In Memoriam: Lynn Margulis (1938–2011)

The Revolutionary Biologist Who Transformed Our Understanding of Life

15

Journal Rejections

2

Billion Years of Evolution Explained

1

Revolutionary Theory

A Scientific Rebel's Journey

In the history of science, few stories are as compelling as that of Lynn Margulis—a brilliant biologist whose revolutionary ideas about the cooperative nature of life were initially rejected by the scientific establishment, only to become foundational principles of modern biology.

When Margulis first proposed that complex cells originated through symbiotic mergers of bacteria, her paper was rejected by fifteen journals before finally seeing publication in 19671 . One reviewer even told her, "Your research is crap"6 . Yet through what Richard Dawkins would later call "sheer courage and stamina," she persevered, ultimately transforming our understanding of how life evolves and interacts1 .

Margulis championed the then-radical idea that evolution progressed as much through cooperation and symbiosis as through competition—a vision that contrasted sharply with the dominant neo-Darwinian perspective of her time6 8 .

1938

Born in Chicago, Illinois

1967

Publishes groundbreaking paper after 15 rejections

1970

Expands theory in "Origin of Eukaryotic Cells"

1983

Elected to National Academy of Sciences

1999

Awarded National Medal of Science

2011

Passes away at age 73

A Revolutionary Idea: Life Through Collaboration

The Endosymbiotic Theory

The Theory

Margulis's most celebrated contribution, the serial endosymbiotic theory (SET), proposed that eukaryotic cells (the complex cells that make up plants, animals, and fungi) originated through a series of symbiotic mergers between different types of bacteria.

She hypothesized that approximately two billion years ago, ancient prokaryotic cells engulfed certain bacteria but didn't digest them—instead, they established permanent, mutually beneficial relationships that evolved into the specialized organelles we recognize in cells today1 .

Key Proposals
  • Mitochondria evolved from oxygen-using proteobacteria3
  • Chloroplasts descended from cyanobacteria3
  • Eukaryotic flagella may have originated from spirochete-like bacteria3
Evidence Mounts

Though initially controversial, Margulis's theory began gaining traction as new genetic evidence emerged. The discovery that mitochondria and chloroplasts contained their own DNA, distinct from the DNA in the cell's nucleus, provided critical support for her idea1 .

Even more convincing was the finding that the DNA in these organelles more closely resembled bacterial DNA than the nuclear DNA of eukaryotic cells7 .

"Thanks to RNA, she proved that the mitochondria of wheat do not resemble wheat: they are genetically alien, but co-opted into the service of wheat"6 .

The Experiment That Changed Everything

Genetic Evidence Validates a Controversial Theory

For more than a decade after Margulis published her revolutionary theory, it remained on the fringes of biological thought. The turning point came in 1978 when Robert Schwartz and Margaret Dayhoff published a groundbreaking study comparing the genetic material of organelles to that of various bacteria1 7 .

Their work provided the first compelling molecular evidence supporting Margulis's once-radical proposal.

Methodology: Tracking Molecular Blueprints

Schwartz and Dayhoff's approach focused on comparing genetic sequences across different organisms. Their experimental procedure followed these key steps:

  1. Isolation of genetic material: DNA was carefully extracted from mitochondria and chloroplasts of various eukaryotic species, as well as from multiple types of bacteria3
  2. Sequence analysis: The researchers analyzed the molecular sequences of the genetic code, paying particular attention to the genes responsible for fundamental cellular processes3
  3. Comparison and alignment: Using emerging bioinformatic techniques, they compared the genetic sequences of organelle DNA with both bacterial DNA and nuclear DNA from eukaryotic cells3
  4. Evolutionary tracking: By identifying similarities and differences in these genetic sequences, the team could reconstruct evolutionary relationships and determine which organisms shared the most recent common ancestors3
Genetic Evidence Supporting Endosymbiotic Theory
Genetic Feature Mitochondrial Similarity to Bacteria Chloroplast Similarity to Cyanobacteria
Ribosomal RNA sequences High similarity to α-proteobacteria High similarity to cyanobacteria
Gene organization More similar to bacterial genomes Nearly identical in some regions
Translation machinery Bacterial-like Bacterial-like
Antibiotic sensitivity Similar to bacteria Similar to cyanobacteria
Timeline of Endosymbiotic Theory Acceptance
Year Event Significance
1967 Margulis publishes "On the Origin of Mitosing Cells" Theory initially rejected or ignored by most scientists
1970 Margulis expands theory in "Origin of Eukaryotic Cells" Presents comprehensive evidence for endosymbiosis
1978 Schwartz & Dayhoff publish genetic evidence Provides first molecular confirmation of theory
Early 1980s Theory gains widespread acceptance Becomes new biological paradigm
1999 Margulis awarded National Medal of Science Official recognition of her contribution

A Scientific Toolkit: Investigating Endosymbiosis

Research into endosymbiotic relationships requires specialized methods and materials

Key Research Tools for Studying Endosymbiosis
Tool/Method Function Application in Endosymbiosis Research
Electron microscopy High-resolution imaging of cellular structures Revealed structural similarities between organelles and bacteria
DNA sequencing Determining genetic code Allowed comparison of organelle and bacterial genomes
PCR amplification Copying specific DNA segments Enabled study of rare genes in organelles
Ribosomal RNA analysis Evolutionary comparison using conserved genes Provided "molecular clock" for dating evolutionary divergences
Antibiotic sensitivity testing Assessing response to bacterial inhibitors Showed organelle translation resembles bacterial systems
Fluorescence in situ hybridization Visualizing specific genetic sequences Confirmed location and organization of organelle DNA

These tools collectively enabled scientists to move beyond morphological observations to the genetic and molecular evidence that ultimately validated Margulis's theory.

"The DNA associated with certain structures had 'evaded detection because it has very little metabolic activity and is difficult to detect by standard techniques,' requiring increasingly sophisticated methods to uncover"5 .

Overcoming Resistance: The Struggle for Acceptance

Challenging the Scientific Orthodoxy

Margulis faced intense opposition throughout much of her career, with her work frequently attracting "intense objections" from leading biologists1 . Her formative paper was rejected by approximately fifteen journals before appearing in the Journal of Theoretical Biology in 19673 .

This resistance stemmed from several factors:

  • Her theory challenged the neo-Darwinian focus on competition as the primary driver of evolution1
  • She proposed major evolutionary transitions through symbiosis, rather than exclusively through gradual accumulation of small mutations6
  • As a young female scientist in a male-dominated field, she faced additional barriers to having her ideas taken seriously6
"I'm not interested in being a 'good sport' if it means compromising what I know to be true"6 .

Recognition and Honors

Despite the initial resistance, Margulis lived to see her ideas become widely accepted and celebrated. Her perseverance earned her some of science's highest honors:

National Medal of Science

Presented by President Bill Clinton in 19991

U.S. National Academy of Sciences

Elected in 19831

Darwin-Wallace Medal

Awarded by the Linnean Society of London in 20081

"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. This is one of the great achievements of twentieth-century evolutionary biology"1 .
Richard Dawkins

A Lasting Legacy: Beyond Endosymbiosis

Margulis's scientific contributions extended far beyond explaining the origin of organelles

The Gaia Hypothesis

In the early 1970s, Margulis began a fruitful collaboration with British chemist James Lovelock, co-developing what would become known as the Gaia hypothesis1 .

This proposition suggested that Earth itself functions as a unified, self-regulating system1 .

Five Kingdoms of Life

Together with biologist Karlene V. Schwartz, Margulis developed a comprehensive five-kingdom classification system that organized life into animals, plants, bacteria (prokaryotes), fungi, and protoctists.

This system rejected the traditional protist kingdom as too general.

Expanding the Symbiotic View

Margulis continued to develop and expand her symbiotic view of evolution throughout her career. She argued that symbiogenesis—the formation of new species through symbiotic mergers—was a major mechanism of evolutionary innovation7 .

This perspective has gained new relevance with recent research into the human microbiome6 .

"We are all of us walking communities of bacteria"

- Lynn Margulis6

A Vision Vindicated

Lynn Margulis left an indelible mark on biology, fundamentally transforming our understanding of life's history and interconnectedness.

Her work replaced a narrative of relentless competition with a more nuanced view that recognized cooperation as a creative force in evolution. By demonstrating that complex cells emerged through symbiotic mergers, she provided a scientific basis for understanding life as fundamentally collaborative—a vision with profound implications for how we see ourselves and our relationship to the natural world.

The journey from dismissed heretic to celebrated scientific hero stands as a testament to her persistence, intellectual courage, and unwavering commitment to evidence. This integrity, combined with her revolutionary insights, cemented her place as one of the most important biologists of the 20th century—one whose vision of a "symbiotic planet" continues to inspire new generations of scientists to see the living world as deeply interconnected and collaborative.

References