Lynn Margulis, 1938-2011

Lynn Margulis died on Tuesday, the 22nd of November.

She is most famous for her theory of endosymbiosis, which says that eukaryotic cells—the complicated cells of protozoa, animals, plants, and fungi—were formed in a series of stages where one organism swallowed another but, instead of digesting it, took it on as a symbiotic partner!

As a young faculty member at Boston University, she wrote a paper on this theory. It was rejected by about fifteen scientific journals, but eventually accepted:

• Lynn Sagan, On the origin of mitosing cells, J. Theor. Bio. 14 (1967), 255–274.

(At the time she was married to the well-known astronomer Carl Sagan; she later wrote a number of books with their son Dorion.)

While Lynn Margulis was not the first to suggest that endosymbiosis played a major role in evolution, her brilliant and endlessly energetic advocacy of this idea helped trigger a scientific revolution. By now, almost all biologists agree that chloroplasts, the little green packages where photosynthesis happens in plant cells, were originally free-living bacteria of their own. Here’s a model of a chloroplast:

As you can see, it’s quite a complex world of its own. Biologists also believe that mitochondria, the ‘powerhouses’ of eukaryotic cells:

were originally bacteria.

Indeed, both chloroplasts and mitochondria still have their own DNA, and they reproduce like bacteria, through fission. This is very useful in research on human populations, since mitochondrial DNA is passed down, not through the sperm cells, but only through the egg cells—which means it’s passed down from mothers to daughters, with sons as a mere sideshow. On the other hand, Y chromosomes go down from father to son.

There’s a lot more evidence for endosymbiosis as the origin of mitochondria and chloroplasts… and in fact, we can see this process going on today in various organisms. There are also many other ways in which organisms pass genes to each other.

This led Margulis to advocate a worldview in which the ‘tree’ of life is replaced by something more like a ‘thicket’:

It also suggests that competition between organisms is part of a bigger story where the formation of partnerships plays a major role. She has written many popular books on this, and they make great reading, even though some of her ideas are still controversial.

In 1995, the famous evolutionary biologist Richard Dawkins wrote:

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.

I’ll end by quoting an excerpt from this essay, which you can read in its entirety online:

• Lynn Margulis, Gaia is a tough bitch, in The Third Culture: Beyond the Scientific Revolution, ed. John Brockman, Simon & Schuster, 1995.

I think it’s a good sample of how she wasn’t afraid to stir up controversy. Indeed, you’ll see her poking Dawkins with a sharp stick at the same time he was complimenting her!

More importantly, you’ll get a sense of how her work helped push evolutionary biology away from neo-Darwinism, also known as the Modern Synthesis. This approach tried to explain all evolution as a process of populations slowly changing through random mutation and natural selection. Now biologists are struggling to formulate an Extended Synthesis which takes many more processes into account.

Her remarks also some contain good warnings for mathematicians and physicists, such as myself, who want to dabble in biology. Physics may be defined as ‘the study of natural systems that can be accurately modeled by beautiful mathematics’. That’s why we can learn a surprising amount about physics just by sitting around scribbling. But living systems always surprise us, since they’re always more rich and complex than our models. So, biology requires more emphasis on experiment, observation, and perhaps a feeling for the organism:

I work in evolutionary biology, but with cells and microorganisms. Richard Dawkins, John Maynard Smith, George Williams, Richard Lewontin, Niles Eldredge, and Stephen Jay Gould all come out of the zoological tradition, which suggests to me that, in the words of our colleague Simon Robson, they deal with a data set some three billion years out of date. Eldredge and Gould and their many colleagues tend to codify an incredible ignorance of where the real action is in evolution, as they limit the domain of interest to animals—including, of course, people. All very interesting, but animals are very tardy on the evolutionary scene, and they give us little real insight into the major sources of evolution’s creativity. It’s as if you wrote a four-volume tome supposedly on world history but beginning in the year 1800 at Fort Dearborn and the founding of Chicago. You might be entirely correct about the nineteenth-century transformation of Fort Dearborn into a thriving lakeside metropolis, but it would hardly be world history.

By “codifying ignorance” I refer in part to the fact that they miss four out of the five kingdoms of life. Animals are only one of these kingdoms. They miss bacteria, protoctista, fungi, and plants. They take a small and interesting chapter in the book of evolution and extrapolate it into the entire encyclopedia of life. Skewed and limited in their perspective, they are not wrong so much as grossly uninformed.

Of what are they ignorant? Chemistry, primarily, because the language of evolutionary biology is the language of chemistry, and most of them ignore chemistry. I don’t want to lump them all together, because, first of all, Gould and Eldredge have found out very clearly that gradual evolutionary changes through time, expected by Darwin to be documented in the fossil record, are not the way it happened. Fossil morphologies persist for long periods of time, and after stasis, discontinuities are observed. I don’t think these observations are even debatable. John Maynard Smith, an engineer by training, knows much of his biology secondhand. He seldom deals with live organisms. He computes and he reads. I suspect that it’s very hard for him to have insight into any group of organisms when he does not deal with them directly. Biologists, especially, need direct sensory communication with the live beings they study and about which they write.

Reconstructing evolutionary history through fossils—paleontology—is a valid approach, in my opinion, but paleontologists must work simultaneously with modern-counterpart organisms and with “neontologists”—that is, biologists. Gould, Eldredge, and Lewontin have made very valuable contributions. But the Dawkins–Williams–Maynard Smith tradition emerges from a history that I doubt they see in its Anglophone social context. Darwin claimed that populations of organisms change gradually through time as their members are weeded out, which is his basic idea of evolution through natural selection. Mendel, who developed the rules for genetic traits passing from one generation to another, made it very clear that while those traits reassort, they don’t change over time. A white flower mated to a red flower has pink offspring, and if that pink flower is crossed with another pink flower the offspring that result are just as red or just as white or just as pink as the original parent or grandparent. Species of organisms, Mendel insisted, don’t change through time. The mixture or blending that produced the pink is superficial. The genes are simply shuffled around to come out in different combinations, but those same combinations generate exactly the same types. Mendel’s observations are incontrovertible.

So J. B. S. Haldane, without a doubt a brilliant person, and R. A. Fisher, a mathematician, generated an entire school of English-speaking evolutionists, as they developed the neo-Darwinist population-genetic analysis to reconcile two unreconcilable views: Darwin’s evolutionary view with Mendel’s pragmatic, anti-evolutionary concept. They invented a language of population genetics in the 1920s to 1950s called neo-Darwinism, to rationalize these two fields. They mathematized their work and began to believe in it, spreading the word widely in Great Britain, the United States, and beyond. France and other countries resisted neo-Darwinism, but some Japanese and other investigators joined in the “explanation” activity.

Both Dawkins and Lewontin, who consider themselves far apart from each other in many respects, belong to this tradition. Lewontin visited an economics class at the University of Massachusetts a few years ago to talk to the students. In a kind of neo-Darwinian jockeying, he said that evolutionary changes are due to the Fisher–Haldane mechanisms: mutation, emigration, immigration, and the like. At the end of the hour, he said that none of the consequences of the details of his analysis had been shown empirically. His elaborate cost-benefit mathematical treatment was devoid of chemistry and biology. I asked him why, if none of it could be shown experimentally or in the field, he was so wedded to presenting a cost-benefit explanation derived from phony human social-economic “theory.” Why, when he himself was pointing to serious flaws related to the fundamental assumptions, did he want to teach this nonsense? His response was that there were two reasons: the first was “P.E.” “P.E.?,” I asked. “What is P.E.? Population explosion? Punctuated equilibrium? Physical education?” “No,” he replied, “P.E. is `physics envy,'” which is a syndrome in which scientists in other disciplines yearn for the mathematically explicit models of physics. His second reason was even more insidious: if he didn’t couch his studies in the neo-Darwinist thought style (archaic and totally inappropriate language, in my opinion), he wouldn’t be able to obtain grant money that was set up to support this kind of work.

The neo-Darwinist population-genetics tradition is reminiscent of phrenology, I think, and is a kind of science that can expect exactly the same fate. It will look ridiculous in retrospect, because it is ridiculous. I’ve always felt that way, even as a more-than-adequate student of population genetics with a superb teacher—James F. Crow, at the University of Wisconsin, Madison. At the very end of the semester, the last week was spent on discussing the actual observational and experimental studies related to the models, but none of the outcomes of the experiments matched the theory.

This passage shows her tough side—these are the top names in evolutionary biology that she’s criticizing here, after all. But when I saw her speak, she was engaging and fun! You can see that yourself in these interviews. Hear how she started as a bad student in 4th grade, why her laboratory budget got cut to $0 in 2004… and get a sense of her career, personality, and ideas.



13 Responses to Lynn Margulis, 1938-2011

  1. John Baez says:

    Here’s another obituary:

    • John Horgan, R.I.P. Lynn Margulis, biological rebel, 24 November 2011.

    An interesting part:

    Toward the end of our interview, I asked Margulis if she minded always being referred to as a provocateur or gadfly, or someone who was “fruitfully wrong,” as one scientist put it. She pressed her lips together, brooding over the question. “It’s kind of dismissive, not serious,” she replied. “I mean, you wouldn’t do this to a serious scientist, would you?” She stared at me, and I finally realized her question was not rhetorical; she really wanted an answer. I agreed that the descriptions seemed somewhat condescending.

    “Yeah, that’s right,” she mused. Such criticism did not bother her, she insisted. “Anyone who makes this kind of ad hominem criticism exposes himself, doesn’t he? I mean, if their argument is just based on provocative adjectives about me rather than the substance of the issue, then…” Her voice trailed off. Like other mavericks I have met, Margulis could not help but yearn, now and then, to be a respected member of the status quo, whose work merely confirmed the prevailing paradigm. But without courageous rebels like her, science would never achieve any progress.

    I’d prefer to say: courageous rebels who are actually correct, like her.

    • David Corfield says:

      But one would hardly write

      But without courageous rebels like her who are actually correct, science would never achieve any progress.

      Even then, perhaps the cowardly mainstream would get there in the end.

      The ‘rebel scientist’ narrative is of course a very powerful one. It would be interesting to have more than an impressionistic sense of the relative advantage of allowing more diversity of thought – Feyerabend’s “Anything goes” or Kuhn’s tight paradigms.

      • John Baez says:

        Okay, maybe something like:

        But without courageous rebels like her (i.e. actually correct, unlike you crackpots out there), science would progress much more slowly.

        It’s worth noting that even if she was a ‘rebel scientist’, she got her paper published, and it instantly made a splash. She said:

        The paper was rejected by about fifteen scientific journals, because it was flawed; also, it was too new and nobody could evaluate it. Finally, James F. Danielli, the editor of The Journal of Theoretical Biology, accepted it and encouraged me. At the time, I was an absolute nobody, and, what was unheard of, this paper received eight hundred reprint requests. Later, at Boston University, it won an award for the year’s best faculty publication. I was only an instructor at the time, so my Biology Department colleagues reacted to the commotion and threw a party.

        So, the ‘unappreciated, misunderstood genius’ trope, beloved of crackpots, does not get much backing from her story. Though she still continued to feel unappreciated and misunderstood:

        But it was more of “Isn’t this cute,” or “It’s so abstruse that I don’t understand it, but others think it worthy of attention.” Even today most scientists still don’t take symbiosis seriously as an evolutionary mechanism. If they were to take symbiogenesis seriously, they’d have to change their behavior. The only way behavior changes in science is that certain people die and differently behaving people take their places.

        And indeed, in that third video, near the end, she said she would die before all her ideas were understood and accepted. I mentioned two of her ideas that are widely accepted now: endosymbiosis as an explanation of chloroplasts and of mitochondria. One of her more controversial theories involves flagella, the whiplike tails that some protozoa have. According to this website:

        “The most [?] controversial claim made by Margulis is that eukaryotic flagella evolved from small, corkscrew-shaped bacteria called spirochetes. Many spirochetes are parasites (the best known, Treponema pallidum, causes syphilis). Others are free-living, found in such exotic environments as the intestines of termites. Regardless of how they live, these unusual bacteria swim with an undulating motion reminiscent of the whiplike movement of eukaryotic flagella.

        Is this similarity evidence for Margulis’s evolutionary claim, or is it simply a coincidence? Why not accept the more orthodox explanation that eukaryotic flagella gradually evolved from the simpler flagella found on many bacteria? Margulis points out that although both types of flagella are used for locomotion, prokaryotic and eukaryotic structures are very different. Prokaryotic flagella consist of a single, hollow filament of protein that spins on its axis like a tiny propeller. Eukaryotic flagella are much larger; they contain a complex arrangement of 11 microtubules, and the entire structure is surrounded by an extension of the cell membrane.

        In contrast to the spinning prokaryotic flagellum, the eukaryotic structure propels the cell by lashing back and forth in a whiplike fashion. Because they are so different in structure, function, and perhaps evolutionary origin, Margulis proposes that the eukaryotic flagellum should be referred to by a different term: undulipodium.”

        That picture doesn’t look like it shows 11 things, but biologists call it a ‘9+2 structure’.

        • David Corfield says:

          I wonder what kinds of consideration would speak for and against this last controversy. She must have had more in mind than their being “so different in structure, function, and perhaps evolutionary origin”. Mind you, I guess it would have helped thinking this way in separating whales and dolphins from fish.

          Ultimately, must it come down to the genetics of flagellum growth and operation?

        • John Baez says:

          The big differences between bacterial flagella and the flagella of eukaryotic cells can serve as evidence that the latter didn’t evolve from the former—but only as rather weak evidence that eukaryotes got their flagella via symbiosis with spirochetes. (Spirochetes are spiral-shaped bacteria that move by twisting around.) Why does Margulis think the flagella came from symbiosis?

          Lynn Margulis has written at length on this, but unfortunately I forget most of what little I once knew, and I’m feeling too lazy tonight to read her papers. I only have the energy to read this:

          Evolution of flagella, Wikipedia.

          Apparently some eukaryotes do use symbiotic spirochetes to help move around—namely, some protozoa called parabasalids that live in the guts of termites. However, the homologies that Margulis proposed between cilia and spirochetes “haven’t stood up to further scrutiny”, according to the doubting masses. And there’s been work on how flagella could have evolved from the ‘spindles’ that eukaryotic cells use to pull apart their nuclei in the process of reproducing.

  2. Frederik De Roo says:

    Now I have learned that her understanding of the Gaia hypothesis is that “the Earth is one very large ecosystem” (instead of organism) the idea starts to sound much more serious to me.

    Also, somehow, but very likely erroneously, I have the vague impression that the dispute between her and the neo Darwinists is partly emotional, due to their different points of view: she wants to emphasize cooperation in life, the others want to stress the competition.

    • I don’t see such a serious difference.

      Her reason to take Gaia as “just” one very large ecosystem seems to be that she doesn’t produce waste, and instead does the recycling herself – unlike any other organism. But Gaia “eats” low entropy solar radiation and “excretes” high entropy radiation into space. Plus, there’s another notable excrement, coal and oil. If you don’t identify the whole planet (incl. the inner) with Gaia, but only the surface, then there’s enough room for an organismic view: The waste products being buried away by sedimentation and recycling being done by plate tectonics.

      One reason for the dispute with neo Darwinists she explains with different perspectives: Her focus is on microbial life – which is a different realm than the megafauna typically studied by evolution biologists. (Her world has e.g. horizontal gene transfer, which has some cooperative flavor. In the realm of animals and plants, genes just compete.)

      • Frederik De Roo says:

        Florifulgurator said:

        Plus, there’s another notable excrement, coal and oil.

        Are they? If I may be sarcastic, perhaps the fact that we currently burn fossil fuels is instead the most efficient method by which Gaia can recycle these otherwise wasteful products and turn them into useful carbon dioxide again which can be eaten by other lifeforms…

        But (seriously) thanks for your other comments, which are clarifying to me.

        • Alas we are re-“cycling” these excrements way too quick. Other cycles can’t catch up, so it’s not efficient. Instead e.g. ocean acidification. Not a productive use of the oceans. What could be plant and plankton food (to the delight of congressman Shimkus) turns out noxious, for it’s administered too fast.

          (This is soo paradigmatic Late Homo S “Sapiens”: Nobody ever was so efficient in non-efficiency. Working hard to annihilate other work. Vernichten von Arbeit mittels Arbeit. wrrr…)

    • John Baez says:

      Frederik wrote:

      I have the vague impression that the dispute between her and the neo Darwinists is partly emotional, due to their different points of view: she wants to emphasize cooperation in life, the others want to stress the competition.

      I think it’s emotional on her side because many people were slow to accept her theories of evolution via symbiosis, and she felt she had to fight to be taken seriously. Meanwhile, she looked at the bigshots of evolutionary biology—the people she’s criticizing in the passage I quoted—and saw many of them talking about evolution as if it were:

      1) always the result of slow incremental changes: random mutations and natural selection,

      2) nicely understood using the mathematical models of Haldane and Fisher,

      3) best studied using techniques of traditional zoology (and botany), which focus on macroscopic animals (and plants) and largely ignore the other kingdoms of life, which she considers more important.

      So, she probably felt that many evolutionary biologists were ignoring really new and interesting phenomena that can only be understood by direct observation, and wasting their time polishing beautiful theories based on insufficient data. As she says: “codifying ignorance”.

      Of course many people had already doubted parts of 1)-3). But the Modern Synthesis had solidified into a bit of a dogma by 1967, when her ground-breaking paper came out, so she probably felt rather oppressed, and probably carried a chip on her shoulder ever since.

      By now the situation is much more fluid, as people are struggling to take lots of new observations and fit them together into an ‘extended synthesis’. It must be an exciting time for anyone working on evolutionary biology! I urge everyone to read this book, to catch some of this excitement:

      • Massimo Pigliucci and Gerd B. Müller, editors, Evolution – The Extended Synthesis, MIT Press, Cambridge, Massachusetts, 2010.

      It will take a while for this new synthesis to come to some sort of resting point, if it ever does.

    • John Baez says:

      In case anyone is too lazy to read the interview with Margulis here, I’ll quote a bit about the Gaia Hypothesis:

      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. He had, by this time, a very good idea of which live organisms were probably “breathing out” the gases in question. These gases were far too abundant in the atmosphere to be formed by chemical and physical processes alone. He argued that the atmosphere was a physiological and not just a chemical system.

      The Gaia hypothesis states that the temperature of the planet, the oxidation state and other chemistry of all of the gases of the lower atmosphere (except helium, argon, and other nonreactive ones) are produced and maintained by the sum of life. We explored how this could be. How could the temperature of the planet be regulated by living beings? How could the atmospheric gas composition — the 20-percent oxygen and the one to two parts per million methane, for example — be actively maintained by living matter?

      It took me days of conversation even to begin to understand Lovelock’s thinking. My first response, just like that of the neo-Darwinists, was “business as usual.” I would say, “Oh, you mean that organisms adapt to their environment.” He would respond, very sweetly, “No, I don’t mean that.” Lovelock kept telling me what he really meant, and it was hard for me to listen. Since his was a new idea, he hadn’t yet developed an appropriate vocabulary. Perhaps I helped him work out his explanations, but I did very little else.

      The Gaia hypothesis is a biological idea, but it’s not human-centered. Those who want Gaia to be an Earth goddess for a cuddly, furry human environment find no solace in it. They tend to be critical or to misunderstand. They can buy into the theory only by misinterpreting it. Some critics are worried that the Gaia hypothesis says the environment will respond to any insults done to it and the natural systems will take care of the problems. This, they maintain, gives industries a license to pollute. Yes, Gaia will take care of itself; yes, environmental excesses will be ameliorated, but it’s likely that such restoration of the environment will occur in a world devoid of people.

      Lovelock would say that Earth is an organism. I disagree with this phraseology. No organism eats its own waste. I prefer to say that Earth is an ecosystem, one continuous enormous ecosystem composed of many component ecosystems. Lovelock’s position is to let the people believe that Earth is an organism, because if they think it is just a pile of rocks they kick it, ignore it, and mistreat it. If they think Earth is an organism, they’ll tend to treat it with respect. To me, this is a helpful cop-out, not science. Yet I do agree with Lovelock when he claims that most of the things scientists do are not science either. And I realize that by taking the stance he does he is more effective than I am in communicating Gaian ideas.

      If science doesn’t fit in with the cultural milieu, people dismiss science, they never reject their cultural milieu! If we are involved in science of which some aspects are not commensurate with the cultural milieu, then we are told that our science is flawed. I suspect that all people have cultural concepts into which science must fit. Although I try to recognize these biases in myself, I’m sure I cannot entirely avoid them. I try to focus on the direct observational aspects of science.

      Gaia is a tough bitch — a system that has worked for over three billion years without people. This planet’s surface and its atmosphere and environment will continue to evolve long after people and prejudice are gone.

  3. P.F. Henshaw says:

    Maybe what Lovelock might have said to give more meaning to his concept is that each living system emerges as an individual whole, and develops within it local “laws of nature” for its parts that change as the whole system evolves over time. That applies to the earth ecology, considered as “Gaia”, an organism of the whole of life, and also applies to each nested whole system that grows as an individual within it.

    Living systems are not the measurements of observers nor the relations between measures that observers find useful for them selves. Instead they are identified by observers as self-defining, by recognizing them as self-contained units of organization that interact with their environment, such as the living unit of organization observed as a storm, a population, a spark, a culture, etc.

  4. Quora says:

    What important ideas were initially ridiculed or rejected by experts?…

    A couple that come to mind are the personal computer and the endosymbiotic theory. Computers: The Personal Computer The personal computer (along with the Internet) is universally acknowledged to have ushered a new era in human history: the information …

You can use Markdown or HTML in your comments. You can also use LaTeX, like this: $latex E = m c^2 $. The word 'latex' comes right after the first dollar sign, with a space after it.

Fill in your details below or click an icon to log in: Logo

You are commenting using your account. Log Out /  Change )

Twitter picture

You are commenting using your Twitter account. Log Out /  Change )

Facebook photo

You are commenting using your Facebook account. Log Out /  Change )

Connecting to %s

This site uses Akismet to reduce spam. Learn how your comment data is processed.