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Stanley Falkow

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Stanley Falkow
Born(1934-01-24)January 24, 1934
DiedMay 5, 2018(2018-05-05)(aged 84)
EducationUniversity of Maine
Brown University
Occupation(s)Technician,Newport Hospital(1956–1957)
Scientist,Walter Reed Army Institute of Research(1961–1966)
Professor,Georgetown University(1967–1972)
Professor,University of Washington(1972–1981)
Professor,Stanford University(1981–2018)
Known forAntibiotic resistance and molecular microbiology research
Partner(s)Rhoda Ostroff (m.1958–1983)
Lucy S. Tompkins(m. 1983–2018)
Children2
Awards
Scientific career
ThesisAn episomic element in a strain of Salmonella typhosa(1960)
Doctoral advisorCharles ( "Doc" ) Arthur Stuart, Brown University
Signature

Stanley "Stan" Falkow(January 24, 1934 – May 5, 2018) was an American microbiologist and a professor ofmicrobiologyatGeorgetown University,University of Washington,andStanford University School of Medicine.Falkow is known as the father of the field of molecular microbial pathogenesis.[1] He formulatedmolecular Koch's postulates,which have guided the study of the microbial determinants of infectious diseases since the late 1980s.[2]Falkow spent over 50 years uncovering molecular mechanisms of how bacteria cause disease and how to disarm them.[1]Falkow also was one of the first scientists to investigate antimicrobial resistance, and presented his research extensively to scientific, government, and lay audiences explaining the spread of resistance from one organism to another, now known ashorizontal gene transfer,and the implications of this phenomenon on our ability to combat infections in the future.[1]

Early life and education[edit]

Falkow was born into a Yiddish-speaking household inAlbany, New York.His father was a shoe salesman and had immigrated from SovietKyiv,Ukraine.[3] His mother came from a family of Jewish immigrants from Poland. She "rented several of their bedrooms and later opened a corset shop".[4]Falkow's family was working-class to poor as he described in his memoir.[1]He was therefore a first-generation Jewish American. Falkow worked at a toy shop in order to collect enough money to buy a model microscope, as well as admitted to stealing textbooks, one a nurse's book on infectious diseases, in order to explore microbiology as a career path despite his family's financial situation.[1]

Falkow attributed his early interest in microbiology to readingMicrobe Huntersin 1943, when he was 11 years old, which he found at the public library after the family had moved toNewport, Rhode Island.The book is a dramatization of microbiological research written by American microbiologistPaul de Kruif.[5] Because of poor grades until his senior year of high school, an adviser suggested military rather than college.[4]Falkow described himself as an average student, and below average at mathematics, which he later rectified by working through workbooks of high school and college level mathematics later in his career, after earning his Ph.D., in order to analyze and design experiments.[1]In his memoir, Falkow gives credit to his teachers and acknowledges the role that the systemic oppression of women in America in the 1940s–1950s played in his life, stating "I now understand that I was the beneficiary of the repression of women by our society. The best and brightest women in many communities were relegated to the one intellectual role deemed suitable for them. They became teachers and many poured their love of language, science, music and art into students like me."[1]

Education and career[edit]

Falkow enrolled in biology at theUniversity of Maine,because of their microbiology department. During the summers he worked in pathology at the hospital in Newport, staining slides and assisting in autopsies. He graduated in 1955.[4] Before he started graduate school atUniversity of Michiganhe developed extreme anxiety. He was later diagnosed withagoraphobia.He avoided movie theaters and crowded places between 1955 and 1983[4]and dropped out of his first attempt at graduate school because of recurring panic attacks.[3][4]He returned to Rhode Island to live with his parents and work at the Newport Hospital as a technician, focusing on bacteriology and processing and examining patient samples.[1]He re-enrolled in graduate school atBrown Universityand graduated with a PhD in 1961.[4]

He finished his work for his Ph.D. thesis at theWalter Reed Army Institute of Research(WRAIR) in the Department of Bacterial Immunology.[1]While at Walter Reed, he worked with African-American microbiologistOthello Washington.Washington was older and more experienced than Falkow, but was assigned to be Stan's technician. Upon meeting, Falkow and Washington decided it would be more appropriate and better for both men to work together as a team. The two men worked together on isolatingmobile genetic elementsand the transfer of genes betweenE. coliandSalmonellaand published a paper together aboutProteusspecies.[1][6]Falkow also worked withShigellaspecies and contracteddysentery,which lead to hospitalization, after being exposed to infected monkey feces flung at his face by an infected monkey.[1]Multiple times through his training and early career more senior scientists, some Nobel prize winners, recommended that Falkow focus more on mechanisms of gene expression and less on pathogens because "nobody cares about typhoid".[1]Falkow remained interested in pathogens despite many warnings from his peers and supervisors that infectious diseases were not interesting and were becoming less frequent in wealthy countries and were therefore a diminishing field of study.[1]

His early work in the 1960s focused on the genetic mechanisms, which enable bacteria to become resistant to antibiotics. He showed that organisms, such asShigella,can possess DNA fragments calledplasmidswhich exist apart from the bacterial chromosome, and that they carry specialized information for survival. Under selective pressure from antibiotics, one species of bacteria can pass its plasmids to another directly without mating, thereby preserving its own specialized survival genes.

In 1966, he joinedGeorgetown University School of Medicineas a professor of microbiology. He later moved toSeattleto join the faculty of the Department of Microbiology and Immunology at theUniversity of Washington School of Medicine.Here he described howmeningitisandgonorrheaorganisms acquire plasmids to become resistant topenicillinand otherantibiotics.[1]

In the 1970s, Falkow shifted his focus to the infection process. During this period, he showed that a life-threatening diarrhea prevalent in many developing countries is caused by a sub-type ofE. coli.He also co-authored (withRoyston C. Clowes,Stanley N. Cohen,Roy CurtissIII,Naomi DattaandRichard Novick) a proposal for uniform nomenclature for bacterial plasmids.[7]

In 1981, he was named chairman of the Department of Medical Microbiology at Stanford University School of Medicine, a position he held until 1985. While at Stanford, Falkow encouragedEsther Lederbergto continue directing the StanfordPlasmid Reference Center,an internationally used registry for plasmids, transposons and insertion sequences.[8])

Personal life and death[edit]

Falkow was married to Rhoda Ostroff in January, 1958 with whom he had two daughters. They divorced in 1983.[4]In December, 1983, he marriedLucy S. Tompkins.[4]

Falkow and Tompkins met while he was a professor at Georgetown University, and she was a medical student who sought Falkow out to work in his lab. At first, Falkow denied the opportunity to Tompkins, claiming that he would not take female graduate students because he was not comfortable having a female student, which was common in the scientific field in the 1960s.[1]Tompkins rebutted hismisogynyfirmly, and explicitly pointed out that he was discriminating against her based on her gender.[1]Her argument forced Stan to reconsider his bias and he then allowed her to enter his laboratory as a trainee.[1]For the remainder of his career Falkow continued to hire women trainees, and by the end of his career most of his trainees were female.[1]Falkow later became an advocate and ally ofgender equalityin science and helped encourage many women to pursue scientific careers.[9]

Falkow had on-going severeanxietythroughout his career. Due to his anxiety early in his career he avoided conferences, airplanes, and giving presentations in general due to the fact that he would shake and sweat noticeably during them.[1]Falkow described his struggle with hismental healthas an "uneasy truce" with his "demons" later on in his career as he learned various coping mechanisms.[1]Because of his anxiety during public speaking, he often employed humor in his teaching and presentations to help cope with the tension of a silent audience, which also lead to most of the audience members finding his talks memorable and enjoyable.[4]Falkow kept his condition a secret from his colleagues during his early career, only privately revealing to Arthur Saz, the chair of microbiology atGeorgetown University School of Medicine,that he received and benefited frompsychoanalysistreatment.[1]Health insurancefor mental health treatment was much better for military employees than academics at the time, and a major reason Falkow continued to work at theWalter Reed Army Institute of Researchfor as long as he did early in his career, but Falkow agreed to a job atGeorgetown Universitybecause of this disclosure, upon which the chair privately organized Falkow's healthcare benefits to cover his psychoanalysis treatments.[1]

In 2004 Falkow was diagnosed withmyelodysplastic syndromeand given a prognosis of two years.[4]He died on May 5, 2018, at the age of 84 at his home inPortola Valley, California,ofmyelodysplastic syndrome.[10]His death was announced and lifetime achievements highlighted in obituaries inThe New York Times,[11]Washington Post,[12]Nature,[13]Science,[9]in a press release from theAmericain Society of Microbiology[14]and in various international news sources.

Contributions[edit]

Molecular microbiology research[edit]

Falkow has been referred to as the "father of molecular microbialpathogenesis",the study of how infectiousmicrobesand host cells interact to cause disease at the molecular level.[15] Falkow adopted the perspective of viewing infection as a process that is ultimately dependent on both the infecting agent and the host.[1][16]He discovered that infectious microbes employgenesthat are activated only inside host cells. His work carries clinical applications, such as a new vaccine forwhooping cough.[citation needed]From enteric pathogens to sexually transmitted diseases to respiratory infections, his influence has left virtually no field untouched.[17]

Falkow and his first graduate student,Richard P. Silver,discovered that episomes (plasmids) are nicked into linear pieces of DNA and transported as linear DNA between two bacterial cells duringconjugation.[18]He then collaborated with Mexican scientistsEmma GalindoandJorge Olarteto investigate new forms of penicillin resistance inShigellaisolates from pediatric patients in Mexico City, which lead to the discovery of new forms of "R factors" - genes responsible for antimicrobial resistance.[19][1]Falkow and traineeJoan Skerman Knappidentified a virulence factor on plasmids ofE. colifound in human feces, demonstrating for the first time that the ability to cause disease may be transferable between bacteria.[20]

Falkow was present at the famous Waikiki beach meeting in 1972 betweenHerb Boyer(who provided the restriction enzyme EcoRI) andStanley Cohen(who provided the plasmid) that led to a plan to developrecombinant DNAtechnology. This led to the first cloning of a bacterial virulence factor, the heat stabletoxinofE. coli,in a seminal 1976 paper withMagdalene Yh So,which heralded the application of molecular genetics to understanding gene transfer, and ultimately led to the creation of the field ofsynthetic biology.

Falkow and his trainees developed many methods that moved biological research forward such as: a method for screening patient samples for enteric pathogens,[21]the identification ofSalmonellaandShigellain patient samples based on lysinemetabolism,[22]the application ofnucleic acid hybridizationto distinguish different bacterial species,[23]the application ofagarose gelsto isolate plasmids of various sizes,[24]isolation and creation of different plasmid backbones now used ascloning vectors,[25]a method for identifying unculturable pathogens based on isolated16S ribosomal RNAsequence,[26]an optimized version ofGFPforflow cytometryapplications,[27]and a fluorescence-based method for the detection of genes expressed by pathogens inside of host cells,[28]among many other techniques.

After studying the DNA content of many different pathogenic microbes, Falkow became one of the first scientists to suggest bacterial taxonomy should be based on DNA composition rather than phenotypic observations.[29][30]

Falkow and others created the universal nomenclature scheme forplasmidsstill in use today.[7]

Falkow established ‘Molecular Koch’s postulates’ for defining virulence determinants,[2]which continues to influence thinking and experimental design in infectious disease research.[17]

Teaching and service[edit]

Falkow valued and enjoyed teaching as a fundamental part of scientific training and believed that all scientists have a duty to teach and train the next generation.[1]In all of his years of employment at academic institutions, he taught students in some capacity between medical, graduate, undergraduate courses and invited lectures for courses whenever he was able to.[1]In 2010, he recorded lectures on microbial pathogenesis for iBiology, and those videos are still publicly available on theiBiology websiteand onYouTube.[16]When Falkow became an emeritus Professor at Stanford University, he still taught students. Indeed, he co-taught withLucy Tompkinsa class open to undergraduate and graduate students on the history of Infectious Disease.[31]

Falkow organized the first international symposium of invited speakers on antimicrobial resistance.[32][1]

Falkow and his studentVickers Herschfieldinvestigated water sources in the D.C. area to find examples of plasmids containing antimicrobial resistance genes. Unexpectedly, they found that thePotomac RiverandRock Creek,considered "clean" water sources used for recreation at the time, were full of fecal waste and alerted many government offices about their findings, which were at first largely ignored.[1]In the summer of 1971, Falkow held a press conference warning D.C. citizens not to let their children play in the Potomac River, stating "one cup of Potomac River water is equivalent to a half gram of human feces".[1][33]

Falkow served on the first NIH Recombinant DNA committee to recommend policy guidelines for the use of recombinant DNA to the United States of America Congress.[1]Many of Falkow's publications are referenced in theRecombinant DNA Regulation Actof 1977.[34]

Falkow worked with theFDAof the United States throughout his career to advocate for the decrease of antimicrobial agents used in livestock feed.[1][35][36]

Awards and honors[edit]

Falkow receiving the National Medal of Science (left), President Barack Obama (middle), military official holding medal about to be awarded (right).

Falkow was elected President of theAmerican Society for Microbiologyfrom July 1997 through June 1998. He was elected to the Institute of Medicine in 1997 and received theMaxwell Finland Awardfrom the National Foundation of Infectious Diseases in 1999. He also received in 1999 an Honorary Doctor of Science,University of Guelph,Guelph,Ontario,and theUniversity of MaineAlumni Career Award. He has received honorary doctorates in Europe and the US. He received numerous awards for his achievements in science, including theBristol-Myers Squibb Awardfor Distinguished Achievement in Infectious Disease Research,[37]the Altemeier Medal from the Surgical Infectious Diseases Society of America, theHoward Taylor RickettsAward Lecture at theUniversity of Chicago,and thePaul Ehrlich and Ludwig Darmstaedter Prize.In 2003, he received the Abbott Lifetime Achievement Award from theAmerican Society for Microbiologyand theSelman A. Waksman Award in Microbiologyfrom theNational Academy of Sciences.[38] He received theRobert Koch Awardin 2000.[39]

Falkow was an elected member of theInstitute of Medicine,the National Academy of Sciences, and the National Academy of Arts and Sciences, an elected Fellow of the American Association for the Advancement of Science. He was also elected into the UK'sRoyal Societyas a Foreign Member.[40]

In September 2008, Falkow was awarded theLasker Awardfor medical research.[41]

In May, 2016, Falkow was awarded by Barack Obama the National Medal of Science for his monumental contributions toward understanding how microbes cause disease and resist the effects of antibiotics, and for his inspiring mentorship that created the field of molecular microbial pathogenesis.[42]

Trainees[edit]

Stanley Falkow is known for training many experts in the field of molecular microbiology who then became prominent professors and scientists all over the world, including;

References[edit]

  1. ^abcdefghijklmnopqrstuvwxyzaaabacadaeafFalkow, Stanley (2020).And what's beyond that?: a memoir by the father of microbial pathogenesis.[United States].ISBN979-8-6873-4014-4.OCLC1225499908.{{cite book}}:CS1 maint: location missing publisher (link)
  2. ^abFalkow S (1988). "Molecular Koch's postulates applied to microbial pathogenicity."Rev Infect Dis10(Suppl 2):S274-S276.
  3. ^abSandeep Ravindran.Curious About EverythingStanford Alumni Magazine, September/October 2011, retrieved June 22 2018
  4. ^abcdefghijHarrison SmithStanley Falkow, microbiologist who studied bacteria and the diseases they cause, dies at 84Washington Post, 12 May 2018, retrieved 21 June 2018
  5. ^"Stanley Falkow, Father of Molecular Microbial Pathogenesis, Dies".The Scientist.RetrievedMay 10,2018.
  6. ^Falkow, Stanley; Ryman, I. R.; Washington, O. (1962)."Deoxyribonucleic Acid Base Composition of Proteus and Providence Organisms".Journal of Bacteriology.83(6): 1318–1321.doi:10.1128/jb.83.6.1318-1321.1962.PMC279453.PMID13891463.
  7. ^abRichard P. Novick et al., "Uniform Nomenclature for Bacterial Plasmids: A Proposal", Bacteriol. Rev., March 1976, pp. 168–189.
  8. ^Seehttp:// esthermlederberg /EImages/Archive/ArchiveIndex.html;click "Special Topics", "Plasmid Reference Center Funding Issues", then "Falkow"
  9. ^abMonack, Denise; Strauss, Evelyn (June 8, 2018)."Stanley Falkow (1934–2018)".Science.360(6393): 1077.Bibcode:2018Sci...360.1077M.doi:10.1126/science.aau2284.S2CID46954621.
  10. ^Kolata, Gina (May 10, 2018)."Stanley Falkow, Who Saw How Bacteria Cause Disease, Dies at 84".The New York Times.Archived fromthe originalon May 11, 2018.RetrievedMay 11,2018.
  11. ^Kolata, Gina (May 10, 2018)."Stanley Falkow, Who Saw How Bacteria Cause Disease, Dies at 84".The New York Times.ISSN0362-4331.RetrievedOctober 8,2021.
  12. ^"Stanley Falkow, microbiologist who studied bacteria and the diseases they cause, dies at 84".Washington Post.ISSN0190-8286.RetrievedOctober 8,2021.
  13. ^Amieva, Manuel R. (June 7, 2018)."Stanley Falkow (1934-2018)".Nature.558(7709): 190.Bibcode:2018Natur.558..190A.doi:10.1038/d41586-018-05377-6.S2CID46958780.
  14. ^"In Memoriam: Falkow, Stanley".ASM.org.RetrievedOctober 8,2021.
  15. ^The Double HelixNFID to Honor Dr. FalkowArchivedSeptember 28, 2007, at theWayback MachineRetrieved on July 4, 2007
  16. ^ab"Host-Pathogen Interactions and Human Disease • iBiology".iBiology.RetrievedOctober 8,2021.
  17. ^ab"Stanley Falkow | Sir William Dunn School of Pathology".path.ox.ac.uk.RetrievedOctober 14,2021.
  18. ^abSilver, Richard P.; Falkow, Stanley (October 1, 1970)."Specific Labeling and Physical Characterization of R-Factor Deoxyribonucleic Acid in Escherichia coli".Journal of Bacteriology.104(1): 331–339.doi:10.1128/jb.104.1.331-339.1970.PMC248218.PMID4919748.
  19. ^Evans, J.; Galindo, E.; Olarte, J.; Falkow, S. (October 1968)."Beta-lactamase of R factors".Journal of Bacteriology.96(4): 1441–1442.doi:10.1128/jb.96.4.1441-1442.1968.ISSN0021-9193.PMC252489.PMID4971890.
  20. ^abSkerman, F. J.; Formal, S. B.; Falkow, Stanley (April 1, 1972)."Plasmid-Associated Enterotoxin Production in a Strain of Escherichia coli Isolated from Humans".Infection and Immunity.5(4): 622–624.doi:10.1128/iai.5.4.622-624.1972.PMC422413.PMID4564682.
  21. ^Falkow, Stanley (July 1, 1957)."A Screening Method for Enteric Organisms, using a Ferric Chloride Test".American Journal of Clinical Pathology.28(1_ts): 99–102.doi:10.1093/ajcp/28.1_ts.99.ISSN0002-9173.PMID13458131.
  22. ^Falkow, S. (May 1958)."Activity of lysine decarboxylase as an aid in the identification of Salmonellae and Shigellae".Technical Bulletin of the Registry of Medical Technologists. American Society of Clinical Pathologists. Registry of Medical Technologists.28(5): 106–108.ISSN0097-0654.PMID13556625.
  23. ^Falkow, Stanley; Rownd, R.; Baron, L. S. (December 1962)."Genetic Homology Between Escherichia Coli K-12 and Salmonella".Journal of Bacteriology.84(6): 1303–1312.doi:10.1128/jb.84.6.1303-1312.1962.ISSN0021-9193.PMC278063.PMID16561983.
  24. ^Meyers, J A; Sanchez, D; Elwell, L P; Falkow, S (September 1976)."Simple agarose gel electrophoretic method for the identification and characterization of plasmid deoxyribonucleic acid".Journal of Bacteriology.127(3): 1529–1537.doi:10.1128/jb.127.3.1529-1537.1976.ISSN0021-9193.PMC232949.PMID821935.
  25. ^So, M.; Gill, R.; Falkow, S. (December 30, 1975)."The generation of a ColE1-Apr cloning vehicle which allows detection of inserted DNA".Molecular & General Genetics.142(3): 239–249.doi:10.1007/BF00425649.ISSN0026-8925.PMID1045010.S2CID38221334.
  26. ^Relman, D. A.; Loutit, J. S.; Schmidt, T. M.; Falkow, S.; Tompkins, L. S. (December 6, 1990)."The agent of bacillary angiomatosis. An approach to the identification of uncultured pathogens".The New England Journal of Medicine.323(23): 1573–1580.doi:10.1056/NEJM199012063232301.ISSN0028-4793.PMID2233945.
  27. ^Cormack, B. P.; Valdivia, R. H.; Falkow, S. (1996)."FACS-optimized mutants of the green fluorescent protein (GFP)".Gene.173(1 Spec No): 33–38.doi:10.1016/0378-1119(95)00685-0.ISSN0378-1119.PMID8707053.
  28. ^Valdivia, R. H.; Falkow, S. (September 26, 1997)."Fluorescence-based isolation of bacterial genes expressed within host cells".Science.277(5334): 2007–2011.doi:10.1126/science.277.5334.2007.ISSN0036-8075.PMID9302299.
  29. ^Marmur, J; Falkow, S; Mandel, M (October 1963)."New Approaches to Bacterial Taxonomy".Annual Review of Microbiology.17(1): 329–372.doi:10.1146/annurev.mi.17.100163.001553.ISSN0066-4227.PMID14147455.
  30. ^Falkow, Stanley (November 1, 1965)."Nucleic acids, genetic exchange and bacterial speciation".The American Journal of Medicine.Symposium on Antibiotics.39(5): 753–765.doi:10.1016/0002-9343(65)90095-1.ISSN0002-9343.PMID5319389.
  31. ^Monack, Denise M; Brodsky, Igor E (April 1, 2020)."Editorial overview: The fortunate students, a tribute to the fortunate professor".Current Opinion in Microbiology.Stanley Falkow Alumni.54:iii–vi.doi:10.1016/j.mib.2020.02.001.ISSN1369-5274.PMID32171689.S2CID212728684.
  32. ^Falkow, Stanley (October 2008)."The Fortunate Professor".Annual Review of Microbiology.62(1): 1–18.doi:10.1146/annurev.micro.62.081307.162931.ISSN0066-4227.PMID18345978.S2CID45328833.
  33. ^Selden, Maury; Llewellyn, Lynn G. (1973).Studies in Environment.U.S. Environmental Protection Agency, Office of Research and Development.
  34. ^Recombinant DNA Regulation Act, 1977.U.S. Government Printing Office. 1977.
  35. ^Guest, Gerald B. (1976)."Status of the FDA's Program on the Use of Antibiotics in Animal Feeds".Food, Drug, Cosmetic Law Journal.31(1): 54–58.ISSN0015-6361.JSTOR26657829.
  36. ^"Inside an Early Attempt to Restrict Antibiotic Use on Farms".FRONTLINE.RetrievedOctober 14,2021.
  37. ^Bristol-Myers SquibbAwards for Distinguished AchievementRetrieved on July 16, 2007
  38. ^"Selman A. Waksman Award in Microbiology".National Academy of Sciences. Archived fromthe originalon December 29, 2010.RetrievedFebruary 15,2011.
  39. ^ISI Highly CitedFalkow, StanleyArchivedSeptember 29, 2007, at theWayback MachineRetrieved on July 12, 2007
  40. ^Ramakrishnan, Lalita; Cormack, Brendan P. (2024)."Stanley Falkow. 24 January 1934—5 May 2018".Biographical Memoirs of Fellows of the Royal Society.76.
  41. ^Altman, Lawrence K. (September 13, 2008)."5 Pioneering Scientists Win Lasker Medical Prizes".New York Times.
  42. ^"Stanley Falkow".
  43. ^abcZonana, Kathy (June 16, 2016)."On mentorship, and how to pay it forward".Scope.RetrievedOctober 8,2021.
  44. ^"In Memoriam of Carleen Collins (1955-2008) | UW Microbiology".microbiology.washington.edu.Archived fromthe originalon October 27, 2021.RetrievedOctober 13,2021.
  45. ^"Biologist Karen Guillemin named as a fellow in the AAAS".Around the O.November 30, 2020.RetrievedOctober 8,2021.
  46. ^Gyles, Carlton; So, Magdalene; Falkow, Stanley (July 1, 1974)."The Enterotoxin Plasmids of Escherichia Coli".The Journal of Infectious Diseases.130(1): 40–49.doi:10.1093/infdis/130.1.40.ISSN0022-1899.PMID4601180.
  47. ^abLeBlanc, Donald J.; Silver, Richard P. (1994),"Molecular Nature, Conjugal Transfer, and Replication of Extrachromosomal Elements, 1961 to 1973",Molecular Genetics of Bacterial Pathogenesis,John Wiley & Sons, Ltd, pp. 3–16,doi:10.1128/9781555818340.ch1,ISBN978-1-68367-274-6,retrievedOctober 12,2021
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  49. ^Vecchio, Riccardo (October 15, 2020)."Virus hunter Peter Piot fights personal battle against COVID-19".Stanford Medicine.RetrievedOctober 13,2021.

External links[edit]