SYMBIOGENESIS. A NEW PRINCIPLE OF EVOLUTION REDISCOVERY OF BORIS MIKHAYLOVICH KOZO-POLYANSKY (1890–1957)

Lynn Margulis

University of Massachusetts

Amherst, USA: celeste@geo. umass. edu

The following is a heavily edited transcript of my illustrated lecture, that included our

14 minute video (with a 2 minute animation model) that shows each step in live organisms

hypothesized in the origin of nucleated cells from bacteria (“eukaryosis”). New observa-tions presented with modern examples of live phenomena make us virtually certain that

B. M. Kozo-Polyansky’s “new principle” (1924) of the importance of symbiogenesis in the

evolutionary process of at least 2000 million years of life on Earth is correct. The widely

touted but undocumented explanation of the origin of evolutionary novelty by “gradual

accumulation of random mutations” will be considered an erroneous early 20th century

hunch proffered primarily by Englishmen, North Americans and other anglophones. They

(Neodarwinist “explanations”) will be replaced by the details of symbiogenesis: genetic

mergers especially speciation by genome acquisition, karyotypic fissions (neocentromere

formation, related chromosome change) and D. I. Williamson’s larval transfer concept for

animals. Although ignored and dismissed in his life time, Kozo-Polyansky’s brilliant work

will be lauded for symbiogenesis in the same style that Gregor Mendel’s studies of inher-itance of “factors” in peas was for recognition of his establishment of diploid organism

genetic principles by the beginning of the 20th century. My talk, photographs and moving

pictures were presented at the Darwin conference, St. Petersburg, on September 23, 2009

introduced by E. Kolchinsky.

Keywords: symbiogenesis, eukaryosis, Kozo-Polyansky, speciation, genetic mergers, ge — nome acquisitions.

In most of these sessions it is customary only at the end of one’s lecture to thank those who do the relevant work or have helped. I prefer to thank my predecessors or colleagues here at the beginning. I want to list the Russian literature that has led me to certain conclusions that I will mention. In my talk I plan to use few words, as I’d rather show you live organisms in color videos and photos, especially electron micrographs. Let us begin with Boris Perfiliev (1891–1969), does anyone know his name? He is the father of microbial ecology, some say. He made capillaries, tiny sedimentary cores, minicores, from square glass capillaries. He used them to study layered sediment, stratified microbial communities. From Perfiliev and his student Galina Dubinina we mention a Academi — cian, Professor Armen Takhtajan, who is still here. The great botanist is 99 years old, as you know. From Takhtajan I continue on to laud a scientist I knew well: Igor Raikov. Rai- kov (1932–1998) was one of the few modern scientists who recognized the importance of the protists for understanding the origin of genetic systems. Raikov was a person who actually did what Charles Darwin had suggested. Darwin admonished, if one wants to understand the evolution of any process or structure one should follow the oddities and

the peculiarities, he called them “oddities and imperfections” in life, because otherwise one may be deceived into thinking that the way life is now is the way it always has been. But the way life forms are now is not the way they have always been. Raikov was excel — lent at reconstruction of ciliate evolution.

From Raikov we come to Liya Nikolaevna Khakhina. It was her book (1992) that led me to Kozo-Polyansky (1924, 2010). Victor Fet, our translator, without whom this work would not have been possible, now lives in West Virginia. He was raised in No — vosibirsk. His extremely literate father taught computers and informatics in Siberia. Unfortunately, Victor could not attend this evolution meeting. But his wife, Dr. Galina Fet, who comes from St. Petersburg, is here now. She has helped us enormously in our visit here. We thank her.

Please, Professor Kolchinsky, ask somebody to translate this for me? Who here knows the work of Dr. Galina Dubinina, microbiology professor from Moscow State University? OK. Nobody! Amazing and unfortunate. Dubinina’s work, along with that of her team and her predecessor, Professor Boris Perfiliev, is crucial to our research, crucially important.

So I want to say, yes, Charles Darwin, in general, was correct. I believe Dr.

the peculiarities, he called them “oddities and imperfections” in life, because otherwise one may be deceived into thinking that the way life is now is the way it always has been. But the way life forms are now is not the way they have always been. Raikov was excel — lent at reconstruction of ciliate evolution.

From Raikov we come to Liya Nikolaevna Khakhina. It was her book (1992) that led me to Kozo-Polyansky (1924, 2010). Victor Fet, our translator, without whom this work would not have been possible, now lives in West Virginia. He was raised in No — vosibirsk. His extremely literate father taught computers and informatics in Siberia. Unfortunately, Victor could not attend this evolution meeting. But his wife, Dr. Galina Fet, who comes from St. Petersburg, is here now. She has helped us enormously in our visit here. We thank her.

Please, Professor Kolchinsky, ask somebody to translate this for me? Who here knows the work of Dr. Galina Dubinina, microbiology professor from Moscow State University? OK. Nobody! Amazing and unfortunate. Dubinina’s work, along with that of her team and her predecessor, Professor Boris Perfiliev, is crucial to our research, crucially important.

So I want to say, yes, Charles Darwin, in general, was correct. I believe Dr. Georgy Levit, who spoke today before me, said that as well. But Neo-Darwinism, fundamen — tally is a religion. It is very bad. It is very limited. It enters realms where it does not belong. It is reductionist. It does not even study life. Perhaps I should simply claim that Neodarwinism took the life out of biology. Somebody please translate that com — ment, as I do want to declare in public that Neodarwinism has had a stultifying effect on the study of the evolution of life on Earth. The Neodarwinists removed life itself, they extracted the life out of “Biology”, the science of life.

Now let us speak about hidden evolution, the arcane narrative of life on Earth through time. We first travel to Voronezh. Who here has been to Voronezh? All of the scenery, the landscape photographs of Voronezh in this presentation have come to me via Victor Fet, whose friend Dr. Viktor Golub sent them to him. Here for example are the old pines of “black earth Russia” (Figure 1).

So what is symbiosis? And what is symbiogenesis? What “symbiosis” is not is “cost — benefit” analysis, or “mutualism”. Such terms are not biology. They are not science, “cost” or “benefit” of life forms can not be measured. But symbiosis and symbiogenesis are sci — ence. “Symbiosis” refers to long term, permanent, sometimes cyclical, for example, sea — sonal, physical association between members of different species or other different taxa in general. To define “symbiosis” one ought to follow the original definition of the German scientist Anton de Bary (1831–1888). Symbiosis refers to a physical association between “differently named” partners, at least two kinds, that endures for most of the life history of the organisms. Or for at least most or all of the life history of one of them. Symbiosis is not a social relationship. Why not? Because social relationships refer to associations be — tween members of the same species. To be symbiotic a relationship must refer to members of different taxa. Symbioses are ecological relationships that, over a long period of time, may become symbiogenesis. In cases where new behaviors, structures or taxa, i. e., new tissues, new organs, new species, new genera, or even new phyla emerge, new relation — ships at many different levels can be identified as direcne cannot privilege one over other of the parts; all parts are required for the system to function. Kozo-Polyansky understood this when, in 1921, he was 30 or 31 years old.

Yes, Konstantin Sergeevich Merezhkovsky and Andrey Sergeevich Famintsyn, and Kozo-Polyansky are recognized as “symbiogeneticists”. These scientists are listed and described in the Russian encyclopedia as symbiogeneticists. They are unknown in the anglophone literature.

In our country we have Ivan Emmanuel Wallin who, of course, did not read Russian (Figure 3). Nearly no North Americans or English people read Russian. Please explain to them what I mean. Wallin (1883–1969) could not read Russian. But he had an immense knowledge of biology, so he invented his own words for the same phenomenon as Kozo — Polyansky’s “new principle”. Wallin came to the same conclusions of the importance of symbiosis in evolution as did Kozo-Polyansky, but entirely independently. And so Ivan Emmanuel Wallin, son of Swedish immigrants to the United States, wrote his wonderful book. He knew nothing about Merezhkovsky, Famintsyn, or Kozo-Polyansky. He knew no or very little Russian literature. Wallin’s idea (1927) he called “symbionticism”. He invented this neologism in the title of his great book on the origins of species. Wallin claimed that mitochondria evolved from symbiotic oxygen-respiring bacteria and that chloroplasts originated from blue-green algae, cyanophytes (called today cyanobacteria). Wallin had the same ideas basically as did Kozo-Polyansky. So these concepts came from two geographically independent sources. The ultimate geographical sources of symbio — genesis concepts came from the study of live organisms in Russia, St. Petersburg, the Mediterranean coast (Villefranche-sur-Mer; Laboratoire Arago; Banyuls-sur-Mer; Stazi — one Zoologica di Napoli), the marine laboratory at Roscoff and the Channel Islands of the

Atlantic, the Marine Biological Laboratory at Woods Hole on Cape Cod, and Wallin’s studies in New York and in the Colorado Rockies and later symbiosis studies on lichens, on corals at the Great Barrier Reef in Australia, and in the Caribbean.

Here we see a shrimp that cleans the teeth of a moray eel (Figure 4). This is symbiosis, but is it symbiogenesis? No. Why? This shrimp picks the teeth, eats debris in the toothy mouth and lives with the eel. Although this particular species of shrimp is not ingested by that eel, the eel eats other kinds of shrimp. It is symbiosis, but not symbiogenesis. Both partners grow and reproduce separately. Both shrimp and eel can live separately. One sees no obvious novelty generated by this symbiosis, i. e., symbiotic physical association. The relationship be — tween the shrimp and the eel is only feeding behavior. Symbiogenesis starts with at least two independent different taxa. Life is difficult, as I don’t have to tell you. Life is probably more difficult in St. Petersburg than it is in Massachusetts. Anyway, it is sometimes exceedingly difficult. Often life is intolerably hard and difficult. And occasionally symbiogenesis provides the solution (Figure 5). Symbiogenesis is evolution, not just ecology.

It is estimated that on Earth today are 250,000 protoctist species. Among them are representatives of the nucleated ancestors of animals, fungi and plants. My thesis, Kozo — Polyansky’s thesis, and Wallin’s thesis, persistently ignored, for the most part, by Anglo — phone biologists, is that your ancestors and mine evolved from bacteria by symbiogenesis in the Proterozoic eon 2,500 to 542 million years ago. Why do we claim that the first nucle — ated cells evolved in the Proterozoic? Because of the presence of fossils called acritarchs, from a few microns to some over 500 micrometers in diameter. No one knows exactly what they are, but everyone who studies these robust microfossils, mainly by palynological tech — niques, agrees that microfossils called acritarchs are eukaryotes. They are fossil nucleated organisms that began to appear in the geological record more than 1,000 million years ago.

Now, this cell, very familiar to all of you, is an animal tissue cell in mitosis. Mitotic cells contain at least one nucleus and the proteinaceous microtubules of the spindle and other cytoskeletal structures. These features are entirely absent in prokaryotes, both bac — teria and so-called “Archaea”. Here we see labeled in blue through fluorescence micros — copy the microtubular “end-binding protein” (Figure 6). The reddish-yellow and yellow signal is from microtubule protein absolutely typical of nucleated cells. This class of pro — tein underlies cell motility seen at the level of the light microscope. Through the optical microscope one observes intracellular movement. Some prokaryotes, that is certain bac — teria may be huge, 80 micrometers in diameter, but inside them one never sees what I call “anima”, “animation”, or any movement like this.

I want to show you now the power of symbiogenesis. We begin with amber, which under certain environmental conditions fossilizes very well. Here is a fossil termite, Mastotermes electrodominicus, that preserved so well that in the amber fossil record Da — vid Grimaldi and his colleagues (American Museum of Natural History, New York City) saw fossil muscle tissue at the light microscopic level! Fossil termites in amber were thin sectioned with a microtome. The investigators visualized the 20-million-year-old muscle tissue of the Mastotermes insect. So, Grimaldi said to us, since we see details of muscle tis — sue, why can we not also seek the termite’s intestinal protists? And we did! We not only see protist fossils in Mastotermes, but we found the oldest spirochetes (and other bacteria, for example bacillus spores) in the fossil record embedded in ancient amber. This had been collected from a mine in Central America, Dominican Republic by the museum director, Dr. Jorge Wagensberg and others (Wier et al., 2002). This specimen I show you here of M. electrodominicus, because it is the best preserved of any mastotermitid in the world, is on

exhibit at the Museum of Science in Barcelona, Spain. However, in less valuable specimens of amber of M. electrodominicus, when cut and imaged the termite microbes are seen.

Now I show you a video of Mixotricha paradoxa, a unique termite protist that lives only inside Mastotermes darwiniensis in the southern hemisphere.

We take a trip to Kakadu Park in northern Australia, near Darwin. The climate there is very hot. The extremely hot climate may have persisted for 15 to 20 million years in this region, continuously unbearably hot. And here we see Mastotermes termites. Inside those termites, in 1956, Professor L. R. Cleveland photographed the organism you see here on 16-mm. black and white film (Cleveland, Grimstone, 1964). He had read that this Mixotricha paradoxa “protozoan” bears both cilia and flagella on its same cell. He did not believe that. After years of study he had never before seen this phenomenon, “cilia and flagella” on one-and-the-same cell. We re-photographed Mixotricha paradoxa many years later. From the 1970s until this century we intermittently studied this paradoxical pro — toctist. And we are now sure that these beating “hairs” are not cilia. They are spirochetes, as Cleveland and Grimstone (1964), first reported on the basis of Grimstone’s superb electron micrographs. The tiny surface spirochetes of Mixotricha are morphologically in — distinguishable from Treponema pallidum, spirochetes of syphilis (Margulis et al., 2009). This, in other words, is a motility symbiosis. The large protist is fringed with 250,000 ec — tobiotic spirochetes embedded in its cortex by attachment sites. Toward the cell’s posterior a different, medium-sized spirochete, is present. It is morphologically indistinguishable from Borrelia burgdorferi. And the cell surface also has the little teeny ones. The mov — ing fringe here, as seen with the electron microscope, on nearly all the anterior surface of Mixotricha, is covered by the treponemes, and by other symbiotic bacteria. There are thus 250,000 treponemes, just after cell division, and before the next division. Perhaps

500,000 treponemes per protist cell. Some half-million attached spirochetes confer motil — ity on the Mixotricha paradoxa. A scanning electron micrograph of Mixotricha’s surface is seen in Figure 7. Five different kinds of spirochetes are physically associated with the protists (Wenzel et al., 2003). In the posterior ingestive area are other endosymbiotic bacteria. Termites can eat wood, but they cannot digest it. Approximately nine different kinds of prokariotic microbes comprising a single protist. One M. paradoxa cell is unites a half a million individuals. Without the symbiotic associates, no termite digests wood. Here you see the large spirochete Canaleparolina, the treponema spirochetes, the protist with a large piece of wood inside. When we say here is “one individual”, you Neodarwin — ists, where is your head? I assume you are not laughing because you don’t understand me? This “single, individual cell” is composed of at least nine different kinds of cells, each with its own genome. This, of course, is a group. And the animal in which this group lives is a termite worker. The termite insect starves within about two weeks if we remove its bacteria and protists. The isopteran dies on the second or third week. But Cleveland showed that if bacteria and protists are injected back into the intestine of the moribund termite the starving insect recovers completely. What, then, is the “individual” here? It is a community. Of course “group selection” occurs!

We have reconstructed the entire evolutionary story of eukaryosis. We have now ob — served every step in the origin of the cilium (Wier et al., 2010). Our hypothesis is that cilia, undulipodia in general, evolved from living spirochetes. Spirochetes donated many genes for crucial metabolic proteins to eukaryotic nuclei (Hall, 2010). The spirochetes form attached round-body (RB) propagules (Brorson et al., 2009). Spirochetes, of course, are gram negative eubacteria (Margulis, Chapman, 2010). They constitute a phylum; they

are pleiomorphic and are totally viable not only in the typical “cork screw” (spiral or heli- cal) swimming stage but also in the slower round-body (RB) form. The RBs (also called vesicular or cystic) are induced by penicillin and other “unfavorable conditions”. Sush as environmental changes i. e., quantity of free oxygen, elevated temperatures, cold, desicca — tion, altered salt concentrations and many chemical or medium alterations (e. g., notably in carbohydrate and other fermentable food source, amino acid composition, antibiotics induce the spirochete RB resistant stage. The Brorsons (2009) proved that RBs are entirely viable and reversible to the more active typical swimming helices. Also the discovery of Spi — rochaeta coccoides from the intestine of the kalotermitid (dry and dampwood-ingesting ter — mite) from Neotermes castaneus (Dröge et al., 2006). S. coccoides feeds osmotrophically, its population maintains and grows in the anoxic or micro-oxic paunch (specialized hindgut) of this subtropical damp wood termite from Florida, Caribbean islands and coasts and other warm, humid North American localities. Nitrogen-fixing bacteria abound in Neotermes cas — taneus, a fact that may be related to the obserbation that Spirochaeta coccoides, in culture does not convert to the typical helical swimmer stage. S. coccoides has not, to our knowl — edge, been reported elsewhere.

Please let us look again at Figures 7 and 8. The epibiotic spirochetes seen here on this trichomonad Mixotricha paradoxa are not cilia even though they look remarkably like cilia and other undulipodia of eukaryotes. Nor are they any other kind of undulipodial homologue. They are analogues as discussed by A. Wier (Wier et al., 2010).

Why are so many termite microbes motile? Because if the microbes do not continu — ously move up the intestine, anteriad, towards the mouth, they will be excreted with the feces. Might you tell them please what I mean, Galina? Many amitochondriates, mostly parabasalids seem to avoid and escape oxygen by remaining centrally located in the in — sect gut. Many must be motile all the time. They must swim up the intestine. They have to either be attached, or they have to swim. Otherwise they will be flushed into the soil. So many, many times, spirochetes alone, spirochetes attached to others, or spirochetes attached to intestinal epithelia evolved the ability to remain inside the insect’s anoxic, warm, wet, nutrient-rich intestinal habitat.

Although this resembles a cilium with a basal kinetosome, it can not be. In this micro — graph the “n” is the nucleoid of the spirochete. There are no missing links here as Kozo — Polyansky said in his 1921 All-Russian Botanical Congress abstract. This is seen as an analogous step in the origin in cilia from spirochetes (Figure 8, see Wier et al., 2010). We have overwhelming evidence that symbiogenesis is the basis of novelty in organisms composed of nucleated cells (Margulis, 1993.) In the earliest symbioses of eukaryotes we hypothesize that intracellular motility, “anima”, was acquired as symbiotic spirochetes by an archaebacterium (Hall, Margulis, 2010).

“Eukaryosis” refers to the process of evolution that formed the earliest eukaryote. The earliest eukaryotes are represented by archaeamoebae, metamonads, and parabasa — lids. None have mitochondria. They still live in anoxic environments today. Most amito — chondriates dwell in environments with low concentrations of oxygen, often with high amounts of sulfur.

A bacterial consortium in sulfur-rich environments was discovered by Boris Perfiliev in the Crimea by 1932. It was later studied by his student Professor Galina Dubinina. She is my age. When first observed Perfiliev called what he thought was one single organ — ism, the genus “Thiodendron”. The name in Greek means “sulfur tree”. Dubinina discov — ered that “Thiodendron” is a spirochete consortium. The spirochete components swim

towards sulfide, or they die. They must have sulfide. These partner organisms produce and release sulfide. They are “sulfidogenic”. They make sulfide. Although described by Perfiliev and his students in work between the 1930s and the 1970s, they never real — ized that they had a consortium of two kinds of bacteria at once. The earliest eukaryotes evolved in anoxic, sulfidic, organic-rich marine or estuarine conditions that were typical of the Proterozoic eon. The “ancestral spirochete” co-descendant is now alive in culture in the Moscow collection, and has recently been deposited in the Braunschweig, Germany culture collection. The multi-authored paper (Dubinina et al., 2010) is finally accepted and published. The first of these spirochetes, attracted by sulfide, is from Staraya Russa, Novgorod Oblast, Russia, is named Spirochaeta perfilievii. The Dubinina team has been working over 20 years on them. They have other related spirochetes to name.

Sulfurous environments are key. The Dubinina spirochetes swim toward sulfide, they are chemotactic. They use O2, ambient oxygen, oxygen in the air, to protect themselves from oxidation damage. They are oxygen-tolerant to the extent that they always need sul — fide. They oxidize the required sulfide to elemental sulfur. I’ve seen them for 30 or 40 years, without knowing what they were; here they are in nature (Figure 9). The white scum is what they first called “Thiodendron latens”. The seaweed is Fucus vesiculosus. The spiro- chete-sulfidogen consortia that Perfiliev discovered is in white in this photograph. Dubi — nina’s team has found them in at least six places: in Pacific Ocean islands, in the White Sea, and the sulfur springs resort, Staraya Russa. This is a place where F. Dostoyevsky lived. Dubinina has found them there, and in other sulfurous environments. They apparently occur all over the world in massive, geochemically important quantities. When Moscow University received their Japanese electron microscope for the first time, Dubinina studied these organisms under strict anoxia, no oxygen: fewer than ten to the minus ten parts oxy — gen. The long stringy filaments broke up. Out swam classical “1:2:1” and “2:4:2” spirochetes. In higher amounts of ambient oxygen (<0.01–2%) this spirochete grows longer and longer and longer with no change in width. Apparently S. perfilievii waits for anoxia in which the sulfide causes the oxygen-induced filament, with no change in width, to break up again to its typical 10–15μm length. This spirochete or its very close relative, if we are correct, shares a common ancestor with the sperm tails of half the people in this room.

Here I show you the animation part of our video called “Eukaryosis”. The atmosphere concentration of oxygen rose because of cyanobacterial photosynthesis. The association be — tween motile spirochetes and sulfidogenic archaebacteria became permanent. A merger oc — curred, and a stable association followed when the eubacterium DNA recombined with the spirochete’s DNA. This “permanent conjugation” of Dubinina’s spirochete occurred with Thermoplasma or Sulfolobus like Dennis Searcy’s thermoacidophilic archaebacterium. The association of the eubacterial sulfide oxidizer and the archaebacterial sulfidogen evolved to become karyomastigont system of eukaryotes called the “last eukaryotic common ances — tor” (LECA), (Margulis et al., 2006). The karyomastigont itself-nucleus “connector” and undulipodia evolved into what became the mitotic spindle. Oxygen concentration was still rising in the mid-Proterozoic eon, and sulfide was decreasing in many local habitats. Mito — chondria were symbiotically acquired only after “eukaryosis”. The karyomastigont-mitotic spindle was already present in anoxic conditions in amitochondriates. The mitochondria begano-spacerun:yes’> are oxygen-tolerant to the extent that they always need sul — fide. They oxidize the required sulfide to elemental sulfur. I’ve seen them for 30 or 40 years, without knowing what they were; here they are in nature (Figure 9). The white scum is what they first called “Thiodendron latens”. The seaweed is Fucus vesiculosus. The spiro- chete-sulfidogen consortia that Perfiliev discovered is in white in this photograph. Dubi — nina’s team has found them in at least six places: in Pacific Ocean islands, in the White Sea, and the sulfur springs resort, Staraya Russa. This is a place where F. Dostoyevsky lived. Dubinina has found them there, and in other sulfurous environments. They apparently occur all over the world in massive, geochemically important quantities. When Moscow University received their Japanese electron microscope for the first time, Dubinina studied these organisms under strict anoxia, no oxygen: fewer than ten to the minus ten parts oxy — gen. The long stringy filaments broke up. Out swam classical “1:2:1” and “2:4:2” spirochetes. In higher amounts of ambient oxygen (<0.01–2%) this spirochete grows longer and longer and longer with no change in width. Apparently S. perfilievii waits for anoxia in which the sulfide causes the oxygen-induced filament, with no change in width, to break up again to its typical 10–15μm length. This spirochete or its very close relative, if we are correct, shares a common ancestor with the sperm tails of half the people in this room.

Here I show you the animation part of our video called “Eukaryosis”. The atmosphere concentration of oxygen rose because of cyanobacterial photosynthesis. The association be — tween motile spirochetes and sulfidogenic archaebacteria became permanent. A merger oc — curred, and a stable association followed when the eubacterium DNA recombined with the spirochete’s DNA. This “permanent conjugation” of Dubinina’s spirochete occurred with Thermoplasma or Sulfolobus like Dennis Searcy’s thermoacidophilic archaebacterium. The association of the eubacterial sulfide oxidizer and the archaebacterial sulfidogen evolved to become karyomastigont system of eukaryotes called the “last eukaryotic common ances — tor” (LECA), (Margulis et al., 2006). The karyomastigont itself-nucleus “connector” and undulipodia evolved into what became the mitotic spindle. Oxygen concentration was still rising in the mid-Proterozoic eon, and sulfide was decreasing in many local habitats. Mito — chondria were symbiotically acquired only after “eukaryosis”. The karyomastigont-mitotic spindle was already present in anoxic conditions in amitochondriates. The mitochondria began as oxygen respiring delta — or alpha-proteobacteria. The cyanobacteria are still laugh — ing. Cyanobacteria can survive nearly everything except they cannot live under acidic con — ditions. They “bleach” as their chlorophylls are destroyed. Our “Eukaryosis” film animation was made by James MacAllister, who is here today to videograph our session.

Now let us look at these cilia to see what these spirochetes have become. We do not know all the details, just most of them. I do not know Y. A. Vinnikov (1910–1987), but he wrote this wonderful book (1982) on the structure of vertebrate sensory systems. A constant in all of cell biology, the nine plus two [9(2)+2] microtubules comprise the undulipodia, they are always 0.25 μm in diameter. Why? Because they are evolutionary homologues. “Undulipodium” is the generic term for the eukaryotic structure that is usu- ally motile. The bacterial flagellum is unrelated. The eukaryotic undulipodium, intrinsi — cally motile, should never be called a “flagellum”. There are many, many different kinds of undulipodia. The flagella are bacterial structures only. They are not ever intrinsically motile. Motility in bacterial flagella is restricted to the basal rotary motor embedded in the cell (plasma) membrane. The bacterial flagellum is passive. The shaft is composed of a single, sometimes two proteins. Undulipodia are intracellular organelles bounded by the eukaryote’s plasma membrane. They are always inside the cell. Undulipodia include all cilia and many other organelles of motility. They are active along their length. They undulate. Intrinsically motile they are composed of at least 350 different proteins, many detailed in the cell biology literature. Perhaps undulipodia have as many as 1,000 pro — teins, not including membrane proteins. Tubulins, tectins, dynein, ATPase and many oth — ers are consistent components of all standard undulipodia. “Undulipodium” is a word from W. Hartmann’s German text book. The word “undulipodium” was used in a crazy A. P. Shmagina’s Russian book (1948). Shmagina opened the throats of dogs, studied their tracheal epithelia to watch their cilia. As the dog died, she watched the ciliary move — ment slow down. I hired a Russian translator who could not find the translation of the term “undulipodium” in any dictionary. We foreigners learned about the term “undulipo — dia” from Polish protozoologists and especially from the California scientist and author: Theodore Jahn. He wrote the popular book on how to identify the “protozoa”.

The ciliated retinal rod cells of the eyes of vertebrates have “nine two plus two” micro- tubular undulipodia, underlain at the base by the “nine three plus zero”, i. e., the [9(3)+0] kinetosomes. The retinal rod ciliary bases, the sperm tails, the epithelial cilia, the algal, the fern and Ginkgo sp. tree sperm tails, all are entirely homologous. They are all examples of undulipodia. They are also present as extensions in cone cells of vertebrate eyes. (The “rods” and “cones” are simply infoldings of the ciliary membranes. The olfactory (smelling) processes are also undulipodia. Here, too, the mechanoreceptor cells of lobster antennules are composed [9(2)+2] pattern of microtubules. The kinocilium of the inner ear and most other sensory cells are modified cilia, as Vinnikov realized. The term undulipodium was introduced into Russian literature but our Anglophone literature refuses to use it. Most people still call these cell sensitive structures and sperm tails “flagella”. Please don’t. They are not flagella. They are undulipodia. “Undulipodium” is singular, undulipodia is originally a mixed Latin and Greek word: undula (wave); podium (foot). The so-called “flagella” of eukaryotes, sperm tails, na — sal epithelial cilia, gravitational sensors, balance organs, taste bud cilia, fish lateral line cells, mechanoreptors of insects and the antennules of lobsters that show microtubules at their dis — tal ends (nine times two=eighteen plus two=20 microtubules in total); these are derived from undulipodia. The antennules evolved as amplification systems based on the microtubules of undulipodia. The sperm tails of aquatic plants such as mosses and liverworts are also [9(2)+2] microtubule motile structures. Scientists agree, based on electron microscopy, that undulipo — dia, limited to eukaryotes, are all evolutionary homologues.

I have hypothesized that all undulipodia are homologues modified from the original ancestral attached spirochete. I continue to say they are evolutionary homologues whose

ancestor was and still is a descendant of a es’> eukaryote we can imagine. It is an has no mitochondria: the medium in which it grows here is anoxic. It has undulipodia. It eukaryote with a single nucleus as part of the karyomastigont system and it grows here in a culture of spirochetes. Why? Because both this tiny mastigote protist and the spirochetes thrive in low oxygen, rich organic mud with an abundance of sulfide gas. We posit that this was the early environment for eukaryotes. Today the same marine habitats support spirochetes and mastigotes similar to this protist. In this tube from sulfide-rich organic mud nearly all swimmers are spiro — chetes of different kinds, except this mastigote. It is a eukaryote, living as a “contaminant” in the mixed spirochete culture. Everywhere in the world 2,000–1,000 million years ago, most likely, this sort of habitat easily could be found.

On the anterior portion of this next protist (Deltatrichonympha, also from masto — termitid termites) are “waving hairs”. They are [9(2)+2] undulipodia, whereas on the posterior portion, here they are spirochetes. Without electron microscopy the details of this Deltatrichonympha cell are hard to discern. The undulipodia and the free-swimming spirochetes are difficult to distinguish. Spirochetes became undulipodia. They began as separate organisms. Sometimes they attach to each other or to different other life forms without modification. Sometimes they are modified. We hypothesized the steps in the at — tachment of spirochetes as they evolved into cilia before we found the analogue (Fig. 10 in Wier et all, 2010). Sometimes Spirochetes may enter cells to which they attach.

Sometimes they are inside other cells permanently. These associations still occur now. Some became permanent attached forms over and over again. Only one type probably is our ancestor. They are fermentors. So each of the steps in evolution exists today in envi — ronments low in oxygen, usually high in sulfide, not always, but always high in organic food. Here we have one of those wood-ingesting insects, termites.

What is happening in this video? Let us look closer at the same insect protist association (Staurojoenina in Neotermes) a little later in the day. We videographed that ten years ago but only recently found in electron micrographic sections that here are five different kinds of spirochetes living inside the protist. We don’t know what they’re doing, but we know that they are spirochetes living inside a vesicle in a single protist cell in an intestine of a termite (Wier et al., 2007). The cilia, née spirochetes evolved to become first sensory cilia after they permanently attached to the sulfidogen in the formation of the karyomastigont. The term “karyomastigont” was invented by С. Janicki in 1915. The karyomastigont comprises the nucleus, the proteinaceous nuclear connector, the kinetosome-centriole and its undulipo- dium. The karyomastigont is an organellar complex. We claim it is the evolutionary prod — uct of the DNA of both sulfidogenic archaebacterium and spirochetal eubacterium partners. The karyomastigont became the mitotic spindle, as it still is in Ochromonas (Figure 10).

Edouard Chatton’s drawings show the karyomastigont with its nucleus, nuclear con — nector, three forward, one trailing undulipodia. During division, the karyomastigont be — comes the mitotic spindle. No one thinks of karyomastigont-to-spindle transformation as an evolutionary phenomenon even though this organellar system permeates the protistological literature. Now, Chatton could not publish these charts, the ones he made for his students. Some are on display in the City Museum of Perpignan in southwest France. The gorgeous charts show too many different types of organisms for a professional journal. But Chatton very well knew the relationship between the karyomastigont nucleus and doubling of the kinetosomes and the formation of the thin mitotic spindle (called a parades mose). Chatton depicted the relationship between the motile undulipodium and mitotic motility as a pro — cess. He knew they were intimately involved and related to the evolution and taxonomy of protist groups. I just added modern terminology to his teaching chart (Figure 11). They are called “course boards” in the literature (Soyer-Gobillard, Schrével, 1986).

The relation between motility, mitosis and symbiogenesis underlies the phyla of the protoctista. We have estimated that there are 36 extant phyla in Kingdom Protoctista (Margulis, Chapman, 2010). That motility, locomotion, and mitosis as cell processes are examples of the same kind of intracellular motility was recognized perfectly well by Chat — ton. A study of these Darwin’s “oddities and imperfections”, not at all typical of animals, show how evolution must be reconstructed from clues. Karl Belar in 1933 in the alga of genus Ochromonas showed how, when the cell divided in mitosis, the karyomastigont became the mitotic spindle.

Here is a surf clam egg. Watch it carefully, please. Here is its “germinal vesicle”, that means “nucleus”. Let us see it again. Germinal vesicle, nucleus, nucleolus, and polar body

(Alliegro, and Alliegro, 2008; Alliegro et al., 2010). Please wait. Nucleus (≡ “germinal

verticle”), nucleolus and the little sphere. The little sphere is the nucleolinus. What is

a nucleolinus?

The “nucleolinus” is an organelle that contains nucleic acids of the kinetosome-cen-triole centrosome system. The nuclear membrane breaks down. The nucleolus in the surf

clam (Spisula) also breaks down periodically. And the nucleolinus is interpreted by Al-liegro (Alliegro et al., 2010) to be the portion of nuclear DNA that in the cytoplasm that

releases centriolar RNA. If we are correct, the legacy should be of the original spirochetes. The recent work on surf clam egg centrosomes by Mark and Mary Anne Alliegro and their colleagues is spectacular (Alliegro et al., 2010).

Each gray triangle represents a symbiotic merger (Figure 12). Some species of amito — chondriates are meiotic. Not many. Mitosis evolved before meiosis. Meiosis evolved as a variation in mitosis probably in response to seasonal environmental cycles. These pro — cesses came after the origin of the karyomastigont, i. e., after eukaryosis. Both mitosis and meiosis evolved at least once before the symbiotic acquisition of mitochondria.

Here I show you here how free-swimming spirochetes act. Just as undulipodia do in amoebomastigote transformation, they actively pull themselves in beneath their mem — branes. Spirochetes produce viable round body (RB) propagules. If we change the growth medium to “unfavorable conditions”, they can form RBs, survive for at least two years in this state, and later revert. First their outer membranes swell. Then they draw in their protoplasmic cylinders. The RBs are totally viable. They reform helical swimmers later when “favorable environmental conditions” are restored. At least twelve different types of spirochetes, free-living and symbiotic, show this behavior (Brorson et al., 2009). Here, once inside, is a spore-like structure produced by Spirosymplokos deltaeiberi. The “spore — like structure” is not heat resistant, but it is desiccation resistant. They are totally viable. Some spirochetes are light sensitive. We hypothesize the cytoskeleton evolved from free — living spirochetes like those of Dubinina (Dubinina et al., 2010). “Cytoskeleton” implies anima, animation, movement, and intracellular movement generally.

Only the bacterial groups that enter the evolutionary story of nucleated cells are de — picted at the bottom of Figure 12. A whole series of protist “oddities and imperfections” in the Proterozoic eon evolved as the oxygen concentration rose. Oxygen-breathing bacte — ria become the mitochondria, and almost all other eukaryotes evolve from mitochondriate protists. The gray triangle, upper right, indicates that photosynthesis that evolved in the bacteria was acquired by algae that evolved into plants. The triangles from bottom to top represent the first symbiotic union, the second symbiotic union, and the third symbiotic fusion. Fusions abounded: undulipodia first, mitochondria next, cyanobacteria precursors to photosynthetic plastids third. Undulipodia confer intracellular motility, mitochondria confer oxygen respiration, plastids provide internal cellular photosynthesis in eukaryotes.

And so where does Kozo-Polyansky come in? The abstract before he wrote his book was shown in Figure 2. He wrote that “nature does make jumps”, because the two or more components of a symbiosis are very different from the combined partnership. Apparently no one read his book. Or maybe they read it but it was not understood. It certainly was not translated. Apparently, from the quote written by Nina Bazilevskaya in 1959 two years after he died, the Russian botanists did not think it was worth translating. He was very young. He went on to become a successful professional botanist. This plant, Andro — sace kozo-polyanskii, is a primula, a primrose. You can see the arctic plant on our Harvard University Press cover of the translation of his book (Figure 13). He did much other work in botany, but as far as I know, he never returned to symbiogenesis after publication of this prescient, unique little book in 1924. Nobody listened to him. Nobody fought for his ideas. Nobody, perhaps in part because he did not write or speak German or English. How can anyone publish about evolution if he knows no English? One cannot pit oneself against all Neodarwinists.

Here we return to Voronezh, to the botanical garden named after Boris Mikhaylovich. Here we see high water in spring. Victor Fet and I explain Kozo-Polyansky’s work but we

didn’t change it. We think he was correct in nearly all his statements. We added a glossary and most importantly we show new photographs of live organisms and new electron mi — crographs of organisms to which he referred. We say it will be obvious to anyone who loves life on Earth that Boris Mikhaylovich Kozo-Polyansky correctly interpreted the evidence for symbiogenesis as source of innovation in evolution. (Figure 13.) The acquisition of genomes, especially of bacteria and other microbes, we show is the most important positive source of evolutionary novelty. The new illustrations show how prescient and correct was his work.

You may wonder how the birches survive the Russian winter. Perhaps bacteria make antifreeze compounds. Kozo-Polyansky did not use the word “prokaryote”. He referred to bacteria by many names: bioblasts, cytodes, flagellated cytodes, nepheloid and consor — tia. Chatton invented the useful term “procariotique”. He did not define it, he used it in a table and placed “cyanophytes” with prokaryotes and algae, including dinoflagellates, under the column “eucariotique”.

However, it was B. M. Kozo-Polyansky who explicity recognized that the tiny life forms were elementary units of the living. He knew well that they have all the properties of life: growth, reproduction, mechano-, chemo-, and photo-sensitivity, motility, photo — synthesis. He understood what today we recognize as the bacterial group. He realized that bacterial elements are recombined and interact. They generate saltations, punctu — ated change in evolutionary lineages. Yes, he claimed, even in his 19 line abstract. Yes, “Nature does make jumps”. Evolution is not gradual as Darwin insisted. Species do not change by accumulation of random mutations as Neodarwinists as insist.

Here, what we don’t see is that in this Voronezh Lake Ugolnoe are the bacteria that run most of the biogeochemical system, as V. I. Vernadsky’s work in his book The Bio — sphere suggests. Thank you all very much.

I am sorry that I can only take questions in English. Please translate the questions for us.

QUESTION: Is symbiogenesis important to prokaryotic evolution, to prokaryotic speciation?

MARGULIS. Fine question. Yes, random mutation and gradualism does occur in prokaryotes. Some symbiogenesis happened, for example in Thioploca. But this was noth — ing like the rampant symbiogenesis in eukaryotes. Speciation itself, as a phenomenon, first evolved in eukaryotes. In prokaryotes overnight one rids the cells of plasmids, and the act of loss changes a species name (e. g., Bacillus anthracis to Bacillus cereus), or even, with the loss of plasmids for nitrogen fixation, induces a name change at the level of genus (e. g., from Rhizobium to Azotobacter!)

Plants, animals, fungi or protoctists don’t change species and genera by overnight treatments in ethidium bromide or a simple increase in temperature. Of course not.

New species in eukaryotes evolve primarily because ancestral ones acquire entire new genomes that, by many steps, integrate over geological periods of time. They generate closely related descendants as new lower taxa radiate. The best contemporary work on this now is by microbiologists and agriculturally-oriented entomologists.

Perhaps you know the older book of Professor Soren Sonea “Une nouvelle bacteriol — ogie”. His new book is called “Prokaryotology” (Sonea, Mathieu, 2000). He’s a Romanian who has lived in Canada, in Quebec, for his entire professional life. He and his colleagues show that the bacterial genetic system is worldwide. When a bacterium is cultured in

the laboratory, one deals with the “terminals” of the worldwide system of bacteria — the “terminals”, not the “computer” itself. In Sonea’s metaphor the computer is outside, the “mainframe computer” dwells in nature.

Bacteria have access to huge numbers of bacterial genes. Indeed they may access any gene of another bacterium by means of transforming-principle DNA extruded into the environment, transduction by phage, conjugation, plasmid transfer and other “small replicon” or “genophore” unidirectional bacterial gene transfer. So we either have one worldwide species of bacteria or no bacterial species at all.

The phenomenon of speciation, even though the words “species” and “speciation” are used as if they meant the same phenomenon, is a process only of eukaryotes. The jumps, the saltations, from prokaryotic components to composite “individuals” is a property of all eukaryotes. All have evolved symbiogenetically. Symbiogenesis is fundamental to all eukaryotic taxa from below species to above phyla (Margulis, Chapman, 2010).

Gradualism, the gradual accumulation of random mutations, ironically is more char — acteristic of prokaryotes. No one has ever shown, in the laboratory, field or fossil record, the veracity of that Neodarwinist mantra. That “gradual accumulation of random muta — tions” results in a passage from one to another, new and different, species has never been documented. By contrast, the phenomenon, rampant in eukaryotes: the symbiogenetic origin of new species has been detailed in much literature (Margulis, Sagan, 2002) ever since the tradition was begun in 1921 and ignored. It is mandated now, that those who wish to understand the evolution of life on Earth, even at this late date, start to read the wise words of the young Boris Mikhlayovich Kozo-Polyansky, native son of Voronezh (Kozo-Polyansky, 2010; Fet, Margulis, 2008). He knew, and we know now, that we live on an Earth more alive and more complicated than the Neodarwinists (or even the more humble of us) can describe or explain. Our world is a prodigy of its grand and profound past, its millions of years of natural history.

Acknowledgements

Drs. Mark A. and Mary Anne Alliegro, Celeste Asikainen, Dr. Andrej Victorovich

Bakhtiarov, Margo Baldwin, Oystein Brorson, Judith Herrick Beard, Dr. Michael J.

Chapman, Kendra Clark, Professor Michael F. Dolan, James MacAllister, Melishia San-tiago, Dr. Galina Fet, Dr. Anna Gorbushina, Professor Ricardo Guerrero, Dr. John Hall,

Dr. Liya N. Khakhina, Professor Eduard I. Kolchinsky, Professor Wolfgang Elisabeth

Krumbein, Professor Jan Sapp, Dr. Bruce Scofield, Dr. M. O. Soyer-Gobillard, Professor

Alfred I. Tauber, Professor Andrew Wier, Dr. Marina V. Loskutova, Professor Michael

L. Williams, and especially Professor Victor Fet (Marshall University, West Virginia).

I express my deepest gratitude for the indispensable (but only partial) financial aid that

came from the Alexander von Humboldt Stiftung (Berlin), The Eastman Distinguished

Professorship (Balliol College of Oxford University, The Tauber Fund, the NASA Plan-etary Biology Internship (Woods Hole, MA) and The Graduate School and the College

of Natural Science at the University of Massachusetts-Amherst.

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Симбиогенез. Новый принцип эволюции:

восстановление приоритета Б. М. Козо-Полянского (1890–1957)

Линн Маргулис

Университет Массачусетса

Амхерст, США: celeste@geo. umass. edu

Публикуемый текст представляет собой отредактированную запись моей лекции, включавшей 14-минутный видеофильм (с двухминутной анимационной моделью) о развитии гипотетических организмов на всех этапах происхождения ядерных клеток от бактерий («эукариоз»). Наши наблюдения, подтвержденные новыми современны — ми примерами, демонстрирующими этапы развития живых организмов, практически наверняка доказывают правоту «нового принципа» Б. М. Козо-Полянского (1924) о важности симбиогенеза в процессе эволюции жизни на Земле в течение не менее двух миллиардов лет. В будущем широко известное, но недокументированное объяснение происхождения эволюционных новшеств путем «постепенного накопления случай — ных мутаций» станет считаться ошибочным предположением начала XX в., которое в основном пропагандировалось англичанами, североамериканцами и другими англо — говорящими учеными. Эти неодарвинистские «объяснения» уступят место подробно — стям симбиогенеза, таким как генетическое слияние, в особенности видообразование через приобретение геномов, кариотипическое расщепление (образование неоцен — тромер и связанное с этим изменение хромосом), концепция ларвального переноса у животных (Д. И. Уильямсон). Работа Козо-Полянского, при его жизни подвергшаяся осмеянию и забвению, приобретет такое же значение, какое имели в начале XX в. рабо — ты Менделя по наследованию признаков у гороха для открытия генетических принци — пов развития диплоидных организмов. Мое выступление, фотографии и фильмы были представлены на Дарвиновской конференции в Санкт-Петербурге 23 сентября 2009 г.

Ключевые слова: симбиогенез, эукариоз, Козо-Полянский, видообразование, гене — тическое слияние, приобретение геномов.

Материал взят из: Чарльз Дарвин и современная биология. Труды Международной научной конференции (21–23 сентября 2009 г., Санкт — Петербург)