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Lynn Margulis

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Lynn Margulis
Margulis in 2005
Born
Lynn Petra Alexander

(1938-03-05)March 5, 1938
Chicago,Illinois,U.S.
DiedNovember 22, 2011(2011-11-22)(aged 73)
Amherst, Massachusetts,U.S.
Alma mater
Known for
Spouses
  • ​
    ​
    (m.1957;div.1965)​
  • Thomas Margulis
    ​
    ​
    (m.1967;div.1980)​
Children4
Awards
Scientific career
FieldsBiology
Institutions
ThesisAn Unusual Pattern of Thymidine Incorporation in Euglena(1965)
Doctoral advisorMax Alfert

Lynn Margulis(bornLynn Petra Alexander;[1][2]March 5, 1938 – November 22, 2011)[3]was an Americanevolutionarybiologist, and was the primary modern proponent for the significance ofsymbiosis in evolution.In particular, Margulis transformed and fundamentally framed current understanding of theevolutionofcells with nucleiby proposing it to have been the result of symbiotic mergers of bacteria. Margulis was also the co-developer of theGaia hypothesiswith the British chemistJames Lovelock,proposing that the Earth functions as a single self-regulating system, and was the principal defender and promulgator of thefive kingdom classificationofRobert Whittaker.

Throughout her career, Margulis' work could arouse intense objections,[4][5]and her formative paper, "On the Origin of Mitosing Cells", appeared in 1967 after being rejected by about fifteen journals.[6]Still a junior faculty member at Boston University at the time, her theory thatcell organellessuch asmitochondriaandchloroplastswere once independentbacteriawas largely ignored for another decade, becoming widely accepted only after it was powerfully substantiated through genetic evidence. Margulis was elected a member of the USNational Academy of Sciencesin 1983. PresidentBill Clintonpresented her theNational Medal of Sciencein 1999. TheLinnean Society of Londonawarded her theDarwin-Wallace Medalin 2008.

Margulis was a strong critic ofneo-Darwinism.[7]Her position sparked lifelong debate with leading neo-Darwinian biologists, includingRichard Dawkins,[8]George C. Williams,andJohn Maynard Smith.[4]: 30, 67, 74–78, 88–92 Margulis' work on symbiosis and herendosymbiotictheory had important predecessors, going back to the mid-19th century – notablyAndreas Franz Wilhelm Schimper,Konstantin Mereschkowski,Boris Kozo-Polyansky,andIvan Wallin– and Margulis not only promoted greater recognition for their contributions, but personally oversaw the first English translation of Kozo-Polyansky'sSymbiogenesis: 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 sonDorion Sagan.

In 2002,Discovermagazine recognized Margulis as one of the 50 most important women in science.[9]

Early life and education

[edit]

Margulis was born inChicago,to aJewishfamily.[10]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.[11]She entered theHyde Park Academy High Schoolin 1952,[12]describing herself as a bad student who frequently had to stand in the corner.[2]

A precocious child, she was accepted at theUniversity of Chicago Laboratory Schools[13]at the age of fifteen.[14][15][16]In 1957, at age 19, she earned a BA from the University of Chicago inLiberal Arts.She joined theUniversity of Wisconsinto study biology underHans Risand Walter Plaut, her supervisor, and graduated in 1960 with anMSin genetics and zoology. (Her first publication, published with Plaut in 1958 in theJournal of Protozoology,was on the genetics ofEuglena,flagellateswhich have features of both animals and plants.)[17]She then pursued research at theUniversity of California, Berkeley,under the zoologist Max Alfert. Before she could complete her dissertation, she was offered research associateship and then lectureship atBrandeis Universityin Massachusetts in 1964. It was while working there that she obtained her PhD from the University of California, Berkeley in 1965. Her thesis wasAn Unusual Pattern of Thymidine Incorporation inEuglena.[18]

Career

[edit]

In 1966 she moved toBoston University,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 theUniversity of Massachusetts at Amherst.She was Distinguished Professor of Biology in 1993. In 1997 she transferred to the Department of Geosciences at UMass Amherst to become Distinguished Professor of Geosciences "with great delight",[19]the post which she held until her death.[20]

Endosymbiosis theory

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Main article:Symbiogenesis
Thechloroplastsofglaucophyteslike thisGlaucocystishave apeptidoglycanlayer, evidence of theirendosymbioticorigin fromcyanobacteria.[21]

In 1966, as a young faculty member atBoston University,Margulis wrote a theoretical paper titled "On the Origin of Mitosing Cells".[22]The paper, however, was "rejected by about fifteen scientific journals," she recalled.[6]It was finally accepted byJournal of Theoretical Biologyand is considered today a landmark in modernendosymbiotic theory.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.[2]The descent of mitochondria from bacteria and of chloroplasts from cyanobacteria was experimentally demonstrated in 1978 byRobert SchwartzandMargaret Dayhoff.[23]This formed the first experimental evidence for the symbiogenesis theory.[2]The endosymbiosis theory of organogenesis became widely accepted in the early 1980s, after the genetic material ofmitochondriaandchloroplastshad been found to be significantly different from that of the symbiont'snuclear DNA.[24]

In 1995, English evolutionary biologistRichard Dawkinshad 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.[6]

Symbiosis as evolutionary force

[edit]
Main article:Symbiosis
See also:Horizontal gene transfer

Margulis opposed competition-oriented views of evolution, stressing the importance of symbiotic or cooperative relationships between species.[25]

She later formulated a theory that proposed symbiotic relationships between organisms of different phyla, or kingdoms, as the driving force ofevolution,and explainedgenetic variationas occurring mainly through transfer of nuclear information betweenbacterial cellsorvirusesandeukaryotic cells.[25]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.[25]

Margulis also held a negative view of certain interpretations ofNeo-Darwinismthat 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."[25] 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 [the slow accrual of mutations by gene-level natural selection], is in a complete funk."[25]

Gaia hypothesis

[edit]
Further information:Gaia hypothesis

Margulis initially sought out the advice ofJames Lovelockfor 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."[6]

Margulis met with Lovelock, who explained his Gaia hypothesis to her, and very soon they began an intense collaborative effort on the concept.[6]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.'"[26]

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 theEarth system,life and its physical environment taken together.[27]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 ").[6]In her 1998 bookSymbiotic Planet,Margulis explored the relationship between Gaia and her work on symbiosis.[28]

Five kingdoms of life

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In 1969, life on earth was classified intofive kingdoms,as introduced byRobert Whittaker.[29]Margulis became the most important supporter, as well as critic[30]– while supporting parts, she was the first to recognize the limitations of Whittaker's classification of microbes.[31]But later discoveries of new organisms, such asarchaea,and emergence of molecular taxonomy challenged the concept.[32]By the mid-2000s, most scientists began to agree that there are more than five kingdoms.[33][34]Margulis became the most important defender of the five kingdom classification. She rejected thethree-domain systemintroduced byCarl Woesein 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).[32]Margulis' concept is given in detail in her bookFive Kingdoms,written with Karlene V. Schwartz.[35]It has been suggested that it is mainly because of Margulis that the five-kingdom system survives.[19]

Metamorphosis theory

[edit]

In 2009, via a then-standard publication-process known as "communicated submission" (which bypassed traditionalpeer review), she was instrumental in getting theProceedings of the National Academy of Sciences(PNAS) to publish a paper byDonald I. Williamsonrejecting "the Darwinian assumption that larvae and their adults evolved from a single common ancestor."[36][37]Williamson's paper provoked immediate response from thescientific community,including a countering paper inPNAS.[36]Conrad Labandeira of theSmithsonian National Museum of Natural Historysaid, "If I was reviewing [Williamson's paper] I 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 [...] [on] the origin of these very radical life cycles." ButDuke Universityinsect developmental biologistFred Nijhoutsaid that the paper was better suited for the "National Enquirerthan the National Academy. "[38]In September it was announced thatPNASwould eliminate communicated submissions in July 2010.PNASstated that the decision had nothing to do with the Williamson controversy.[37]

AIDS/HIV theory

[edit]

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'?"[39]which states that, "Detailed research that correlates life histories of symbioticspirochetesto changes in the immune system of associated vertebrates is sorely needed ", and urging the" reinvestigation of the natural history of mammalian,tick-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 ".[39]

In aDiscover Magazineinterview, Margulis explained her reason for interest in the topic of the 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) and the spirochete which causes Lyme disease (Borrelia) only retain about 20% of the genes they would need to live freely, outside of their human hosts.[7]

However, in theDiscover Magazineinterview Margulis said that "the set of symptoms, or syndrome, presented by syphilitics overlaps completely with another syndrome: AIDS", and also noted thatKary Mullis[a]said that "he went looking for a reference substantiating that HIV causes AIDS and discovered, 'There is no such document' ".[7]

This provoked a widespread supposition that Margulis had been an "AIDS denialist".Jerry Coyne reacted on hisWhy Evolution is Trueblog against his interpretation that Margulis believed "that AIDS is really syphilis, not viral in origin at all."[40]Seth Kalichman,a social psychologist who studies behavioral and social aspects of AIDS, cited her [Margulis] 2009 paper as an example of AIDS denialism "flourishing",[41]and asserted that her [Margulis] "endorsement of HIV/AIDS denialism defies understanding".[42]

Reception

[edit]

HistorianJan Sapphas said that "Lynn Margulis's name is as synonymous with symbiosis asCharles Darwin's is with evolution. "[4]She has been called "science's unruly earth mother",[43]a "vindicated heretic",[44]or a scientific "rebel",[45]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.[4]

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,"[46]and in the commemorative collection of essaysLynn Margulis: The Life and Legacy of a Scientific Rebel,commentators again and again depict her as a modern embodiment of the "scientific rebel",[4]akin toFreeman Dyson's 1995 essayThe Scientist as Rebel,a tradition Dyson saw embodied inBenjamin Franklin,and which Dyson believed to be essential to good science.[47]

Awards and recognitions

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Personal life

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Margulis married astronomerCarl Saganin 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, acrystallographer.They had a son named Zachary Margulis-Ohnuma, a New York City criminal defense lawyer, and a daughter Jennifer Margulis, teacher and author.[64][65]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."[65]

In the 2000s she had a relationship with fellow biologist Ricardo Guerrero.[12]

Margulis argued that theSeptember 11 attackswere a "false-flag operation,which has been used to justify the wars in Afghanistan and Iraq as well as unprecedented assaults on [...] civil liberties. "She wrote that there was" overwhelming evidence that the three buildings [of the World Trade Center] collapsed by controlled demolition. "[4]

She was a religiousagnostic,[12]and a staunchevolutionist,but rejected themodern evolutionary synthesis,[7]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."[6]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.[7]

Margulis died on November 22, 2011, at home inAmherst,Massachusetts,five days after suffering ahemorrhagic stroke.[1][2][65][66]As her wish, she was cremated and her ashes were scattered in her favorite research areas, near her home.[67]

Works

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Books

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  • Margulis, Lynn (1970).Origin of Eukaryotic Cells,Yale University Press,ISBN0-300-01353-1
  • Margulis, Lynn (1982).Early Life,Science Books International,ISBN0-86720-005-7
  • Margulis, Lynn, and Dorion Sagan (1986).Origins of Sex: Three Billion Years of Genetic Recombination,Yale University Press,ISBN0-300-03340-0
  • Margulis, Lynn, and Dorion Sagan (1987).Microcosmos: Four Billion Years of Evolution from Our Microbial Ancestors,HarperCollins,ISBN0-04-570015-X
  • Margulis, Lynn, and Dorion Sagan (1991).Mystery Dance: On the Evolution of Human Sexuality,Summit Books,ISBN0-671-63341-4
  • Margulis, Lynn, ed. (1991).Symbiosis as a Source of Evolutionary Innovation: Speciation and Morphogenesis,The MIT Press,ISBN0-262-13269-9
  • Margulis, Lynn (1991). "Symbiosis in Evolution: Origins of Cell Motility". In Osawa, Syozo; Honzo, Tasuku (eds.).Evolution of Life.Japan: Springer. pp. 305–324.doi:10.1007/978-4-431-68302-5_19.ISBN978-4-431-68304-9.
  • Margulis, Lynn (1992).Symbiosis in Cell Evolution: Microbial Communities in the Archean and Proterozoic Eons,W.H. Freeman,ISBN0-7167-7028-8
  • Sagan, Dorion, and Margulis, Lynn (1993).The Garden of Microbial Delights: A Practical Guide to the Subvisible World,Kendall/Hunt,ISBN0-8403-8529-3
  • Margulis, Lynn, Dorion Sagan andNiles Eldredge(1995)What Is Life?,Simon and Schuster,ISBN978-0684810874
  • Margulis, Lynn, and Dorion Sagan (1997).Slanted Truths: Essays on Gaia, Symbiosis, and Evolution,Copernicus Books,ISBN0-387-94927-5
  • Margulis, Lynn, and Dorion Sagan (1997).What Is Sex?,Simon and Schuster,ISBN0-684-82691-7
  • Margulis, Lynn, and Karlene V. Schwartz (1997).Five Kingdoms: An Illustrated Guide to the Phyla of Life on Earth,W.H. Freeman & Company,ISBN0-613-92338-3
  • Margulis, Lynn (1998).Symbiotic Planet: A New Look at Evolution,Basic Books,ISBN0-465-07271-2
  • Margulis, Lynn,et al.(2002).The Ice Chronicles: The Quest to Understand Global Climate Change,University of New Hampshire,ISBN1-58465-062-1
  • Margulis, Lynn, and Dorion Sagan (2002).Acquiring Genomes: A Theory of the Origins of Species,Perseus Books Group,ISBN0-465-04391-7
  • Margulis, Lynn (2007).Luminous Fish: Tales of Science and Love,Sciencewriters Books,ISBN978-1-933392-33-2
  • Margulis, Lynn, andEduardo Punset,eds. (2007).Mind, Life and Universe: Conversations with Great Scientists of Our Time,Sciencewriters Books,ISBN978-1-933392-61-5
  • Margulis, Lynn, and Dorion Sagan (2007).Dazzle Gradually: Reflections on the Nature of Nature,Sciencewriters Books,ISBN978-1-933392-31-8
  • Margulis, Lynn (2009). "Genome Acquisition in Horizontal Gene Transfer: Symbiogenesis and Macromolecular Sequence Analysis". In Gogarten, Maria Boekels;Gogarten, Johann Peter;Olendzenski, Lorraine C. (eds.).Horizontal Gene Transfer.Methods in Molecular Biology. Vol. 532. Humana Press. pp. 181–191.doi:10.1007/978-1-60327-853-9_10.ISBN978-1-60327-852-2.PMID19271185.

Journals

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Explanatory notes

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  1. ^Kary Mulliswon the 1993Nobel Prizefor thepolymerase chain reaction,and was known for his unconventional scientific views.

References

[edit]
  1. ^abWeber, Bruce (November 24, 2011)."Lynn Margulis, evolution theorist, dies at 73".The New York Times.RetrievedJuly 25,2014.
  2. ^abcdefLake, James A.(2011)."Lynn Margulis (1938–2011)".Nature.480(7378): 458.Bibcode:2011Natur.480..458L.doi:10.1038/480458a.PMID22193092.S2CID205069081.
  3. ^Schaechter, M (2012). "Lynn Margulis (1938–2011)".Science.335(6066): 302.Bibcode:2012Sci...335..302S.doi:10.1126/science.1218027.PMID22267805.S2CID36800637.
  4. ^abcdefSagan, Dorion, ed. (2012).Lynn Margulis: The Life and Legacy of a Scientific Rebel.White River Junction: Chelsea Green.ISBN978-1603584470.
  5. ^"Lynn Margulis".The Telegraph.December 13, 2011.Archivedfrom the original on January 12, 2022.RetrievedMarch 9,2021.
  6. ^abcdefgMargulis, Lynn,Gaia Is a Tough BitchArchivedNovember 22, 2017, at theWayback Machine.Chapter 7 in The Third Culture: Beyond the Scientific Revolution by John Brockman (Simon & Schuster, 1995)[dead link]
  7. ^abcdeTeresi, Dick(June 17, 2011)."Lynn Margulis says she's not controversial, she's right".Discover Magazine.Discover Interview. No. April 2011.RetrievedJune 22,2023. [Broken link]
  8. ^Gilbert, Scott F.;Sapp, Jan;Tauber, Alfred I.(2012)."A Symbiotic View of Life: We have never been individuals".The Quarterly Review of Biology.87(4): 325–341.doi:10.1086/668166.PMID23397797.S2CID14279096.
  9. ^Svitil, Kathy (November 13, 2002)."The 50 Most Important Women in Science".Discover Magazine.RetrievedMay 1,2019.
  10. ^Goldman, Jason."Ad Memoriam: Lynn Margulis (5.03.1938 - 22.11.2011)"(PDF).Jason G. Goldman.RetrievedSeptember 14,2015.
  11. ^Oakes, Elizabeth H. (2007).Encyclopedia of World Scientists(Revised ed.). New York: Facts on File. p. 484.ISBN978-1-4381-1882-6.
  12. ^abc"Lynn Margulis".NNDB.Soylent Communications.RetrievedDecember 18,2014.
  13. ^di Properzio, James (February 1, 2004)."Lynn Margulis: Full speed ahead".University of Chicago Magazine.Archived fromthe originalon July 23, 2014.RetrievedJuly 25,2014.
  14. ^Scoville, Heather."Lynn Margulis".About.com.Archived fromthe originalon December 18, 2014.RetrievedDecember 18,2014.
  15. ^"Lynn Margulis".Encyclopedia of World Biography.2004.RetrievedDecember 18,2014.
  16. ^A Life With...(interview). Series 5. BBC Radio 4. July 16, 2009.
  17. ^Archibald, John (2014).One Plus One Equals One: Symbiosis and the evolution of complex life.Oxford: Oxford University Press. p. 50.ISBN978-0-19-966059-9.
  18. ^abcMargulis, Lynn (2002).Una revolución en la evolución: Escritos seleccionados(in Spanish). Valencia: Universitat de Valencia. pp. 45–48.ISBN978-8-437-05494-0.
  19. ^abcdYount, Lisa (2003).A to Z of biologists.New York, NY: Facts on File. p. 198.ISBN978-1-4381-0917-6.
  20. ^abcdHaskett, Dorothy Regan."Lynn Petra Alexander Sagan Margulis (1938–2011)".The Embryo Project Encyclopedia.Arizona Board of Regents, Arizona State University.RetrievedDecember 18,2014.
  21. ^Keeling, Patrick J. (2004)."Diversity and evolutionary history of plastids and their hosts".American Journal of Botany.91(10): 1481–1493.doi:10.3732/ajb.91.10.1481.PMID21652304.
  22. ^Sagan, Lynn (1967). "On the origin of mitosing cells".Journal of Theoretical Biology.14(3): 225–274.Bibcode:1967JThBi..14..225S.doi:10.1016/0022-5193(67)90079-3.PMID11541392.
  23. ^Schwartz, R.; Dayhoff, M. (1978). "Origins of prokaryotes, eukaryotes, mitochondria, and chloroplasts".Science.199(4327): 395–403.Bibcode:1978Sci...199..395S.doi:10.1126/science.202030.PMID202030.
  24. ^Gillham, Nicholas W. (January 14, 2014)."Chloroplasts and Mitochondria".In Reeve, Eric C.R. (ed.).Encyclopedia of Genetics.Routledge. pp. 721–735.ISBN978-1-134-26350-9.
  25. ^abcdeMann, C (1991). "Lynn Margulis: Science's unruly Earth mother".Science.252(5004): 378–381.Bibcode:1991Sci...252..378M.doi:10.1126/science.252.5004.378.PMID17740930.
  26. ^Lovelock, J.E.; Margulis, L. (1974)."Atmospheric homeostasis by and for the biosphere: the gaia hypothesis".Tellus A.26(1–2): 2–10.Bibcode:1974Tell...26....2L.doi:10.3402/tellusa.v26i1-2.9731.S2CID129803613.
  27. ^Lovelock, James (1988).The Ages of Gaia: A Biography of Our Living Earth.New York: W.W.Norton & Co.
  28. ^Margulis, Lynn (1998).Symbiotic Planet.New York, NY: Basic Books.
  29. ^Whittaker, R.H. (January 1969). "New concepts of kingdoms or organisms. Evolutionary relations are better represented by new classifications than by the traditional two kingdoms".Science.163(3863): 150–60.Bibcode:1969Sci...163..150W.CiteSeerX10.1.1.403.5430.doi:10.1126/science.163.3863.150.PMID5762760.
  30. ^Margulis, Lynn (1974). "Five-Kingdom Classification and the Origin and Evolution of Cells".Evolutionary Biology.Vol. 7. pp. 45–78.doi:10.1007/978-1-4615-6944-2_2.ISBN978-1-4615-6946-6.PMC1847511.PMID17376230.
  31. ^Margulis, Lynn (1971). "Whittaker's Five Kingdoms of Organisms: Minor Revisions Suggested by Considerations of the Origin of Mitosis".Evolution.25(1): 242–245.doi:10.2307/2406516.JSTOR2406516.PMID28562945.
  32. ^abHagen, Joel B. (2012)."Five Kingdoms, More or Less: Robert Whittaker and the Broad Classification of Organisms".BioScience.62(1): 67–74.doi:10.1525/bio.2012.62.1.11.S2CID86253586.
  33. ^Simpson, Alastair G.B. & Roger, Andrew J. (2004)."The real 'kingdoms' of eukaryotes".Current Biology.14(17): R693–6.Bibcode:2004CBio...14.R693S.doi:10.1016/j.cub.2004.08.038.PMID15341755.S2CID207051421.
  34. ^Adl, SM; Simpson, AG; Farmer, MA; Andersen, RA; Anderson, OR; Barta, JR; Bowser, SS; Brugerolle, G; et al. (2005)."The new higher level classification of eukaryotes with emphasis on the taxonomy of protists"(PDF).The Journal of Eukaryotic Microbiology.52(5): 399–451.doi:10.1111/j.1550-7408.2005.00053.x.PMID16248873.S2CID8060916.
  35. ^Tao, Amy (October 22, 2013)."Lynn Margulis".Encyclopædia Britannica.RetrievedDecember 18,2014.
  36. ^abWilliamson, D. I. (2009)."Caterpillars evolved from onychophorans by hybridogenesis".Proceedings of the National Academy of Sciences.106(47): 19901–19905.Bibcode:2009PNAS..10619901W.doi:10.1073/pnas.0908357106.PMC2785264.PMID19717430.
  37. ^abControversial caterpillar-evolution study formally rebutted,Scientific AmericanOnline
  38. ^Borrell, Brendan."National Academy as National Enquirer? PNAS Publishes Theory That Caterpillars Originated from Interspecies Sex".Scientific American.RetrievedNovember 23,2011.
  39. ^abMargulis, Lynn; Maniotis, Andrew; MacAllister, James; Scythes, John; Brorson, Oystein; Hall, John; Krumbein, Wolfgang E.; Chapman, Michael J. (2009)."Syphilis, Lyme disease & AIDS: Resurgence of" the great imitator "?"(PDF).Symbiosis.47(1): 51–58.doi:10.1007/BF03179970.S2CID25177964.
  40. ^Jerry Coyne (April 12, 2011)."Lynn Margulis disses evolution in Discover Magazine – Embarrasses both herself and the field".Why Evolution is True.
  41. ^Kalichman, S. C.; Eaton, L.; Cherry, C. (2010).""There is no Proof that HIV Causes AIDS": AIDS denialism beliefs among people living with HIV/AIDS ".Journal of Behavioral Medicine.33(6): 432–440.doi:10.1007/s10865-010-9275-7.PMC3015095.PMID20571892.
  42. ^Seth C. Kalichman(January 16, 2009).Denying AIDS: Conspiracy Theories, Pseudoscience, and Human Tragedy.Springer Science & Business Media. pp. 181–82.ISBN978-0-387-79476-1.
  43. ^Mann, C (1991). "Lynn Margulis: Science's unruly Earth mother".Science.252(5004): 378–381.Bibcode:1991Sci...252..378M.doi:10.1126/science.252.5004.378.PMID17740930.
  44. ^Barlow, Connie (1992).From Gaia to Selfish Genes: Selected writings in the life sciences(1st MIT Press paperback ed.). Cambridge, MA: MIT Press. p. 47.ISBN978-0-262-52178-9.
  45. ^Fiveash, Kelly (November 24, 2011)."'Rebel' biologist and neo-Darwinian skeptic Lynn Margulis dies ".The Register.RetrievedDecember 19,2014.
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