Ernst W. Mayr
Ernst W. Mayr |
![Ernst W. Mayr](/2009-wikipedia_en_wp1-0.7_2009-05/I/225px_Ernst_Mayr_PLoS.jpg)
Ernst W. Mayr
|
Born |
July 5, 1904
Kempten, Germany |
Died |
February 3, 2005 (aged 100)
Bedford, Massachusetts
|
Nationality |
German |
Fields |
evolutionary biology |
Notable awards |
Linnean Society of London's Darwin-Wallace Medal in 1958. |
Ernst Walter Mayr (July 5, 1904, Kempten, Germany – February 3, 2005, Bedford, Massachusetts U.S.), was one of the 20th century's leading evolutionary biologists. He was also a renowned taxonomist, tropical explorer, ornithologist, historian of science, and naturalist. His work contributed to the conceptual revolution that led to the modern evolutionary synthesis of Mendelian genetics, systematics, and Darwinian evolution, and to the development of the biological species concept.
Neither Darwin nor anyone else in his time knew the answer to the species problem: how multiple species could evolve from a single common ancestor. Ernst Mayr approached the problem with a new definition for the concept species. In his book Systematics and the Origin of Species (1942) he wrote that a species is not just a group of morphologically similar individuals, but a group that can breed only among themselves, excluding all others. When populations of organisms get isolated, the sub-populations will start to differ by genetic drift and natural selection over a period of time, and thereby evolve into new species. The most significant and rapid genetic reorganization occurs in extremely small populations that have been isolated (as on islands).
His theory of peripatric speciation (a more precise form of allopatric speciation which he advanced) based on his work on birds, is still considered a leading mode of speciation, and was the theoretical underpinning for the theory of punctuated equilibrium. Mayr is generally credited with inventing the modern philosophy of biology, particularly of evolutionary biology, which he distinguished from physics, for its introduction of (natural) history into science.
Biography
Mayr was the second son of Helene Pusinelli and Dr. Otto Mayr. His father was a jurist (District Prosecuting Attorney at Würzburg)[1] but took an interest in natural history and took the children out on field trips. He learnt all the local birds in Würzburg from his elder brother Otto. He also had access to a natural history magazine for amateurs, Kosmos. His father died just before he was thirteen. The family then moved to Dresden and he studied at the Staatsgymnasium (“Royal Gymnasium” until 1918) in Dresden-Neustadt and completed his high school education there. In April 1922, while still in high school, he joined the newly founded Saxony Ornithologists’ Association. Here he met Rudolf Zimmermann who was his ornithological mentor during this time. He passed his high school examination (Abitur), in February 1923, and Mayr's mother rewarded him with a pair of binoculars.[2] On 23 March 1923 on the lakes of Moritzburg, the Frauenteich, he spotted what he identified as a Red-crested Pochard. The species had not been seen in Saxony since 1845 and the local club argued about the identity. Raimund Schelcher (1891–1979) of the club then suggested that Mayr visit his classmate Erwin Stresemann on his way to Greifswald where Mayr was to begin his medical studies.[2] After a tough interrogation, Stresemann accepted and published the sighting as authentic. Stresemann was very impressed and suggested that Mayr could work as a volunteer between semesters in the ornithological section of the museum. Mayr wrote about this event It was as if someone had given me the key to heaven.[2] He entered the University of Greifswald in 1923 and, according to Mayr himself, "took the medical curriculum (to satisfy a family tradition)."[3] Mayr was endlessly interested in ornithology and "chose Greifswald at the Baltic for my studies for no other reason than that...it was situated in the ornithologically most interesting area."[3] Although he ostensibly planned to become a physician, he was "first and foremost an ornithologist."[3] During the first semester break Stresemann gave him a test to identify treecreepers and Mayr was able to identify most of the specimens correctly. Stresemann declared that Mayr 'was a born systematist'.[4] In 1925 Stresemann suggested that he give up his medical studies and join the Berlin Museum with the prospect of bird-collecting trips to the tropics on the condition that he completed his doctoral studies in 16 months. Mayr completed his doctorate in ornithology at the University of Berlin under Dr. Carl Zimmer, who was a full professor (Ordentlicher Professor), on 24 June 1926 at the age of 21. On 1 July he accepted the position offered to him at the Museum for a monthly salary of 330.54 Reichsmark.[5]
At the International Zoological Congress at Budapest in 1927, Mayr was introduced by Stresemann to banker and naturalist Walter Rothschild, who asked him to undertake an expedition to New Guinea on behalf of himself and the American Museum of Natural History in New York. In New Guinea Mayr collected several thousands bird skins (he named 26 new bird species during his lifetime) and, in the process also named 38 new orchid species. During his stay in New Guinea, he was invited to accompany the Whitney South Seas Expedition to the Solomon Islands.
He returned to Germany in 1930 and in 1931 he accepted a curatorial position at the American Museum of Natural History, where he played the important role of brokering and acquiring the Walter Rothschild collection of bird skins, which was being sold in order to pay off a blackmailer. During his time at the museum he produced numerous publications on bird taxonomy, and in 1942 his first book, Systematics and the Origin of Species, which completed the evolutionary synthesis started by Darwin.
After Mayr was appointed at the American Museum of Natural History, he influenced American ornithological research by cultivating mentoring relationships with young birdwatchers. Mayr organized a monthly seminar under the auspices of the Linnean Society of New York. This society, under the influence of J. A. Allen, Frank Chapman and Jonathan Dwight concentrated on taxonomy and later became a clearing house for bird banding and sight records. There were a group of eight young birdwatchers from the Bronx and later became the Bronx County Bird Club and they were led by Ludlow Griscom. Mayr was surprised at the differences between American and German Birding Societies. He noted that the German society was "far more scientific, far more interested in life histories and breeding bird species, as well as in reports on recent literature." Mayr also encouraged his Linnaean Society seminar participants to take up a specific research project of their own. "Everyone should have a problem" was the way one Bronx County Bird Club member recalled Mayr's refrain. One of Mayr's seminar participants was Joseph Hickey and under Mayr's influence went on to write A Guide to Birdwatching (1943). Hickey remembered later –"Mayr was our age and invited on all our field trips. The heckling of this German foreigner was tremendous, but he gave tit for tat, and any modern picture of Dr E. Mayr as a very formal person does not square with my memory of the 1930s. He held his own." Mayr's said of his own involvement with the local birdwatchers: "In those early years in New York when I was a stranger in a big city, it was the companionship and later friendship which I was offered in the Linnean Society that was the most important thing in my life."[6]
Another person that Mayr greatly influenced was Margaret Morse Nice. Mayr encouraged her to correspond with the European ornithologists of the time, and helped her in her landmark study on Song Sparrows. Nice wrote to Joseph Grinnell in 1932 trying to get foreign literature reviewed in the Condor: "Too many American ornithologists have despised the study of the living bird; the magazines and books that deal with the subject abound in careless statements, anthropomorphic interpretations, repetition of ancient errors, and sweeping conclusions from a pitiful array of facts. ... in Europe the study of the living bird is taken seriously. We could learn a great deal from their writing." Mayr ensured that Nice could publish her two volume Studies in the Life History of the Song Sparrow, finding her a publisher, and her book was reviewed by Aldo Leopold, Grinnell, Jean Delacour. Nice dedicated her book to "My Friend Ernst Mayr."[6]
Mayr joined the faculty of Harvard University in 1953, where he also served as director of the Museum of Comparative Zoology from 1961 to 1970. He retired in 1975 as emeritus professor of zoology, showered with honors. Following his retirement, he went on to publish more than 200 articles, in a variety of journals—more than some reputable scientists publish in their entire careers; 14 of his 25 books were published after he was 65. Even as a centenarian, he continued to write books. On his 100th birthday, he was interviewed by Scientific American magazine.
He received awards including the National Medal of Science, the Balzan Prize, the Sarton Medal of the History of Science Society, the International Prize for Biology, the Loye and Alden Miller Research Award, and the Lewis Thomas Prize for Writing about Science. In 1939 he was elected a Corresponding Member of the Royal Australasian Ornithologists Union. He was awarded the Linnean Society of London's prestigious Darwin-Wallace Medal in 1958. He was never awarded a Nobel Prize, but he noted that there is no Prize for evolutionary biology, and that Darwin would not have received one, either. Mayr did win a 1999 Craford Prize. That prize honors basic research in fields that do not qualify for Nobel Prizes and is administered by the same organization as the Nobel Prize.
Mayr was co-author of six global reviews of bird species new to science (listed below).
He was also an atheist, stating "there is nothing that supports the idea of a personal God".[7]
Mayr's ideas
As a traditionally trained biologist with little mathematical experience, Mayr was often highly critical of early mathematical approaches to evolution such as those of J.B.S. Haldane, famously calling in 1959 such approaches "beanbag genetics". He maintained that factors such as reproductive isolation had to be taken into account. In a similar fashion, Mayr was also quite critical of molecular evolutionary studies such as those of Carl Woese.
In many of his writings, Mayr rejected reductionism in evolutionary biology, arguing that evolutionary pressures act on the whole organism, not on single genes, and that genes can have different effects depending on the other genes present. He advocated a study of the whole genome rather than of isolated genes only. Current molecular studies in evolution and speciation indicate that although allopatric speciation seems to be the norm in groups (possibly those with greater mobility) such as the birds, there are numerous cases of sympatric speciation in many invertebrates (especially in the insects).
After articulating the biological species concept in 1942, Mayr played a central role in the species problem debate over what was the best species concept. He staunchly defended the biological species concept against the many definitions of "species" that others proposed.
Mayr was an outspoken defender of the scientific method, and one known to sharply critique science on the edge. As a notable recent example, he criticized the search for aliens as conducted by fellow Harvard professor Paul Horowitz as being a waste of university and student resources, for its inability to address and answer a scientific question.
Mayr rejected the idea of a gene-centered view of evolution and starkly but politely criticized Richard Dawkins' ideas:
The funny thing is if in England, you ask a man in the street who the greatest living Darwinian is, he will say Richard Dawkins. And indeed, Dawkins has done a marvelous job of popularizing Darwinism. But Dawkins' basic theory of the gene being the object of evolution is totally non-Darwinian. I would not call him the greatest Darwinian.[8]
Mayr insisted throughout his career that the gene as the target of selection cannot and should not be considered a valid idea in modern evolutionary thought.
“The idea that a few people have about the gene being the target of selection is completely impractical; a gene is never visible to natural selection, and in the genotype, it is always in the context with other genes, and the interaction with those other genes make a particular gene either more favorable or less favorable. In fact, Dobzhanksy, for instance, worked quite a bit on so-called lethal chromosomes which are highly successful in one combination, and lethal in another. Therefore people like Dawkins in England who still think the gene is the target of selection are evidently wrong. In the 30's and 40's, it was widely accepted that genes were the target of selection, because that was the only way they could be made accessible to mathematics, but now we know that it is really the whole genotype of the individual, not the gene. Except for that slight revision, the basic Darwinian theory hasn't changed in the last 50 years.”[8]
See also
Bibliography
Books
- Mayr, Ernst (1942). Systematics and the origin of species, from the viewpoint of a zoologist. Cambridge: Harvard University Press. ISBN 0-674-86250-3.
- Mayr, Ernst (1966). Animal Species and Evolution. Cambridge, Mass: Belknap Press. ISBN 0-674-03750-2.
- Mayr, Ernst. Populations, Species, and Evolution : An Abridgment of Animal Species and Evolution. Cambridge, Mass: Belknap Press. ISBN 0-674-69013-3.
- Mayr, Ernst. Evolution and the Diversity of Life : Selected Essays. Cambridge, Mass: Belknap Press. ISBN 0-674-27105-X.
- Mayr, Ernst. The Growth of Biological Thought: Diversity, Evolution, and Inheritance. Cambridge, Mass: Belknap Press. ISBN 0-674-36446-5.
- Mayr, Ernst. Toward a New Philosophy of Biology: Observations of an Evolutionist. Cambridge: Harvard University Press. ISBN 0-674-89666-1.
- Ashlock, Peter D.; Mayr, Ernst (1991). Principles of systematic zoology. New York: McGraw-Hill. ISBN 0-07-041144-1.
- Mayr, Ernst. One Long Argument: Charles Darwin and the Genesis of Modern Evolutionary Thought (Questions of Science). Cambridge: Harvard University Press. ISBN 0-674-63906-5.
- Mayr, Ernst (1997). This is biology: the science of the living world. Cambridge: Belknap Press of Harvard University Press. ISBN 0-674-88469-8.
- Diamond, Jared M.; Mayr, Ernst (2001). The birds of northern Melanesia: speciation, ecology & biogeography. Oxford [Oxfordshire]: Oxford University Press. ISBN 0-19-514170-9.
- Mayr, Ernst (2001). What evolution is. New York: Basic Books. ISBN 0-465-04426-3.
- Mayr, Ernst (2004). What makes biology unique?: considerations on the autonomy of a scientific discipline. Cambridge, UK: Cambridge University Press. ISBN 0-521-84114-3.
Global reviews of species new to science
- Zimmer, J. T. & E. Mayr (1943) New species of birds described from 1938 to 1941 The Auk Vol. 60 pp. 249-262
- Mayr, E. (1957) New species of birds described from 1941 to 1955 Journal for Ornithology Vol. 98 pp. 22-35
- Mayr, E. (1971) New species of birds described from 1956 to 1965 Jour. f. Ornith. Vol. 112 pp. 302-316
- Mayr, E. & F. Vuilleumier (1983) New species of birds described from 1966 to 1975 Jour. f. Ornith. Vol. 124 pp. 217-232
- Vuilleumier, F. & E. Mayr (1987) New species of birds described from 1976 to 1980 Jour. f. Ornith. Vol. 128 pp. 137-150
- Vuilleumier, François, Mary LeCroy & Ernst Mayr (1992) New species of birds described from 1981 to 1990 Bulletin of the British Ornithologists' Club Vol. 112A pp. 267-309
Other notable publications
- 1923 "Die Kolbenente (Nyroca rufina) auf dem Durchzuge in Sachsen". Ornithologische Monatsberichte 31:135-136
- 1923 "Der Zwergfliegenschnäpper bei Greifswald". Ornithologische Monatsberichte 31:136
- 1926 "Die Ausbreitung des Girlitz (Serinus canaria serinus L.) Ein Beitrag zur Tiergeographie". J. für Ornithologie 74:571-671
- 1927 "Die Schneefinken (Gattungen Montifringilla und Leucosticte)" J. für Ornithologie 75:596-619
- 1929 with W Meise. Zeitschriftenverzeichnis des Museums für Naturkunde Mitteilungen aus dem Zoologischen Museum in Berlin 14:1-187
- 1930 (by Ernst Hartert) "List of birds collected by Ernst Mayr". Ornithologische Monatsberichte 36:27-128
- 1930 "My Dutch New Guinea Expedition". 1928. Ornithologische Monatsberichte 36:20-26
- 1931 Die Vögel des Saruwaged und Herzoggebirges (NO Neuginea) Mitteilungen aus dem Zoologischen Museum in Berlin 17:639-723
- 1931 "Birds collected during the Whitney South Sea Expedition. XII Notes on Halcyon chloris and some of its subspecies". American Museum Novitates no 469
- 1932 "A tenderfoot explorer in New Guinea" Natural History 32:83-97
- 1935 "Bernard Altum and the territory theory". Proceedings of the Linnaean Society of New York 45, 46:24-38
- 1940 "Speciation phenomena in birds". American Naturalist 74:249-278
- 1941 "Borders and subdivision of the Polynesian region as based on our knowledge of the distribution of birds". Proceedings of the 6th Pacific Scientific Congress 4:191-195
- 1941 "The origin and history of the bird fauna of Polynesia". Proceedings of the 6th Pacific Scientific Congress 4:197-216
- 1943 "A journey to the Solomons". Natural History 52:30-37,48
- 1944 "Wallace's Line in the light of recent zoogeographics studies". Quarterly Review of Biology 19:1-14
- 1944 "The birds of Timor and Sumba". Bulletin of the American Museum of Natural History 83:123-194
- 1944 "Timor and the colonization of Australia by birds". Emu 44:113-130
- 1946 "History of the North American bird fauna" Wilson Bulletin 58:3-41
- 1946 "The naturalist in Leidy's time and today". Proceedings of the Academy of Natural Sciences of Philadelphia 98:271-276
- 1947 "Ecological factors in speciation". Evolution 1:263-288
- 1948 "The new Sanford Hall". Natural History 57:248-254
- 1950 The role of the antennae in the mating behavior of female Drosophila. Evolution 4:149-154
- 1951 Introduction and Conclusion. Pages 85,255-258 in The problem of land connections across the South Atlantic with special reference to the Mesozoic. Bulletin of the American Museum of Natural History 99:79-258
- 1951 with Dean Amadon, "A classification of recent birds". American Museum Novitates no. 1496
- 1953 with E G Linsley and R L Usinger. Methods and Principles of Systematica Zoology. McGraw-Hill, New York.
- 1954 "Changes in genetic environment and evolution". Pages 157-180 in Evolution as a Process (J Huxley, A C Hardy and E B Ford Eds) Allen and Unwin. London
- 1955 "Karl Jordan's contribution to current concepts in systematics and evolution". Transactions of the Royal Entomological Society of London 107:45-66
- 1956 with C B Rosen. "Geographic variation and hybridization in populations of Bahama snails (Cerion)". American Museum Novitates no 1806.
- 1957 "Species concepts and definitions". Pages 371-388 in The Species Problem (E. Mayr ed). AAAS, Washington DC.
- 1959 "The emergence of evolutionary novelties". Pages 349-380 in The Evolution of Life: Evolution after Darwin, vol 1 (S. Tax, ed) University of Chicago.
- 1959 "Darwin and the evolutionary theory in Biology". Pages 1-10 in Evolution and Anthropology: A Centennial Appraisal (B J Meggers, Ed) The Anthropological Society of Washington, Washington DC.
- 1959 "Agassiz, Darwin, and Evolution". Harvard Library Bulletin. 13:165-194
- 1961 "Cause and effect in biology: Kinds of causes, predictability, and teleology are viewed by a practicing biologist". Science 134:1501-1506
- 1962 "Accident or design: The paradox of evolution". Pages 1-14 in The Evolution of Living Organisms (G W Leeper, Ed) Melbourne University Press.
- 1964 Introduction, Bibliography and Subject Pages vii-xxviii, 491-513 in On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life, by Charles Darwin. A Facsimile of the First Edition. Harvard University Press.
- 1965 Comments. In Proceedings of the Boston Colloguium for the Philosophy of Science, 1962-1964. Boston Studies in the Philosophy of Science 2:151-156
- 1969 Discussion: Footnotes on the philosophy of biology. Philosophy of Science 36:197-202
- 1972 Continental drift and the history of the Australian bird fauna. Emu 72:26-28
- 1972 Geography and ecology as faunal determinants. Pages 549-561 in Proceedings XVth International Ornithological Congress (K H Voous, Ed) E J Brill, Leiden, The Netherlands.
- 1972 Lamarck revisited. Journal of the History of Biology. 5:55-94
- 1974 Teleological and teleonomic: A new analysis. Boston studies in the Philosophy of Science 14:91-117
- 1978 Tenure: A sacred cow? Science 199:1293
- 1980 How I became a Darwinian, Pages 413-423 in The Evolutionary Synthesis (E Mayr and W Provine, Eds) Harvard University Press, Cambridge, Massachusetts.
- 1980 with W B Provine, Eds. The Evolutionary Synthesis. Harvard University Press.
- 1981 Evolutionary biology. Pages 147-162 in The Joys of Research (W. Shripshire Jr, Ed.) Smithsonian Institution Press.
- 1984 Evolution and ethics. Pages 35-46 in Darwin, Mars and Freud: Their influence on Moral Theory (A L Caplan and B Jennings, Eds.) Plenum Press, New York.
- 1985. Darwin's five theories of evolution. In D. Kohn, ed., The Darwinian Heritage, Princeton NJ: Princeton University Press, pp. 755-772.
- 1985. How biology differs from the physical sciences. In D. J. Depew and B H Weber, eds., Evolution at a Crossroads: The New Biology and the New Philosophy of Science, Cambridge MA: The MIT Press, pp. 43-63.
- 1988. The why and how of species. Biology and Philosophy 3:431-441
- 1992. The idea of teleology. Journal of the History of Ideas 53:117-135
- 1994. with W.J. Bock. Provisional classifications v. standard avian sequences: heuristics and communication in ornithology. Ibis 136:12-18
- 1996. What is a species, and what is not? Philosophy of Science 63 (June): 262-277.
- 1996. The autonomy of biology: the position of biology among the sciences. Quarterly Review of Biology 71:97-106
- 1997. The objects of selection Proc. Natl. Acad. Sci. USA 94 (March): 2091-94.
- 1999. Darwin's influence on modern thought Crafoord Prize lecture, September 23, 1999.
- 2000. Biology in the Twenty-First Century Bioscience 50 (Oct. 2000): 895-897.
- 2001. The philosophical foundations of Darwinism Proceedings of the American Philosophical Society 145:488-495
- 2002. with Walter J Bock. Classifications and other ordering systems. Zeitschrift Zool. Syst. Evolut-Forsch. 40:1-25
Notes
References
- Coyne, Jerry. 2005. Ernst Mayr (1904-2005). Science 307:1212-1213.
- Diamond, Jared. 2005. Obituary: Ernst Mayr (1904−2005). Nature 433:700-701.
- Haffer, Jürgen 2007. Ornithology, Evolution, and Philosophy: The Life and Science of Ernst Mayr 1904–2005. Springer.
- Milner, Richard. 1990. The Encyclopedia of Evolution Facts on File, New York. ISBN 0-8160-1472-8
- Schilthuizen, Menno. 2001. Frogs, Flies, and Dandelions: Speciation-The Evolution of New Species. Oxford ISBN 0-19-850393-8
- Kutschera, Ulrich. 2006. Dogma, not faith, is the barrier to scientific enquiry Nature 443, 26.
- Mayr, E. & William B. Provine (Eds) 1998. The Evolutionary Synthesis: Perspectives on the Unification of Biology. Harvard University Press.
- Meyer, Axel On the Importance of Being Ernst Mayr PLoS Biology Vol. 3, No. 5, e152 DOI:10.1371/journal.pbio.0030152
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1970s
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1970: Robert H. Dicke · Allan R. Sandage · John C. Slater · John A. Wheeler · Saul Winstein 1973: Carl Djerassi · Maurice Ewing · Arie Jan Haagen-Smit · Vladimir Haensel · Frederick Seitz · Robert Rathbun Wilson 1974: Nicolaas Bloembergen · Paul Flory · William Alfred Fowler · Linus Carl Pauling · Kenneth Sanborn Pitzer 1975: Hans A. Bethe · Joseph Hirschfelder · Lewis Sarett · E. Bright Wilson · Chien-Shiung Wu 1976: Samuel Goudsmit · Herbert S. Gutowsky · Frederick Rossini · Verner Suomi · Henry Taube · George Uhlenbeck 1979: Richard P. Feynman · Herman Mark · Edward M. Purcell · John Sinfelt · Lyman Spitzer · Victor F. Weisskopf
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1980s
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1982: Philip W. Anderson · Yoichiro Nambu · Edward Teller · Charles H. Townes 1983: E. Margaret Burbidge · Maurice Goldhaber · Helmut Landsberg · Walter Munk · Frederick Reines · Bruno B. Rossi · J. Robert Schrieffer 1986: Solomon Buchsbaum · Horace Crane · Herman Feshbach · Robert Hofstadter · Chen Ning Yang 1987: Philip Abelson · Walter Elsasser · Paul C. Lauterbur · George Pake · James A. Van Allen 1988: D. Allan Bromley · Paul Ching-Wu Chu · Walter Kohn · Norman F. Ramsey · Jack Steinberger 1989: Arnold O. Beckman · Eugene Parker · Robert Sharp · Henry Stommel
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1990s
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1990: Allan M. Cormack · Edwin M. McMillan · Robert Pound · Roger Revelle 1991: Arthur L. Schawlow · Ed Stone · Steven Weinberg 1992: Eugene M. Shoemaker 1993: Val Fitch · Vera Rubin 1994: Albert Overhauser · Frank Press 1995: Hans Dehmelt · Peter Goldreich 1996: Wallace S. Broecker 1997: Marshall Rosenbluth · Martin Schwarzschild · George Wetherill 1998: Don L. Anderson · John N. Bahcall 1999: James Cronin · Leo Kadanoff
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2000s
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2000: Willis E. Lamb · Jeremiah P. Ostriker · Gilbert F. White 2001: Marvin L. Cohen · Raymond Davis Jr. · Charles Keeling 2002: Richard Garwin · W. Jason Morgan · Edward Witten 2003: G. Brent Dalrymple · Riccardo Giacconi 2004: Robert N. Clayton 2005: Ralph A. Alpher · Lonnie Thompson 2006: Daniel Kleppner
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