols. (1929; reprinted terns: An Atypical Autobiography (Melbourne, 1968); F. J. Fenner, "Frank Macfarlane Burnet, 1899-1985," BiographicalMemoirsof Fellows of the Royal Society 33 (1987): 100-62; and ChristopherSexton, TheSeeds of Time:TheLife of Sir MacfarlaneBurnet(Melbourne,1991). Burnetalso recalled (ChangingPatterns,75-6) the influenceof SinclairLewis, Arrowsmith (London, 1925).
29 Alfred W.

Burnet also read Julian Huxley's Essays of a Biologist, and all of The Science of Life,

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WARWICK ANDERSON

me recalled,Wells, Huxley,andWells "brought into biological science andprobably gave thatecological slantto the studyof humandisease which I thinkis characterisH. tic of whatI have written."35 G. Wells andhis coauthors providedan extensiveaccount of the new science of ecology, which they definedas "theobservationof aniand and malsin theirpropersurroundings of theirnormalinteraction ways of life, and what one may call field physiology."36 They described the gradualtransitionon the fromdevastation the "richness tropicalclimax vegetation," importo of Krakatoa tance of parasitechains and food chains, the life cycle of lemmings,the biological of deer in New controlof pricklypearin Australia,and the proliferation introduced for man and aniZealand.Accordingto TheScience of Life, "unrestrained breeding, malsalike,whethertheyaremice, lemmings,locusts,Italians,Hindoos,or Chinamen, evil of is biologicallya thoroughly thing." Burnetwouldtaketo heartexplanations the for Life,""thetangledweb of interrelationships," the dangersof overand "struggle Wells warnedthat crowdingandhumanmobility.37
but freedomof intercourse communication and bothtradeandthought; it gives stimulates disease-germsnew facilitiesfor rapidspreading .... Thusthe growthof civilizationhas been markedby a trailof plagues, more explosive and more widespreadthan anything which primitivemancan haveexperienced.38

Yetin its treatment healthanddisease, TheScience of Lifemostlypresenteda conof ventionalversionof bacteriology. Buret musthave been disappointed, the book but him to thinkmoreaboutthe ecology of the viruseshe was cultivating wouldstimulate in the WalterandEliza Hall Instituteduringthe 1930s. At the Hall Institutein Melbourne,Burnetcompleted his studies of the interactions of bacteriaand bacteriophage,stimulatedduringan earliervisit to the Lister Institutein London,and examinedthe behaviorof viruses, especially the influenza virus, in the chick embryo. In 1935, an opportunityto investigatepsittacosis confirmedhis growing interestin biological aspects of infectious disease. Burnetwas able to demonstratethat asymptomaticinfection with psittacosis was common among wild parrots,but when the birds were confined and stressed, the illness would become manifest.39Such latent, inapparentinfection drew Burnet into a broaderconsiderationof disease ecology and immunity.The small scale of the medicalresearchenterprisein Australiawould frequentlycompel the Hall Institute to take on a microbiologicalservice role for government,much of which involved field studies of disease outbreakssuch as psittacosis. As a consequence, Burnet, was neverable to limit his researchto thougha brilliantlaboratory experimentalist, bench work. His presence in a "peripheral" settler society-a place preoccupied with populationproblemsand its fragile environment-and the occasional need to
35 F M. Bumet, "Life's Complexities:Misgivings aboutModels,"AustralianAnnals of Medicine4 (1969): 363-7, on 364. William B. Provine argues that "Huxley's discussion of evolution was the of single most encompassingpresentation a neo-Darwinianviewpoint availablein 1930,"and he suggests that "theinfluence of The Science of Life on scientists and the educatedpublic deserves careful in studyby historians" ("England," TheEvolutionarySynthesis:Perspectiveson the Unificationof BiMass., 1980], 332). ology, ed. Erst MayrandWilliam B. Provine [Cambridge, 36 Wells, Huxley, andWells, Science of Life (cit. n. 34), 1:22. 37 Ibid., 2:974, 1012, 1010, 1012. 38 Ibid., 2:1016. 39 F M. Bumet, "Enzootic Psittacosis Journal of Hygiene 35 amongst Wild AustralianParrots," (1935): 412-20.

NATURALHISTORIESOF INFECTIOUSDISEASE

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undertakefield microbiology led him repeatedlytowardan ecological perspective on disease.

In 1937, Buret began to write Biological Aspects of Infectious Disease, a book de-

He that disease reader. wantedto demonstrate "infectious signedfor the nonspecialist canbe thoughtof withprofitalongecologicallines as a strugglefor existencebetween manandmicro-organisms." is, he would arguethattherewas a natural That
conflictbetweenman andhis parasiteswhich, in a constantenvironment, would tendto result in a virtualequilibrium,in which both species would surviveindefinitely.Man, however,lives in an environment constantlybeingchangedby his own activities,andfew of his diseaseshave attainedsuch an equilibrium.40

Buret regarded workas a combination epidemiology immunology his of and viewed from a wide biological perspective.Examininga numberof common diseases, he of soughtto providean evolutionary explanation the relationsbetweenhumanpopulations andtheirparasites. Infectiousdiseasewas, he claimed,nothingmoreor less than of "a manifestation the interaction living beings"in a changingenvironment. of Processes suchas migration, and increase thuswouldlead urbanization, generalpopulation of to redistribution old diseasesandthe emergenceof "new"diseases."Wars, internal andexternal,financial and he of depressions labourtroubles," wrote,"areall breeders infectiousdisease,andthefuture diseasewill dependon theessentiallyfortuitous of circumstances which will let loose or withhold these calamities."41 conclusion, Buret In

warned(perhaps of the of presciently) the dangersof biologicalwarfare, deployment germssuchas anthrax. Invitedto give the EdwardK. DunhamLecturesat Harvard 1944, Burnetdistinin guished his recentwork from those disciplines "whichdeal essentiallywith the orchemicaland ganismin isolationandwhich are chiefly concernedwith its structure, and with its functioningas a single unit."Rather,he associatedhis morphological, own researchinterestswith "thesciences whose objectis to interpret controlthe and of living things as they are, in such environments the presenceof naas phenomena tureand the activitiesof mankindhave allottedthem."Burnetwantedto understand how organismsare "distributed space and time"and the "long-term in historicalasof the interaction betweenorganismandenvironment."42 hadbegunto view He pects himself as a contributor the new evolutionarysynthesis that Julian Huxley deto scribedin the early 1940s. The synthesiswas initiallya combination Mendelism, of chromosometheory,mathematical studiesof heredityin populations, natural seand lectionism;a consequenceof the revivalof Darwinismin the 1930s, it broughtexperimentalgeneticiststogetherwith naturalists who studiedevolutionin the field.43 a In
sense, then, Burnet was extending the synthesis to microbiology and immunology.
Buret, Biological Aspects of Infectious Disease (cit. n. 3), 3, 23. See also idem, "Changesin Twenty-FiveYears in Outlook on Infectious Disease," The Medical Journal of Australia (hereafter MJA)2 (1939): 23-8; and idem, "CharlesMackayLecture:Biological Approachesto InfectiousDisease,"MJA2 (1941): 607-12. 41 Buret, BiologicalAspects of InfectiousDisease (cit. n. 3), 4, 307. 42 F. M. Bumet, Virus Organism:Evolutionaryand EcologicalAspectsof SomeHumanVirus as Diseases (Cambridge,Mass., 1945), 3. 43 MayrandProvine,EvolutionarySynthesis(cit. n. 35). See JulianHuxley,Evolution:TheModern Synthesis(London, 1942); R. A. Fisher,The Genetical Theoryof Natural Selection (1930; reprinted New York, 1958); and Theodosius Dobzhansky,Genetics and the Origin of Species (1937; reprinted New York, 1951).
40

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WARWICK ANDERSON

Huxley recognized Burnet'scontributionin the second edition of Evolution:The ModernSynthesis,thoughhe emphasizedmostly the researchon microbialgenetics rather thanthe morespeculativeecological modelingof disandantibodyproduction ease processes.44 to moderBurnet'santipathy facile reductionism his discomfortwith industrial and in nity became more pronounced latereditions of his surveyof disease ecology, retitled the Natural History of Infectious Disease. In 1953, he lamented that "the older

as of who weremostlytrained medicalmenhasnow almost generation bacteriologists The had as been replacedby workerstrainedprimarily biochemists." new generation little or no interestin ecological aspects of infectiousdisease. The adventof antibithe otics had furthernarrowed focus of medicalresearch,and simplifiedthe clinical encounter.In this and later editions, Buret, an admirerof Malthus,amplifiedhis and numbers, a declinein population quality,as warningsof an increasein population infectiousdisease became a less significantsocial factor.He was also more worried duringthe cold war that the great powers would resortto biological warfare.The was, he wrote,"astrangeandgloomy endingof this account prospectof germwarfare of infectiousdisease."45 of the natural history In 1972, David O. White,the professorof microbiologyat the Universityof Melbourne,joined Buret as the authorof the fourthedition of the NaturalHistory of InfectiousDisease, but the conceptualframeworkhardlyaltered.This last edition includedsome additionalintroductory chapterson susceptibilityand resistance,the of transmission infection,evolutionandthe survivalof host andparasite,antibiotics, andhospitalinfectionsandiatrogenicdisease.The antireductionist themecontinued, with the disparagement genetics andthe new molecularbiology. "Thefasciof along nationwith molecularbiology and its implicationsat all levels of biology has persisted,"Buret andWhite wrote, "andamongstbacteriologistsand virologiststhere has been a trendaway from what may be called the ecological aspectsof infectious disease."In the fourthedition, ecology was again definedas "theinteractionof orand ganismswith theirenvironment especially with otherorganismsof theirown or 46 differentspecies in thatenvironment." More thanever before, the text was strewn with termssuch as "climaxstate," and "niche," "virgin-soil epidemics," "ecosystem." Thus "everyorganismis itself a productof evolutionand ecologically relatedto almost every otherorganismin the ecosystem."Urbanization, and overcrowding, humanmobilitywere causingever-greater to suchecosystems.All the same, disruption antibioticsseemed in the 1970s to have controlledinfectiousdiseases, whose future looked dull. "Theremay be some wholly unexpectedemergenceof a new and dangerous infectious disease, but nothing of the sort has marked the last fifty years."47

Buret andWhiteremainedconcerned,however,thatnuclearor bacteriological warfare was imminentand thatmoderncivilizationwas environmentally unsustainable. "Thereis unease everywhereamongstresearchmicrobiologistswith a social conJulianHuxley,Evolution:TheModernSynthesis,2d ed. (London, 1963), xxx-xxxi, 1. F. M. Buret, NaturalHistory of InfectiousDisease, 2d ed. (Cambridge,1953), 3, 351. The third edition, publishedin 1962, was little altered. 46 F. M. Burnet and David O. White, Natural History of Infectious Disease, 4th ed. (Cambridge, 1972), 3, 4. 47 Ibid., 138, 263. During the 1960s, Burnetalso explained that "communicabledisease has more than a human significance;it is also a manifestationof the way of life of anotherorganism,the responsiblepathogen"("TheNaturalHistoryof InfectiousDisease,"in TheTheoryand Practice of Public Health, 2d. ed., ed. W. Hobson [London, 1965], 121).
44

45

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science,"they noted.It seemedthat"theadvanceof science is allowingthe developmentof wholly unnatural modes of humandominationandmass destruction."48 Buret andWhitepointedto immunologyas one of the few branchesof "medical" researchto embraceecological approaches. "Themedicalaspectsof immunityhave movedawayfromthe centreof scientificinterestas in the last ten yearsimmunology has begunto developits full statusas partof fundamental the biology."49 Apparently, stimulusto this development Burnet'sclonal selectiontheoryof antibodyformawas of tion, andhis explanation immunologicaltolerance,which hadearnedhim and Sir PeterMedawarthe 1960 Nobel Prize for physiology or medicine.In the late 1950s, Bumet had encounteredNiels Jere's "selective"hypothesis,which stated that an antigenwould combinewith the best fit amongthe organism'sdiversenatural globulins andtransport to antibody-producing it cells, readyto makemultiplecopies of the this presented globulin.Buret reconfigured idea, arguingthatit wouldmakesense if cells produceda patternof globulinfor genetic reasonsand the arrivalof the correcausedthe proliferation the appropriate of spondingantigenicdeterminant globulin. The theoryimplieda geneticpolymorphism the lymphocytes,with a varietyof surof face receptorsand coding for antibodyglobulins arisingfrom somatic mutationor from some otherprocess occurringin differentiation development.50 "bioand This LinusPauling'searlier"instructive" whichsuglogical"conceptreplaced hypothesis, gestedthatthe antigenacteddirectlyas a templatefor antibody production-this idea hadbeen fartoo "chemical" Burnetto countenance The clonal selectiontheory for it. would come to represent culmination Burnet's"ecological"thinking. the of "Inmore senses thanone I have always been a humanbiologist,"Buret declared in 1966, "andI knowthatto look at manas a product evolution,as partof the whole of has given me some useful insightinto my own characteristics diffiand living world, net's lexicon "biological," "ecological,"signaledattentionto an integrative, or Darwinian conceptualframework;a biological approachdid not thereforeimply the abandonment reductionist of methodsso muchas theirintegration a dynamicand into interactive modelof therelationsof organism environment. and Buret generallyused his interpretation prewarecology as a metaphoric of resourceratherthanan analytic tool. AfterBuret readWells andHuxleyandstudiedwithAgar,his understanding of a form.Despitetheoccasionalreference ecology hadacquired moreorless permanent
to "ecosystem" in the fourth edition of the Natural History of Infectious Disease, he
Buret andWhite, Natural History of InfectiousDisease, 4th ed. (cit. n. 46), 266, 267. Buret in particular hopedthatethology mightprovidesome insightinto humanaggressionandsuggest a means of channelingit into more constructiveactivities. He communicatedextensively with KonradLorenz andfrequentlyvisited him. 49BurnetandWhite, NaturalHistory of InfectiousDisease (cit. n. 46), 31. 50F. M. Bumet, "A Modificationof Jere's Theory of Antibody ProductionUsing the Concept of Clonal Selection,"AustralianJournal of Science 20 (1957): 67-9; and idem, The Clonal Selection (Nashville, 1959). For an earlierversion of the theory,see F M. Buret TheoryofAcquiredImmunity andF. J. Fenner,TheProductionofAntibodies,2d ed. (Melbourne,1949). See also JoshuaLederberg, "The Ontogeny of the Clonal Selection Theory of Antibody Formation:Reflections on Darwin and Annals of the New York Ehrlich," Academyof Sciences 546 (1988): 175-87; ArthurSilverstein,His(San Diego, 1989);Anne-MarieMoulin, Le derier langage de la medecine:History of Immunology toire de l'immunologiede Pasteurau Sida (Paris, 1991); andEileen CrustandA. I. Tauber, "Selfhood, Immunity,and the Biological Imagination:The Thoughtof FrankMacfarlaneBuret," Biology and Philosophy 15 (2000): 509-33. 51 F M. Burnet,Biology and theAppreciationof Life. Boyer Lectures 1966 (Sydney, 1966), 1.
48

culties, and added a special intensity to my appreciation of the world as it is."51In Bur-

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the appearsto have disregarded mathematical modelingand systems theoryof postwar ecology.52 Indeed,althoughmanyof the scholarswho shapedtwentieth-century with them.53 ecology workedin Australia,thereis no evidencethatBuret interacted ForBurnet,the term"ecology"continuedto suggesta generalappreciation natural of of processes,the transcendence merephysicochemical complexityand evolutionary mechanism;he would use some older ecological insights,but he did not embarkon systematicecological research.
RENE DUBOSAND MANADAPTING

In 1943, Rene Dubos, the professorof comparative had pathologyat Harvard, recommendedBuret as the next Dunhamlecturer the medicalschool. Dubos,though at he had not met the Australianscientist,was interestedin Buret's views on disease After doctoralstudiesin bacteriologyat Rutgerswith SelmanWaksman, ecology.54 Dubos had investigatedthe propertiesof soil microorganisms, on concentrating the role of competition,and its biochemicalmediators,in limitingpopulations.55 Later, settledatthe RockefellerInstitute, extracted bacteriostatic he a substance, gramicidin, fromBacillus brevis,anddemonstrated use in the treatment externalinfections. its of Such investigationsof soil microbiologyand bacterialcompetitionquickly attuned Dubos to an interactive ecological vision of humanhealthanddisease.56 and Dubos even found supportfor his emergingecological views from Surprisingly, 0. T. Avery,his mentorat the RockefellerInstitute,who otherwisepreferred conto ductcloisteredlaboratory studiesof immunochemistry bacterialtransformation. and Dubos thoughtthatAvery'sworkhad little influenceon medicalpractice;rather, he to contributed "theunderstanding biological phenomena." of "WhenAverybecame in interested a biologicalphenomenon, firstobservedit for the sheerfun of it, as a he naturalist."57 studyof microbiologyshouldbe morethana meremedicalinstruThe ment;it shouldalso help to illuminatea wide rangeof biological problems.The scientist should thereforeacquirea broadbiological perspectiveon disease processes. between Accordingto Dubos, his seniorcolleague was convincedthat"theinterplay the life processesof the host andthose of the parasitewas the way of the future."58
52 See, e.g., W. C. Allee et al., Principles of Animal Ecology (Philadelphia, 1949); and Eugene Odum,Fundamentalsof Ecology (Philadelphia,1953). For more on the historyof ecology, see Gregg and Mitman,TheState of Nature:Ecology, Community, AmericanSocial Thought,1900-1950 (Chicago, 1992). 53See A. J. Nicholson, "TheBalance of Animal Populations," JournalofAnimal Ecology 2 (1933): and 551-98; and H. G. Andrewartha L. C. Birch, The Distributionand Abundanceof Animals (Chicago, 1954). The leading Australianecologists are discussed in SharonE. Kingsland,Modeling Nature:Episodes in theHistoryof PopulationEcology (Chicago, 1985), 116-23, 171-2; andMartinMulligan and StuartHill, Ecological Pioneers: A Social History of AustralianEcological Thoughtand Action (Cambridge,2001), chap. 7. 54 AlexanderN. Zabusky,"Ecological Odyssey: The IntellectualDevelopment of Rend J. Dubos" (seniorhonorsthesis, Harvard University,1986). Dubos was at Harvard only between 1942 and 1944; otherwisehe workedat the RockefellerInstitute. 55BernardD. Davis describesWaksman,who was awardeda Nobel Prize for his discoveryof strepa historianof the soil, cataloguingthe micro-organisms foundthere,and tomycin,as "primarily natural focusing on theirtaxonomyandtheirecological effects" ("TwoPerspectives:On RendDubos, and on AntibioticActions,"Perspectivesin Biology and Medicine 35 [1991]: 37-48, on 40). 56 Barbara GutmannRosenkrantz, "ReneJules Dubos,"AmericanNational BiographyOnline,Feb. 2000, http://www.anb.org/articles/12/12-01795.html. 57 Dubos, TheProfessor,the Institute,and DNA (cit. n. 2), 89, 113. 58Ibid., 100.

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In the early 1940s, Dubos beganto formulatea more complex biological account of host-parasite relationships, drawingcertainlyon TheobaldSmith'spreviouswork andprobably Bumet'srecentmonograph. TheBacterialCell,publishedin 1945, on In Dubos described
the organization those moleculargroupingswhich, on accountof theirchemical acof unitandin its tivity,conditionthebehaviorof the cell bothas an independent functioning of relationto the environment. ultimateunderstanding the natural The historyof infectious diseases, and the rationaldevelopmentof methodsfor theircontrol,dependupon this knowledge.59

It was an effortto place bacteriain an evolutionary time frame,reporton bacterial and mutation,and indicatethe biochemicalmanifestations organismic of variability Dubos claimed that in the past, microbiologistshad narrowlystudied competition. 'events'rather thanbacteria themselves." Theseresearchers were"al"bacteriological mostuninfluenced the doctrines methodsof classicalbiology." and Dubossoughtto by he remedythis deficiency."Itis necessary," declared,"to abandonthe anthropomorwhich characterized attitude earlierefforts;bacteriamustbe studied,not only in phic theeffectswhichtheyhaveon practical human but problems, also forwhattheyareand whatthey do as independent living organisms."60 and Dubos,however,was gradually movingaway fromlaboratory investigation rehimselfas a popular In writerandcommentator. a seriesof books,he would fashioning socialandbiologicalunderstanding human of disease.61 His arguefor a moreintegrative dominant themewas that"thestatesof healthanddiseasearetheexpressions the sucof cess or failureof theorganism its effortsto respond in to chaladaptively environmental Like Bumet,he believed"organismic environmental and neededas lenges." biology" muchattention "physicochemical as He withthe enorbiology." wrote,"Incomparison mous effortdevotedto the components the body machine,living as a processhas of beenstudied scientific In methods."62 bookssuchasMirageof HealthandMan hardly by Dubossoughtto explore"thecomplexinter-relationships betweenmanandhis Adapting, and wouldhavebeen physicochemical biologicalenvironment."63 of his arguments Many familiarto readersof Bumet's work:the "interplay" betweenorganismsreachingan the betweenparasitism predation, determinants bacterial the and of equilibrium, balance
virulence and host resistance, the impact of increasing population density, and the general evolution of microbial diseases. However, Dubos, unlike Buret, would increasingly focus on more direct physical influences on human health, referringback to Hippocrates and medical geography and pointing to the dangers of environmentalpollution.
59Ren6 J. Dubos, The Bacterial Cell: In its Relation to Problems of Virulence,Immunity,and Mass., 1945), 17. The book was based on a series of lecturesdeliveredin Chemotherapy (Cambridge, 1944 at the Lowell Institutein Boston. 60 Ibid., 339, 342. See also Rene J. Dubos, "Utilizationof Selective MicrobialAgents in the Study of Biological Problems," Bulletinof the N.Y.Academyof Medicine 17 (1941): 405-22; idem, "Microin biology,"AnnualReviewof Biochemistry11 (1942): 659-78; andidem, "Trends the StudyandControl of InfectiousDiseases,"Proceedings of theAmericanPhilosophical Society 88 (1944): 208-13. 61 The firstof these was Ren6Dubos andJean Dubos, TheWhitePlague: Tuberculosis, Man, and SoNew Brunswick,N.J., 1987), which describesthe impactof povertyand waron ciety (1952; reprinted the incidence of this "social disease." 62 Dubos, ManAdapting(cit. n. 3), xvii, xix, xx, 333. 63 Ibid., xxi. See also Ren6 J. Dubos, Mirage of Health: Utopias, Progress,and Biological Change (London, 1959).

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Although some of Dubos's and Buret's interestsand approachescontinuedto overlap,theirpathsbeganto divergeas well, as Dubos emphasizeda critiqueof capitalism and civilization.In his work, monitorycommentoften overcameecological traditional: scruples.His message was becomingever-more
To a surprising unaltered bodily constitution, the extent,modem manhas retained physfromhis Paleolithicaniological responses,andemotionaldriveswhichhe has inherited cestors.Yethe lives in a mechanized, and difworldradically air-conditioned, regimented ferentfromthe one in which he evolved.64

Dubos's condemnationof "the pathology of urban and suburbanlife, of antiresphysiological leisure in a mechanized,automatedand crowdedenvironment," onatedwith the sensibilityof the 1960s protestmovements,and he soon became a of He prophet the "counterculture."65 beganto talkmoreaboutthe moralcost of adaptationto degraded conditions."Alltoo often,"he lamented,"thebiologicalandsocial to changesthatenablemankind overcomethethreats posedby the modemworldmust be eventuallypaidfor at a cruelpricein termsof humanvalues." thereAdaptability, fore, was "oftena passive acceptanceof conditionswhichreallyarenot desirablefor mankind." This led Dubos to arguethat "thebiological view of adaptation inadeis that quatefor humanlife"-a statement Burnetwas unlikelyto utter.66 Dubos continuedin much the same vein throughout 1960s and the 1970s. In the 1968, he warned,"[M]anwill ultimatelydestroyhimself if he thoughtlesslyeliminatesthe organismsthatconstituteessentiallinks in the complexanddelicateweb of life of which he is a part." he himself as an heir of the HipIncreasingly, represented time, good medicalcare implies attenpocratictradition: "today,as in Hippocrates' tion not only to the body butto the whole personandto his total environment."67 Alto of disease,Dubos thoughhe was now callingattention his "holistic" understanding still arguedthatthis generalconceptualframework to be informedby preciselabhad deteroratoryknowledge.He triedto resist, not always successfully,environmental minismandto hold on to a moreinteractive, model. "Allnatural ecological phenomena,"he wrote, "arethe result of complex inter-relationships; manifestations all of humandisease are the consequencesof the interplaybetweenbody, mind, and environment."68 increasingly, also wantedto condemnthe damageindustrial But he capitalismwas doing to "human radicalhumanism his ecologito values,"counterposing cal sensibility."Medicalproblemsposed by the environmental stimuliandinsultsof modem civilizationhave acquireda criticalurgency," asserted.Indeed,his main he worryhad become "thethreatto mankindposed by technologiesderivedfrommodem physicochemicalandbiological sciences."69
FRANKFENNERAND THE ECOLOGYOF VIRUSERADICATION

FrankFenner,an Australianmicrobiologist,worked in Dubos's laboratoryat the RockefellerInstitute 1948 and 1949.Therehe studiedtheproduction BCG, since in of
64Dubos, ManAdapting(cit. n. 3), xviii. See Rosenberg,"Civilizationas Risk"(cit. n. 15).
65 66

Dubos, ManAdapting(cit. n. 3), 252. Ibid., 275, 279. 67 Ren6 J. Dubos, Man, Medicine,and Environment (New York, 1968), 9, 61. 68 Ibid., 61.
69

Ibid., 88, 111.

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Dubos was thinkingmoregenerallyabouttuberculosis the time.70 afterWorld at Just WarII, Fennerhad gone to the Hall Institutein Melbourneto investigatethe experimentalepidemiologyof infectiousectromeliaof mice and to conductresearchmore generallyon poxviruses.It was Buret who suggestedthatFennermight appreciate to workingwith Dubos. Fennerwas, in a sense, preadapted this intellectualterrain. While studyingmedicine at the Universityof Adelaide, he attendedthe lecturesof J. B. Cleland,the professorof pathologyandlaterauthorof an essay on the naturalist in medicine,andC. StantonHicks, the professorof physiology and a pioneerecoloAs himselfto the SouthAustralian MuFennereven attached gist.71 a medicalstudent, seum as its firstandlast "craniologist"-a joke title, he laterrecalled.Workingin an of oldertradition the natural historyof man,Fennerhadused his skullmeasurements to challengeprevailingtheoriesof Aboriginalhomogeneity, insteada hypostulating bridorigin.As therewas no futurein craniology,duringthe warhe retrained a mias a diplomaof tropicalmedicine,and then treatingmalaria, crobiologist,completing scrubtyphus,and denguein PapuaNew Guinea.Throughout subsequentcareer, his Fennerwouldretainhis interestin natural andin the relationsof humansand history theirenvironment, environment.7 especiallythe animated In 1949, Fennertookup the foundation chairof microbiologyat the newAustralian NationalUniversity, thoughhe continuedfor a few yearsto workat the Hall Institute on Mycobacterium ulceransandits peculiarterrestrial relations.In the 1950s,as myxa poxvirus,was introduced the rabbit into of omatosis, Australia, population southern Fennerdecidedto investigateevolutionary in the organism'svirulenceandin change the resistanceof rabbitsto it. It was, he believed,"thebest natural on experiment the co-evolutionof viral virulenceand host resistanceavailablefor a disease of verteAt brates."73 first,99 percentof rabbitsdied from infection with the myxoma virus. The virus,however,actedas a powerfulselectionpressureandthe next generation of rabbitswas far less susceptible.In this researchproject,Fennercollaboratedwith FrancisRatcliffe,a zoologist at the CSIRO.Ratcliffehad trainedwith CharlesElton andJulianHuxleyin Englandbeforeventuring Australia to whereinitiallyhe studied the distribution the fruit-eating of and "flying-fox" laterthe problemof soil erosionin the outback.74 Fenner'secological orientationwas confirmedand enrichedthrough his associationwithRatcliffe,who since the 1940shadbeen seekingbiologicalmeans to controlrabbitpopulations.75
70Rene J. Dubos, F. J. of Fenner,and C. H. Pierce, "Properties a Cultureof BCG Grownin Liquid Media ContainingTween 80 and the Filtrateof HeatedSerum," AmericanReviewof Tuberculosis 61 (1950): 66-76; and F. J. Fenner,"Bacteriologicaland ImmunologicalAspects of BCG Vaccination," Advances in Tuberculosis Research4 (1951): 112-86. 71 On the environmentalist and sometimesneo-Lamarckian of the Adelaide medical school, see cast WarwickAnderson, The Cultivationof Whiteness:Science, Health, and Racial Destiny in Australia Refer(New York,2003), chap. 7. See also J. B. Cleland,"TheNaturalistin Medicine, with Particular ence to Australia," MJA 1 (1950): 549-65; and C. StantonHicks, Soil, Food and Life (Melbourne, of 1945). Fenner'sfather,Charles,a geographerwho earnedhis living as superintendent educationin SouthAustralia,was perhapsa more importantinfluence. 72 F J. Fenner,"Nature,Nurture,and My Experience with Smallpox Eradication: CareerInfluA enced by ChanceEvents," MJA171 (1999): 638-41. Whenhe was at the Hall Institute,Fennerbecame thejunior author,with Bumet, of the second edition of TheProductionof Antibodies(cit n. 50). 73 Fenner,"Nature,Nurtureand My Experience with Smallpox Eradication" (cit. n. 72), 639. See also idem, ed., History of Microbiology in Australia (Canberra, 1990).
74 F. N. Ratcliffe,Flying Foxand DriftingSand:TheAdventures a Biologist inAustralia(London, of to 1938). JulianHuxley wrote the introduction this book. 75 F. J. Fennerand F. N. Ratcliffe, Myxomatosis(Cambridge,1965).

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administrative dutiestook Duringthe 1960s,the writingof booksandaccumulating Fennerawayfromthe laboratory The bench.76 Biology of AnimalViruses,a majorrevision of Buret's Principles of Animal Virology, was Fenner'sfirst attemptto describein detail the "broader In biological principlesof animalvirology."77 a chapter on the transmission viruses, he arguedthat "theepidemiologyof infectious disof eases is a branchof ecology concernedwith the spreadand survivalof infectious on He agentsin nature." reported routesof entryintothe body andexit fromit, the differencesbetween single-hostand multiple-hostviruses, variationsin immunity,the influenceof weatherconditionson disease manifestation, the importance exand of Fenner viruses," (for which he cited Greenwood)."Animal perimental epidemiology In concluded,"survivein natureas partof an ecosystem."78 a laterpartof the same he consideredthe evolution of virulence and host response. Focusing on chapter, myxomatosis,influenza,and dengue,he emphasizedthe processes of naturalselecand tion, adaptation, genetic drift. Even in 1968, Fennerwas concernedthatincreasinghumanmobilityandenvironmentalalterationwould lead to the emergenceof new diseases. His warningswere amplifiedin the secondeditionof TheBiology of AnimalViruses,publishedin 1974. "Man'sdisturbance the biosphere,and some medicalinnovations," of wrote Fenner andhis coauthors,"havecreated'new' viraldiseases for manhimself andfor his doIt mesticatedanimals." was clearby thenthat"thegreatincreasein the numbers huof manbeings, theircrowdinginto ever largercities, andthe increasingcommunication world by menbetweenthesecities all overthe world,aretendingto makethe 'human' into a single ecological unit."79 or previouslyunknown,microbesmight proRare, liferatewith rapidityin these circumstances. an especially prescientpassage-in In view of the outbreak bovinespongiform of in encephalopathy Britaina decadelaterFennerand his colleagues suggestedthat "thegreatconcentrations livestock that of characterize'industrial in some Westerncountries,and the widespread agriculture' of shipmentand aggregation largenumbersof animalsfor fattening,providegreatly increasedopportunities the spreadof 'rare'virusesthroughlargenumbersof anifor mals andfor the emergenceof viraldiseasesof livestock."80 the 1970s, Buret may In have thoughtthatthe outlookfor infectiousdisease was still reassuringly but "dull," Fennerwas increasinglypessimistic. The prospectsfor smallpox eradication,however,did seem to improve.Dubos's distastefor humanintervention the environment led him to dismiss effortsto in had eradicatediseases;he urged,rather, theircontrol,by which he meanta "proper, skilful handlingof the ecological situation." assertedin 1965: "Eradication microHe of bial disease is a will-o'-the-wisp;pursuingit leads into a morassof hazy biological This vain pursuitencouraged"the illusion that man can concepts and half-truths."
76 In 1967 Fennerbecame deanof the JohnCurtinSchool of MedicalResearchat theAustralianNational University.From 1973 until 1979, he was directorof the Centrefor Resourceand Environment Studiesthere. 77 E J. Fenner,TheBiology ofAnimal Viruses,2 vols. (New York,1968), 2:v. See also F M. Burnet, 2d Principles ofAnimal Virology, ed. (New York, 1960). Elsewhere,Fennerclaimed thatBumet's BiologicalAspects of InfectiousDisease "shareswith TheobaldSmith'sParasitismand Disease the distinction of being the first attemptto apply ecological principles to infectious diseases" ("Brahma, Shiva,andVishnu:ThreeFaces of Science,"AustralianAnnalsof Medicine4 [ 1969]: 351-60, on 351). 78Fenner,Biology of AnimalViruses(cit. n. 77), 2:760, 758. 79 E J. Fenner,B. R. McAuslan,C. A. Mims et al., TheBiology ofAnimal Viruses, ed. (New York, 2d 1974), 618, 635. 80Ibid., 635.

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to controlhis responsesto stimuliand can makeadjustments new ways of life without havingto pay for these adaptivechanges.""Havingstudiedbiological effortsto eradicate rabbits,Fennerwas not so surethatcompletefailurewas inevitable,though he, too, could be doubtful.Because the germsof malariaand yellow fever had multhese diseases was "biologically tiple hosts, Fennerbelievedany attemptto eradicate unsound." could, however,conceive of the biological possibility of eradicating He smallpox, since the virus was monotypicand had no animalhost, the disease was readilyidentified,immunityafterinfection was life long, and vaccinationwas relaSmallpoxEradication tively easy. In 1968, the yearafterthe launchof the Intensified he still thought"technicaldifficulties"stood in the way of the complete Campaign, His eradication smallpox,butby 1974 he was morehopeful.82 expertisein poxvirus of had of research,as well as his understanding the ecological dynamicsof vaccination, of attracted attention D. A. Henderson, chief of the SmallpoxEradication the Unit the of theWorldHealthOrganization, invitedFennerto serveon the committeeoverwho the eradication of Later,as chairman the GlobalCommissionfor the seeing program. of in 1979, was able to confirmthe disCertification SmallpoxEradication, Fenner, of appearance the disease since 1977.83 In 1968, S. V. Boyden, an immunologistwho had workedwith Dubos and then in at Fenner'sdepartment theANU, organized conferencein Canberra civilization's a on on humanbiology. The meetingbroughtBumet, Fenner,and Dubos together impact again.Boyden,the headof a new "urban biology"group,haddevelopedan interestin the social andbiologicalcausesof humanmaladjustment. was not surprising he It that Dubos in condemning costs of culturaladaptation an unnatural to the civilizajoined tion.Accordingto Boyden,"[W]hen conditionsof life of an animalpopulation the deviatefromthoseto whichit hasbecome,through natural selection,geneticallyadapted, some signs of biological maladjustment almostinevitable."84 are Dubos Predictably, asserted "inapplyingthe conceptof adaptation the humanspecies,it is ... necthat to essary to use criteriadifferentfrom those used in Darwinianpopulationtheory." are Instead,he believed,humansneededto "decidewhatkindsof adaptation compatible with the maintenance desirablehumanvalues."85 of to Buret, in his introduction the symposium,avoidedsuch talk of "human values,"focusing on the changingdistributionof humangenotypes secondaryto civilization.His old enthusiasmfor euincreasedtheir genics neverfarbelow the surface,Bumet worriedthatas populations of quality was declining; the "overbreeding" American"Negroes"especially dismayedhim.86 Fenner provided what was perhapsthe most thoroughly"ecological,"or least of anthropocentric, the papers."Fromthe point of view of infectious diseases,"he featuresof man'sculturaldevelopment the size of the are wrote,"themost important
81 Dubos, ManAdapting(cit. n. 3), 381. See SocratesLitsios, "Ren6Dubos andFredL. Soper:Their Views on VectorandDisease Eradication," Perspect.Biol. Med.41 (1997): 138-49. Soper, Contrasting with the RockefellerFoundation,led the campaignto eradicatemalariain the 1950s and 1960s. 82 Fenner,Biology of AnimalViruses(cit. n. 77), 2:784. 83 F. J. Fenner,D. A. Henderson,I. Aritaet al., Smallpoxand its Eradication(Geneva, 1988). 84 to S. V. Boyden, introduction TheImpactof Civilisationof the Biology of Man, ed. S. V. Boyden (Toronto,1970), xiii-xiv, on xiii. 85 Rene J. Dubos, "TheBiology of Civilisation-With in Emphasison PerinatalInfluences," Boyden, Impactof Civilisation(cit. n. 84), 219-29, on 220, 222. 86 F. M. Bumet, "HumanBiology as the Study of HumanDifferences,"in Boyden, Impactof Civilisation (cit. n. 84), xv-xx.

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of individualcommunities men, the numberandproximityof such communities, and theextentof movementandinterchange betweenthem."87 Fennerdiscussedchangesin in interactions malaria, salmonellosis, cholera,measles,smallpox,yellow host-parasite New viraldiseaseshadbeen"recognized fever,andpoliomyelitis. relatively commonly most but not all of these have been due to humanintervention in duringthis century: some natural or human situation, to changesin the socialhabitsof man." Urbanization, and of the colonization, airtravelseemedespeciallyto promote spread disease.In Fenner'sopinion,the best solutionto the problemof disease was "theeradication the of sourcesof infectionby theelimination poverty."88 discussingFenner's In of S. paper, D. of at warnedof selectionfor antibiotic Rubbo,the professor microbiology Melbourne, resistance of of in amongmicrobes."Theappearance thesenew strains bacteria the affluentsocietiesillustrates unusualeffect of the changingsocial organization the an on In infectiousdiseasesof man."89 response,Fennerdrewhis audience's attention backto the emergenceof "new"diseases."I am surewe will witnessthe appearance antiof novel virusesin the nextfiftyyears,andI do notmeanonly influenza viruses; genically of possiblywe mayalsowitnesstheappearance virusesof novelpathogenic potential."90
CONCLUSION

While some othermedicalinvestigators the twentiethcenturyhadturnedto sociolin ogy to gain a widerperspectiveon healthand disease, advocatesof disease ecology drewon theirknowledgeof biologicalprocesses.Forthem,even the social mightbecome a figmentof biology. Manyof these scientistshadtrainedin tropicalmedicine, science and were thus preadapted seeing the to veterinary pathology,or agricultural broaderbiological determinants apparently of specific or idiosyncraticpathological events.91 Commonly,they had acquiredin youth an enthusiasmfor naturalhistory, whichcontinuedto inflecttheirmedicalstudies;andtheproblemstheyinvestigatedTexas cattle fever, psittacosis,soil microbiology,myxomatosis-tended to confirm theirecologicalinclinations. Residentin settlersocieties suchas the UnitedStatesand to Australia, pioneerdisease ecologists were especiallyattuned the persistingimpact of colonialdevelopment policies, to the lastingeffects of agricultural changeandhumanresettlement. neverlost his interestin the qualityandquanBurnet,in particular, fostered by the intense debate in the tity of populations,a eugenic preoccupation 1920s overAustralian Disease ecology populationpolicy-or "carrying capacity."92
87 F. J. Fenner,"TheEffects of ChangingSocial Organisation the InfectiousDiseases of Man,"in on Boyden, Impactof Civilisation(cit. n. 84), 48-76, on 48. 88 Ibid., on 63, 66. 89 S. D. Rubbo,quotedin ibid., 69. 90Ibid., 76. 91 Experienceof tropicalpracticeclearly influencedthe ideas of JacquesM. May, a disease ecologist who mixed mostly with geographersbut was also on the marginsof the group I describe. In the 1940s and 1950s, with the supportof theAmericanGeographical Society,May triedto refashionmedical geographyas disease ecology. See his "MedicalGeography:Its Methods and Objectives,"GeographicalReview40 (1950): 9-41. On May's tropicalcareer,see JacquesM. May, SiamDoctor (Garden City,N.Y., 1949). 92 I discuss this debate in The Cultivationof Whiteness(cit. n. 71). Interestingly,Buret seems to have followed a trajectory similarto thatof RaymondPearl.In Buret's case, it led him away fromthe "Australia Unlimited"partisansof the Melbournemedical school, towarda concernwith "overpopulation."On Pearl,see GarlandE. Allen, "OldWine in New Bottles:FromEugenics to PopulationControl in the Workof RaymondPearl," TheExpansionofAmericanBiology,ed. KeithR. Benson, Jane in Maienschein,and RonaldRainger(New Brunswick,N.J., 1991), 231-61.

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For thusemergedas a legacy of settlercolonial anxieties.93 Buret andothersit would a cogent explanation disease patterns a worldshapedby humanmobilof in provide towardthe modernity coloniality; it also allowedthemto expresstheirambivalence madeglobal. ism had apparently In the 1980s, furtherchanges in the naturaland conceptualenvironments would a largerniche for ecological reasoningin medical science. Above all, the provide emergenceof a global epidemicof HIV/AIDSshook those previously"nonecological"scientistswho hadbecome complacentaboutinfectiousdisease in the developed world.Justas epidemiologistssuch as FlexnerandGreenwoodhadcalled for a more of complex understanding disease transmissionin the wake of the influenzapandemic of 1918-1919, manymicrobiologistsnow soughta moreintegrative explanation for the emergenceof AIDS and otherdiseases, includinglegionnaires'disease, disLyme disease, dengue hemorrhagicfever, and new variantCreutzfeldt-Jacob a matterof old diseases conquering ease. Of course, disease "emergence," usually was new territories, hardlya novel occurrence, Fennercould havetold them.Since as the late nineteenthcentury,many "new"diseases had spreadwidely, among them nontyphoidsalmonellosis,poliomyelitis, kuru,and even the pandemicform of influenza.All of these emergentconditionshad at the time stimulatedspeculationon theirunderlyingbiological or social causes,butAIDS wouldexertan even moreprofoundimpacton biomedicalscience, as it struckaftera long periodof complacency, when the futureof infectiousdiseases, in Burnet'sopinion,looked exceedingly dull in the developed world. In the developing world, however, there had never been much cause for contentment, therethe impactof economic developmenton mifor crobialabundance distribution still demonstrated and was daily.94 Ecological insight was rarelyabsentfromtropicalmedicine;thus,in a sense, "mainstream" biomedical science was simply catchingup, recognizingthatdisease even in Europeand North America might be the outcome of dynamic processes in a global ecosystem. The complacencyshatteredin the 1980s was not so much overconfidencein the global controlof infectiousdisease, as the conventionalassumptionthatEuropeandNorth Americahad managedsomehow to remove themselves from the naturalprocesses thatdisease ecologists described.
for the emergenceof a generalenvironmental 93 This echoes RichardGrove's argument consciousness from concerns about colonial environmentaldegradation: his Green Imperialism:Colonial see 1600-1860 (Cambridge, Expansion, TropicalIsland Edens, and the Origins of Environmentalism, in 1995). See also Libby Robin, "Ecology:A Science of Empire," Ecology and Empire:Environmental Historyof SettlerSocieties, ed. TomGriffithsandLibbyRobin (Melboure, 1997), 63-75; Thomas R. Dunlap, "Ecology and Environmentalism the Anglo Settler Colonies,"in Griffithsand Robin, in Ecology and Empire,76-86; JohnMackenzie, "Empireand the Ecological Apocalypse:The Historiin ographyof the ImperialEnvironment," GriffithsandRobin,Ecology and Empire,215-28; andPeder Orderin the BritishEmpire,1895-1945 (Cambridge, Anker,ImperialEcology: Environmental Mass., Ankermeans Oxfordand SouthAfrica. 2001). By "Britishempire," 94 Much of this work has focused on schistosomiasisand in trypanosomiasis Africa. See CharlesC. Hughes and JohnM. Hunter,"Disease and 'Development'in Africa,"Social Science and Medicine 3 Man-MadeEnvironments, (1970): 443-93; andDuncanPedersen,"DiseaseEcology at a Crossroads: Human Rights, and Perpetual Development Utopias," Social Science and Medicine 43 (1996): 745-58. Helen Tilley has discussed the persistingecological perspectiveof Africantropicalmedicine in more detail in "Ecologies of Complexity"(cit. n. 18). See also StephenJ. Kunitz,Disease and Social Diversity:TheEuropeanImpacton the Health of Non-Europeans(New York,1994). In the 1960s and 1970s, medical anthropology an to frequentlydemonstrated "eco-social"approach understanding disease in the "Third World." MarciaC. Inhom andPeterJ. Brown,TheAnthropology nfectious See of Disease: InternationalHealth Perspectives(Amsterdam,1997).

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Fenner's prediction in the 1960s of the emergence of "new" diseases in the developed world was fulfilled in the 1980s. Joshua Lederberg was among those who finally read the ecological lesson. A colleague of Dubos's at the Rockefeller University, Lederberg had worked with Buret at the Hall Institute in the 1950s, so their work was familiar to him. In 1993, though, Lederberg complained that "the historiography of epidemic disease is one of the last refuges of the concept of special creationism"Now, however, many scipioneering ecological approaches had largely been ignored.95 entists were prepared to admit that evolutionary processes operating on a global scale were responsible for the emergence of "new" diseases. As environments changed, as urbanization, deforestation, and human mobility increased, so, too, did disease patterns alter, with natural selection promoting the proliferation of microbes in new niches. But pessimism should not overwhelm us. "We recall that since Frank Macfarlane Buret, Theobald Smith, and others,"wrote Lederberg, "we have understood that evolutionary equilibrium favors mutualistic rather than parasitic or unilaterally destructive interactions. Natural selection, in the long run, favors host resistance, on the one hand, and temperate virulence and immunogenic masking on the parasite's part on the other."All the same, Lederberg, echoing Dubos, remained concerned that too good a human adaptation to an increasingly degraded environment might yet be detrimental to human values. "In a biological sense," he mused, "we may achieve new genomic equilibria with these radically altered environments; but the price of naturalselection is so high that I doubt we would find it ethically acceptable: it conflicts violently with the nominally infinite worth that we place on every individual."96 During the 1990s, amplified concern about emerging infectious diseases, along with fears of increasing antibiotic resistance and the health effects of climate change, would boost interest in disease ecology. Stephen S. Morse, a virologist and immunologist at the Rockefeller University, joined Lederberg in arguing that since "most 'new' or 'emerging' viruses are the result of changes in trafficpatternsthat give viruses new highInways," we need "a science of trafficpatterns,partbiology and part social science."97 terest in the emergence of "new" diseases soon led to a proliferationof conferences and symposia; it gave rise to numerous reports and to popular books, such as Laurie Garrett's The Coming Plague.98The journals Emerging Infectious Diseases and Ecosystem Health were launched in the mid-1990s.99At the end of the century, clinicians and scientists were also coming to recognize antibiotic resistance as a growing problem, and
95 JoshuaLederberg, "VirusesandHumanKind:Intracellular Symbiosis and EvolutionaryCompetition,"in EmergingViruses,ed. Stephen S. Morse (New York, 1993), 3-9, on 3. This book derived froma 1989 conferenceon emergingviruses supported the RockefellerUniversityandthe National by Institutesof Health,andattendedby D. A. Henderson,FrankFenner,WilliamH. McNeill, andRobert M. May. See also JoshuaLederberg,R. E. Shope, and S. C. Oaks Jr.,eds., EmergingInfections:Microbial Threatsto Health in the UnitedStates (Washington, D.C., 1992). Like Burneta generationearlier, Lederbergwas profoundlyshapedby Wells and Huxley: "TheScience of Life was the most influentialsourceof my perspectiveon biology andman'splace in the cosmos, seen as evolutionary drama" ("GeneticRecombinationin Bacteria:A Discovery Account,"in The Excitementand Fascinationof Science, ed. JoshuaLederberg[PaloAlto, Calif., 1990], vol. 3, part 1, 893-915, on 895). 96Lederberg,"Virusesand HumanKind"(cit. n. 95), on 8, 4. 97 Viruses:Defining the Rules for ViralTraffic," StephenS. Morse, "Emerging Perspect.Biol. Med. 34 (1991): 387-409, on 388, 404. Morse thanksFrankFennerfor discussing this paperwith him. 98 LaurieGarrett, The ComingPlague: Newly EmergingDiseases in a WorldOut of Balance (London, 1994). 99See Nicolas B. King, "TheScale Politics of EmergingDiseases" (this volume). King relatesfears of biological warfare-a problemthathauntedBurneteven in 1940-to the growinginterestin emerging diseases.

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it they attributed to evolutionary processes.Accordingto S. B. Levy,a microbiologist at Tufts,profligate antibiotic had delivereda selectionpressure microbes"unuse on in "We he precedented the historyof evolution."'?? must somehow," wrote, "finda in and meansto reversethe ecologicalimbalancethathas occurred termsof resistant Others usreaping ecologicalwhirlwind climatechange. saw the of susceptible strains."'01 Alterationsin the abundanceand distribution microorganisms their vectors of and to a large extent,mediatethe influenceof climatechange and otherphysical might, Thus as mosquitoesextendedtheir range so, too, would malaria, transfigurations. andothersupposedly In Environments, dengue "tropical" pathogens. HumanFrontiers, andDisease,A. J. McMichael, who occupiedGreenwood's chairat theLondonSchool of HygieneandTropical in Medicine,declaredthat"ashumanintervention the global and environment its life processesintensifies,we need betterunderstanding the poof tentialconsequences of... ecologicaldisruptions healthanddisease."'02 for In form andfunction,the understanding disease ecology emergingat the end of of the twentiethcenturyoften departed theories,and quiteradicallyfromthe antecedent of Smith,Buret, andDubos.However,therewas still a commetaphoric borrowings, mon interestin mitigatingfacile reductionismin medical research,a desire to assemble a more complex and integrative framework disease patterns. for explanatory of an ecological perspectiveon infectiousdiseases soughta meansto reProponents late microbiological or processesto largerenvironmental biological forces, a way to describethe interactive, betweenhost andparasiteandphysidynamicrelationships cal milieu. In so doing, they conventionally invokedan evolutionary time frameand a global compass.Indeed,it is hardto imagine the developmentof disease ecology withoutthe concomitanteconomic and political globalizationoccurringduringthe twentiethcentury.Now, as before,the leadinginstigators wide-ranging of ecological thattheirviews are marginal biomedicalresearch. they conin Yet argue approaches tinue to occupy key positions at elite researchinstitutions,and they continueto acquireNobel Prizes at an enviablerate.Despite claims of marginality, speculationon disease ecology has often functionedto distinguishscientistsfrommeretechnicians or fromthe allegedreductionism routinism juniorcolleaguesandcompetitors. and of But theoretical of disspeculation this sort,once littlemorethana markof intellectual tinction,a flashybit of plumage,would eventuallybecome a majorselection advanand of tage in the rapidlychanging,andperplexing,natural conceptualenvironments the late twentiethcentury.
100 B. Levy, "Antibiotic S. Resistance:An Ecological Imbalance," AntibioticResistance:Origins, in Evolution,Selectionand Spread,CibaFoundationSymposium207 (Chichester, U.K,, 1997), 1-14, on 2. See also idem, TheAntibioticParadox:How MiracleDrugsAre Destroyingthe Miracle (New York, 1992). Dubos predictedselectionfor antibioticresistanceas earlyas 1942. See CarolL. Moberg,"Rene Dubos:A Harbinger MicrobialResistanceto Antibiotics," of Perspect.Biol. Med. 42 (1999): 559-80. 101 Resistance"(cit. n. 100), 8. Levy, "Antibiotic 102A. J. and McMichael, Human Frontiers,Environments, Disease: Past Patterns, UncertainFutures (Cambridge,2001), xiv. The WHO opened an office to study the health implicationsof climate to since the late 1970s of mathematical changein 1990. It is also important recognize the contribution modeling of host-parasiteinteractionsand populationdynamics. See R. M. May, "Ecology and the Evolutionof Host-VirusInteractions," Morse, EmergingViruses(cit. n. 95), 58-68; R. M. Anderin son andR. M. May, "Population Biology of InfectiousDiseases,"Nature280 (1979): 361-7,455-61; and R. M. May and R. M. Anderson, "The TransmissionDynamics of Human Immunodeficiency in Virus," TheEpidemiologyand Ecology of InfectiousDisease Agents, ed. R. M. AndersonandJ. M. Thresh(London, 1988), 239-81. May regardedFrankFenneras the "realhero"of the biological analysis of disease because of his classic mathematicalanalysis of myxomatosis (May, "Ecology and the Evolutionof Host-VirusInteractions," 63).