Editing Carl Woese

{{Short description|American microbiologist (1928–2012)}}
{{Use mdy dates|date=November 2011}}
{{Infobox scientist
|name              = Carl Woese
|image             = Carl Woese.jpg
|image_size        = 
|caption           = Woese in 2004
|birth_date        = {{birth date|1928|07|15}}
|birth_place       = [[Syracuse, New York]], U.S.
|death_date        = {{death date and age|2012|12|30|1928|07|15}}
|death_place       = [[Urbana, Illinois]], U.S.
|ethnicity         =
|field             = [[Microbiology]]
|work_institutions = [[University of Illinois Urbana–Champaign]]
|education         = {{unbulleted list|[[Amherst College]]|[[Yale University]]}}
| thesis_title = Physical Studies on Animal viruses
| thesis_url = https://www.proquest.com/docview/302030453/
| thesis_year = 1953
|doctoral_advisor  = [[Ernest C. Pollard]]<ref name="pnas2012"/>
|doctoral_students =
|notable_students = [[David Stahl (biologist)|David Stahl]]<ref>{{cite web |url=https://mcb.illinois.edu/departments/microbiology/downloads/history_of_microbiology_at_illinois.pdf |archive-url=https://ghostarchive.org/archive/20221009/https://mcb.illinois.edu/departments/microbiology/downloads/history_of_microbiology_at_illinois.pdf |archive-date=2022-10-09 |url-status=live |title=History of the Department of Microbiology |publisher=University of Illinois Urbana–Champaign |date=2015-06-01 |access-date=2017-03-09}}</ref>
|known_for         = Recognition of [[Archaea]] as a domain of life
|influences        =
|influenced        =
|prizes            = {{Plainlist|
* [[Leeuwenhoek Medal]] (1992)
* [[Selman A. Waksman Award in Microbiology|Selman A. Waksman Award]] (1995)
* [[National Medal of Science]] (2000)
* [[Crafoord Prize]] (2003)
}}
|religion          =
|footnotes         =
|signature         =
}}
'''Carl Richard Woese''' ({{IPAc-en|w|oʊ|z}} {{respell|WOHZ}};<ref name="sayhow">{{cite web | editor-last = Hagen | editor-first = Ray | title = Say How? A Pronunciation Guide to Names of Public Figures | work = National Library Service for the Blind and Physically Handicapped | date = August 2012 | url = https://www.loc.gov/nls/about/organization/standards-guidelines/uvwx/#w }}</ref> July 15, 1928 – December 30, 2012) was an American [[microbiologist]] and [[biophysicist]]. Woese is famous for defining the [[Archaea]] (a new [[domain (biology)|domain]] of life) in 1977 through a pioneering [[phylogenetic]] [[taxonomy (biology)|taxonomy]] of [[16S ribosomal RNA]], a technique that has revolutionized microbiology.<ref name="Woese_1990">{{cite journal | last1 = Woese | first1 = Carl R.| author-link1 = Carl Woese | last2 =  Kandler | first2 = O | last3 = Wheelis | first3= M | title = Towards a natural system of organisms: proposal for the domains Archaea, Bacteria, and Eucarya | journal = Proc Natl Acad Sci USA | volume = 87 | issue = 12 | pages = 4576–9 | year = 1990 | pmid = 2112744 | doi = 10.1073/pnas.87.12.4576 | pmc = 54159 | bibcode=1990PNAS...87.4576W| author2-link = Otto Kandler| doi-access = free}}</ref><ref name="woese1978">{{cite journal | last1 = Woese | first1 = C.R.| author-link1 = Carl Woese | last2 = Magrum | first2 = L.J. | last3 = Fox | first3 = G.E.| author-link3 = George E. Fox | title = Archaebacteria | journal = J Mol Evol | volume = 11 | issue = 3 | pages = 245–51 | year = 1978 | pmid = 691075 | doi = 10.1007/BF01734485| bibcode = 1978JMolE..11..245W | s2cid = 260611975}}</ref><ref name="woese1977">{{cite journal | issn = 0027-8424| volume = 74| issue = 11| pages = 5088–5090| last1 = Woese| first1 = C. R.| author-link1 = Carl Woese| author2 = G. E. Fox| author-link2 = George E. Fox| title = Phylogenetic structure of the prokaryotic domain: The primary kingdoms| journal = Proceedings of the National Academy of Sciences| date = 1977-11-01| pmid = 270744 | pmc = 432104| doi = 10.1073/pnas.74.11.5088| bibcode = 1977PNAS...74.5088W| doi-access = free}}{{open access}}</ref><ref name="morell1997"/> He also originated the [[RNA world hypothesis]] in 1967, although not by that name.<ref name="woese1967">{{Cite book | publisher = Harper & Row | last = Woese | first = Carl | title = The Genetic Code: the Molecular basis for Genetic Expression | location = New York | year = 1967 | author-link = Carl Woese}}</ref> Woese held the [[Stanley O. Ikenberry]] Chair and was professor of microbiology at the [[University of Illinois Urbana–Champaign]].<ref name="natureobit">{{Cite journal | last1 = Noller | first1 = H. | author-link = Harry F. Noller| title = Carl Woese (1928–2012) Discoverer of life's third domain, the Archaea| doi = 10.1038/493610a | journal = Nature | volume = 493 | issue = 7434 | page = 610 | year = 2013 | pmid =  23364736|bibcode = 2013Natur.493..610N | s2cid = 205076152 | doi-access = free }}</ref><ref>{{Cite journal 
| last1 = Goldenfeld | first1 = N. 
| last2 = Pace | first2 = N. R. 
| author-link2 = Norman R. Pace
| doi = 10.1126/science.1235219 
| title = Retrospective: Carl R. Woese (1928-2012) 
| journal = Science 
| volume = 339 
| issue = 6120 
| page = 661 
| year = 2013 
| pmid = 23393257 
|bibcode = 2013Sci...339..661G | s2cid = 36566952 
}}</ref><ref name="uofi2006">{{cite web
 |url         = http://news.illinois.edu/NEWS/06/0519woese.html
 |title       = U. of I. microbiologist Carl Woese elected to Royal Society
 |access-date  = 2009-03-02
 |work        = News Bureau, University of Illinois Urbana–Champaign
 |date        = 2006-05-19
 |url-status     = dead
 |archive-url  = https://web.archive.org/web/20120213181353/http://news.illinois.edu/NEWS/06/0519woese.html
 |archive-date = February 13, 2012
 |df          = mdy-all
}}</ref>
{{TOC limit|4}}

==Life and education==
Woese was born in Syracuse, New York on July 15, 1928. His family was [[German American]]. Woese attended [[Deerfield Academy]] in [[Massachusetts]]. He received a bachelor's degree in [[mathematics]] and [[physics]] from [[Amherst College]] in 1950. During his time at Amherst, Woese took only one biology course ([[Biochemistry]], in his senior year) and had "no scientific interest in plants and animals" until advised by [[William M. Fairbank]], then an assistant professor of physics at Amherst, to pursue [[biophysics]] at [[Yale]].<ref name="woese2005qa">{{Cite journal | last1 = Woese | first1 = C. R. | author-link = Carl Woese| title = Q & A | doi = 10.1016/j.cub.2005.02.003 | journal = Current Biology | volume = 15 | issue = 4 | pages = R111–R112 | year = 2005 | pmid =  15723774| s2cid = 45434594 | doi-access = free | bibcode = 2005CBio...15.R111W }}</ref>

In 1953, he completed a [[PhD]] in [[biophysics]] at [[Yale University]], where his doctoral research focused on the inactivation of [[virus]]es by heat and [[ionizing radiation]].<ref name=mcb>{{cite web |url=http://mcb.illinois.edu/faculty/profile/1204 |title=Carl R Woese, Professor of Microbiology |publisher=University of Illinois Urbana–Champaign |access-date=February 16, 2010 |url-status=dead |archive-url=https://web.archive.org/web/20100213210457/http://mcb.illinois.edu/faculty/profile/1204 |archive-date=February 13, 2010 |df=mdy }}</ref><ref name="sapp2009"/> He studied medicine at the [[University of Rochester]] for two years.<ref name="sapp2009">{{Cite book
| publisher = Oxford University Press
| isbn = 978-0-199-73438-2
| last = Sapp
| first = Jan A.
| title = The new foundations of evolution: on the tree of life
| location = New York
| year = 2009
}}</ref> Then he became a postdoctoral researcher in biophysics at Yale University investigating bacterial spores.<ref name="woese1960rad">{{Cite journal 
| last1 = Woese | first1 = C. R. 
| author-link1 = Carl Woese
| title = Phage induction in germinating spores of Bacillus megaterium 
| journal = Radiation Research 
| volume = 13 
| issue = 6 
| pages = 871–878 
| year = 1960 
| pmid = 13786177
 | doi=10.2307/3570863
| bibcode = 1960RadR...13..871W| jstor = 3570863 
}}</ref> From 1960 to 1963, he worked as a biophysicist at the [[General Electric Research Laboratory]] in [[Schenectady, New York]].<ref name=mcb /><ref name=newsgazette>{{cite web
|url=http://www.news-gazette.com/news/university-illinois/2012-12-30/visionary-ui-biologist-carl-woese-84-dies.html
|title=Visionary UI biologist Carl Woese, 84, dies
|access-date=2012-12-31
|work=The News-Gazette: Serving East Central Illinois
|date=2012-12-30
|archive-date=February 16, 2013
|archive-url=https://web.archive.org/web/20130216170841/http://www.news-gazette.com/news/university-illinois/2012-12-30/visionary-ui-biologist-carl-woese-84-dies.html
|url-status=dead
}}</ref> In 1964, Woese joined the microbiology faculty of the University of Illinois Urbana–Champaign, where he focused on Archaea, genomics, and [[molecular evolution]] as his areas of expertise.<ref name="uofi2006"/><ref name=mcb /><ref name=newsgazette /> He became a professor at the University of Illinois Urbana–Champaign's [[Carl R. Woese Institute for Genomic Biology]], which was renamed in his honor in 2015, after his death.<ref name=newsgazette />

Woese died on December 30, 2012, following complications from [[pancreatic cancer]], leaving as survivors his wife Gabriella and a son and daughter.<ref>{{cite news|url=https://www.washingtonpost.com/local/obituaries/carl-woese-dies-evolutionary-biologist-was-84/2013/01/19/a91a051c-61e9-11e2-b05a-605528f6b712_story.html |title=Carl Woese dies; evolutionary biologist was 84 |newspaper=The Washington Post |date=2013-01-19 |accessdate=2022-02-16}}</ref><ref name="uofiigb2012">{{cite web
 |url         = http://www.igb.illinois.edu/news/carl-r-woese-1928-%E2%80%93-2012
 |title       = Carl R. Woese: 1928 – 2012
 |access-date  = 2012-12-30
 |work        = News, The Institute for Genomic Biology, University of Illinois Urbana–Champaign
 |date        = 2012-12-30
 |url-status     = dead
 |archive-url  = https://web.archive.org/web/20130102225304/http://www.igb.illinois.edu/news/carl-r-woese-1928-%E2%80%93-2012
 |archive-date = January 2, 2013
 |df          = mdy-all
}}</ref><ref name=nytobit>{{cite news |title=Carl Woese Dies at 84. Discovered Life's 'Third Domain' |url=https://www.nytimes.com/2013/01/01/science/carl-woese-dies-discovered-lifes-third-domain.html |quote=Carl Woese, a biophysicist and evolutionary microbiologist whose discovery 35 years ago of a “third domain” of life in the vast realm of micro-organisms altered scientific understanding of evolution, died on Sunday at his home in Urbana, Ill. He was 84. ... |newspaper=[[The New York Times]] |date=December 31, 2012 |access-date=2013-01-04 }}</ref>

==Work and discoveries==
=== Early work on the genetic code ===
Woese turned his attention to the [[genetic code]] while setting up his lab at [[General Electric]]'s [[GE Global Research|Knolls Laboratory]] in the fall of 1960.<ref name="sapp2009"/> Interest among physicists and molecular biologists had begun to coalesce around deciphering the correspondence between the twenty [[amino acids]] and the four letter alphabet of [[Nucleobase|nucleic acid bases]] in the decade following [[James D. Watson]], [[Francis Crick]], and [[Rosalind Franklin]]'s discovery of the structure of DNA in 1953.<ref name="pnas2012"/> Woese published a series of papers on the topic. In one, he deduced a correspondence table between what was then known as "soluble RNA" and DNA based upon their respective [[base pair]] ratios.<ref name="woese1961ratio">{{Cite journal 
| doi = 10.1038/189920a0 
| last1 = Woese | first1 = C. R. 
| author-link1 = Carl Woese
| title = Composition of various ribonucleic acid fractions from micro-organisms of different deoxyribonucleic acid composition 
| journal = Nature 
| volume = 189 
| issue = 4768 
| pages = 920–921 
| year = 1961 
| pmid = 13786175
|bibcode = 1961Natur.189..920W | s2cid = 4201322 }}</ref> He then re-evaluated experimental data associated with the hypothesis that viruses used one base, rather than a triplet, to encode each amino acid, and suggested 18 codons, correctly predicting one for [[proline]].<ref name="sapp2009"/><ref name="woese1961viruses">{{Cite journal 
| doi = 10.1038/190697a0 
| last1 = Woese | first1 = C. R. 
| author-link1 = Carl Woese
| title = Coding ratio for the ribonucleic acid viruses 
| journal = Nature 
| volume = 190 
| issue = 4777 
| pages = 697–698 
| year = 1961 
| pmid = 13786174
|bibcode = 1961Natur.190..697W | s2cid = 4221490 }}</ref> Other work established the mechanistic basis of protein translation, but in Woese's view, largely overlooked the genetic code's evolutionary origins as an afterthought.<ref name="pnas2012"/>

In 1962, Woese spent several months as a visiting researcher at the [[Pasteur Institute]] in [[Paris]], a locus of intense activity on the molecular biology of gene expression and gene regulation.<ref name="sapp2009"/> While in Paris, he met [[Sol Spiegelman]], who invited Woese to visit the [[University of Illinois]] after hearing his research goals; at this visit Spiegelman offered Woese a position with immediate [[tenure]] beginning in the fall of 1964.<ref name="sapp2009"/> With the freedom to patiently pursue more speculative threads of inquiry outside the mainstream of biological research, Woese began to consider the genetic code in evolutionary terms, asking how the codon assignments and their translation into an amino acid sequence might have evolved.<ref name="sapp2009"/><ref name="woese1964">{{Cite journal | last1 = Woese | first1 = C. R. | author-link1 = Carl Woese| last2 = Hinegardner | first2 = R. T. | last3 = Engelberg | first3 = J. | doi = 10.1126/science.144.3621.1030 | title = Universality in the Genetic Code | journal = Science | volume = 144 | issue = 3621 | pages = 1030–1031 | year = 1964 | pmid =  14137944|bibcode = 1964Sci...144.1030W | doi-access =  }}</ref>

=== Discovery of the third domain ===
For much of the 20th century, prokaryotes were regarded as a single group of organisms and classified based on their [[biochemistry]], [[morphology (biology)|morphology]] and [[metabolism]]. In a highly influential 1962 paper, [[Roger Stanier]] and [[C. B. van Niel]] first established the division of cellular organization into [[prokaryote]]s and [[eukaryote]]s, defining prokaryotes as those organisms lacking a [[cell nucleus]].<ref name=stanier1962>{{Cite journal | doi = 10.1007/BF00425185 | last1 = Stanier | first1 = R. Y. | author-link1 = Roger Stanier| last2 = Van Niel | first2 = C. B. | author-link2 = C. B. van Niel| title = The concept of a bacterium | journal = Archiv für Mikrobiologie | volume = 42 | pages = 17–35 | year = 1962 | issue = 1 | pmid = 13916221| bibcode = 1962ArMic..42...17S | s2cid = 29859498 }}</ref><ref name=pace2009>{{Cite journal | last1 = Pace | first1 = N. R. | author-link1 = Norman R. Pace| title = Problems with "Procaryote" | doi = 10.1128/JB.01224-08 | journal = Journal of Bacteriology | volume = 191 | issue = 7 | pages = 2008–2010; discussion 2010 | year = 2009 | pmid =  19168605| pmc =2655486 }}</ref> Adapted from [[Édouard Chatton]]'s generalization, Stanier and Van Niel's concept was quickly accepted as the most important distinction among organisms; yet they were nevertheless skeptical of microbiologists' attempts to construct a natural [[phylogenetic]] classification of bacteria.<ref name=sapp2005>{{Cite journal | last1 = Sapp | first1 = J. | author-link1 = Jan Sapp| title = The Prokaryote-Eukaryote Dichotomy: Meanings and Mythology | doi = 10.1128/MMBR.69.2.292-305.2005 | journal = Microbiology and Molecular Biology Reviews | volume = 69 | issue = 2 | pages = 292–305 | year = 2005 | pmid =  15944457| pmc =1197417 }}</ref> However, it became generally assumed that all life shared a common prokaryotic (implied by the [[Greek language|Greek]] root πρό (pro-), before, in front of) ancestor.<ref name="pace2009" /><ref name=oren2010>{{Cite book| chapter = Concepts About Phylogeny of Microorganisms–an Historical Perspective| publisher = Caister Academic Press| isbn = 9781904455677| pages = 1–22|editor1=Aharon Oren |editor2=R. Thane Papke 
| last = Oren| first = Aharon| title = Molecular Phylogeny of Microorganisms| location = Norfolk, UK| date = 2010-07-01| chapter-url = https://books.google.com/books?id=a5t9DYZ-wccC&q=concepts+Aharon+Oren+phylogeny&pg=PR7}}</ref>

In 1977, Woese and [[George E. Fox]] experimentally disproved this universally held hypothesis about the basic structure of the [[tree of life (biology)|tree of life]].<ref name="pace2011pnas" /> Woese and Fox discovered a kind of microbial life which they called the “archaebacteria” ([[Archaea]]).<ref name="woese1977" /> They reported that the archaebacteria comprised "a third kingdom" of life as distinct from bacteria as plants and animals.<ref name="woese1977" /> Having defined Archaea as a new "urkingdom" (later [[domain (biology)|domain]]) which were neither bacteria nor eukaryotes, Woese redrew the [[Taxonomy (biology)|taxonomic]] tree. His [[three-domain system]], based on phylogenetic relationships rather than obvious morphological similarities, divided life into 23 main divisions, incorporated within three domains: [[Bacteria]], [[Archaea]], and [[Eukaryote|Eucarya]].<ref name="Woese_1990" />

[[File:PhylogeneticTree, Woese 1990.PNG|thumb|left|450px|Phylogenetic tree based on Woese et al. rRNA analysis. The vertical line at bottom represents the [[last universal common ancestor]] (LUCA).<ref name="Woese_1990"/>]]

Acceptance of the validity of Woese's phylogenetically valid classification was a slow process. Prominent biologists including [[Salvador Luria]] and [[Ernst Mayr]] objected to his division of the prokaryotes.<ref>{{Cite journal | title = Two empires or three?| doi = 10.1073/pnas.95.17.9720| volume = 95
| issue = 17| pages = 9720–9723| last = Mayr| first = Ernst| author-link = Ernst Mayr| journal = Proceedings of the National Academy of Sciences |year = 1998| bibcode=1998PNAS...95.9720M| pmid=9707542| pmc=33883| doi-access = free}}</ref><ref name="sapp2007">{{Cite journal| title = The structure of microbial evolutionary theory| doi = 10.1016/j.shpsc.2007.09.011| pmid = 18053933|  volume = 38| issue = 4| pages = 780–95 | last = Sapp| first = Jan A.| author-link = Jan Sapp| journal = Studies in History and Philosophy of Science Part C: Studies in History and Philosophy of Biological and Biomedical Sciences |date=December 2007}}</ref> Not all criticism of him was restricted to the scientific level. A decade of labor-intensive [[oligonucleotide]] cataloging left him with a reputation as "a crank," and Woese would go on to be dubbed as "Microbiology's Scarred Revolutionary" by a news article printed in the journal ''[[science (journal)|Science]]''.<ref name="morell1997">{{Cite journal| title = Microbiology's scarred revolutionary| doi = 10.1126/science.276.5313.699| issn = 0036-8075| volume = 276| issue = 5313| pages = 699–702| last = Morell| first = V.| journal = Science| date = 1997-05-02| pmid = 9157549| s2cid = 84866217}}</ref> The growing body of supporting data led the [[scientific community]] to accept the Archaea by the mid-1980s.<ref name="sapp2009"/> Today, few scientists cling to the idea of a unified Prokarya.

Woese's work on Archaea is also significant in its implications for the search for life on other planets. Before the discovery by Woese and Fox, scientists thought that Archaea were extreme organisms that evolved from the microorganisms more familiar to us. Now, most believe they are ancient, and may have robust evolutionary connections to the first organisms on Earth.<ref>{{Cite journal| title = Archaeal phylogenomics provides evidence in support of a methanogenic origin of the Archaea and a thaumarchaeal origin for the eukaryotes| doi = 10.1098/rspb.2010.1427| volume = 278| issue = 1708| pages = 1009–1018| last = Kelly| first = S.|author2=B. Wickstead |author3=K. Gull | journal = Proceedings of the Royal Society B: Biological Sciences| date = 2010-09-29 | pmid=20880885 | pmc=3049024}}</ref> Organisms similar to those archaea that exist in extreme environments may have developed on other planets, some of which harbor conditions conducive to [[extremophile]] life.<ref name="stetter2006">{{Cite journal| title = Hyperthermophiles in the history of life| doi = 10.1098/rstb.2006.1907| volume = 361| issue = 1474| pages = 1837–1843| last = Stetter| first = Karl O. | journal = Philosophical Transactions of the Royal Society B: Biological Sciences| date = 2006-10-29| pmc = 1664684| pmid=17008222}}</ref>

Notably, Woese's elucidation of the [[Tree of life (biology)|tree of life]] shows the overwhelming diversity of microbial lineages: single-celled organisms represent the vast majority of the biosphere's genetic, metabolic, and ecologic niche diversity.<ref name=woese2006prok>{{Cite book | last1 = Woese | first1 = C. R. | chapter = How We Do, Don’t and Should Look at Bacteria and Bacteriology | title = The Prokaryotes | pages = 3–4 | year = 2006 | isbn = 978-0-387-25476-0 | doi = 10.1007/0-387-30741-9_1}}</ref> As microbes are crucial for many [[biogeochemical cycle]]s and to the continued function of the biosphere, Woese's efforts to clarify the evolution and diversity of microbes provided an invaluable service to [[ecologists]] and [[conservationists]]. It was a major contribution to the theory of [[evolution]] and to our knowledge of the history of life.<ref name=pnas2012>{{cite journal | doi = 10.1073/pnas.1120749109 | issn = 1091-6490 | volume = 109 | issue = 4 | pages = 1019–1021 | last = Nair | first = Prashant | title = Woese and Fox: Life, rearranged | journal = Proceedings of the National Academy of Sciences | date = 2012-01-17 | bibcode=  2012PNAS..109.1019N | pmid=22308527 | pmc=3268309| doi-access = free }}</ref>

Woese wrote, "My evolutionary concerns center on the bacteria and the archaea, whose evolutions cover most of the planet's 4.5-billion-year history. Using ribosomal RNA sequence as an evolutionary measure, my laboratory has reconstructed the phylogeny of both groups, and thereby provided a phylogenetically valid system of classification for prokaryotes. The discovery of the archaea was in fact a product of these studies".<ref name=mcb />

=== Evolution of primary cell types ===
Woese also speculated about an era of rapid evolution in which considerable [[horizontal gene transfer]] occurred between organisms.<ref name="pace2011pnas">{{Cite journal
| title = Phylogeny and beyond: Scientific, historical, and conceptual significance of the first tree of life
| doi = 10.1073/pnas.1109716109
| issn = 1091-6490 
| volume = 109
| issue = 4
| pages = 1011–1018
| last1 = Pace
| first1 = Norman R.
| author-link1 = Norman R. Pace
| first2 = Jan 
| last2 = Sapp
| author-link2 = Jan Sapp
| first3 = Nigel
| last3 = Goldenfeld
| journal = Proceedings of the National Academy of Sciences
| date = 2012-01-24
|bibcode = 2012PNAS..109.1011P
| pmid=22308526
| pmc=3268332| doi-access = free
}}</ref><ref name="woese2002cells">{{Cite journal
| title = On the evolution of cells
| doi = 10.1073/pnas.132266999
| volume = 99
| issue = 13
| pages = 8742–8747
| last = Woese
| first = Carl R.
| journal = Proceedings of the National Academy of Sciences of the United States of America
| date = 2002-06-25
| pmid = 12077305
| pmc = 124369
| bibcode = 2002PNAS...99.8742W
| doi-access = free
}}</ref> First described by Woese and Fox in a 1977 paper and explored further with microbiologist [[Jane Gibson]] in a 1980 paper, these organisms, or ''[[Progenote|progenotes]]'', were imagined as protocells with very low complexity due to their error-prone translation apparatus ("noisy genetic transmission channel"), which produced high mutation rates that limited the specificity of cellular interaction and the size of the genome.<ref>{{Cite journal 
| last1 = Woese | first1 = C. R. 
| last2 = Fox | first2 = G. E. 
| title = The concept of cellular evolution 
| journal = Journal of Molecular Evolution 
| volume = 10 
| issue = 1 
| pages = 1–6 
| year = 1977 
| pmid = 903983
 | doi=10.1007/bf01796132
| bibcode = 1977JMolE..10....1W| s2cid = 24613906 
}}</ref><ref>{{Cite journal|last1=Woese|first1=Carl R.|last2=Gibson|first2=Jane|last3=Fox|first3=George E.|date=January 1980|title=Do genealogical patterns in purple photosynthetic bacteria reflect interspecific gene transfer?|url=https://www.nature.com/articles/283212a0|journal=Nature|language=en|volume=283|issue=5743|pages=212–214|doi=10.1038/283212a0|pmid=6243180|bibcode=1980Natur.283..212W|s2cid=4243875|issn=1476-4687}}</ref> This early translation apparatus would have produced a group of structurally similar, functionally equivalent proteins, rather than a single protein.<ref name="pace2011pnas"/> Furthermore, because of this reduced specificity, all cellular components were susceptible to horizontal gene transfer, and rapid evolution occurred at the level of the ecosystem.<ref name="woese2002cells"/><ref name="buchanan2010">{{Cite news|title=Evolution, but not as we know it|last=Buchanan|first=Mark|date=2010-01-23|work=New Scientist|issue=2744|volume=205|pages=34–37|issn=0262-4079}}</ref>

The transition to modern cells (the "[[Darwinian threshold|Darwinian Threshold]]") occurred when organisms evolved translation mechanisms with modern levels of fidelity: improved performance allowed cellular organization to reach a level of complexity and connectedness that made genes from other organisms much less able to displace an individual's own genes.<ref name="woese2002cells"/>

In later years, Woese's work concentrated on genomic analysis to elucidate the significance of horizontal gene transfer (HGT) for evolution.<ref name="woese2005hgt">{{Cite book
| title = Microbial Phylogeny and Evolution:Concepts and Controversies: Concepts and Controversies
| chapter = Evolving biological organization
| publisher = Oxford University Press
| pages = 99–117
|editor1= Jan Sapp
| last = Woese
| first = Carl R.
| author-link = Carl Woese
| access-date = 2013-01-04
| year = 2005
| isbn = 9780198037774
| chapter-url = https://books.google.com/books?id=SMvLpiK-fgsC&q=Evolving+Biological+Organization&pg=PA99
}}</ref> He worked on detailed analyses of the phylogenies of the aminoacyl-tRNA synthetases and on the effect of horizontal gene transfer on the distribution of those key enzymes among organisms.<ref name="woese2008">{{Cite journal 
| last1 = Woese | first1 = C. R. 
| author-link1 = Carl Woese
| last2 = Olsen | first2 = G. J. 
| last3 = Ibba | first3 = M. 
| last4 = Söll | first4 = D. 
| title = Aminoacyl-tRNA synthetases, the genetic code, and the evolutionary process 
| journal = Microbiology and Molecular Biology Reviews 
| volume = 64 
| issue = 1 
| pages = 202–236 
| year = 2000 
| pmid = 10704480 
| pmc = 98992
| doi = 10.1128/MMBR.64.1.202-236.2000
}}</ref> The goal of the research was to explain how the primary cell types (the archaeal, eubacterial, and eukaryotic) evolved from an ancestral state in the [[RNA world]].<ref name=mcb />

== Perspectives on biology ==
Woese shared his thoughts on the past, present, and future of biology in ''[[Current Biology]]'':<ref name="woese2005qa"/>

<blockquote>The "important questions" that 21st century biology faces all stem from a single question, the nature and generation of [[biological organization]]. . . . Yes, Darwin is back, but in the company of . . . scientists who can see much further into the depths of biology than was possible heretofore. It is no longer a "10,000 species of birds" view of evolution—evolution seen as a procession of forms. The concern is now with the process of evolution itself.<ref name="woese2005qa"/></blockquote>

<blockquote>I see the question of biological organization taking two prominent directions today. The first is the evolution of (proteinaceous) cellular organization, which includes sub-questions such as the evolution of the translation apparatus and the genetic code, and the origin and nature of the hierarchies of control that fine-tune and precisely interrelate the panoply of cellular processes that constitute cells. It also includes the question of the number of different basic cell types that exist on earth today: did all modern cells come from a single ancestral cellular organization?<ref name="woese2005qa"/></blockquote>

<blockquote>The second major direction involves the nature of the global ecosystem. . . . Bacteria are the major organisms on this planet—in numbers, in total mass, in importance to the global balances. Thus, it is [[microbial ecology]] that . . . is most in need of development, both in terms of facts needed to understand it, and in terms of the framework in which to interpret them.<ref name="woese2005qa"/></blockquote>

Woese considered biology to have an "all-important" role in society. In his view, biology should serve a broader purpose than the pursuit of "an engineered environment":<ref name="woese2005qa"/>

<blockquote>What was formally recognized in physics needs now to be recognized in biology: science serves a dual function. On the one hand it is society's servant, attacking the applied problems posed by society. On the other hand, it functions as society's teacher, helping the latter to understand its world and itself. It is the latter function that is effectively missing today.<ref name="woese2005qa"/></blockquote>

== Honors and scientific legacy ==
Woese was a [[MacArthur Fellowship|MacArthur Fellow]] in 1984, was made a member of the [[United States National Academy of Sciences|National Academy of Sciences]] in 1988, received the [[Leeuwenhoek Medal]] (microbiology's highest honor) in 1992, the [[Selman A. Waksman Award in Microbiology]] in 1995 from the [[United States National Academy of Sciences|National Academy of Sciences]],<ref name=Waksman>{{cite web|title=Selman A. Waksman Award in Microbiology|url=http://www.nasonline.org/site/PageServer?pagename=AWARDS_waksman|publisher=National Academy of Sciences|access-date=2011-02-27|url-status=dead|archive-url=https://web.archive.org/web/20110112174319/http://www.nasonline.org/site/PageServer?pagename=AWARDS_waksman|archive-date=January 12, 2011|df=mdy-all}}</ref> and was a [[National Medal of Science]] recipient in 2000. In 2003, he received the [[Crafoord Prize]] from the [[Royal Swedish Academy of Sciences]] "for his discovery of a third domain of life".<ref>{{cite web |url=http://astrobiology.nasa.gov/articles/carl-woese-and-new-perspectives-on-evolution/ |title=Carl Woese and New Perspectives on Evolution |last=Morrison |first=David |date=December 10, 2003 |work=Astrobiology: Life in the Universe |publisher=NASA |access-date=February 16, 2010 |url-status=dead |archive-url=https://web.archive.org/web/20100224074131/http://astrobiology.nasa.gov/articles/carl-woese-and-new-perspectives-on-evolution |archive-date=February 24, 2010 |df=mdy }}</ref><ref name="crafoord2003">{{cite web | last = Huss | first = Erik | title = The Crafoord Prize 2003 – Crafoordprize | work = The Crafoord Prize | format = Press Release | access-date = 2013-01-03 | date = 2003-02-12 | url = http://www.crafoordprize.se/press/arkivpressreleases/thecrafoordprize2003.5.32d4db7210df50fec2d800018585.html | archive-date = October 31, 2020 | archive-url = https://web.archive.org/web/20201031171837/https://www.crafoordprize.se/press/arkivpressreleases/thecrafoordprize2003.5.32d4db7210df50fec2d800018585.html | url-status = dead }}</ref>  He was elected to the [[American Philosophical Society]] in 2004.<ref>{{cite web|title=APS Member History|url=https://search.amphilsoc.org/memhist/search?creator=Carl+R.+Woese&title=&subject=&subdiv=&mem=&year=&year-max=&dead=&keyword=&smode=advanced|access-date=2021-06-09|website=search.amphilsoc.org}}</ref> In 2006, he was made a [[foreign member of the Royal Society]].<ref name="uofi2006"/>

Many microbial species, such as ''[[Pyrococcus woesei]]'',<ref name="Zillig1987">{{cite journal |doi=10.1016/S0723-2020(87)80057-7 |title=Pyrococcus woesei, sp. Nov., an ultra-thermophilic marine archaebacterium, representing a novel order, Thermococcales |year=1987 |last1=Zillig |first1=Wolfram |last2=Holz |first2=Ingelore |last3=Klenk |first3=Hans-Peter |last4=Trent |first4=Jonathan |last5=Wunderl |first5=Simon |last6=Janekovic |first6=Davorin |last7=Imsel |first7=Erwin |last8=Haas |first8=Birgit |journal=Systematic and Applied Microbiology |volume=9 |issue=1–2 |pages=62–70|bibcode=1987SyApM...9...62Z }}</ref> ''[[Methanobrevibacter woesei]]'',<ref name="Miller2002">{{cite journal |doi=10.1099/ijs.0.02022-0 |title=Description of ''Methanobrevibacter gottschalkii'' sp. nov., ''Methanobrevibacter thaueri'' sp. nov., ''Methanobrevibacter woesei'' sp. nov. And ''Methanobrevibacter wolinii'' sp. nov |year=2002 |last1=Miller |first1=T. L. |journal=International Journal of Systematic and Evolutionary Microbiology |volume=52 |issue=3 |pages=819–822|pmid=12054244 }}</ref> and ''Conexibacter woesei'',<ref name="conexibacter">{{cite journal |doi=10.1099/ijs.0.02400-0 |title=''Conexibacter woesei'' gen. nov., sp. nov., a novel representative of a deep evolutionary line of descent within the class Actinobacteria |year=2003 |last1=Monciardini |first1=P. |journal=International Journal of Systematic and Evolutionary Microbiology |volume=53 |issue=2 |pages=569–576 |pmid=12710628|doi-access=free }}</ref> are named in his honor.

Microbiologist Justin Sonnenburg of [[Stanford University]] said "The 1977 paper is one of the most influential in microbiology and arguably, all of biology. It ranks with the works of [[James D. Watson|Watson]] and [[Francis Crick|Crick]] and Darwin, providing an evolutionary framework for the incredible diversity of the microbial world".<ref name=pnas2012 />

With regard to Woese's work on horizontal gene transfer as a primary evolutionary process, Professor [[Norman R. Pace]] of the [[University of Colorado at Boulder]] said, "I think Woese has done more for biology [[Wiktionary:writ large|writ large]] than any biologist in history, including [[Charles Darwin|Darwin]]... There's a lot more to learn, and he's been interpreting the emerging story brilliantly".<ref>[[Mark Buchanan]], [https://www.newscientist.com/article/mg20527441.500-horizontal-and-vertical-the-evolution-of-evolution.html?full=true&print=true Horizontal and vertical: The evolution of evolution], ''[[New Scientist]]'', January 26, 2010</ref>

== Selected publications==
===Books===
* {{Cite book
| last = Woese
| first = Carl
| title = The Genetic Code: the Molecular Basis for Genetic Expression
| location = New York
| publisher = Harper & Row
| year = 1967
| oclc = 293697
}}

===Selected articles===
* {{Cite journal
| title = Phylogenetic structure of the prokaryotic domain: the primary kingdoms
| issn = 0027-8424
| volume = 74
| issue = 11
| pages = 5088–5090
| last = Woese
| first = Carl R.
|author2=George E. Fox
 | journal = Proceedings of the National Academy of Sciences of the United States of America
| year = 1977
| bibcode = 1977PNAS...74.5088W |doi = 10.1073/pnas.74.11.5088
| pmid = 270744
| pmc = 432104
| doi-access = free
}}
* {{Cite journal
| title = Bacterial evolution
| volume = 51
| issue = 2
| pages = 221–271
| last = Woese
| first = Carl R.
| journal = Microbiological Reviews
| date = 1987-06-01
| pmid = 2439888
| pmc = 373105
| doi = 10.1128/MMBR.51.2.221-271.1987
}}
* {{Cite journal
| title = Towards a natural system of organisms: proposal for the domains Archaea, Bacteria, and Eucarya
| issn = 0027-8424
| volume = 87
| issue = 12
| pages = 4576–4579
| last = Woese
| first = Carl R.
|author2=O Kandler | author3-link = Mark Wheelis
|author3=M L Wheelis
| journal = Proceedings of the National Academy of Sciences of the United States of America
| year = 1990
| doi = 10.1073/pnas.87.12.4576
| pmid = 2112744
| pmc = 54159
| bibcode = 1990PNAS...87.4576W
| author2-link = Otto Kandler
| doi-access = free
}}
* {{Cite journal
| title = A New Biology for a New Century
| doi = 10.1128/MMBR.68.2.173-186.2004
| issn = 1098-5557
| volume = 68
| issue = 2
| pages = 173–186
| last = Woese
| first = Carl R.
| journal = Microbiology and Molecular Biology Reviews
| date = 2004-06-01
| pmid=15187180
| pmc=419918
}}
* {{Cite book
| title = Microbial Phylogeny and Evolution:Concepts and Controversies: Concepts and Controversies
| chapter = Evolving biological organization
| publisher = Oxford University Press
| pages = 99–117
|editor1= Jan Sapp
| last = Woese
| first = Carl R.
| access-date = 2013-01-04
| year = 2005
| isbn = 9780198037774
| chapter-url = https://books.google.com/books?id=SMvLpiK-fgsC&q=Evolving+Biological+Organization&pg=PA99
}}
* {{Cite book
| title = The Prokaryotes
| chapter = How We Do, Don't and Should Look at Bacteria and Bacteriology
| pages = 3–23
| last = Woese
| first = Carl R.
| year = 2006
| doi=10.1007/0-387-30741-9_1
|isbn=978-0-387-30741-1
}}
* {{Cite journal
| title = How the Microbial World Saved Evolution from the Scylla of Molecular Biology and the Charybdis of the Modern Synthesis
| doi = 10.1128/MMBR.00002-09
| volume = 73
| issue = 1
| pages = 14–21
| last = Woese
| first = Carl R.
|author2=Nigel Goldenfeld
 | journal = Microbiology and Molecular Biology Reviews
| year = 2009
| pmid = 19258530
| pmc = 2650883
| author2-link = Nigel Goldenfeld
}}

==See also==
* [[Archaea]]
* [[Bacterial phyla]], the major lineages of Bacteria
* [[George E. Fox]]
* [[Karl Stetter]]
* [[Norman R. Pace]]
* [[Otto Kandler]]
* [[Phylogenetics]]
* [[Tree of life (biology)]]
* [[16S ribosomal RNA]]
* [[Woeseian revolution]]
* [[Woese's dogma]]

==References==
{{reflist}}

==External links==
*{{PubMedAuthorSearch|Woese|CR}}
* [http://archives.library.illinois.edu/archon/?p=collections/controlcard&id=11138 Carl Woese papers] at the University of Illinois, Champaign
* [http://www.igb.illinois.edu/ The Carl R. Woese Institute for Genomic Biology, University of Illinois]
* [http://www.igb.illinois.edu/people/archaea Woese's Homepage, Carl R. Woese Institute for Genomic Biology], November 30, 2017
* [http://www.igb.illinois.edu/woese-guest-book Carl R. Woese Guestbook, Carl R. Woese Institute for Genomic Biology], November 30, 2017
* [http://www.microbeworld.org/index.php?Itemid=195&catid=54%3Aprogram-one-qthe-tree-of-lifeq&id=261%3Asolving-the-puzzle-part-2-of-10&option=com_content&view=article Excerpts from a documentary on Woese's Tree of Life]
*{{cite journal |doi=10.1016/j.cub.2005.02.003 |title=Q & A |year=2005 |last1=Woese |first1=Carl R. |journal=Current Biology |volume=15 |issue=4 |pages=R111–2 |pmid=15723774|s2cid=45434594 |doi-access=free |bibcode=2005CBio...15.R111W }}
{{Winners of the National Medal of Science|biological}}
{{FRS 2006}}
{{Authority control}}

{{DEFAULTSORT:Woese, Carl}}
[[Category:Carl Woese| ]]
[[Category:1928 births]]
[[Category:2012 deaths]]
[[Category:American microbiologists]]
[[Category:Amherst College alumni]]
[[Category:Deerfield Academy alumni]]
[[Category:Foreign members of the Royal Society]]
[[Category:MacArthur Fellows]]
[[Category:Members of the United States National Academy of Sciences]]
[[Category:National Medal of Science laureates]]
[[Category:Scientists from Syracuse, New York]]
[[Category:University of Illinois Urbana-Champaign faculty]]
[[Category:Leeuwenhoek Medal winners]]
[[Category:Deaths from pancreatic cancer]]
[[Category:Members of the American Philosophical Society]]
[[Category:Deaths from cancer in Illinois]]
[[Category:History of genetics]]