A brief history of the endosymbiotic theory

By Constanza Loreto Di Giammarco

Earth’s life history is substantiated by evolution nevertheless this force is everything but
homogenous it encompasses multiple events at different paces, which leaves behind the
fixated idea of gradualism, introduced in Charles Darwin’s On the Origin of Species
(Darwin, 1859). Indeed life itself has undergone a process characterized by ancestry and
diversity, the latter having notable events, like the Cambric Explosion, 540 million years
ago, nonetheless if a definitive and crucial event should be suggested, it would be life’s
most notable division: 2.000 million years ago Earth only harbored bacteria and archaea,
and in a matter of an instant, powered by specific physical and chemical conditions,
symbiosis occurred and a complex, nucleated, doubled membrane cell appeared, called
eukarya (Sampedro, 2010, 2015).
The term symbiosis originated in Russia in a very specific social climate, characterized by
the influence of the works of Charles Darwin and Ernst Haeckel, apart from any Boshevik
influence, Russian scientists view Darwin’s natural selection as a theory of mutual aid
rather than mere competition (Todes, 1989)(Lazcano and Peretó, 2017). In this complex
scientific context, the role of symbiosis in the evolution of nucleated cells was discussed by
Konstantin Merezhkovsky, Andrei Famintsyn and Boris Kozo-Polyanski (Sapp, 2009)
(Lazcano and Peretó, 2017), although these figures were motivated by anti-mendelism,
their contributions were the first steps towards a more open perspective regarding the ways
of evolution. Merezhkovsky and his peers remained in the dark, and when instigated their
ideas were credited as interesting but not vital to the understanding of heredability and
evolution, in the words of Thomas H. Morgan, renowned scientist and winner of the Nobel
prize of physiology and medicine: “In a word the cytoplasm may be ignored genetically”
(Morgan, 1926).
Symbiosis remained a latent idea, until the re-emergence of molecular biology and electron
microscopy, demonstrating the ultra-structural similarity of a cell organelle and free-living
organisms (Ris and Plaut, 1962). Hans Ris, an American cytologist and pioneer electron
microscopist, introduced the symbiotic theory to the then undergraduate Lynn Margulis, in
her own words: “when Ris read to us ‘More recently Wallin (1922) has maintained that
chondriosomes may be regarded as symbiotic bacteria whose associations with other
cytoplasmic components may have arisen in the earliest stages of evolution … To many, no
doubt, such consideration may appear too fantastic to mention in polite scientific society;
nevertheless, it is in the range of possibility that they may someday call for more serious
consideration…’ the course of my professional life was set forever!” (Margulis, 2005;
Lazcano and Peretó, 2017).
Lynn Margulis was critical in the positioning and validation of the symbiotic theory, in her
article On the Origin of Mitosing Cells (Sagan, 1967) she accomplished the exceptional
task of finding and connecting morphological threads to support the symbiotic ancestry of
life on earth; she hypothesized that the mitochondria, photosynthetic plastids and basal
bodies of flagella were once free-living prokaryotic cells, stablishing the first unified theory
of eukaryogenesis (Gray, 2017). Margulis’s ideas initiated an interesting debate between
proponents of autogenous origin and exogenous origin theories of organelle evolution
(Gray, 2017) along with the rise of microevolutionary biology and the constant idea that
communities prevail over individuals.

Darwin, C. (1859) On the Origin of Species, On the Origin of Species, 1859. Available at:
https://doi.org/10.4324/9780203509104.

Gray, M.W. (2017) ‘Lynn Margulis and the endosymbiont hypothesis: 50 years later’, Molecular
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Lazcano, A. and Peretó, J. (2017) ‘On the origin of mitosing cells: A historical appraisal of Lynn
Margulis endosymbiotic theory’, Journal of Theoretical Biology, 434, pp. 80–87. Available at:
https://doi.org/10.1016/j.jtbi.2017.06.036.

Margulis, L. (2005) ‘Hans Ris (1914-2004): Genophore, chromosomes and the bacterial origin of
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http://scielo.isciii.es/scielo.php?script=sci_arttext&pid=S1139-67092005000200011&nrm=iso.

Morgan, T.H. (1926) ‘Genetics and the physiology of development’, The American Naturalist ,
10(671), pp. 489–515.

Ris, H. and Plaut, W. (1962) Ultrastructure of DNA-containing areas in the chloroplast of

chlamydomonas. Available at: http://rupress.org/jcb/article-pdf/13/3/383/1479103/383.pdf.

Sagan, L. (1967) On the Origin of Mitosing Cells, J. Theoret. Biol.

Sampedro, J. (2010) Deconstruyendo a Darwin. Los enigmas de la evolución a la luz de la nueva

genética, Ciencias. Available at:
https://www.revistas.unam.mx/index.php/cns/article/view/18905.

Sampedro, J. (2015) ‘La genómica da la razón a Lynn Margulis’, El País, 20 August. Available at:
https://elpais.com/elpais/2015/08/19/ciencia/1440001134_537219.html?
event=go&event_log=go&prod=REGCRARTCIENCIA&o=cerrciencia.

Sapp, J. (2009) The New Foundations of Evolution: On the Tree of Life. . Oxford: Oxford University
Press.

Todes, D.P. (1989) Darwin Without Malthus: The Struggle for Existence in Russian Evolutionary
Thought., Russian History. New York: Oxford University Press. Available at:
https://doi.org/10.1163/187633190x00877.