Lynn Margulis
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Lynn Margulis (born Lynn Petra Alexander; March 5, 1938 – November 22, 2011) was an American
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
ary biologist, and was the primary modern proponent for the significance of symbiosis in evolution. Historian
Jan Sapp Jan Anthony Sapp (born June 12, 1954) is a professor in the Department of Biology, York University, Canada. His writings focus especially on evolutionary biology beyond the classical neo-Darwinian framework, and emphasize the fundamental importanc ...
has said that "Lynn Margulis's name is as synonymous with symbiosis as
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended fr ...
's is with evolution." In particular, Margulis transformed and fundamentally framed current understanding of the
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
of cells with nuclei – an event
Ernst Mayr Ernst Walter Mayr (; 5 July 1904 – 3 February 2005) was one of the 20th century's leading evolutionary biologists. He was also a renowned Taxonomy (biology), taxonomist, tropical explorer, ornithologist, Philosophy of biology, philosopher o ...
called "perhaps the most important and dramatic event in the history of life" – by proposing it to have been the result of symbiotic mergers of bacteria. Margulis was also the co-developer of the
Gaia hypothesis The Gaia hypothesis (), also known as the Gaia theory, Gaia paradigm, or the Gaia principle, proposes that living organisms interact with their inorganic surroundings on Earth to form a synergistic and self-regulating, complex system that help ...
with the British chemist
James Lovelock James Ephraim Lovelock (26 July 1919 – 26 July 2022) was an English independent scientist, environmentalist and futurist. He is best known for proposing the Gaia hypothesis, which postulates that the Earth functions as a self-regulating sys ...
, proposing that the Earth functions as a single self-regulating system, and was the principal defender and promulgator of the five kingdom classification of Robert Whittaker. Throughout her career, Margulis' work could arouse intense objection (one grant application elicited the response, "Your research is crap. Don't ever bother to apply again.") and her formative paper, "On the Origin of Mitosing Cells", appeared in 1967 after being rejected by about fifteen journals. Still a junior faculty member at Boston University at the time, her theory that cell organelles such as
mitochondria A mitochondrion (; ) is an organelle found in the Cell (biology), cells of most Eukaryotes, such as animals, plants and Fungus, fungi. Mitochondria have a double lipid bilayer, membrane structure and use aerobic respiration to generate adenosi ...
and
chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s were once independent
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
was largely ignored for another decade, becoming widely accepted only after it was powerfully substantiated through genetic evidence. Margulis was elected a member of the US
National Academy of Sciences The National Academy of Sciences (NAS) is a United States nonprofit, non-governmental organization. NAS is part of the National Academies of Sciences, Engineering, and Medicine, along with the National Academy of Engineering (NAE) and the Nati ...
in 1983. President
Bill Clinton William Jefferson Clinton ( né Blythe III; born August 19, 1946) is an American politician who served as the 42nd president of the United States from 1993 to 2001. He previously served as governor of Arkansas from 1979 to 1981 and agai ...
presented her the
National Medal of Science The National Medal of Science is an honor bestowed by the President of the United States to individuals in science and engineering who have made important contributions to the advancement of knowledge in the fields of behavioral and social scienc ...
in 1999. The
Linnean Society of London The Linnean Society of London is a learned society dedicated to the study and dissemination of information concerning natural history, evolution, and taxonomy. It possesses several important biological specimen, manuscript and literature colle ...
awarded her the Darwin-Wallace Medal in 2008. Called "science's unruly earth mother", a "vindicated heretic", or a scientific "rebel", Margulis was a strong critic of
neo-Darwinism Neo-Darwinism is generally used to describe any integration of Charles Darwin's theory of evolution by natural selection with Gregor Mendel's theory of genetics. It mostly refers to evolutionary theory from either 1895 (for the combinations of Dar ...
. Her position sparked lifelong debate with leading neo-Darwinian biologists, including
Richard Dawkins Richard Dawkins (born 26 March 1941) is a British evolutionary biologist and author. He is an emeritus fellow of New College, Oxford and was Professor for Public Understanding of Science in the University of Oxford from 1995 to 2008. An ath ...
, George C. Williams, and
John Maynard Smith John Maynard Smith (6 January 1920 – 19 April 2004) was a British theoretical and mathematical evolutionary biologist and geneticist. Originally an aeronautical engineer during the Second World War, he took a second degree in genetics und ...
. Margulis' work on symbiosis and her
endosymbiotic An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within ...
theory had important predecessors, going back to the mid-19th century – notably
Andreas Franz Wilhelm Schimper Andreas Franz Wilhelm Schimper (12 May 1856 – 9 September 1901) was a German botanist and phytogeographer who made major contributions in the fields of histology, ecology and plant geography. He travelled to South East Asia and the Caribbea ...
,
Konstantin Mereschkowski Konstantin Sergeevich Mereschkowski ( rus, Константи́н Серге́евич Мережко́вский, p=mʲɪrʲɪˈʂkofskʲɪj; – 9 January 1921) was a prominent Russian biologist and botanist, active mainly around Kazan, Tat ...
,
Boris Kozo-Polyansky Boris Mikhailovich Kozo-Polyansky (; 20 January 1890 – 21 April 1957) was a Soviet and Russian botanist and evolutionary biologist, best known for his seminal work, ''Symbiogenesis: A New Principle of Evolution'', which was the first work to pl ...
, and
Ivan Wallin Ivan Emanuel Wallin (22 January 1883 – 6 March 1969) was an American biologist who made the first experimental works on endosymbiotic theory. Nicknamed the "Mitochondria Man", he claimed that mitochondria, which are cell organelles, were once i ...
– and Margulis, not only promoted greater recognition for their contributions, but personally oversaw the first English translation of Kozo-Polyansky's ''Symbiogenesis: A New Principle of Evolution'', which appeared the year before her death. Many of her major works, particularly those intended for a general readership, were collaboratively written with her son Dorion Sagan. In 2002, ''
Discover Discover may refer to: Art, entertainment, and media * ''Discover'' (album), a Cactus Jack album * ''Discover'' (magazine), an American science magazine Businesses and brands * DISCover, the ''Digital Interactive Systems Corporation'' * D ...
'' magazine recognized Margulis as one of the 50 most important women in science.


Biography

Lynn Margulis was born in
Chicago (''City in a Garden''); I Will , image_map = , map_caption = Interactive Map of Chicago , coordinates = , coordinates_footnotes = , subdivision_type = Country , subdivision_name ...
, to a
Jewish Jews ( he, יְהוּדִים, , ) or Jewish people are an ethnoreligious group and nation originating from the Israelites Israelite origins and kingdom: "The first act in the long drama of Jewish history is the age of the Israelites""The ...
,
Zionist Zionism ( he, צִיּוֹנוּת ''Tsiyyonut'' after ''Zion'') is a nationalist movement that espouses the establishment of, and support for a homeland for the Jewish people centered in the area roughly corresponding to what is known in Je ...
family. Her parents were Morris Alexander and Leona Wise Alexander. She was the eldest of four daughters. Her father was an attorney who also ran a company that made road paints. Her mother operated a travel agency. She entered the
Hyde Park Academy High School Hyde Park Academy High School (formerly known as Hyde Park High School and Hyde Park Career Academy) is a public 4–year high school located in the Woodlawn neighborhood on the south side of Chicago, Illinois, United States. Opened in 1863, Hyd ...
in 1952, describing herself as a bad student who frequently had to stand in the corner. A precocious child, she was accepted at the
University of Chicago Laboratory Schools The University of Chicago Laboratory Schools (also known as Lab or Lab Schools and abbreviated as UCLS though the high school is nicknamed U-High) is a Private school, private, co-educational Day school, day Early childhood education, Pre-K and K ...
at the age of fifteen. In 1957, at age 19, she earned a BA from the University of Chicago in
Liberal Arts Liberal arts education (from Latin "free" and "art or principled practice") is the traditional academic course in Western higher education. ''Liberal arts'' takes the term ''art'' in the sense of a learned skill rather than specifically the ...
. She joined the
University of Wisconsin A university () is an institution of higher (or tertiary) education and research which awards academic degrees in several academic disciplines. Universities typically offer both undergraduate and postgraduate programs. In the United States, t ...
to study biology under
Hans Ris Hans Ris (June 15, 1914 – November 19, 2004) was an American cytologist and pioneer electron microscopy, electron microscopist. His studies of chromosome structure revealed the importance of non-histone proteins, and along with evolutionary biol ...
and Walter Plaut, her supervisor, and graduated in 1960 with an MS in genetics and zoology. (Her first publication, published with Plaut in 1958 in the ''Journal of Protozoology'', was on the genetics of ''
Euglena ''Euglena'' is a genus of Unicellular organism, single cell flagellate eukaryotes. It is the best known and most widely studied member of the class Euglenoidea, a diverse group containing some 54 genera and at least 200 species. Species of ''Eug ...
'',
flagellate A flagellate is a cell or organism with one or more whip-like appendages called flagella. The word ''flagellate'' also describes a particular construction (or level of organization) characteristic of many prokaryotes and eukaryotes and their ...
s which have features of both animals and plants.) She then pursued research at the
University of California, Berkeley The University of California, Berkeley (UC Berkeley, Berkeley, Cal, or California) is a public land-grant research university in Berkeley, California. Established in 1868 as the University of California, it is the state's first land-grant u ...
, under the zoologist Max Alfert. Before she could complete her dissertation, she was offered research associateship and then lectureship at
Brandeis University , mottoeng = "Truth even unto its innermost parts" , established = , type = Private research university , accreditation = NECHE , president = Ronald D. Liebowitz , pro ...
in Massachusetts in 1964. It was while working there that she obtained her PhD from the University of California, Berkeley in 1965. Her thesis was ''An Unusual Pattern of Thymidine Incorporation in ''Euglena''.'' In 1966 she moved to
Boston University Boston University (BU) is a private research university in Boston, Massachusetts. The university is nonsectarian, but has a historical affiliation with the United Methodist Church. It was founded in 1839 by Methodists with its original campu ...
, where she taught biology for twenty-two years. She was initially an Adjunct Assistant Professor, then was appointed to Assistant Professor in 1967. She was promoted to Associate Professor in 1971, to full Professor in 1977, and to University Professor in 1986. In 1988 she was appointed Distinguished Professor of Botany at the
University of Massachusetts at Amherst The University of Massachusetts Amherst (UMass Amherst, UMass) is a public research university in Amherst, Massachusetts and the sole public land-grant university in Commonwealth of Massachusetts. Founded in 1863 as an agricultural college, it ...
. She was Distinguished Professor of Biology in 1993. In 1997 she transferred to the Department of Geosciences at Amherst to become Distinguished Professor of Geosciences "with great delight", the post which she held until her death.


Personal life

Margulis married astronomer
Carl Sagan Carl Edward Sagan (; ; November 9, 1934December 20, 1996) was an American astronomer, planetary scientist, cosmologist, astrophysicist, astrobiologist, author, and science communicator. His best known scientific contribution is research on ext ...
in 1957 soon after she got her bachelor's degree. Sagan was then a graduate student in physics at the University of Chicago. Their marriage ended in 1964, just before she completed her PhD. They had two sons, Dorion Sagan, who later became a popular science writer and her collaborator, and Jeremy Sagan, software developer and founder of Sagan Technology. In 1967, she married Thomas N. Margulis, a crystallographer. They had a son named Zachary Margulis-Ohnuma, a New York City criminal defense lawyer, and a daughter Jennifer Margulis, teacher and author. They divorced in 1980. She commented, "I quit my job as a wife twice," and, "it's not humanly possible to be a good wife, a good mother, and a first-class scientist. No one can do it — something has to go." In the 2000s she had a relationship with fellow biologist Ricardo Guerrero. Her sister Joan Alexander married Nobel Laureate
Sheldon Glashow Sheldon Lee Glashow (, ; born December 5, 1932) is a Nobel Prize-winning American theoretical physicist. He is the Metcalf Professor of Mathematics and Physics at Boston University and Eugene Higgins Professor of Physics, Emeritus, at Harvard U ...
; another sister, Sharon, married mathematician
Daniel Kleitman Daniel J. Kleitman (born October 4, 1934)article availableon Douglas West's web page, University of Illinois at Urbana–Champaign)."Kleitman, Daniel J.," in: ''Who's Who in Frontier Science and Technology'', 1, 1984, p. 396. is an American mathe ...
. She was a religious
agnostic Agnosticism is the view or belief that the existence of God, of the divine or the supernatural is unknown or unknowable. (page 56 in 1967 edition) Another definition provided is the view that "human reason is incapable of providing sufficient ...
, and a staunch
evolutionist Evolutionism is a term used (often derogatorily) to denote the theory of evolution. Its exact meaning has changed over time as the study of evolution has progressed. In the 19th century, it was used to describe the belief that organisms deliberate ...
, but rejected the
modern evolutionary synthesis Modern synthesis or modern evolutionary synthesis refers to several perspectives on evolutionary biology, namely: * Modern synthesis (20th century), the term coined by Julian Huxley in 1942 to denote the synthesis between Mendelian genetics and s ...
, and said: "I remember waking up one day with an epiphanous revelation: I am not a neo-Darwinist! I recalled an earlier experience, when I realized that I wasn't a humanistic Jew. Although I greatly admire Darwin's contributions and agree with most of his theoretical analysis and I am a Darwinist, I am not a neo-Darwinist." She argued that "Natural selection eliminates and maybe maintains, but it doesn't create", and maintained that symbiosis was the major driver of evolutionary change. In 2013, Margulis was listed as having been a member of the Advisory Council of the
National Center for Science Education The National Center for Science Education (NCSE) is a not-for-profit membership organization in the United States whose stated mission is to educate the press and the public on the scientific and educational aspects of controversies surrounding t ...
. Margulis died on 22 November 2011 at home in Amherst,
Massachusetts Massachusetts (Massachusett language, Massachusett: ''Muhsachuweesut assachusett writing systems, məhswatʃəwiːsət'' English: , ), officially the Commonwealth of Massachusetts, is the most populous U.S. state, state in the New England ...
, five days after suffering a
hemorrhagic stroke A stroke is a disease, medical condition in which poor cerebral circulation, blood flow to the brain causes cell death. There are two main types of stroke: brain ischemia, ischemic, due to lack of blood flow, and intracranial hemorrhage, hemorr ...
. As her wish, she was cremated and her ashes were scattered in her favorite research areas, near her home.


Contributions


Endosymbiosis theory

In 1966, as a young faculty member at
Boston University Boston University (BU) is a private research university in Boston, Massachusetts. The university is nonsectarian, but has a historical affiliation with the United Methodist Church. It was founded in 1839 by Methodists with its original campu ...
, Margulis wrote a theoretical paper titled "On the Origin of Mitosing Cells". The paper, however, was "rejected by about fifteen scientific journals," she recalled.Margulis, Lynn
Gaia Is a Tough Bitch
. Chapter 7 in The Third Culture: Beyond the Scientific Revolution by John Brockman (Simon & Schuster, 1995)
It was finally accepted by ''
Journal of Theoretical Biology The ''Journal of Theoretical Biology'' is a biweekly peer-reviewed scientific journal covering theoretical biology, as well as mathematical, computational, and statistical aspects of biology. Some research areas covered by the journal include cell ...
'' and is considered today a landmark in modern
endosymbiotic theory Symbiogenesis (endosymbiotic theory, or serial endosymbiotic theory,) is the leading evolutionary theory of the origin of eukaryotic cells from prokaryotic organisms. The theory holds that mitochondria, plastids such as chloroplasts, and possibl ...
. Weathering constant criticism of her ideas for decades, Margulis was famous for her tenacity in pushing her theory forward, despite the opposition she faced at the time. The descent of mitochondria from bacteria and of chloroplasts from cyanobacteria was experimentally demonstrated in 1978 by Robert Schwartz and
Margaret Dayhoff Margaret Belle (Oakley) Dayhoff (March 11, 1925 – February 5, 1983) was an American physical chemist and a pioneer in the field of bioinformatics. Dayhoff was a professor at Georgetown University Medical Center and a noted research biochem ...
. This formed the first experimental evidence for the symbiogenesis theory. The endosymbiosis theory of organogenesis became widely accepted in the early 1980s, after the genetic material of
mitochondria A mitochondrion (; ) is an organelle found in the Cell (biology), cells of most Eukaryotes, such as animals, plants and Fungus, fungi. Mitochondria have a double lipid bilayer, membrane structure and use aerobic respiration to generate adenosi ...
and
chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s had been found to be significantly different from that of the symbiont's
nuclear DNA Nuclear DNA (nDNA), or nuclear deoxyribonucleic acid, is the DNA contained within each cell nucleus of a eukaryotic organism. It encodes for the majority of the genome in eukaryotes, with mitochondrial DNA and plastid DNA coding for the rest. It ...
. In 1995, English evolutionary biologist
Richard Dawkins Richard Dawkins (born 26 March 1941) is a British evolutionary biologist and author. He is an emeritus fellow of New College, Oxford and was Professor for Public Understanding of Science in the University of Oxford from 1995 to 2008. An ath ...
had this to say about Lynn Margulis and her work:
I greatly admire Lynn Margulis's sheer courage and stamina in sticking by the endosymbiosis theory, and carrying it through from being an unorthodoxy to an orthodoxy. I'm referring to the theory that the eukaryotic cell is a symbiotic union of primitive prokaryotic cells. This is one of the great achievements of twentieth-century evolutionary biology, and I greatly admire her for it.


Symbiosis as evolutionary force

Margulis opposed competition-oriented views of evolution, stressing the importance of symbiotic or cooperative relationships between species. She later formulated a theory that proposed symbiotic relationships between organisms of different phyla, or kingdoms, as the driving force of
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
, and explained
genetic variation Genetic variation is the difference in DNA among individuals or the differences between populations. The multiple sources of genetic variation include mutation and genetic recombination. Mutations are the ultimate sources of genetic variation, ...
as occurring mainly through transfer of nuclear information between bacterial cells or
virus A virus is a submicroscopic infectious agent that replicates only inside the living cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea. Since Dmitri Ivanovsky's 1 ...
es and
eukaryotic cell Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacter ...
s. Her organelle genesis ideas are now widely accepted, but the proposal that symbiotic relationships explain most genetic variation is still something of a fringe idea. Margulis also held a negative view of certain interpretations of
Neo-Darwinism Neo-Darwinism is generally used to describe any integration of Charles Darwin's theory of evolution by natural selection with Gregor Mendel's theory of genetics. It mostly refers to evolutionary theory from either 1895 (for the combinations of Dar ...
that she felt were excessively focused on competition between organisms, as she believed that history will ultimately judge them as comprising "a minor twentieth-century religious sect within the sprawling religious persuasion of Anglo-Saxon Biology." She wrote that proponents of the standard theory "wallow in their zoological, capitalistic, competitive, cost-benefit interpretation of Darwin – having mistaken him ... Neo-Darwinism, which insists on he slow accrual of mutations by gene-level natural selection is in a complete funk."


Gaia hypothesis

Margulis initially sought out the advice of
James Lovelock James Ephraim Lovelock (26 July 1919 – 26 July 2022) was an English independent scientist, environmentalist and futurist. He is best known for proposing the Gaia hypothesis, which postulates that the Earth functions as a self-regulating sys ...
for her own research: she explained that, "In the early seventies, I was trying to align bacteria by their metabolic pathways. I noticed that all kinds of bacteria produced gases. Oxygen, hydrogen sulfide, carbon dioxide, nitrogen, ammonia—more than thirty different gases are given off by the bacteria whose evolutionary history I was keen to reconstruct. Why did every scientist I asked believe that atmospheric oxygen was a biological product but the other atmospheric gases—nitrogen, methane, sulfur, and so on—were not? 'Go talk to Lovelock,' at least four different scientists suggested. Lovelock believed that the gases in the atmosphere were biological." Margulis met with Lovelock, who explained his Gaia hypothesis to her, and very soon they began an intense collaborative effort on the concept. One of the earliest significant publications on Gaia was a 1974 paper co-authored by Lovelock and Margulis, which succinctly defined the hypothesis as follows: "The notion of the biosphere as an active adaptive control system able to maintain the Earth in homeostasis we are calling the 'Gaia hypothesis.'" Like other early presentations of Lovelock's idea, the Lovelock-Margulis 1974 paper seemed to give living organisms complete agency in creating planetary self-regulation, whereas later, as the idea matured, this planetary-scale self-regulation was recognized as an emergent property of the
Earth system Earth system science (ESS) is the application of systems science to the Earth. In particular, it considers interactions and 'feedbacks', through material and energy fluxes, between the Earth's sub-systems' cycles, processes and "spheres"— atmo ...
, life and its physical environment taken together. When climatologist Stephen Schneider convened the 1989 American Geophysical Union Chapman Conference around the issue of Gaia, the idea of "strong Gaia" and "weak Gaia" was introduced by James Kirchner, after which Margulis was sometimes associated with the idea of "weak Gaia", incorrectly (her essay "''Gaia is a Tough Bitch''" dates from 1995 – and it stated her own distinction from Lovelock as she saw it, which was primarily that she did not like the metaphor of Earth as a single organism, because, she said, "No organism eats its own waste"). In her 1998 book ''Symbiotic Planet'', Margulis explored the relationship between Gaia and her work on symbiosis.


Five kingdoms of life

In 1969, life on earth was classified into
five kingdoms In biology, a kingdom is the second highest taxonomic rank, just below domain. Kingdoms are divided into smaller groups called phyla. Traditionally, some textbooks from the United States and Canada used a system of six kingdoms (Animalia, Pla ...
, as introduced by Robert Whittaker. Margulis became the most important supporter, as well as critic – while supporting parts, she was the first to recognize the limitations of Whittaker's classification of microbes. But later discoveries of new organisms, such as
archaea Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaebac ...
, and emergence of molecular taxonomy challenged the concept. By the mid-2000s, most scientists began to agree that there are more than five kingdoms. Margulis became the most important defender of the five kingdom classification. She rejected the
three-domain system The three-domain system is a biological classification introduced by Carl Woese, Otto Kandler, and Mark Wheelis in 1990 that divides cellular life forms into three domains, namely Archaea, Bacteria, and Eukaryota or Eukarya. The key difference fr ...
introduced by
Carl Woese Carl Richard Woese (; July 15, 1928 – December 30, 2012) was an American microbiologist and biophysicist. Woese is famous for defining the Archaea (a new domain of life) in 1977 through a pioneering phylogenetic taxonomy of 16S ribosomal RNA, ...
in 1990, which gained wide acceptance. She introduced a modified classification by which all life forms, including the newly discovered, could be integrated into the classical five kingdoms. According to Margulis, the main problem, archaea, falls under the kingdom Prokaryotae alongside bacteria (in contrast to the three-domain system, which treats archaea as a higher taxon than kingdom, or the six-kingdom system, which holds that it is a separate kingdom). Margulis' concept is given in detail in her book ''Five Kingdoms'', written with Karlene V. Schwartz. It has been suggested that it is mainly because of Margulis that the five-kingdom system survives.


Controversies

It has been suggested that initial rejection of Margulis' work on the endosymbiotic theory, and the controversial nature of it as well as Gaia theory, made her identify throughout her career with scientific mavericks, outsiders, and unaccepted theories generally. In the last decade of her life, while key components of her life's work began to be understood as fundamental to a modern scientific viewpoint – the widespread adoption of Earth System Science and the incorporation of key parts of endosymbiotic theory into biology curricula worldwide – Margulis if anything became more embroiled in controversy, not less. Journalist John Wilson explained this by saying that Lynn Margulis "defined herself by oppositional science," and in the commemorative collection of essays ''Lynn Margulis: The Life and Legacy of a Scientific Rebel'', commentators again and again depict her as a modern embodiment of the "scientific rebel", akin to
Freeman Dyson Freeman John Dyson (15 December 1923 – 28 February 2020) was an English-American theoretical physicist and mathematician known for his works in quantum field theory, astrophysics, random matrices, mathematical formulation of quantum m ...
's 1995 essay, ''The Scientist as Rebel'', a tradition Dyson saw embodied in
Benjamin Franklin Benjamin Franklin ( April 17, 1790) was an American polymath who was active as a writer, scientist, inventor, statesman, diplomat, printer, publisher, and political philosopher. Encyclopædia Britannica, Wood, 2021 Among the leading inte ...
, and which Dyson believed to be essential to good science. At times, Margulis could make highly provocative comments in interviews that appeared to support her most strident critics' condemnation. The following describes three of those controversies.


Metamorphosis theory

In 2009, via a then-standard publication-process known as "communicated submission" (which bypassed traditional
peer review Peer review is the evaluation of work by one or more people with similar competencies as the producers of the work (peers). It functions as a form of self-regulation by qualified members of a profession within the relevant field. Peer review ...
), she was instrumental in getting the ''
Proceedings of the National Academy of Sciences ''Proceedings of the National Academy of Sciences of the United States of America'' (often abbreviated ''PNAS'' or ''PNAS USA'') is a peer-reviewed multidisciplinary scientific journal. It is the official journal of the National Academy of Sci ...
'' (''PNAS'') to publish a paper by Donald I. Williamson rejecting "the Darwinian assumption that larvae and their adults evolved from a single common ancestor."Controversial caterpillar-evolution study formally rebutted
''
Scientific American ''Scientific American'', informally abbreviated ''SciAm'' or sometimes ''SA'', is an American popular science magazine. Many famous scientists, including Albert Einstein and Nikola Tesla, have contributed articles to it. In print since 1845, it i ...
Online''
Williamson's paper provoked immediate response from the
scientific community The scientific community is a diverse network of interacting scientists. It includes many " sub-communities" working on particular scientific fields, and within particular institutions; interdisciplinary and cross-institutional activities are als ...
, including a countering paper in ''PNAS''. Conrad Labandeira of the
Smithsonian National Museum of Natural History The National Museum of Natural History is a natural history museum administered by the Smithsonian Institution, located on the National Mall in Washington, D.C., United States. It has free admission and is open 364 days a year. In 2021, with 7 ...
said, "If I was reviewing illiamson's paperI would probably opt to reject it," he says, "but I'm not saying it's a bad thing that this is published. What it may do is broaden the discussion on how metamorphosis works and .. nthe origin of these very radical life cycles." But
Duke University Duke University is a private research university in Durham, North Carolina. Founded by Methodists and Quakers in the present-day city of Trinity in 1838, the school moved to Durham in 1892. In 1924, tobacco and electric power industrialist James ...
insect developmental biologist Fred Nijhout said that the paper was better suited for the "''National Enquirer'' than the National Academy." In September it was announced that ''PNAS'' would eliminate communicated submissions in July 2010. ''PNAS'' stated that the decision had nothing to do with the Williamson controversy.


AIDS/HIV theory

In 2009 Margulis and seven others authored a position paper concerning research on the viability of round body forms of some spirochetes, "Syphilis, Lyme disease, & AIDS: Resurgence of 'the great imitator'?" which states that, "Detailed research that correlates life histories of symbiotic
spirochete A spirochaete () or spirochete is a member of the phylum Spirochaetota (), (synonym Spirochaetes) which contains distinctive diderm (double-membrane) gram-negative bacteria, most of which have long, helically coiled (corkscrew-shaped or s ...
s to changes in the immune system of associated vertebrates is sorely needed", and urging the "reinvestigation of the natural history of mammalian,
tick Ticks (order Ixodida) are parasitic arachnids that are part of the mite superorder Parasitiformes. Adult ticks are approximately 3 to 5 mm in length depending on age, sex, species, and "fullness". Ticks are external parasites, living by ...
-borne, and venereal transmission of spirochetes in relation to impairment of the human immune system". The paper went on to suggest "that the possible direct causal involvement of spirochetes and their round bodies to symptoms of immune deficiency be carefully and vigorously investigated". In a ''
Discover Magazine ''Discover'' is an American general audience science magazine launched in October 1980 by Time Inc. It has been owned by Kalmbach Publishing since 2010. History Founding ''Discover'' was created primarily through the efforts of ''Time'' mag ...
'' interview, which was published less than six months before her death, Margulis explained to writer Dick Teresi her reason for interest in the topic of 2009 "AIDS" paper: "I'm interested in spirochetes only because of our ancestry. I'm not interested in the diseases", and stated that she had called them "symbionts" because both the spirochete which causes syphilis (''
Treponema ''Treponema'' is a genus of spiral-shaped bacteria. The major treponeme species of human pathogens is ''Treponema pallidum'', whose subspecies are responsible for diseases such as syphilis, bejel, and yaws. ''Treponema carateum'' is the cause o ...
'') and the spirochete which causes Lyme disease (''
Borrelia ''Borrelia'' is a genus of bacteria of the spirochete phylum. Several species cause Lyme disease, also called Lyme borreliosis, a zoonotic, vector-borne disease transmitted by ticks. Other species of ''Borrelia'' cause relapsing fever, and are tr ...
'') only retain about 20% of the genes they would need to live freely, outside of their human hosts. However, in the ''Discover Magazine'' interview Margulis said that "the set of symptoms, or syndrome, presented by syphilitics overlaps completely with another syndrome: AIDS", and also noted that
Kary Mullis Kary Banks Mullis (December 28, 1944August 7, 2019) was an American biochemist. In recognition of his role in the invention of the polymerase chain reaction (PCR) technique, he shared the 1993 Nobel Prize in Chemistry with Michael Smith and wa ...
said that "he went looking for a reference substantiating that HIV causes AIDS and discovered, 'There is no such document' ". This provoked a widespread supposition that Margulis had been an "
AIDS denialist HIV/AIDS denialism is the belief, despite conclusive evidence to the contrary, that the human immunodeficiency virus (HIV) does not cause acquired immune deficiency syndrome (AIDS). Some of its proponents reject the existence of HIV, while oth ...
". Notably Jerry Coyne reacted on his ''Why Evolution is True'' blog against his interpretation that Margulis believed "that AIDS is really syphilis, not viral in origin at all."
Seth Kalichman Seth C. Kalichman is an American clinical community psychologist and professor of social psychology at the University of Connecticut, known for his research into HIV/AIDS treatment and HIV/AIDS denialism. Kalichman is also the director of the So ...
, a social psychologist who studies behavioral and social aspects of AIDS, cited her urgulis2009 paper as an example of AIDS denialism "flourishing", and asserted that her argulis"endorsement of HIV/AIDS denialism defies understanding".


9/11 "Truth"

Margulis argued that the
September 11 attacks The September 11 attacks, commonly known as 9/11, were four coordinated suicide terrorist attacks carried out by al-Qaeda against the United States on Tuesday, September 11, 2001. That morning, nineteen terrorists hijacked four commercia ...
were a "
false-flag operation A false flag operation is an act committed with the intent of disguising the actual source of responsibility and pinning blame on another party. The term "false flag" originated in the 16th century as an expression meaning an intentional misr ...
, which has been used to justify the wars in Afghanistan and Iraq as well as unprecedented assaults on ..civil liberties." She claimed that there was "overwhelming evidence that the three buildings f the World Trade Centercollapsed by controlled demolition."


Awards and recognitions

* Elected Fellow of the
American Association for the Advancement of Science The American Association for the Advancement of Science (AAAS) is an American international non-profit organization with the stated goals of promoting cooperation among scientists, defending scientific freedom, encouraging scientific respons ...
in 1975. *
Guggenheim Fellowship Guggenheim Fellowships are grants that have been awarded annually since by the John Simon Guggenheim Memorial Foundation to those "who have demonstrated exceptional capacity for productive scholarship or exceptional creative ability in the ar ...
in 1978. * Elected to the
National Academy of Sciences The National Academy of Sciences (NAS) is a United States nonprofit, non-governmental organization. NAS is part of the National Academies of Sciences, Engineering, and Medicine, along with the National Academy of Engineering (NAE) and the Nati ...
in 1983. * Guest Hagey Lecturer, University of Waterloo, 1985 * Miescher-Ishida Prize in 1986. * 1989, conferred the Commandeur de l’Ordre des Palmes Académiques de France. * Has her papers permanently archived in the
Library of Congress The Library of Congress (LOC) is the research library that officially serves the United States Congress and is the ''de facto'' national library of the United States. It is the oldest federal cultural institution in the country. The library is ...
,
Washington, DC ) , image_skyline = , image_caption = Clockwise from top left: the Washington Monument and Lincoln Memorial on the National Mall, United States Capitol, Logan Circle, Jefferson Memorial, White House, Adams Morgan ...
. * 1992, recipient of Chancellor's Medal for Distinguished Faculty of the University of Massachusetts at Amherst. * 1995, elected Fellow of the
World Academy of Art and Science The World Academy of Art and Science (WAAS), founded in 1960, is an international non-governmental scientific organization and global network of more than 800 scientists, artists, and scholars in more than 90 countries. It serves as a forum for s ...
. * 1997, elected to the
Russian Academy of Natural Sciences The Russian Academy of Natural Sciences ( Russian: Российская академия естественных наук) is a Russian non-governmental organization founded on August 31 1990 in Moscow in the former Soviet Union, following a de ...
. * 1998, recipient of the Distinguished Service Award of the
American Institute of Biological Sciences The American Institute of Biological Sciences (AIBS) is a nonprofit scientific charity. The organization’s mission is to promote the use of science to inform decision-making and advance biology for the benefit of science and society. Overvie ...
. * 1998, elected Fellow of the
American Academy of Arts and Sciences The American Academy of Arts and Sciences (abbreviation: AAA&S) is one of the oldest learned societies in the United States. It was founded in 1780 during the American Revolution by John Adams, John Hancock, James Bowdoin, Andrew Oliver, and ...
. * 1999, recipient of the
William Procter Prize for Scientific Achievement The William Procter Prize for Scientific Achievement is an award given by Sigma Xi, a scientific-research honor society. The Procter Prize is presented annually to a scientist who has made an outstanding contribution to scientific research and has d ...
. * 1999, recipient of the
National Medal of Science The National Medal of Science is an honor bestowed by the President of the United States to individuals in science and engineering who have made important contributions to the advancement of knowledge in the fields of behavioral and social scienc ...
, awarded by President William J. Clinton. * 2001, Golden Plate Award of the
American Academy of Achievement The American Academy of Achievement, colloquially known as the Academy of Achievement, is a non-profit educational organization that recognizes some of the highest achieving individuals in diverse fields and gives them the opportunity to meet o ...
* 2002–05, Alexander von Humboldt Prize. * 2005, elected President of
Sigma Xi Sigma Xi, The Scientific Research Honor Society () is a highly prestigious, non-profit honor society for scientists and engineers. Sigma Xi was founded at Cornell University by a junior faculty member and a small group of graduate students in 1886 ...
, The Scientific Research Society. * Profiled in ''Visionaries: The 20th Century's 100 Most Important Inspirational Leaders'', published in 2007. * Founded Sciencewriters Books in 2006 with her son Dorion. * Was one of thirteen recipients in 2008 of the Darwin-Wallace Medal, heretofore bestowed every 50 years, by the
Linnean Society of London The Linnean Society of London is a learned society dedicated to the study and dissemination of information concerning natural history, evolution, and taxonomy. It possesses several important biological specimen, manuscript and literature colle ...
. * 2009, speaker at the Biological Evolution Facts and Theories Conference, held at the
Pontifical Gregorian University The Pontifical Gregorian University ( it, Pontificia Università Gregoriana; also known as the Gregorian or Gregoriana,) is a higher education ecclesiastical school ( pontifical university) located in Rome, Italy. The Gregorian originated as ...
,
Rome , established_title = Founded , established_date = 753 BC , founder = King Romulus (legendary) , image_map = Map of comune of Rome (metropolitan city of Capital Rome, region Lazio, Italy).svg , map_caption ...
aimed at promoting dialogue between
evolutionary biology Evolutionary biology is the subfield of biology that studies the evolutionary processes (natural selection, common descent, speciation) that produced the diversity of life on Earth. It is also defined as the study of the history of life fo ...
and
Christianity Christianity is an Abrahamic monotheistic religion based on the life and teachings of Jesus of Nazareth. It is the world's largest and most widespread religion with roughly 2.38 billion followers representing one-third of the global pop ...
. * 2010, inductee into the Leonardo da Vinci Society of Thinking at the
University of Advancing Technology University of Advancing Technology (UAT) is a private for-profit university in Tempe, Arizona. Founded in 1983, UAT integrates technology into its general education requirements. The institution offers core classes (e.g., Legal Issues in Technol ...
in
Tempe, Arizona , settlement_type = City , named_for = Vale of Tempe , image_skyline = Tempeskyline3.jpg , imagesize = 260px , image_caption = Tempe skyline as se ...
. * 2010, NASA Public Service Award for Astrobiology. * 2012
Lynn Margulis Symposium: Celebrating a Life in Science
University of Massachusetts, Amherst, March 23–25, 2012 * 2017, the ''Journal of Theoretical Biology'' 434, 1–114 commemorated the 50th anniversary of "The origin of mitosing cells" with a special issue * Honorary doctorate from 15 universities.


Works


Books

* Margulis, Lynn (1970). ''Origin of Eukaryotic Cells'', Yale University Press, * Margulis, Lynn (1982). ''Early Life'', Science Books International, * Margulis, Lynn, and Dorion Sagan (1986). ''Origins of Sex : Three Billion Years of Genetic Recombination'', Yale University Press, * Margulis, Lynn, and Dorion Sagan (1987). ''Microcosmos: Four Billion Years of Evolution from Our Microbial Ancestors'', HarperCollins, * Margulis, Lynn, and Dorion Sagan (1991). ''Mystery Dance: On the Evolution of Human Sexuality'', Summit Books, * Margulis, Lynn, ed. (1991). ''Symbiosis as a Source of Evolutionary Innovation: Speciation and Morphogenesis'', The MIT Press, * * Margulis, Lynn (1992). ''Symbiosis in Cell Evolution: Microbial Communities in the Archean and Proterozoic Eons'', W.H. Freeman, * Sagan, Dorion, and Margulis, Lynn (1993). ''The Garden of Microbial Delights: A Practical Guide to the Subvisible World'', Kendall/Hunt, * Margulis, Lynn, Dorion Sagan and
Niles Eldredge Niles Eldredge (; born August 25, 1943) is an American biologist and paleontologist, who, along with Stephen Jay Gould, proposed the theory of punctuated equilibrium in 1972. Education Eldredge began his undergraduate studies in Latin at Columb ...
(1995
''What Is Life?''
Simon and Schuster, * Margulis, Lynn, and Dorion Sagan (1997). ''Slanted Truths: Essays on Gaia, Symbiosis, and Evolution'', Copernicus Books, * Margulis, Lynn, and Dorion Sagan (1997). ''What Is Sex?'', Simon and Schuster, * Margulis, Lynn, and Karlene V. Schwartz (1997). ''Five Kingdoms: An Illustrated Guide to the Phyla of Life on Earth'', W.H. Freeman & Company, * Margulis, Lynn (1998). ''Symbiotic Planet : A New Look at Evolution'', Basic Books, * Margulis, Lynn, ''et al.'' (2002). ''The Ice Chronicles: The Quest to Understand Global Climate Change'', University of New Hampshire, * Margulis, Lynn, and Dorion Sagan (2002). ''Acquiring Genomes: A Theory of the Origins of Species'', Perseus Books Group, * Margulis, Lynn (2007). ''Luminous Fish: Tales of Science and Love'', Sciencewriters Books, * Margulis, Lynn, and
Eduardo Punset Eduard Punset i Casals (; 9 November 1936 – 22 May 2019) was a Spanish politician, lawyer, economist, and science popularizer. He held a degree in Law from the Universidad Complutense de Madrid and a Master's in Economic Sciences from the Uni ...
, eds. (2007). ''Mind, Life and Universe: Conversations with Great Scientists of Our Time'', Sciencewriters Books, * Margulis, Lynn, and Dorion Sagan (2007). ''Dazzle Gradually: Reflections on the Nature of Nature'', Sciencewriters Books, *


Journals

* * * * * * * * * * * * * * * * * * * *


Notes


References


External links

* * * ** ** ** * * * * {{DEFAULTSORT:Margulis, Lynn American biologists Evolutionary biologists Theoretical biologists Symbiosis 1938 births 2011 deaths American women biologists Women evolutionary biologists Women zoologists Lyme disease researchers Members of the United States National Academy of Sciences National Medal of Science laureates Boston University faculty University of Massachusetts Amherst faculty University of California, Berkeley alumni University of Chicago Laboratory Schools alumni University of Wisconsin–Madison College of Letters and Science alumni American people of Jewish descent American agnostics Jewish women scientists Sagan family Carl Sagan Scientists from Massachusetts Symbiogenesis researchers 20th-century American zoologists 21st-century American zoologists 20th-century American biologists 21st-century biologists 20th-century American women scientists 21st-century American women scientists