341:
184:, a frequently fatal disease in the early 20th century. He found that multiple types—some virulent and some non-virulent—were often present over the course of a clinical case of pneumonia, and thought that one type might change into another (rather than simply multiple types being present all along). In testing that possibility, he found that transformation could occur when dead bacteria of a virulent type and live bacteria of a non-virulent type were both injected in mice: the mice would develop a fatal infection (normally only caused by live bacteria of the virulent type) and die, and virulent bacteria could be isolated from such infected mice.
321:
110:
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531:, it was soon widely accepted that DNA was the material. Despite the much less precise experimental results (they found a not-insignificant amount of protein entering the cells as well as DNA), the Hershey–Chase experiment was not subject to the same degree of challenge. Its influence was boosted by the growing network of the phage group and, the following year, by the publicity surrounding the DNA structure proposed by
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20:
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genetics research, in part because it made little difference for classical genetics experiments in which genes were defined by their behavior in breeding experiments rather than their chemical makeup. H. J. Muller, while interested, was focused more on physical rather than chemical studies of the gene, as were most of the members of the
410:, challenged Avery's finding that the transforming principle was pure DNA, suggesting that protein contaminants were instead responsible. Although transformation occurred in some kinds of bacteria, it could not be replicated in other bacteria (nor in any higher organisms), and its significance seemed limited primarily to medicine.
494:
Despite the significant number of citations to the paper and positive responses it received in the years following publication, Avery's work was largely neglected by much of the scientific community. Although received positively by many scientists, the experiment did not seriously affect mainstream
518:
was an influential member of the phage group. Erwin
Chargaff had shown that the base composition of DNA varies by species (contrary to the tetranucleotide hypothesis), and in 1952 Rollin Hotchkiss published his experimental evidence both confirming Chargaff's work and demonstrating the absence of
440:
A few microbiologists and geneticists had taken an interest in the physical and chemical nature of genes before 1944, but the Avery–MacLeod–McCarty experiment brought renewed and wider interest in the subject. While the original publication did not mention genetics specifically, Avery as well as
460:
Between 1944 and 1954, the paper was cited at least 239 times (with citations spread evenly through those years), mostly in papers on microbiology, immunochemistry, and biochemistry. In addition to the follow-up work by McCarty and others at the
Rockefeller Institute in response to Mirsky's
364:
The experimental findings of the Avery–MacLeod–McCarty experiment were quickly confirmed, and extended to other hereditary characteristics besides polysaccharide capsules. However, there was considerable reluctance to accept the conclusion that DNA was the genetic material. According to
1080:
Boivin; Boivin, André; Vendrely, Roger; Lehoult, Yvonne (1945). "L'acide thymonucléique hautement polymerise, principe capable de conditioner la spécificité sériologique et l'équipement enzymatique des Bactéries. Conséquences pour la biochemie de l'hérédité".
291:(enzymes that break apart proteins or RNA) did not affect it, but an enzyme preparation of "deoxyribonucleodepolymerase" (a crude preparation, obtainable from a number of animal sources, that could break down DNA) destroyed the extract's transforming power.
180:, reported in 1928, identified that some "transforming principle" in pneumococcal bacteria could transform them from one type to another. Griffith, a British medical officer, had spent years applying serological typing to cases of
270:
Chemical analysis showed that the proportions of carbon, hydrogen, nitrogen, and phosphorus in this active portion were consistent with the chemical composition of DNA. To show that it was DNA rather than some small amount of
405:
in the 1950s, were dismissive of DNA as the genetic material (and were inclined to avoid the "messy" biochemical approaches of Avery and his colleagues). Some biologists, including fellow
Rockefeller Institute Fellow
199:
at the
Rockefeller Institute. A series of Rockefeller Institute researchers continued to study transformation in the years that followed. With Richard H. P. Sia, Dawson developed a method of transforming bacteria
461:
criticisms, the experiment spurred considerable work in microbiology, where it shed new light on the analogies between bacterial heredity and the genetics of sexually-reproducing organisms. French microbiologist
385:, and the parallels among viruses, genes, and enzymes; many biologists thought genes might be a sort of "super-enzyme", and viruses were shown according to Stanley to be proteins and to share the property of
564:"Studies on the Chemical Nature of the Substance Inducing Transformation of Pneumococcal Types: Induction of Transformation by a Deoxyribonucleic Acid Fraction Isolated from Pneumococcus Type III"
90:
Studies on the
Chemical Nature of the Substance Inducing Transformation of Pneumococcal Types: Induction of Transformation by a Desoxyribonucleic Acid Fraction Isolated from Pneumococcus Type III
449:
and others praised the result as establishing the biological specificity of DNA and as having important implications for genetics if DNA played a similar role in higher organisms. In 1945, the
263:. An immunological precipitation caused by type-specific antibodies was used to verify the complete destruction of the capsules. Then, the active portion was precipitated out by alcohol
809:
220:
of the transforming principle by 1933. Colin MacLeod worked to purify such solutions from 1934 to 1937, and the work was continued in 1940 and completed by Maclyn McCarty.
523:
was quickly becoming established, and biologists were more inclined to think of heredity in the same terms for bacteria and higher organisms. After
Hershey and Chase used
98:, Avery and his colleagues suggest that DNA, rather than protein as widely believed at the time, may be the hereditary material of bacteria, and could be analogous to
88:
strain type III-S, when injected along with living but non-virulent type II-R pneumococci, resulted in a deadly infection of type III-S pneumococci. In their paper "
373:", DNA consisted of repeating units of the four nucleotide bases and had little biological specificity. DNA was therefore thought to be the structural component of
377:, whereas the genes were thought likely to be made of the protein component of chromosomes. This line of thinking was reinforced by the 1935 crystallization of
441:
many of the geneticists who read the paper were aware of the genetic implications—that Avery may have isolated the gene itself as pure DNA. Biochemist
955:
539:(Watson was also a member of the phage group). Only in retrospect, however, did either experiment definitively prove that DNA is the genetic material.
279:, or some other cell component that was responsible for transformation, Avery and his colleagues used a number of biochemical tests. They found that
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and showed that genetics could apply to bacteria, even if Avery's specific method of transformation was not general. Avery's work also motivated
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Scientists looking back on the Avery–MacLeod–McCarty experiment have disagreed about just how influential it was in the 1940s and early 1950s.
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Hotchkiss, Roland D. "The role of deoxyribonucleotides in bacterial transformations". In W. D. McElroy; B. Glass (eds.).
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to purify and characterize the "transforming principle" responsible for the transformation phenomenon first described in
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studies of DNA, even as he faced pressure from funders to focus his research on whole cells, rather than biomolecules.
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with many enzymes. Furthermore, few biologists thought that genetics could be applied to bacteria, since they lacked
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174:'s studies bacteriologists believed that the types were fixed and unchangeable from one generation to the next.
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Chargaff E (June 1950). "Chemical specificity of nucleic acids and mechanism of their enzymatic degradation".
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Follow-up work in response to criticism and challenges included the purification and crystallization, by
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1410:"The transformation of genetics by DNA: an anniversary celebration of Avery, MacLeod and McCarty (1944)"
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Fry, Michael (2016) Landmark
Experiments in Molecular Biology; Elsevier-Academic Press, United States,
370:
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On the intersecting theories of viruses, genes and enzymes in this period, see: Creager, Angela N. H.
756:"The transformation of pneumococcal types: II. The interconvertibility of type-specific S pneumococci"
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to show that it was primarily DNA, rather than protein, that entered bacteria upon infection with
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The purification procedure Avery undertook consisted of first killing the bacteria with heat and
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Deichmann, UTE (2004). "Early responses to Avery et al.'s paper on DNA as hereditary material".
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170:, a German bacteriologist, had discovered the pneumococcal types and serological typing; until
166:. The antibodies will react with other bacteria of the same type as the original inoculation.
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Explorers of the Body: Dramatic
Breakthroughs in Medicine from Ancient Times to Modern Science
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formation; the different types are classified according to their immunological specificity.
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suggested that it was largely ignored, and only celebrated afterwards—similarly to
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956:"Isolation of a Crystalline Protein Possessing the Properties of Tobacco-Mosaic Virus"
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took up the attempt to extend
Griffith's findings, resulting in the extraction of
70:). It was the culmination of research in the 1930s and early 20th century at the
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showing that virtually all the detected nitrogen in the purified DNA came from
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The Life of a Virus: Tobacco Mosaic Virus as an
Experimental Model, 1930–1965
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that served the function of carrying genetic information (with the very word
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such as this, precipitated from solutions of cell components, to perform
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Lehrer, Steven. Explorers of the Body. 2nd edition. iuniverse 2006 p 46
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protein in Avery's transforming principle. Furthermore, the field of
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The findings of Griffith's experiment were soon confirmed, first by
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The transforming principle: discovering that genes are made of DNA
860:. U.S. National Library of Medicine. Accessed February 25, 2009.
397:. In particular, many of the geneticists known informally as the
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Avery and his colleagues showed that DNA was the key component of
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Shifting Focus: Early Work on Bacterial Transformation, 1928–1940
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Avery, Oswald T.; Colin M. MacLeod; Maclyn McCarty (1944-02-01).
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Proteins, enzymes, genes: the interplay of chemistry and biology
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Proceedings of the Society for Experimental Biology and Medicine
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130:, medical researchers were able to sort bacteria into different
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claimed to extend Avery's bacterial transformation findings to
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containing the antibodies can then be extracted and applied to
19:
1245:. Baltimore: Johns Hopkins University Press. pp. 426–36.
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Hyder, Avery, MacLeod and McCarty used strands of purified
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Historical Studies in the Physical and Biological Sciences
401:, which would become influential in the new discipline of
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colonies which have a polysaccharide capsule that induces
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as Griffith had done). After Dawson's departure in 1930,
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itself coined to indicate a belief that its function was
58:, in an era when it had been widely believed that it was
51:
24:
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457:, in part for his work on bacterial transformation.
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Profiles in Science: The Oswald T. Avery Collection
805:"The Transformation of Pneumococcal Types In Vitro"
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1100:Lederberg, Joshua; Edward L. Tatum (1946-10-19).
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92:", published in the February 1944 issue of the
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1029:. University of Chicago Press: Chicago, 2002.
803:Dawson, Martin H.; Sia, Richard H. P. (1930).
1770:History of the creation-evolution controversy
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731:"Beitrage zur Variabilitat der Pneumokokken"
247:-soluble components. Next, the protein was
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1333:. New Haven, Conn: Yale University Press.
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881:"Remembering Rollin Hotchkiss (1911–2004)"
499:. Avery's work was also neglected by the
72:Rockefeller Institute for Medical Research
1800:Relationship between religion and science
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729:Neufeld, Fred; Levinthal, Walter (1928).
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138:. When a person or test animal (e.g., a
633:"The Significance of Pneumococcal Types"
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1355:. Cambridge: Harvard University Press.
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255:and the polysaccharide capsules were
38:was an experimental demonstration by
760:The Journal of Experimental Medicine
631:Griffith, Frederick (January 1928).
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736:Zeitschrift für Immunitätsforschung
421:'s work decades before the rise of
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1677:Central dogma of molecular biology
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954:Stanley, Wendell M. (1935-06-28).
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1262:The Journal of General Physiology
852:The Oswald T. Avery Collection: "
228:Pneumococcus is characterized by
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1256:Hershey AD, Chase M (May 1952).
1188:Fruton (1999), pp. 440–442
944:Fruton (1999), pp. 440–441
610:Fruton (1999), pp. 438–440
568:Journal of Experimental Medicine
298:in 1948, of a DNA depolymerase (
95:Journal of Experimental Medicine
36:Avery–MacLeod–McCarty experiment
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1353:A history of molecular biology
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688:Dawes, Heather (August 2004).
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1408:Lederberg J (February 1994).
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754:Dawson, MH (1 January 1930).
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154:that react specifically with
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54:is the substance that causes
50:that, in 1944, reported that
1376:. United States: iUniverse.
1160:Deichmann, pp. 227–231
1070:Deichmann, pp. 215–220
1061:Deichmann, pp. 207–209
1052:Deichmann, pp. 220–222
16:1944 microbiology experiment
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146:with a particular type, an
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1426:10.1093/genetics/136.2.423
1351:; Morange, Michel (1998).
1329:Fruton, Joseph S. (1999).
1322:10.1525/hsps.2004.34.2.207
371:tetranucleotide hypothesis
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879:Witkin EM (August 2005).
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649:10.1017/S0022172400031879
29:bacterial transformations
1470:Stegenga, Jacob (2011).
1451:McCarty, Maclyn (1986).
823:10.3181/00379727-27-5078
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512:Hershey–Chase experiment
510:By the time of the 1952
126:With the development of
81:Streptococcus pneumoniae
56:bacterial transformation
1370:Lehrer, Steven (2006).
1102:"Gene Recombination in
302:), and precise work by
1722:Spontaneous generation
1672:Germ theory of disease
1649:Zoology (through 1859)
690:"The quiet revolution"
637:The Journal of Hygiene
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1743:Philosophy of biology
1243:Phosphorus Metabolism
1008:on September 27, 2006
869:Fruton (1999), p. 439
843:Fruton (1999), p. 438
477:bacterial conjugation
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178:Griffith's experiment
115:Griffith's experiment
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106:in higher organisms.
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1847:Genetics experiments
1775:Human Genome Project
1687:Great chain of being
1654:Zoology (since 1859)
1589:Evolutionary thought
1559:Agricultural science
1455:. New York: Norton.
772:10.1084/jem.51.1.123
580:10.1084/jem.79.2.137
525:radioactive isotopes
379:tobacco mosaic virus
360:Reception and legacy
1852:Biology experiments
1780:Humboldtian science
1717:Sequence hypothesis
1624:Molecular evolution
1274:10.1085/jgp.36.1.39
1122:1946Natur.158..558L
975:1935Sci....81..644S
858:Profiles in Science
425:. Others, such as
395:sexual reproduction
300:deoxyribonuclease I
197:Martin Henry Dawson
150:ensues, generating
1790:Natural philosophy
1738:History of science
1538:History of biology
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521:bacterial genetics
453:awarded Avery the
435:molecular genetics
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172:Frederick Griffith
158:on the bacteria.
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1362:978-0-674-00169-5
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1349:Cobb, Matthew
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1316:(2): 207–32.
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1116:(4016): 558.
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529:bacteriophage
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475:demonstrated
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451:Royal Society
448:
445:, geneticist
444:
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419:Gregor Mendel
416:
415:Gunther Stent
411:
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408:Alfred Mirsky
404:
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387:autocatalysis
384:
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354:
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346:
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335:Colin MacLeod
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265:fractionation
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214:James Alloway
211:
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206:(rather than
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44:Colin MacLeod
41:
37:
30:
26:
21:
1828:
1629:Paleontology
1569:Biochemistry
1479:
1475:
1452:
1420:(2): 423–6.
1417:
1413:
1372:
1352:
1330:
1313:
1309:
1268:(1): 39–56.
1265:
1261:
1251:
1242:
1236:
1206:(6): 201–9.
1203:
1199:
1193:
1156:
1113:
1109:
1103:
1095:
1086:
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1075:
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1021:
1010:. Retrieved
1003:the original
966:
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626:
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571:
567:
509:
493:
487:to continue
480:
473:Edward Tatum
466:
463:André Boivin
459:
455:Copley Medal
447:H. J. Muller
439:
412:
363:
325:Oswald Avery
296:Moses Kunitz
293:
289:ribonuclease
285:chymotrypsin
269:
249:precipitated
238:
229:
227:
207:
201:
189:Fred Neufeld
186:
176:
168:Fred Neufeld
135:
125:
93:
89:
79:
67:
63:
40:Oswald Avery
35:
33:
1755:Ethnobotany
1644:RNA biology
1552:disciplines
1200:Experientia
505:Nobel Prize
497:phage group
399:phage group
391:chromosomes
375:chromosomes
160:Blood serum
1836:Categories
1697:Lamarckism
1604:Immunology
1302:References
1089:: 646–648.
1012:2009-02-26
743:: 324–340.
257:hydrolyzed
253:chloroform
251:out using
241:extracting
152:antibodies
144:inoculated
122:Background
1765:Dysgenics
1748:Teleology
1712:RNA world
1707:Protocell
1682:Darwinism
1663:Theories,
1634:Phycology
182:pneumonia
1818:Category
1760:Eugenics
1665:concepts
1609:Medicine
1594:Genetics
1542:timeline
1488:21789957
1414:Genetics
1292:12981234
1220:15421335
1140:21001945
999:17743301
915:16144981
885:Genetics
831:84395600
790:19869670
716:15296771
675:20474956
598:19871359
423:genetics
259:with an
234:antibody
203:in vitro
156:antigens
86:virulent
60:proteins
1731:Related
1599:Geology
1584:Ecology
1564:Anatomy
1550:Fields,
1444:8150273
1435:1205797
1283:2147348
1228:2522535
1148:1826960
1118:Bibcode
991:1658941
971:Bibcode
963:Science
906:1449782
781:2131805
666:2167760
657:4626734
589:2135445
481:E. coli
315:adenine
308:glycine
281:trypsin
277:protein
209:in vivo
195:and by
191:at the
132:strains
104:viruses
102:and/or
84:of the
68:primary
64:protein
1579:Botany
1486:
1459:
1442:
1432:
1394:
1380:
1359:
1337:
1290:
1280:
1226:
1218:
1146:
1138:
1110:Nature
1033:
997:
989:
913:
903:
829:
788:
778:
714:
673:
663:
655:
596:
586:
533:Watson
349:Watson
347:(with
261:enzyme
245:saline
230:smooth
46:, and
1224:S2CID
1144:S2CID
1006:(PDF)
987:JSTOR
959:(PDF)
827:S2CID
653:JSTOR
543:Notes
537:Crick
353:Crick
142:) is
140:mouse
136:types
134:, or
100:genes
1484:PMID
1457:ISBN
1440:PMID
1392:ISBN
1378:ISBN
1357:ISBN
1335:ISBN
1288:PMID
1216:PMID
1136:PMID
1031:ISBN
995:PMID
911:PMID
856:."
786:PMID
712:PMID
671:PMID
594:PMID
535:and
429:and
393:and
351:and
287:and
243:the
34:The
1430:PMC
1422:doi
1418:136
1318:doi
1278:PMC
1270:doi
1208:doi
1126:doi
1114:158
1087:221
979:doi
901:PMC
893:doi
889:170
819:doi
776:PMC
768:doi
702:doi
661:PMC
645:doi
584:PMC
576:doi
479:in
381:by
273:RNA
52:DNA
25:DNA
1838::
1480:33
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Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
