Once Upon a Time Lynn Margulis by EstudiPuche

In memory of Lynn Margulis

(1938-2011)

[Previous page] Puffer's Pond, in North Amherst (Massachusetts), was one of Lynn's favorite places.

The cover composition tries to evoke Lynn Margulis, absent, in an environment close to her. Puffer's Pond, where she used to swim during most of the year, even in cold temperatures. Her dog, who used to accompany her. The bike, her favorite transport and in the background, the extraordinary hues of autumn in New England. On the back cover, the same environment with a beaver that could often be seen around.

Mercè Piqueras Wendy Ran Sandra Young Graphic design

Jordi Rabascall Madrid (Estudi Puche S. L.) © of this edition Editorial Septimus c/ Rajoleria, 3 (08472) Campins First edition: October 2013

ISBN: 978-84-940179-3-3 D.L.: B.21068-2013 Printing

Vanguard Gràfic S.A. c/ Joan Miró, 19 08930 Sant Adrià de Besòs, Barcelona This work is licensed under a

Creative Commons Attribution-NoDerivs 3.0 Unported

contents

Thinking about Lynn. Moselio Schaechter . . . . . . . . . . . . . . . . . . . . . . In search of truth. Ricardo Guerrero . . . . . . . . . . . . . . . . . . . . . . . . . . . Teaching for science, teaching for life. Montse Vallmitjana . . . . . . . Beyond the sea. Marta Estrada . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Looking at microbes. Isabel Esteve . . . . . . . . . . . . . . . . . . . . . . . . . . . . Engaged to science, married to life. Ricardo Amils . . . . . . . . . . . . . . Life is symbiosis. Eva Barreno . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A world of microbes. Ramon Folch . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lynn Margulis, my inspiration. Mercè Piqueras . . . . . . . . . . . . . . . . Let’s listen to the universe. Eduard Punset . . . . . . . . . . . . . . . . . . . . . . Nature and nurture. Mónica Solé Rojo . . . . . . . . . . . . . . . . . . . . . . . Teaching in the rain. Martí Domínguez . . . . . . . . . . . . . . . . . . . . . . . Swimming with bacteria. Núria Gaju . . . . . . . . . . . . . . . . . . . . . . . . . . Bitten by the science bug. Laura Villanueva . . . . . . . . . . . . . . . . . . . . . A woman who left her mark. Rubén López and Concepción Ronda . . Not gone with the wind. Begoña Vendrell . . . . . . . . . . . . . . . . . . . . . . . Sharing knowledge, sharing life. Jordi Mas-Castellà . . . . . . . . . . . . A groundbreaking. Balbina Nogales . . . . . . . . . . . . . . . . . . . . . . . . . . . A woman for all seasons. Carles Pedrós-Alió . . . . . . . . . . . . . . . . . . . . The living experience of natural history museums. Alícia Duró i Sans Gaia and more. Héctor Ruiz . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . My dear Lynn. A humble and late eulogy. Xavier Maymó-Gatell . . Let’s go to Ithaca. Gemma Reguera . . . . . . . . . . . . . . . . . . . . . . . . . . . . Members of all the kingdoms. Juli Peretó . . . . . . . . . . . . . . . . . . . . . . The art of storytelling. Nicole Skinner . . . . . . . . . . . . . . . . . . . . . . . . . The ins and outs of life. Carmen Chica . . . . . . . . . . . . . . . . . . . . . . . . .

9 13 29 35 43 49 55 61 67 73 81 87 93 99 105 111 115 121 127 131 137 143 151 161 169 175

About the authors and collaborators . . . . . . . . . . . . . . . . . . . . . . . . . . . .

187

thinking about lynn Moselio Schaechter

San Diego State University, San Diego, CA, USA

Las honestas palabras nos dan un claro indicio de la honestidad del que las pronuncia o las escribe.

Honest words give us a clear indication of the honesty of the one who pronounces or writes them.

Miguel de Cervantes (1547-1616)

Lynn Margulis changed the way we think about evolution, our cells, ourselves. To her we owe the now universal acceptance of the symbiotic origin of the eukaryotic cell. Seldom even among the scientific notables do we find an insight of such weightiness. Notice that I call her achievement an insight, not a discovery or an invention. It was not the result of direct experimentation, rather a realization that emerged from her deep thinking and erudition, an understanding that she then put forward vehemently. And therein lies the tale. How did this come about? If you seek historical details and specifics, I would refer you to Jan Sappâ&#x20AC;&#x2122;s chapter in the book on Lynn edited by Dorion Sagan. Here I will sketch a brief picture of the unique endosymbiosis that brought about this achievement, a symbiosis between Lynnâ&#x20AC;&#x2122;s excellence as a geneticist/cytologist and her fervent personality. Lynn trained under a distinguished cytologist at the University of Wisconsin, Hans Ris, whose career was anchored in both the 19th century era of hugely detailed protist cytology and [ 9 ]

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the modern biological revolution. In such an atmosphere, and Lynn being Lynn, she became steeped in cytological learnedness, both of the old and the new. She gravitated to the elaborate and beautiful world of protist biology and became an expert in its intricacies. From early on, she was aware of the proposal of Ris and others that organelles such as mitochondria and plastids were of bacterial origin. However, in their advocacy they did not fully use the tools that conveyed the implications of this process for evolution, one could say its evolutionary immanence. The notion that the eukaryotic cell, replete with its organelles, arose through symbioses required such a huge mental leap that most people were refractory to its acceptance. Somehow, people needed a proverbial hit over the head, and Lynn delivered just that with her classic 1967 paper in the Journal of Theoretical Biology. Its message took some time to penetrate, but penetrate it did, and with a vengeance. Lynn did not stop here. She reached further and saw symbiosis â&#x20AC;&#x201D; previously treated as an interesting eccentricity of Nature pertinent to lichens, the roots of legumes, or the rumens of cowsâ&#x20AC;&#x201D; as the main force in evolution. At the time, this was blasphemy against the neoDarwinian dogma that evolution proceeded by the stepwise selection of single mutations. In Lynnâ&#x20AC;&#x2122;s powerful vision, symbiosis provided that alternative. Nowadays, a testament to her impact, it is common place to consider evolution as modular, with important changes in a lineage arising (probably most of them early in evolution) by the acquisition of packets of genes. Such a packet has on occasion arrived, as she envisioned, in the form of the entire genome of a symbiont; more frequently, via other mechanisms that deliver smaller clusters of genes. Genes travel via viruses, through sex, or simply as naked DNA. However, so imbued was she with the symbiogenesis idea that [ 10 ]

thinking about lynn | moselio schaechter

she had little time for those other ways to transfer genes en masse. This passionate espousal of her view was also reflected in the ire with which she savaged Neo-Darwinism and its adherents.

Photo by Lluís Pou at his home in L’Astort, near Barcelon, Sept 12, 1992. From left to right Carl Rustenholz, Lorraine Olendzenski, Lynn Margulis, Ricardo Guerrero, Moselio Schaechter, Carmen Chica.

Now let me turn to Lynn as the person I knew, a friend with whom I had the opportunity to talk at length on numerous occasions through the years. Thinking about Lynn is not a simple matter. She was probably the most unusual person I have known, and she presented puzzling contradictions. For example, consider her understanding of science. On the one hand, she made a monumental contribution to biology, while on the other hand she championed views that were unjustifiable by the evidence, e.g., that AIDS is caused by a spirochete. How can achievements of gleaming clarity coexist in one brain with ideas that seem inexplicable to most? ¿Que pasa? [ 11 ]

once upon a time lynn margulis

I wish I knew. Lynn left us with only hints, elusive inklings. One trait that does stand out was the impassioned intensity of her convictions. Talking with her meant listening to her. One contradicted at one’s peril. On the rare occasions when I did, my comments elicited an avalanche of arguments and scientific facts, often peppered by pointed remarks about those who held views contrary to hers. It was not that she was uncaring or disrespectful. Rather, she was visibly carried away by the force of her argument. What deeper meaning can we read in her intensity? Perhaps, because her brain was so focused on developing and then defending her own primary insight that other ideas escaped close scrutiny. Lynn’s enchantment with the poet Emily Dickinson has been commented on by many. I believe that Lynn was a poet at heart. I contrast to the character in Molière’s Le Bourgeois Gentilhomme who realized that he had been speaking in prose all his life, Lynn may have known that she spoke in a poetic language. Her writings are handsome in a poetic sense, sometimes lyrical, sometimes dramatic. But, beyond that, her writing was imbued with candor and utter integrity. As in the words of Cervantes, the honesty of her writing gave proof to her inner honesty. I thank Merry Youle for her skillful and insightful editing of this contribution. 3

[ 12 ]

in search of truth Ricardo Guerrero

Institute for Catalan Studies, Barcelona, Catalonia

“The story of science, a continuous joint narrative, is never the personal tale of any scientist”, writes Lynn Margulis at the end of the prologue to her only fiction book, Luminous Fish. Tales of Science and Love (Chelsea Green Pub. Co., White River Junction, VT, 2007; originally published in Spanish in 2002 by Tusquets Ed., Barcelona). Since her death on November 22, 2011, several tributes have been paid in honor of Lynn’s work and surely there will be many more such events in the future. The contributions here aim to put into words the recognition and admiration of a few of her Spanish colleagues and friends, who consider themselves fortunate to have been part of the Spanish experience of this incredible woman. Certainly, Lynn was neither a perfect scientist nor a perfect woman, but she is a role model for all of us. Perfection does not exist and surely she would not have wanted to be perfect. She was tenacious in defending her ideas, provocative in the way she did so, and unorthodox in almost every aspect of her life. In my twenty-eight years of “coexistence” with her, both in professional and personal matters, what I certainly can add to this description is the integrity, the scientific convictions, and the overwhelming personality of this unique woman, housewife, mother, writer, teacher,... who was Lynn Petra Alexander (Chicago, IL, 1938; Amherst, MA, 2011). In the history and development of science, there are milestones linked to one or more scientists. Besides being the product of the [ 13 ]

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wit and personal work of someone, they also incorporate ideas and discoveries that others have previously achieved. Science is a cooperative task (â&#x20AC;&#x153;a continuous joint narrativeâ&#x20AC;?) that builds and modifies ideas, and leaves doors open to other new ideas to move forward. Lynn followed that way and left us a huge scientific production, her ideas and her commitment to what she believed and defended. The short texts collected here by Carmen Chica are contributions by colleagues and friends, many of them leading biology researchers, particularly in the field of microbiology, and to a lesser extent in other disciplines. All of them met Lynn and have tried to highlight the mark she left in all of them. So, there are occasionally repetitions in the texts because while authors (either senior or young) recount their work in a particular field of science, they also dive into their first meeting with Lynn. For the youngest, getting to know her was also a way to look towards a future devoted to science and to consider what that commitment meant. In this regard, Lynn highlighted the influence that some of her own teachers had had in her training. With them, the students worked, commented on articles of researchers and criticized them directly; it was a way to participate in their own education under the guidance of vocational teachers. So, the task of good teachers will never be overemphasized. A good teacher, the one that we will always remember, is the one that infected us with the contagious virus of curiosity. In the individual contributions of this book, there are also coincidences. This is because all authors had the opportunity to experience with Lynn the best of her: her communication skills; her extreme generosity (her house has been home for almost all of them); her perseverance to go forward despite the difficulties; her tireless way to [ 14 ]

in search of truth | ricardo guerrero

work (until exhaustion of herself and their collaborators); her sense of humor (using amusing puns both in English and in Spanish), her laboriousness (it was impossible to get bored with her). Although briefly, because this is not a scientific book, other frequent topics are the origin of the eukaryotic cell by serial endosymbiosis, the Gaia theory developed by James E. Lovelock, the origin and evolution of life. And more. Either reading her writings or listening from her lips, the long and arduous process that eventually led to the establishment of these theories was a unique experience. Lynn expressed the difficulty of the scientific community to accept changes; she expressed this feeling in a book in which she developed the idea of symbiosis leading to the evolution of nucleated cells and then to other organisms (fungi, plant, animals) when she wrote “many circumstances conspire to extinguish scientific discoveries, especially those that cause discomfort about our culture’s norms.” (The Symbiotic Planet. A New Look at Evolution, Weindenfeld & Nicolson, London, 1998). In the same sense are Thomas S. Kuhn’s words: “Concerned with scientific development, the historian then appears to have two main tasks. On the one hand, he must determine by what man and at what point in time each contemporary scentific fact, law, and theory was discovered or invented. On the other hand, he must describe and explain the congeries of error, myth, and superstition that have inhibited the more rapid accumulation of the constituents of the modern science.” (The Structure of Scientific Revolutions, The University of Chicago Press, Chicago, 1962). Apart from some explanation about the problem with changes in science, surely neither Lynn nor any other modern scientist (either woman or man) would consider politically correct to atribute discoveries and inventions only to men. But this has been a predominant idea for a long [ 15 ]

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time, and it does not detract from the great value of a book that in depth analyses science and its philosophical and historical foundations. In another part of Khun’s book we can read: “Normal science, for example, often suppresses fundamental novelties because they are necessarility subversive of its basic commitments. Nevertheless, so long as those commitments research ensures that novelty shall not be suppressed for very long.” Lynn never hesitated to mention the scepticism shown towards the theory of endosymbiosis by recognized scientists, including Niles Eldredge and Stephen Jay Gould (1941-2002), both of them good friends of hers. Perhaps she thought that the main problem with those and other relevant scientists was their misconception of seeing microorganisms mainly as pathogenic agents. For this reason, she admired and loved Lewis Thomas (1913-1993), the North American physician, essayist, educator, policy advisor, and researcher. In fact, Thomas, despite being a physician, with a real implication in clinical research, was willing to recognize the existence and the role of many non-pathogenic microorganisms. In his delightful book (which can be considered his autobiography) The Youngest Science. Notes of a Medicine-Watcher (The Viking Press, New York, 1983), Thomas stated (refering to some criticism he received on several articles): “Some of the evolutionary biologists criticize the suggestion, running through many of the essays, that the earth’s body represents a kind of organism, displaying so many instances of interdependency and connectedness as to resemble an enormous embryo still in the process of developing. This notion has seemed reasonable enough to me, considering the plain paleontological fact that the life on the earth was, for nearly 75 per cent of its existence made up entirely of separate, prokaryote, microbial cells, themselves the progeny of what may have [ 16 ]

in search of truth | ricardo guerrero

been, long ago, the single first cell […] The Gaia hypothesis, proposed by Lovelock and supported by Margulis, goes a step further to postulate that the conjoined life of the planet not only comprises a sort of organism but succeeds in regulating itself.” An intrinsic property of any scientific assertion is that it can be invalidated by the discovery of new facts or the change in some ideas. Carl Sagan wrote: “In science it often happens that scientists say, ‘You know that’s a really good argument; my position is mistaken,’ and then they would actually change their minds and you never hear that old view from them again. They really do it. It doesn’t happen as often as it should, because scientists are human and change is sometimes painful. But it happens every day. I cannot recall the last time something like that happened in politics or religion.” Turning again to Symbiotic Planet, we find Lynn comments on what being a woman firmly convinced to follow a scientific career implies. In 1957, when she was 19, she married astrophysicist Carl Sagan and she mothered with him two children (Dorion, born in 1959, and Jeremy, born in 1960). They got divorced and she later married chemist Thomas Margulis, with whom she had two more children (Zachary, born in 1967, and Jennifer, born in 1969). Nevertheless, in no way was she willing to adopt the only role of a housewife and to attend social activities consisting of playing cards, being a member of social clubs or participating in contests of apple pies. “I preferred—she wrote—the company of babies, mud, trees, fossils, puppies and microbes, to the normal world of adults.” In 1958, when she was 20, she published her first scientific article, a short one: “Incorporation of Thymidine in the Cytoplasm of Amoeba proteus,” by W. Plaut and L. Alexander Sagan (J. Biophysic. Biochem. Cytol., 1958, 4:843-846). Walter Plaut was an out[ 17 ]

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standing scientist that, four years later, in 1962, along with Hans Ris, his close colleague at the Zoology Department of the University of Wisconsin, Madison, reported a major discovery on cell structure. They (Plaut and Ris) found DNA in the chloroplast of Chlamydomonas, establishing for the first time that organelles might contain their own genetic information. Lynn had the opportunity to know both of them and surely became contaminated with their passion for science. It was while working in Plaut’s lab and earning a master’s degree in zoology-genetics that she was intrigued by the curiously independent nature of mitochondria, and later became a proponent of endosymbiosis—a theory first introduced in the 1920s by several Russian scientists—which would have a profound influence on all of her later research. At that time, she stated “I owe my passion for working on Saturday nights to the utter necessity of amoebae feeding and bowl cleaning every day.” It is well known—in fact she herself used to tell it—that in 1966 several editors of different journals to whom she submitted the manuscript of “The origin of mitosing eukaryotic cells”, noted the novelty of that hypothesis... but rejected it. She later admitted that it had been poorly presented and not correctly written. Finally the work appeared in the Journal of Theoretical Biology (14:225-274, 1967), under her name then: Lynn Sagan. Also the publication of an expanded version of her hyphotesis as a book suffered uncomfortable situations until it was published, in 1970, by Yale University Press, under the title Origin of the Eukaryotic Cell. Here Lynn discusses her early work on the symbiotic origin of cellular organelles, mainly mitocondria and chloroplasts. The preface, written by the worldwide-known ecologist G. Evelyn Hutchinson (1903-1991), contributed indeed to the quick impact of the book. [ 18 ]

in search of truth | ricardo guerrero

Lynn loved spirochetes and took them as a matter of study for a long time. As an example, in an interview, she said that her favorite organism was a huge spirochete identified for the first time at the Ebro Delta. She called it her “Catalan spirochete”, referring to Spirosymplokos deltaeiberi, that was described in an article published in 1993 in Archives of Microbiology (R. Guerrero, J. Ashen, M. Solé and L. Margulis: “Spirosymplokos deltaeiberi nov. gen., nov. sp.: variable-diameter composite spirochete from microbial mats.” 160: 461-470). Later on, in 1998, after having acquired more data about this spirochete, the authors were interested in establishing its possible distribution in microbial mats from other locations. The results of that work were published in an article by L. Margulis, A. Navarrete and M. Solé: “Cosmopolitan distribution of the large composite microbial mat spirochete Spirosymplokos deltaeiberi” (Int. Microbiol. 1: 27-34, 1998). The publication, International Microbiology, the official journal of the Spanish Society for Microbiology, was founded by Lynn and myself in that very year, and has nowadays a healthy life of sixteen years. These unusual spirochetes, previously reported only from the Alfacs Peninsula at the delta of the Ebro River in northeast Spain, contain striking arrays of cytoplasmic granules packed into their protoplasmic cylinders. Both in summer and in winter, the huge spirochetes were found also in mats growing in the Sippewissett salt marsh at Woods Hole, Massachusetts, as well as in microbial mats at North Pond (Laguna Figueroa, Baja California Norte, Mexico). The identity of these spirochetes was confirmed by electron microscopy: number and disposition of flagella, composite structure, measurements of their distinctive cytoplasmic granules. The granules, larger, more conspicuous and present in addition to ribosomes, are hypothesized to contain ATPases. As culture conditions worsen, these spi[ 19 ]

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rochetes retract into membrane-bounded round bodies in which they form refractile inclusions. From morphology and behavior, we concluded that the North American spirochetes from both Atlantic and Pacific intertidal microbial mats were indistinguishable from those at the Ebro Delta. We concluded a cosmopolitan distribution for Spirosymplokos deltaeiberi. Besides, the observation that spirochetes could resist long periods of desiccation (or other â&#x20AC;&#x153;lethalâ&#x20AC;? conditions) was a clue to a potential explanation of the intermitent life-course and strange behavior of a very well known bacterium, Treponema pallidum, the agent of syphilis. This book contains several friendly opinions on Lynn. In some cases, with Lynnâ&#x20AC;&#x2122;s own words from talks with friends, interviews, or excerpts of texts. In other cases, with comments on her, her work, her ideas, her personality, and the evolution of her own thoughts in her basic conception of life. In no case, however, we believe this can distort the meaning and significance of her message. Lynn was able to clearly express her thoughts and opinions about many thing in just a few words. On the many occasions on which she was interviewed for newspapers and magazines, Lynn clearly expressed their ideas and thoughts on microorganisms, evolution and related issues. For instance, she used to explain that evolution came from the recombination of microorganisms that formed a new living being. All living beings are made up of bacteria, included we humans. Ten per cent of our dry weight consists of bacteria. Homo sapiens is nothing special. Biologically, not very different from a chimpanzee. Although many people think that bacteria are bugs that we have to fight, there are many and different kinds of bacteria, and very few of them are pathogenic. Even these act only under certain environmental conditions. When [ 20 ]

in search of truth | ricardo guerrero

the interviewer tells Lynn that she speaks with more fascination about microorganisms than about humans, she answers: “…I am not at all interested in human biology. We behave like a mammal pest, trying to destroy the environment. Fortunately, nature has proven much more intelligent than us [...] And even though humans are applied to pollution and annihilation, the Earth, which is an ecosystem with its own physiology—a kind of living organism—has sufficient resources to defend itself against these attacks.” 1 In another occasion Lynn said: “I refuse to do what people tell me to do. That’s always been my problem. The critics say I need proof by molecular biology… but classical biology is perfectly good science, just obscure. To make a contribution to evolution, one has to be broad. People tried to tell me what science to do, and I don’t do what they say.” 2 The Catalan journalist Núria Escur asked her once what was the most beautiful thing she had seen through a microscope, and Lynn’s answer was: “Microbes!, but people do not understand this and they make products to kill them. And by doing this, they are killing themselves…” 3 In another interview and in relation to the spread of intelligent design (ID) or creationism, Lynn said, “evolution is a fact as proven as gravity. As much as I want to deny it, science is always more powerful than any dogma.” 4 Finally, I remember that, in a lecture at the Institute for Catalan Studies and referring to the “hidden evolution”, she said (in Spanish): “Yes, I am a Darwinist, but I am against neo-Darwinism, which is a limited synthesis of the ideas of the Agustinian monk Mendel with the theory of evolution of “daddy” Charles. It is a vision of Dar1

El Periódico, March 29, 2001, interviewed by Elena Hevia. 2 The Scientist vol. 17, No. 13, 2003.

[ 21 ]

La Vanguardia, March 19, 2006. El Mundo, March 20, 2006, interviewed by Pablo Jauregui.

once upon a time lynn margulis

winism that has found its maxim prophet in the idea of the selfish gene. Neo-Darwinism is a reductionist theory because it only considers biological aspects, which are just a small part of the issues that play a role in evolution.” 5 In the last years of her life, Lynn used to remember the quote by the North American physicist David Bohm (1917–1992): “Science is the search for truth... whether we like it or not.” Living intensely is laborious and difficult, because it requires a great effort, but also provides great satisfaction. I shared with Lynn a love for science and life. Science that seeks the truth, and life that shares that love, which I find no better way to express than through the words of the Spanish poet Francisco de Quevedo. 3

La Vanguardia, July 13, 2008, interviewed by Salvador Llopart.

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in search of truth | ricardo guerrero

Amor constante más allá de la muerte

Love constant beyond death

Cerrar podrá mis ojos la postrera

The last shadow a cloudless day

Y podrá desatar esta alma mía

And this, my soul, may be freed by

Sombra que me llevare el blanco día,

May cast on me, could close my eyes;

Hora a su afán ansioso lisonjera;

An hour eager to flatter its ardor:

Mas no, de esotra parte, en la ribera,

But on that far shore it will not

Nadar sabe mi llama el agua fría,

My flame can swim frigid water

Dejará la memoria, en donde ardía:

Forsake the memory where it burned;

Y perder el respeto a ley severa.

And will flaunt so cruel a law.

Alma a quien todo un dios prisión ha sido, Soul, long imprisoned by a god, Venas que humor a tanto fuego han dado,

Veins, fuel you gave to the blaze,

Su cuerpo dejará no su cuidado;

It will leave its body, not its cares;

Polvo serán, mas polvo enamorado.

Dust they will be, but dust in love.

Medulas que han gloriosamente ardido,

Serán ceniza, mas tendrá sentido;

Marrow, gloriously you burned;

They will be ashes, but still will feel: Francisco de Quevedo (1580-1645)

Lynn and Ricardo in Barcelona, June 1983. Photo by B. Dyer.

[ 23 ]

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Cuba, October 1983. Photo courtesy of R. Guerrero.

[ 24 ]

in search of truth | ricardo guerrero

Olga California, Ricardo and Lynn in Amherst (Lynnâ&#x20AC;&#x2122;s home), December 1990. Photo courtesy of R. Guerrero.

Zach, Lynn, Dany, and Ricardo near Marrakech, January 1993. Photo courtesy of R. Guerrero.

[ 25 ]

once upon a time lynn margulis

A picnic with J.F. Crow in Amherst, June 1993. Photo courtesy of R. Guerrero.

Sampling the microbial mats at the Ebro Delta. December 1997. Photo courtesy of R. Guerrero.

[ 26 ]

in search of truth | ricardo guerrero

Brooklin Botanical Garden, at Zachâ&#x20AC;&#x2122;s wedding, August 1997. Photo courtesy of R. Guerrero.

Pedralbes Monastery, Barcelona, November 2003. Photo by M. Piqueras.

[ 27 ]

once upon a time lynn margulis

A windy day at the Giza Pyramids, Cairo, Egypt, January 2004. Photo by M. Piqueras.

Awaiting the departure, Madrid, September 2009. Photo courtesy of R. Guerrero.

[ 28 ]

teaching for science , teaching for life Montse Vallmitjana

High school teacher, Barcelona

“For most of the history of life on this planet, the living landscape resembled a time-forgotten seashore.

Although inconspicuous, life in the form of bacteria and their diverse communities changed forever the surface

and atmosphere of the planet. Although tiny, early

life was complex and original. In mudflats, evaporite expanses, fens, and ponds, microbes evolved innovations that we now associate with animals and plants”

Lynn Margulis, Early Life

A young Lynn Margulis first came to Barcelona in 1973, to attend the Fourth International Conference on the Origin of Life. For her, the conference “had nothing to do with the city,” but “everyone was very nice and it was a lot of fun.” In 1982, she participated, along with other leading scientists, in a conference on Darwinism, organized to commemorate the centenary of Darwin’s death and the opening of the new building housing the Faculty of Biology of the University of Barcelona. But it was at a dinner party held at the home of Joan Senén-Josa and his wife, Catherine, to which Antonio Lazcano, Marta Estrada, Montse Ponsà, Lynn, and I were invited, that I discovered her to be not only a highly intelligent sci[ 29 ]

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entist and communicator but also a warm person with a wonderful sense of humor. Getting to know her was a great privilege and the beginning of a long friendship. During that visit to Barcelona, I interviewed Lynn for CiĂ¨ncia, which at the time was the first and only popular science magazine published in Catalan, edited by Joan SenĂŠn-Josa. In that interview 30 years ago, Lynn talked about her research, the results of which

Lynn Margulis interviewed by Montse Vallmitjana for the magazine CiĂ¨ncia. The interview was conducted in 1982 but published in 1983. Photo by G. Serra.

[ 30 ]

teaching for science, teaching for life | montse vallmitjana

would later be expanded upon and supported: the theory of the origin of the eukaryotic cell by endosymbiosis; the emergence of life; the five kingdoms—four of them defined by cytology, ecology, and life cycle studies, and the fifth, the Protoctista. Fortunately, in the following years Lynn would often come to Barcelona, to continue her collaboration with the Microbial Ecology Group at the Autonomous University of Barcelona. The group’s director, Ricardo Guerrero, always made sure Lynn was aware of the attendance of many high school teachers at her lectures. Afterwards, in her humorous and affectionate way she always asked me “Cómo está papá Vallmitjana?” She was asking about my father, Lluis Vallmitjana, professor at the Faculty of Biology of the University of Barcelona. Lynn was quite familiar with his research on mitochondria and other organelles and on protozoa. Through the powerful arguments laid out in Lynn’s books, the scientific community became well acquainted with the undulipodia, Warrawoona, fossil bacterial stromatolites, and especially Protoctista, which we had previously referred to as protists (unicellular eukaryotes). From her we learned about microbial mats, depicted in photos of those formations at Shark Bay, Australia, and later on about the “Catalan spirochete” (Spirosymplokos deltaiberi), discovered at the Ebro Delta. We were captivated by the image that later would provide the cover of Five Kingdoms, a hand showing the Earth with a kingdom on each finger. We watched films showing animals in symbiosis with green algae, among them the flatworm Convoluta roscoffensis, which contains the alga Platymonas in all of its cells. We talked about planets, reducing atmospheres, bacterial photosynthesis, and so much more. Many of her lectures began by asking us to consider the Earth [ 31 ]

once upon a time lynn margulis

as seen from the Moon, or to imagine the biosphere, life, and time: three billion years of only bacterial life. We not so-young high school teachers had been taught that life was about 600 million years old. At university we familiarized ourselves with the many amazing chemical reactions performed by bacteria, a classroom blackboard full of biochemical formulae. But all that information was without any further connection. Little information could be found in books about the many forms of bacteria (cocci, bacilli, vibrios, spirilla) or the diseases they caused. Algae, including the Cyanophyta (blue-green algae) were studied as such; living organisms were classified into three kingdoms (Monera, Plants, and Animals), and we were told that they had appeared and survived on Earth because it was a planet with oxygen in its atmosphere! Now,

LluĂs Vallmitjana (center) at the Faculty of Biology of the University of Barcelona, with Lynn and Ricardo Guerrero. Photo courtesy of M. Vallmitjana.

[ 32 ]

teaching for science, teaching for life | montse vallmitjana

and in large part because of Lynnâ&#x20AC;&#x2122;s work and her advocacy of that of others, we know that the oxygen in the Earthâ&#x20AC;&#x2122;s atmosphere is a byproduct of bacterial metabolism. Recently, a friend noted that we will always be indebted to Lynn for providing us with an overview of life broad enough to encompass its history, microbial ecology, and endosymbiosis. Life had developed and it proceeded through evolutionary processes. Millions of years of life, allowing the diversification of bacterial metabolism, the appearance of oxygen in the atmosphere, giving rise to the eukaryotic cell and, finally, to multicellular beings. Over the last three decades, Lynn Margulisâ&#x20AC;&#x2122; ideas on endosymbiosis and the relationship of the biosphere to the environment (later included in the Gaia theory), the classification of living organisms, and the processes of speciation (symbiogenesis) have

Lynn Margulis during a lecture in Barcelona. Photo by M. Vallmitjana.

[ 33 ]

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become part of the standard high school curriculum. Within a continuing education program, a group of teachers from the Catalan Society for Biology has been regularly organizing a series of lectures. We had invited Lynn to give a lecture in 2012. Sadly, it was not possible. I will remember her showing one of her videos, passionately discussing spirochetes, or reciting a poem by Emily Dickinson in her distinct voice. But I imagine her on a remote beach, in the Proterozoic, walking in the sea, barefoot. 3

[ 34 ]

beyond the sea Marta Estrada

Institute of Marine Sciences, CSIC, Barcelona

El meu amic el mar

My friend the sea

quan la meva nau busca recer

when my ship seeks shelter

té la calma d'un déu adormit, a l'illa del seu pit.

has the calm of a sleeping god, on the island of its chest.

El meu amic el mar

My friend the sea

i quan s'omple d'aire el meu velam

and when air fills my sails

té el coratge d'un déu exaltat, seguim un joc incert.

has the courage of an exalted god, we play an uncertain game.

El meu amic el mar

My friend the sea

i en el seu va-i-vé de so i color

and in its coming and going

és l'immens bressol de tots els blaus, aprenc el poc que tinc.

is the immense cradle of all blues, of sound and color

I realize how little I have.

És per això que mai

That's why I will never

i fidel viuré amarinat

and I shall live faithfully in the sea

no em podré allunyar del seu batec, fins acabar el vent.

manage to escape its heartbeat, until the wind is no more.

Lluis Llach (songwriter)

[ 35 ]

once upon a time lynn margulis

Throughout the 1970s I made several visits to various research centers in the United States. One of them was to the Marine Biological Laboratory, which, together with the Woods Hole Oceanographic Institute and the National Marine Fisheries Service Laboratory, was located in the vacation town of Woods Hole, Massachusetts. As a center of global importance for oceanographic research, Woods Hole receives many visitors and along with its famous summer courses is the site of many conferences and workshops. I was fortunate to participate in some of them and I remember with pleasure the lessons, field work, and the many discussions with peers and teachers. At the time, Lynn was a professor at Boston University, which maintained a marine studies program at Woods Hole. It was there that I met her and a lifelong friendship began. That stay at Woods Hole was a major impetus for my research, which has been carried out mainly at the Institute of Marine Sciences in Barcelona, part of the Spanish Research Council. My researchâ&#x20AC;&#x201D;and I can say my passionâ&#x20AC;&#x201D;focuses on the ecology of phytoplankton and its interactions in the global marine ecosystem. Over the years I have developed this work, exploring the Mediterranean, Antarctica, and other marine regions of the world within programs of the Institute of Marine Sciences. With my colleagues, I have also extended my interests to include the control mechanisms of harmful algal blooms. In my work as a researcher, in the training of personnel, and in performing management tasks I seek to encourage an enthusiasm for science, especially among young people. It is important for them to realize that scientific work, regardless of the chosen area, is worth doing. In the case of marine science, there is the satisfaction of contributing to the [ 36 ]

beyond the sea | marta estrada

knowledge of an important part of the world that surrounds us, the sea. In this regard, Lynn was certainly an expert. Lynn spoke fluent Spanish, which she had learned during her stays in various Latin American countries, especially Mexico (she also spoke French and Italian). I was impressed by the enthusiasm that she conveyed in her lessons and lectures, dedicated to topics as diverse as cellular evolution, the origin of life, and the influence of the biosphere on the planet. In 1978, I was a guest at her home in Newton, near Boston, going swimming with her in the local public pool and enjoying the company of her family. Zach, her third son, was at that time very young but he was a wonderful guide during my visits to downtown Boston. Lynn Margulis leaves us a great body of scientific work, full of innovative ideas. When she proposed that eukaryotic cells had evolved from a symbiotic community of prokaryotic cells, her theory seemed quite eccentric. Now, of course, the endosymbiotic origin of mitochondria and plastids is well accepted. In marine phytoplankton, to which I have devoted many years of research, the important role of endosymbiotic evolutionary events, not only primary, but also secondary and tertiary ones, can be readily appreciated. Lynn cited Russian biologists as originators of the theory of symbiosis, but it was she who developed it, guiding it away from its early rejection to its full adoption by the scientific community. Lynn was a revolutionary thinker as well as a passionate observer of the microscopic world. I was fascinated by her documented presentations on the strange consortia of microorganisms that truly exist but seem to belong to the world of science fiction. Among her many other scientific contributions, we also have to thank her for her role in developing and staunchly supporting James Lovelockâ&#x20AC;&#x2122;s Gaia theory. [ 37 ]

once upon a time lynn margulis

Sampling in the Indian Ocean during the Malaspina cruise (March 2011). The 24 oceanographic bottles, with a CTD probe (conductivity, temperature, depth) in the middle, were submerged to a depth of 4000 m to record temperature, salinity and other variables, and to take water samples at selected depths. Photo by M. Estrada.

[ 38 ]

beyond the sea | marta estrada

Bow of the icebreaker the CCGS Amundsen, a Canadian ship used for scientific research, during the Circumpolar Flaw Lead cruise in the Arctic Ocean (March 2008). Photos by M. Estrada.

[ 39 ]

once upon a time lynn margulis

In 1973, Lynn came to Barcelona to attend the first meeting of the International Society for the Study of the Origin of Life (ISSOL). We also met in 1982, when she was invited to the opening of the new building of the Faculty of Biology of the University of Barcelona. In 1988, I had the pleasure to see her in Perpignan, at a meeting held to honor Edouard Chatton, the first person to propose the division between prokaryotes and eukaryotes. Beginning in 1982, her visits not only to Barcelona but to all of Spain became very frequent. She participated in projects and research collaborations, was invited by universities and research centers, and gave courses, lectures, and presentations. She was awarded Doctor Honoris Causa degrees by twenty universities around the world, including several from Spain (Autonomous University of Madrid, University of Valencia, University of Vigo and Autonomous University of Barcelona), evidence of the great esteem in which she was held. Lynn Margulis had a strong relationship to Catalonia, one that was not limited to giving lectures or participating in conferences. She collaborated in the preparation of several monographs, for instance issue 39 of Treballs de la Societat Catalana de Biologia devoted to the origin of life. In addition, she collaborated in research projects, including the one that led to the identification and description of the spirochete Spirosymplokos deltaiberi, which was isolated from the microbial mats of the Ebro Delta. With her sense of humor, she used to call it the â&#x20AC;&#x153;Catalan spirochete.â&#x20AC;? In November 2011, Lynn was still a highly active and dynamic researcher; thus, the news of her death was unexpected and very sad. Among Lynnâ&#x20AC;&#x2122;s many qualities I remember her kindness and her engagement with others. I can recall, for example, how enthusiastically she spoke about a group of people in Boston dedicated to promoting [ 40 ]

beyond the sea | marta estrada

womenâ&#x20AC;&#x2122;s health. She had a keen interest in popularizing science and she contributed to it through publications and in collaborations with the media. It is a pleasure to re-read her interviews, some of which, like the one published by the magazine MĂ¨tode, can be found on the web [http://metode.cat/es/Revistas/Entrevista/Lynn-Margulis] I consider myself fortunate to have met Lynn Margulis. If I had to define her in three keywords, I would say revolutionary, enthusiastic, committed. She leaves us a legacy of scientific ideas and an example of a scientist and a woman who will never be forgotten. 3

[ 41 ]

looking at microbes Isabel Esteve

Autonomous University of Barcelona, Bellaterra

â&#x20AC;&#x153;Faithâ&#x20AC;? is a fine invention

When Gentlemen can see â&#x20AC;&#x201C;

But Microscopes are prudent In an Emergency

Emily Dickinson (1830-1886)

Scientific facts are based on observations and experimental results, both of which benefit from the progressive and, as has lately been the case, accelerated development of technological tools. Among these tools, the one that undoubtedly has contributed the most to our understanding of the vital unit, the cell, has been the electron microscope. Indeed, in biology, electron microscopy has played a role comparable to that of the elementary particle accelerator in revealing the intimate structure of matter. Molecular genetics, nanotechnology, and other life-science-related disciplines, including medicine, have profited enormously and effectively from scanning and transmission electron microscopy observations of the structure of the cell. The resulting progress in the field of cell biology has in turn promoted further scientific and technological advances that have improved our understanding of the functioning of prokaryotic and eukaryotic cells. The first electron microscopes were built by Knoll and Ruska between 1930 and 1933. Although first marketed in 1939, the electron [ 43 ]

once upon a time lynn margulis

microscope was not used to study the structure of the eukaryotic cell until 1952; its application to the field of microbial ecology would not occur until the 1970s. Since then, much of what we know about the identity and distribution of microorganisms in nature has been aided by electron microscopy analyses. The identification of organisms according to their distinctive ultrastructural features is among the major achievements resulting from the application of electron microscopy to the knowledge of microbial diversity in natural environments. Also relevant to classification efforts is an understanding of the metabolic processes performed by the various groups of microorganisms. This knowledge is enhanced by relating ultrastructural to chemical changes inside the cell. Of special interest is the role of electron microscopy in revealing the importance of microbial diversity in extreme environments and the mechanisms developed by microorganisms, mainly bacteria, to survive in life-limiting conditions, such as high and low temperatures, changing solar radiation, and variations in pH and salinity. The use of extracellular polymeric substances and the structuring and aggregation of microbial communities as defenses against adverse environmental conditions are now well-defined strategies, thanks to the great resolving power of modern electron microscopes. Recent studies have focused on the partnerships formed between different organisms, which are relevant in phenomena ranging from the development of infectious diseases (biofilm formation) to the bioremediation of contaminated ecosystems in natural environments. Our Microbial Ecology group at the Autonomous University of Barcelona (UAB), has used high-resolution microscopy techniques for many years, mainly to study two types of environments: the Cis贸 Lagoon in Banyoles (Girona) and the microbial mats of the Ebro [ 44 ]

looking at microbes | isabel esteve

Delta (Tarragona). We have specialized in the study of phototrophic microorganisms, both oxygenic and anoxygenic. In recent years, we have worked on the role of microorganisms in detoxifying metalpolluted ecosystems. Confocal laser scanning microscopy, scanning electron microscopy, and transmission electron microscopy, the latter two coupled to an X-ray dispersive energy system, have allowed us to evaluate the ability of various microorganisms, especially cyanobacteria, to capture metals, as an indicator of their potential to bioremediate polluted environments. Very little is as yet known about the biodiversity of ecosystems and the interactions of the resident organisms, among themselves and in their adaptations to environmental conditions. Continued technological advances in microscopy and multidisciplinary approaches using biochemical and molecular methods will guide the future of these and other studies in microbial ecology. It has been a long road of research, work, and life since I first met Lynn Margulis. I have to thank her for the great interest that she always showed in our work and for her friendship. I first met Lynn when she came to the UAB in 1982, invited by Ricardo Guerrero. It was impossible not to be caught up in her vitality and enthusiasm. Her open and unceasingly curious mind were obvious to anyone in her presence. At the same time, she impressed us with her closeness although by that time she was already a well-known scientist. During that first encounter, I was working, in collaboration with Nuria Gaju, on the antagonistic relationship between the predatory bacterium Daptobacter, isolated in our laboratory, and Chromatium, a photosynthetic bacterium that Daptobacter attacked by penetrating its cytoplasm and causing cellular lysis. Lynn was immediately fascinated by this work, which led to a long-lasting collaboration and, [ 45 ]

once upon a time lynn margulis

equally important, to our close friendship. We also collaborated in exploring other aspects of symbiosis, first the relationship between the ciliated Coleps hirtus and Cryptomonas phaseolus and, later on, the microorganisms that form the microbial mats of the Ebro Delta. More recently, by applying methods of high resolution microscopy, our group â&#x20AC;&#x201D;in a collaborative research projectâ&#x20AC;&#x201D; has been able to improve toxicity studies in marine sediment microorganisms. And also to validate the effectiveness of high-resolution microscopy techniques to determine the toxic effect of heavy metals at different concentrations in photosynthetic microorganisms inhabiting microbial mats of the Ebro delta. We have characterized a novel microalga isolated from this ecosystem, called DE2009, which has a greater capacity to sequester lead than other microorganisms studied so far.

The microalga Scenedesmus DE2009 capturing lead (indicated by arrows) from a metal-polluted culture. The image was obtained using transmission electron microscopy coupled to an energy dispersive X-ray detector. Image courtesy of J. Maldonado and I. Esteve.

[ 46 ]

looking at microbes | isabel esteve

In 1987, I went to Boston for a stay in Lynnâ&#x20AC;&#x2122;s laboratory at Boston University. Of course, she refused to allow me to live anywhere else other than in her own home. She did the same with students and researchers from all over the world. Her hospitality was well known and much appreciated. During my stay, we went to Laguna Figueroa (Baja California, Mexico) to collect samples of microbial mats. She also invited me to deliver lectures at Boston University and at Woods Hole and shared with me an impressive file of electron microscopy images of different microorganisms taken by David Chase. Her generosity had no limits. For instance, she organized a Thanksgiving Day feast a week before the actual date so that I could participate in the celebration before my return to Spain. I owe her the two most intense months of my life.

Lynn Margulis during the ceremony that awarded her an Honoris Causa Doctorate from the Autonomous University of Barcelona, June 6, 2007. Photo courtesy of UAB-I. Esteve.

[ 47 ]

once upon a time lynn margulis

In 2007, the UAB honored Lynn Margulisâ&#x20AC;&#x2122; long scientific career and her educational work by awarding her an Honoris Causa Doctorate in a ceremony in which I proudly acted as madrina (godmother). It was appropriate recognition of a woman with a passion for life and for science, an excellent professor, and a generous friend. 3

[ 48 ]

engaged to science , married to life Ricardo Amils

Autonomous University of Madrid

Yo tengo tantos hermanos que no los puedo contar

En el valle, la montaĂąa, en la pampa y en el mar.

Cada cual con sus trabajos, con sus sueĂąos cada cual. Con la esperanza delante, con los recuerdos detrĂĄs. Yo tengo tantos hermanos, que no los puedo contar

y una hermana muy hermosa, que se llama libertad. I have so many brothers I cannot count them all

In the valley, in the mountains, on the plain and in the sea. Each with his work, with his dreams every one. With hope ahead, with memories behind.

I have so many brothers I cannot count them all

And a very beautiful sister whose name is liberty.

Atahualpa Yupanqui (1908-1992, singer-songwriter)

The Rio Tinto (Red River) is located in southwestern Spain, in Huelva province. The red color of its waters is due to the high concentrations of ferric iron but also to the sulfuric acid produced by chemolithotrophic microorganisms that thrive in the high concentrations of metallic sulfides, mainly pyrite, characteristic of the Iberian Pyrite Belt (IPB). [ 49 ]

once upon a time lynn margulis

Today we know that what was initially considered to be industrial pollution is the product of microbial metabolism. From a paleontological point of view it is well established that microbial activity has been ongoing in the Río Tinto for several million years, reaching back well before mining activities started in the area some 5000 years ago. Consequently, the Rio Tinto is an excellent model to study primitive life systems based on the anaerobic respiration of reduced minerals and to suggest alternative explanations for the massive removal of iron from Archean oceans. All metallic sulfidic mines have acid drainage problems. What makes Río Tinto unique is its size (92 km long), its constant acidity (around pH 2.3), and its high level of microbial diversity, all of which are due to the nature of the water table, the high concentration of metal sulfides of the IPB, and the temperature. In the presence of water, a generous amount of substrate, and a suitable temperature, chemolithotrophic microorganisms are able to thrive, producing large amounts of the oxidation products of pyrite, ferric iron, and sulfuric acid, the main constituents of Río Tinto’s water column. The biotope of Río Tinto is rather complex. Two main extreme characteristics are the acidity of the system, as already mentioned, with a pH of about 2.3, and the high concentrations of highly toxic heavy metals (copper, arsenic, nickel, and cadmium). Given these extreme conditions, it is surprising to find an enormous eukaryotic diversity, including algae, filamentous fungi, yeasts, protozoa, and amoebae, in a place where nobody could rationally have imagined it. Our group, in collaboration with experts in protists, has isolated heterotrophic microorganisms, amoebae, and photosynthetic algae, growing them in the lab in order to study their properties. These organisms likely take part in well-established symbiotic relation[ 50 ]

engaged to science, married to life | ricardo amils

ships that have allowed them to adapt to the extreme conditions of their habitat. Thus, the questions we have posed are why and how they have managed to succeed, and what is the advantage, if any, in inhabiting this nasty environment. The same questions fascinated Lynn Margulis, who from the beginning of her career as a scientist stubbornly believed in the importance of symbiotic relationships, such as those established in this peculiar ecosystem. During our conversations her enthusiasm for pursuing these and other such biological challenges could be seen in the sparkle in her eyes. RĂo Tinto provides us with a model based on iron. Iron is an extremely important element for life. Free access to iron, a limiting factor at the neutral pH in which most organisms develop on Earth,

Runway intermediate zone of Rio Tinto (Berrocal); the development of acidophilic algae can be seen. Photo courtesy of J. Segura.

[ 51 ]

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would have supported protists living in Río Tinto. More than 100 years ago, microorganisms were found in extreme halophilic conditions. However, it was not until the late 1960s that Thomas Brock and his collaborators isolated the first hyperthermophile, from Yellowstone National Park. This organism, Thermus aquaticus, transformed previous conceptions of the bacterial limits of life. Among the large number of articles and books published by Brock, surely the most interesting is the autobiographical text which appeared in the Annual Review of Microbiology in 1995. In “The road to Yellowstone–and beyond,” Brock describes the work of his group that led to those discoveries and their implications. In that same article, he essentially establishes microbial ecology both as an analytical approach and as a field of study. Recognition of Thermus aquaticus ended the long-held notion that life could develop only under “normal” conditions of temperature, pH, pressure, and water activity. But extremophiles not only extend the limits of life, they are also of great help in studying its origins and in guiding the search for life on other planets. It is no wonder then that Lynn was deeply interested in our work, as all her life she had sought answers to questions regarding the origin of life in the Universe and its evolution here on Earth. I remember her fascination with the amoebas detected along the banks of the Río Tinto and the adaptation of these organisms to extreme conditions probably not much different than those that characterized primitive Earth. Although we have learned a lot about the origins of life (in part thanks to Lynn), there is still much more to discover. Even the definition of life is complicated. Those with great minds, Lynn Margulis among them, have had to conclude that although we can describe the properties of living things, list them, and obtain images of them, [ 52 ]

engaged to science, married to life | ricardo amils

we cannot convincingly define the essence of life, what it is all about. This inability prevents us, for example, from understanding whether viruses are alive. It hinders not only biology students, but also, e.g., astrobiologists, who are engaged in searching for life outside our planet. How can we detect something if we are not quite sure what it is? It was in July 1986, at the symposium of the International Society for the Study of the Origin of Life, held in Berkeley, California, that I have met Lynn for the first time. The president of that meeting was Stanley Miller and the four major questions addressed at the gathering were: What were conditions like on primitive Earth? What is â&#x20AC;&#x153;primitiveâ&#x20AC;? in biology? How did bioenergetic systems develop in the prebiotic milieu? What were the first self-replicating systems? Among the things that impressed me the most about Lynn were her passionate drive as a scientist and her humanity as a person. After that, we met many times, discussing science, mainly microbes, but also often laughing at many things. Her strongest influence on me as a scientist was in deepening my appreciation of the role of microorganisms in shaping our blue planet; on a personal level, she is responsible for many of the good friends that I made, based on our shared friendship with Lynn. Now, in my daily work, with the joys and deceptions that are the nature of a life in science, I frequently think back on my encounters with this remarkable, intelligent and devoted woman, Lynn Margulis, and the deep friendship we enjoyed. 3

[ 53 ]

life is symbiosis Eva Barreno

University of Valencia

Lichens, the Earth slowest telegramâ&#x20AC;Ś The lichen describes itself inscribes itself, writes

a verbose silence: in encoded script Graphis scripta

Hans Magnus Enzensberger, Flechtenkunde

Lichens represent major radiations of ascomycetes in a symbiotic stage that is characterized by a unique symbiogenetic phenotype of specific biological organization in the lichen thallus. The cyclical associations of lichens involve at least two very different organisms, a heterotrophic fungus (mycobiont) and a photosynthetic (photobiont) cyanobacterium (cyanobiont) or unicellular green alga (phycobiont). Recently, the coexistence of several phycobionts in some lichen thalli has been demonstrated. These specific and selective forms of symbiotic associations result in positive interactions among potential competitors, leading to complex but stable entities. The important functional interactions between photobionts and mycobionts, and possibly other symbionts such as specific bacterial communities, suggest that these partners evolved simultaneously. The evolutionary success of holobionts is evidenced by the fact that more than 15,000 lichen species are known thus far. [ 55 ]

once upon a time lynn margulis

Lichenization allows the partners to thrive in habitats that would otherwise be unavailable to either one on its own. A predominant characteristic of lichens is their desiccation tolerance. Lichen thalli are found in almost all terrestrial habitats, from the poles to the tropics, including extreme environments such as deserts and high mountains. Indeed, they are the only organisms able to survive for prolonged periods in outer space. Lichens are also essential to ecological succession, by enhancing the weathering of rock surfaces. The study of lichen symbioses is one of the most exciting fields in biology, as it makes use of a broad spectrum of methodologies, from molecular to ecological. Interdisciplinary approaches are essential to the study of evolutionary processes but also for efforts aimed at biodiversity protection. Most of my professional life has been devoted to lichens. At the University of Valencia, our group investigates several aspects of lichens, including their taxonomy, biogeography, ecophysiology, physiology, genomics, proteomics, and the role of the symbiotic algae Trebouxia sp.pl. We have identified the coexistence of multiple phycobionts in lichen thalli and examined how changes in fungal genomes are accompanied by changes in the algal genome, resulting in concerted changes in three genomes. The physiological results have been used to develop tools in which lichens serve as bioindicators of atmospheric pollution and environmental changes, both under field (biomonitoring networks) and controlled laboratory conditions. Our current efforts are aimed at determining whether lichen phycobionts are of biotechnological interest. It was my activity in the world of lichens that sparked my interest in the work of Lynn Margulis; it led to many discussions with her over the years and to our deep friendship. As soon as Lynn learned that I was a lichenologist, lichen symbioses were a constant [ 56 ]

life is symbiosis | eva barreno

Parmelia barrenoae Divakar, MC. Molina & A. Crespo. This foliose lichen was originally described from the Iberian Peninsula even though it is now widely known in Europe, where it was often confused with P. sulcata Taylor. Recently, it was reported in North America (California, Idaho, Oregon, and Washington) and in Africa (Middle Atlas, Morocco). Courtesy of E. Barreno, at Sierra del Toro, Castellón, Spain

subject of our talks. At a gathering in Valencia in 1996 she presented the Spanish edition of her wonderful book What is Life? The following year I invited her to give the opening lecture of the 12th Symposium on Cryptogamic Botany, held at the University of Valencia (September 17–20, 1997). The title she chose for her talk was “Photosynthetic life: Symbiogenesis and the origins of algae.” All who attended that meeting—including myself, of course— were greatly impressed by Lynn’s originality and brilliance. She greatly enjoyed the event and the opportunity to interact with specialists in the biology of algae, bryophytes, fungi, lichens, and ferns. Interdisciplinary meetings, in which experts in several fields interact, were [ 57 ]

once upon a time lynn margulis

rare, she noted, and she expressed her hope that this aspect of the Symposium would be preserved for as long as possible. In the year 2000, the University of Valencia celebrated its 500th anniversary. Among the various activities organized to commemorate the occasion, Lynn collaborated with us in organizing the “GAIA 2000: Second Chapman Conference on the Gaia Hypothesis.” With the support of the American Geophysical Union, she invited some of the foremost researchers in evolutionary biology, geophysics, meteorology, microbiology, and paleontology. Even James E. Lovelock himself participated. Lynn’s efforts were consistent with her well-known generosity. Later on she asked me to co-author a paper for the journal BioScience, as it was her wish to share what was known about lichens with an extended audience on the occasion of the publication of Lichens of North America (Brodo et al. 2001). In our article for BioScience we wrote: “Lichens exemplify the details of complex individuality. The relations between syntrophic metabolism and morphogenesis in the emergence of novelty through physical association is made obvious in these colourful creatures, and so too the contribution of symbiogenesis to speciation and taxonomy in them is manifest and their capacity for revival from severe desiccation is impressive.” At the time, Lynn had suggested that I should search for more phycobionts and bacteria in the cyclical symbiosis of lichens. And it was following this wise advice that I decided to change my line of research on lichens. Now we know of the coexistence of several species of algae and different bacteria. We have discovered new algal species, including one that is found in North America and in Europe. This microalga will be named Trebouxia lynnae in her honor. I am indebted to her for so much intellectual encouragement. [ 58 ]

life is symbiosis | eva barreno

On June 8, 2001, Lynn was distinguished as Doctor Honoris Causa by the University of Valencia and I was honored to be her academic companion. During the ceremony, Lynn gave the lectio “Bacteria in the origin of species: neo-Darwinian paradigm death.” The text was included in the book Una revolución en la evolución (A revolution in evolution), published by the University of Valencia in 2002, within the Honoris Causa collection. The book also contained biographical data and a selection of Lynn’s articles and other writings. She was a devoted mother, a sparkling personality, a good friend, and a passionate woman full of life. She was tireless, brilliant, tenacious, and completely determined to pursue her ideas and defend her convictions. 3

[ 59 ]

a world of microbes Ramon Folch

President of ERF (Estudi Ramon Folch & Associates), Barcelona

It took hundreds of millions of years to produce the life

that now inhabits the earth —eons of time in which that developing and evolving and diversifying life reached a

state of adjustment and balance with its surroundings. The environment, rigorously shaping and directing the life it supported contained elements that were hostile as well as supporting[…] Given time —time not in years but in millennia— life adjusts, and a balance has been reached.

For time is the essential ingredient; but in the modern world there is no time.

Rachel Carson (1907-1964), Silent Spring

What a long time ago it was when we humans believed that the Earth was the center of the universe, with ourselves as the monarchs of creation! It took many years of effort and study to discover otherwise and enormous bravery to dare say so, in a time when knowledge was nurtured and supported by religious ideology. With the overpowering strength of its arguments, science revealed Earth’s astronomical condition and its position: a small planet in a modest stellar system in a secondary galaxy. It also became obvious that the biological dimension of humans was modest; as practically newlyarrived beings with a very minor relevance in ecological terms. [ 61 ]

once upon a time lynn margulis

We are eukaryotes; our cells are extremely complex, with a differentiated nucleus and packed with structures whose functions we are only slowly beginning to understand. Initially, our complexity was thought to confer us with superiority over the insignificant bacteria, those trifling unicellular beings with a simple anatomy. But even that presumption has turned out to be illusory, because we now know that bacterial physiology is prodigiously diverse, much more efficient than ours, and prepared to exploit every imaginable environment. Indeed, bacteria thrive and proliferate in environments where no other living being could survive. Lynn Margulis was among those who quickly recognized the value of bacteria and granted them their due respect. She understood their function on our planet and the role they played in the appearance and evolution of life. This native Chicagoan converted Bostonian grasped the transcendence of bacterial cells and shook up the classification of living beings by constructing a five-branch tree, with four branches containing eukaryotes, i.e., animals, vegetables, fungi and protoctista (formerly, protozoa and algae) and the fifth, prokaryotes (bacteria). She proposed the theory of serial endosymbiosis which explains the eukaryote cell as a symbiontic integration of diverse bacterial cells, and formulated, together with James Lovelock, the Gaia hypothesis (Earth as a macro-organism). Her achievements in her chosen field and outside of it were impressive. Besides being good friends, she contributed decisively to the success of Biosfera, the collectively written encyclopedia whose compilation I directed in the 1990s and of which she was an enthusiastic supporter from the moment she learned of it. She was an active collaborator and co-author of some of its eleven volumes, especially the first. She also helped to publicize the work, acting as intermediary [ 62 ]

a world of microbes | ramon folch

for a version in English, which became possible. There is also a German version and the Japanese edition is in the works. The socioecologic focus of Biosfera was what pleased Lynn the most. Biosfera is an encyclopedic work, set up according to the principles of the UNESCO MAB (Man and Biosphere) Program, and with its patronage. The work was published by EnciclĂ˛pedia Catalana and contains the contributions of nearly 200 scientists from all over the world, experts in different fields. Of importance to Lynn

Invitation to the presentation of Biosfera, March 22, 1993, at the Science Museum in Barcelona, with the participation as a lecturer of Lynn Margulis, co-author of the first volume. As a curiosity, it can be noted the similarity of this graphism to the cover of Google Earth twenty years later.

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was that although the presence of macroscopic living beings often masks the role and function of microorganisms in many ecosystems, bacterial presence is manifest in all media. In coastal environments (treated in her contribution to Biosfera), microorganisms have retained much of their physiognomy from the remotest eras. Lynn was a frequent visitor in Catalonia, where she had many friends, collaborators, and admirers-followers of her work. She was enchanted by the microbial mats of the Ebro Delta and other places in Spain and shared her fascination for these ecosystems, in which we find important clues to the origin of life. As I am both an ecologist and a botanist, I always appreciated Lynnâ&#x20AC;&#x2122;s defense of the environment. She was a woman who loved Nature dearly and who tread upon it cautiously. Yet she was also realistic and did not believe that human intervention can be absolutely innocuous. Faithful to the premise of â&#x20AC;&#x153;think globally, act locally,â&#x20AC;? outside her house in Amherst there was, prominently, an area for composting. Defense of the environment needs a sustainable focus. Sustainability is a word lately more often heard than understood. Understanding, however, is essential to successful environmental management and in efforts to conciliate the various interests, whether of ecological advocates or of conventional socioeconomic sectors. Ecology, education, and development are basic elements that will guarantee our respect of the environment. They require the clear communication of environmental goals if we are to implement the necessary actions efficiently. For young people who choose to follow a scientific career in ecology it is important that they appreciate the many other possibilities, besides academic ecology or ecological research, that surely would satisfy their desire to contribute to the preservation of our environment. In my case, I was fortunate to be able to put into practice all of that I [ 64 ]

a world of microbes | ramon folch

learned into the ecology of human systems. The application of ecological principles and concepts are particularly relevant in urbanism, such as the design of new buildings. The use of available elements like natural light, the heat of the sun, or shaded areas are among the many aspects that contribute to our welfare and economy, in this case through the benefits of artificial-energy savings. At the same time, we must be aware that nearly all of the things that make our lives easier contribute to pollution, mainly in the form of CO2. So what are we willing to give up? At what cost? What level of emissions will we consider to be acceptable? We must answer these questions realistically and honestly. Perhaps more difficult but equally important is the need for governments to realize, based on accurate information and careful predictions, that without a global view they will hardly be in a position to tackle the planetâ&#x20AC;&#x2122;s problems. This global view is what must steer scientistsâ&#x20AC;&#x2122; tasks. I have followed the principles of ecology in my professional activities since I get my doctorate in this matter, that was firstly established in Spain at the University of Barcelona in the 1960s. These words are also meant as a short call to consider ecology not only with respect to the natural environment but also in its urban dimension. The two are, of course, intimately related and ensure the preservation of biodiversity. They include the avoidance of both aquifer depletion and pollution of the seas and the need to monitor the health of forests and rivers. Words and actions; with these tools Lynn Margulis contributed to our understanding of biology and to making it understandable, to pointing out the value of microbes and their function, to practicing the principles of ecology, and to supporting efforts to conserve the resources of our planet, with responsibility and knowledge. Thank you, Lynn. We miss you, but your work and your memory remain. 3 [ 65 ]

lynn margulis, my inspiration Mercè Piqueras

International Microbiology, Barcelona

No tornaràs mai més, però perdures en les coses i en mi de tal manera

que em costa imaginar-te absent per sempre. You will never return, but you will last

In both things and myself in such a way That it’s hard for me to imagine you’ll be absent for ever.

Miquel Martí i Pol (1929-2003), Llibre d'absències

In 1995, during a talk with an undergraduate biology student, I recommended him the Spanish edition of Lyn Margulis’ Microcosmos, which had just been released. “Is that Lynn Margulis the same person that explained the origin of eukaryotic cells by means of serial endosymbiosis?”, he asked me. “Yes, she is”, I replied. “I learnt serial endosymbiosis in the first year of university, but… why nobody told me that Margulis was a woman and… that she was still alive!?” Had that conversation taken place today, unfortunately, the student would no longer be able to say “and… that she was still alive!” It is true that Lynn is no longer physically with us, but her memory will be always alive in the people that got to know her, and her ideas have already a place in the history of biology. Indeed, Lynn, it is hard for me to imagine you will be absent for ever. [ 67 ]

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I first met Lynn on November 1987, during one of her frequent visits to Barcelona. She gave a lecture in a meeting on Gaia that Prof. Ricardo Guerrero had organized at the Barcelona Science Museum (currently CosmoCaixa). Reading Early Life and later on Microcosmos—both books by Lynn—offered me a new insight in the history of life on Earth. One that was quiet different from what I had learnt at the University in the late 1960s. The explanation Lynn gave to the origin of the eukaryotic cell by serial endosymbiosis and to changes in the Earth atmosphere throughout evolution due to the interaction between the Earth’s biotic and abiotic components seemed obvious, yet nobody had realized before that it had to be that way. Only one imaginative, curageous person, with a great mind, could propose those theories, and that was Lynn. It is true that—as she herself repeated many times—Lynn had rescued the ideas from several Russian researchers that had hypothesized about the origin of cellular organelles, but she must be credited for having developed and spread those ideas, and for having encouraged historians of science to delve into Russian scientific literature to find out more about those pioneers of symbiogenesis. Lynn would come often to Barcelona, where she had her second home, and always accepted to lecture to any kind of audiences: either to university researchers or students, or to lay publics. To all of them, she could communicate her theories at the appropriate, comprehensible level. However, I think that students were those that became more fascinated listening to her and surely now there are biology researchers that owe to Lynn Margulis their decision of following a career in biology. In fact, throughout the years I have seen, in the faces of students attending Lynn’s lectures, the same fascination and wonder that I felt when I first met her and [ 68 ]

lynn margulis, my inspiration | mercĂ¨ piqueras

read Early Life and Microcosmos. She indeed conveyed her enthusiasm and love for microbes to her audiences and to her readers. When I first met Lynn I was a bit confused about my future in the world of biology. A few months earlier I had started my collaboration with Ricardo Guerreroâ&#x20AC;&#x2122;s team and I was catching up with biology. I did not have the intention to start a doctorate; I was aware I had already missed the boat. In fact, I had been seventeen years far from the world of science, devoted only to my family, growing up three children, and I felt overwhelmed by the changes in subjects such as genetics, cell biology, and evolution, mostly due to the development of molecular biology. My collaboration with Ricardo Guerrero was not in the lab, but in the office; I was assisting in the English correspondence, preparation of articles and reports. I also did bibliographical research and collaborated in the organization of several scientific international meetings. I had always had a liking for language and writing, and started doing other activities on my own, in which I could develop my skills in those fields. So I translated Lynnâ&#x20AC;&#x2122;s book Microcosmos into Spanish and then began writing popular science. It took more than seven years for the Spanish Microcosmos to be published. The translation was ready, but the publisher that had the rights for the Spanish edition was specialized in university textbooks and finally thought that a popular science book would not fit in its catalogue. Fortunately, a few years later, another publisher (Editorial Tusquets) became interested in the book and bought the rights. My first popular-science story was about the American biologist and science writer Rachel Carson and the impact of her book Silent spring; I wrote it for the weekly science supplement of La Vanguardia in 1994. Since then I have published many articles and stories, [ 69 ]

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A view of the kitchen of Lynn Margulisâ&#x20AC;&#x2122; house. Here she used to prepare meals either for her and for their frequent visitors, family, colleagues and students. And also a nice place to look outside while reading, studying or writing. Photo by M.Piqueras.

as well as several books, and currently science communication in various ways is my main activity. Both Lynn Margulis and Ricardo Guerrero encouraged me to follow that path, which has allowed me to be involved in both science and writing, the two things I like the most. Being a freelancer science writer, science editor and translator has many advantages, but has also disadvantages. Anyway, at my age, when many of my university fellows are retired, I do not even think of it. I have stepped down from some duties (I was in the Board of the Catalan Association for Science Communication for eleven years, five of which as the President; I was also Vice-President of the Catalan Association for the History of Science and Technology for two years), but keep being Associate Editor of International Microbiologyâ&#x20AC;&#x201D;the official journal of the Spanish Society for Microbiology, [ 70 ]

lynn margulis, my inspiration | mercè piqueras

which was founded by Ricardo Guerrero with the collaboration of Lynn Margulis—doing some translations, and writing popular science either in press or in my personal blog La lectora corrent (The common reader, http://lectoracorrent.blogspot.com). A recent project in which I was involved was the current permanent exhibition of the Natural History Museum of Barcelona at its new facilities: the Museu Blau (Blue Museum; the name comes from the indigo-blue color of its building). The external curator of the project was Ricardo Guerrero, and he envisaged an exhibition that provided a comprehensive vision of the Earth and life on it, showing the interactions between both the biotic and abiotic components of the planet, and stressing the role that microbes had had throughout the history of life. That approach was based on the ideas that the British chemist James Lovelock, with the collaboration of Lynn Margulis, expressed in his Gaia theory, or the science of

Some things packed in Lynn Margulis’ office after she died. Photo by C. Puche.

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Earthâ&#x20AC;&#x2122;s physiology. When Lynn died, she left her scientific legacy to Ricardo Guerrero, who thought that the Natural History Museum of Barcelona would be the most suitable place to host it. In March 2012, when the University of Massachusetts-Amherst organized a symposium in the memory of Lynn Margulis, nine people from Barcelona attended it. Had not been for the long, expensive trip, surely many more would have gone to Amherst. A great part of Lynnâ&#x20AC;&#x2122;s own scientific library, many specimens including geological samples from biological origin, as well as videos, article reprints and drawings are already in Barcelona and in the future will be available to students and scholars at the Natural History Museum. This will contribute to keep her memory alive in a city that Lynn loved so much and that also loved her. 3

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let s listen to the universe Eduard Punset

Grupo Punset Producciones, Barcelona

There are more things in Heaven and Earth, Horatio, than are dreamt of in your philosophy. William Shakespeare (1564-1616) Hamlet I.5

Those who have followed the program Redes on the Spanish TVE and/or have read any of the books I have published over my long career know my passion for science. I am passionate about all of its branches, aspects, and specialties and about everything it can give us, including progress and intellectual strength. What I also find exciting is communicating science, making it available to the public in a meaningful, engaging way, and not only to those already equipped with a basic knowledge of science but also to those who are simply curious and motivated by the desire to learn. In science communication, the pursuit of quality is essential. It is what awakens the interest of both experts and lay people in discussions of any topic, whether presented in the form of an interview, a print article, or a documentary. Quality, respect, and an engaging presentation are essential ingredients in science communication. Of the many scientific questions that have captivated me, the origin of life is perhaps at the top of the list. As is often the case when we think about life, the first image that comes to us is our own, that of humans. If we think of our beginnings, we might imagine ourselves as primates, hairy and with a clumsy gait, but [ 73 ]

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Eduard Punset working in his office in Pineda (2012). Photo courtesy of E. Punset.

little else. We know, however, that we are not the only chapter in the story of life; rather, that life encompasses infinitely more than us and that we are not even the most important beings. Many scientists have helped us to understand this. Among them is a woman, an extraordinary woman, whom I had the honor and pleasure to get to know, not only professionally on the various occasions that I interviewed her, but also as a friend. She was Lynn Margulis, who said goodbye to us on a November afternoon in 2011, leaving behind a scientific legacy that continues to allow us to learn and to know a little more every day. In the following, I have provided two excerpts from interviews and conversations with Lynn, translated into English from their original Spanish versions, about things that she not only knew well but also was uniquely able to effectively communicate them. On [ 74 ]

let’s listen to the universe | eduard punset

one of those occasions, I mentioned that there had been two key moments in the history of mankind: first, the Copernican revolution, which put an end to the idea that our planet, Earth, was the center of the universe, and second, the Darwinian revolution, which showed that mankind was neither the oldest form nor the center of life. And when I asked Lynn if she thought that humans were truly convinced that they are not the most important thing on Earth, she answered: Margulis: Of course not; there is a huge disconnect, because what scientists learn does not reach the general public, or even many other scientists. A large majority of scientists still believe

in their hearts that it is obvious that humans are the center, that we are the smartest, that we run the system, that we are

independent. This is a very political word, but people are not independent because we completely depend on plants and oth-

er organisms for food and for our air supply. Yes, I believe that there is a huge disconnect between what most people believe and what science has proven.

In another interview (number 348), which took place in 2007, we discussed almost exclusively the question “What is life?” Here is an excerpt of that discussion: Punset: Lynn, for a few thousand years we have wondered “What

is life?” If our audience does not learn today, after listening to you, what life is, they never will.

Margulis: In all likelihood they will not, because this is a question that must be answered in words and life is more than any [ 75 ]

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words, can describe; so it’s difficult. A possible definition is

that life is matter, because life surely is matter, made of carbon and hydrogen, etc., matter that comes from the stars, supernovas.... and is a flow of energy. Many people believe that life is nothing more than a recipe; you take water, amino acids, minerals, etc., mix them, and out comes life.

Punset: In one of your books, What is Life?, which I recom-

mend, even if it is not very recent, you say: “I do not really know whether life is matter surrounded by energy or energy immersed in a broth of matter.”

Margulis: We think that life started at least three thousand mil-

lion years ago and there is direct evidence of this. We wonder when life enters a fetus, but the truth is that it has never not

been there; that is, life is continuous, it is a thermodynamic

phenomenon, but a phenomenon that has never stopped from the time it started. It is a physical phenomenon as well.

Punset: You say that in the history of evolution, there is more cooperation than competition, than aggression...

Margulis: Those are words that I never use. Life is diverse in the sense that the waste of one life form is the food of another. For

example, urine is full of nitrogen, which other organisms consume. So I don’t talk about competition or cooperation because

it doesn’t make sense. When we pee, do we think we are helping the bug that uses urine [as a nitrogen source]? No. This is why I don’t refer to cooperation.

Punset: One question that interests many people is that there are

photosynthetic organisms that use the sun’s energy to produce

nutrients essential for life. Yet, others cannot carry out photosynthesis and instead must eat those who do. Why are we un[ 76 ]

letâ&#x20AC;&#x2122;s listen to the universe | eduard punset

able to [perform photosynthesis], at what moment in evolution [was it ruled out]....?

Margulis: From the viewpoint of metabolism, a human is almost the same as a mouse, a rat, or a dog. Some scientists removed

cells from a mouse and introduced plastids, chloroplasts from plants, into those cells which then became photosynthetic. But

they survived only a few weeks. And so the question is: why canâ&#x20AC;&#x2122;t we photosynthesize? Because in mammals, in animals, the selection pressure is to flee and look for food. However, in

algae and plants, which ultimately are photosynthetic bacteria, there is pressure to look for light. So if an animal stands still searching for light, it ends up being food for other beings.

Punset: So what was really lost over all those years was the possibility of a non-predatory evolutionary path?

Margulis: Yes, but algae of many kinds survived, as did plants, with this kind of non-predatory behavior opening up this path.

Punset: We are aware of lifeless rocks nearly four-billion-year-old here on the Earth.

Margulis: Yes, for example in Greenland, but it is impossible to say whether there was ever life there. In Canada, in the North,

there are rocks from more than three and a half thousand million years ago, and there is no evidence of life in those rocks.

Punset: Well, but, how is it possible that from more than three-

million-year-old rocks with no evidence of life there is now so much life? What happened?

Margulis: First we must understand that life happens at the level of the cell, which is a micron in size. Life is nothing but a

growing cell. Inside the membrane, the growth of the genetic

code begins, and so does protein synthesis, etc. All this hap[ 77 ]

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pens from the beginning within the membrane, carried out by

the system. The minimal cell has to produce its own membrane and its own proteins... and there is no cell less complex than

that. The truth is that nobody knows exactly when the single step in which rocks, minerals, and water became life took place, but we have some clues.

Punset: On the one hand, you biologists venture deeply into the

chemical origins of life and the reliance on minerals, while on the other hand, geologists, for whom the planet and celestial bodies are inert, have started to say that those bodies look as if they are alive, that they harbor certain processes.

Margulis: The first person to observe this was Lovelock, but he had not read Vernadsky, for whom life was living matter, and

who also said that it was animated water. Vernadsky said that independence is a political idea and has nothing to do with

science, because any living being independent of the biosphere dies. He had enormous vision. Lovelock says that the Earth is a living being, Vernadsky said that life is a geological

force. Within each cell there is a large pool of calcium and

phosphate, for reasons related to physiology; without control of those pools, we would eventually have stones, of calcium

phosphate, within our cells. For example, all teeth are made of calcium phosphate. Lovelock discovered why we donâ&#x20AC;&#x2122;t have

stones in all our cells, for example, in our heads. The reason is that we have cells that produce mucus and this inhibits

the production of stones inside the cell. When we remove

this mucus under very controlled conditions, we can make calcium scales, teeth, and many other things because we have removed the inhibitor.

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let’s listen to the universe | eduard punset

Punset: It’s awesome, but you’re leaving us without borders....

Margulis: It is also art. We believe that art is the work of mankind but that’s nonsense, because art began much earlier, with

many organisms, such as the design of diatoms, among others; it is so beautiful, so detailed, and it is the work of a cell. Art is a part of life that spans far beyond the limits of human life.

Punset: To the extent to which symbiosis occurs, the mixture of earth and life, of mineral and bacteria, is it easier or more difficult to define what life is?

Margulis: The notion of life has improved thanks to experiments and observations but the work still left to do will take a very

long time and is very complicated. I would never say what you said about symbiosis with rocks. Symbiosis is very simple to

define: you have living beings with different names, that is, different types that can be identified, and when they live in

physical contact, that is symbiosis. You have mites in your eye-

brows, the palm is a desert but in the armpit there is a jungle of organisms, and there are more bacteria inhabiting our mouths than people in New York City. Yes, many more, but there are

also different types. When different types of organisms merge

and create new cells by fusion or create new and different organs, when they do that, that’s symbiogenesis.

Punset: And when changes take place over time, evolving, you

also say it is not so much a genetic history of genetic changes…

Margulis: These are genetic changes that result from interactions, from permanent or temporary mergers. Genes in themselves are nothing, just chemicals.

Punset: After all this, if someday your grandchildren ask you what life is, what are you going to tell them? [ 79 ]

once upon a time lynn margulis

Margulis: Life is matter that you can handle, that you can choose, that you can move, that has motility inside it. We donâ&#x20AC;&#x2122;t think

of plants as motile, but there is a lot of motility inside them....it is matter that can change its own environment.

*** Nothing that happens today in our minds, in our lives, and in the universe can be explained without going back to the origins. This woman, who is no longer physically with us, immersed herself, and us by extension, in the origins of life in order to understand life. For this, Lynn, we owe you our gratitude! 3

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nature and nurture Mónica Solé Rojo

Ph.D., microbiologist, Madrid

I still find each day too short for all the thoughts I want to think, all the walks I want to take, all the books I want to read, and all the friends I want to see. John Burroughs (1837-1921)

We will never know whether a scientist is born as a scientist or after birth becomes one; it’s a difficult question to answer. In fact, it would be a waste of time if it were not for the fact that Lynn Margulis, one of the greatest scientific thinkers of our time, often declared, Ever since I was a child I have always wanted to be an explorer and writer, but I did not know what to explore. But now I find myself exploring the world of bacteria and writing about that. Today I cannot imagine doing anything else. She was fortunate, as she was educated at the University of Chicago, her native city, where the intellectual atmosphere and educational methods were unique at that time, in the late 1950s. It was the Great Books era at the University of Chicago, with a curriculum in which students read original works, following the development of ideas rather than academic disciplines. Science was much more a way of thinking than a body of knowledge, an intellectual approach that Lynn continued to embrace for the rest of her life. In fact, during her studies for her Master’s degree at the [ 81 ]

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University of Wisconsin, she read the cell biology and genetics literature of the late 19th and early 20th centuries. It was from this source that she developed her “crackpot” idea that the genes in the cytoplasm and those in the nucleus had different origins in evolution, a hypothesis that her advisor, the cell biologist Hans Ris, forbade her to pursue—although he eventually did so himself, after Lynn had left his group, and indeed confirmed its validity. Today the distinct evolutionary history of mitochondrial and nuclear genes is considered a scientific fact and is described in biology textbooks in use throughout the world. All of us remember those teachers and professors who awakened in us a passion for knowledge, understanding, and inquiry—those who definitively influenced us, and led us to our chosen vocation. As Carl Sagan once said: In every person is a kind of circuit which resonates to intellectual discovery—and the idea is to make that resonance work. Lynn was someone who inspired that kind of response not only in students but also in well established scientists. My time as a graduate student in her group was my first research experience. Her laboratory was always a center of activity, filled with books, scientific papers, fossils, movies, living samples, microscopes, students from different countries, discussions, lectures, poems, some of her family members, and even her dog. Life as a member of Lynn’s group was intense and very stimulating intellectually. It was an opportunity to learn not only about the scientific subjects on which we were working but also how to observe, think, formulate hypotheses, set up experiments, search for and read the right papers, write our own papers, think with an open mind, defy scientific dogmas, find colleagues to learn from, defend our ideas, etc. It was intense, exhausting, and exhilarating. [ 82 ]

nature and nurture | mónica solé rojo

All that training and experience ultimately gave me the personal skills and tools required for research and for communicating science, whether to the general public or specifically to school children, undertakings to which Lynn attached great importance. So, we combined “hard” research on spirochetes, microbial mats, protists, etc., with hands-on workshops on trash and garbage, scales of time and space, fossils in the sand, fermentation, symbiosis, etc. Lynn instilled in me a similar passion for teaching science to a non-specialized audience.

Lynn with Mónica Solé teaching a practical course in Oviedo, Spain. Photo courtesy of M. Solé

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Lynn was not only my scientific mentor and a loyal friend but also an older sister, as she used to tell me in those days back in the early 1990s. I was only 25 when I first arrived at her lab at the University of Massachusetts Amherst. I had recently graduated with a degree in Biology from the Autonomous University of Madrid. It was there that I fell in love with her books, with her ideas. A bedroom in her home, a plate at her table, and a bicycle were always ready for me, and for any of her students or anyone else in need. Unbridled generosity. Emily Dickinson’s poems, discoveries regarding rare protoctists were among the wide-ranging topics of conversation in her home. She was the most demanding professor and, at the same time, the warmest “mother” to us, her students. Always restless, she organized her life so that there would be time for an intense and fruitful scientific career but also to be attentive to the needs of others, whether human, animal, or plant.

The author collecting larvae for the production of queen bees. Photo courtesy of M. Solé

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nature and nurture | mónica solé rojo

The depth and breadth of Lynn’s knowledge were unsurpassable. I remember when she and Stephen J. Gould were co-lecturers at the opening session of a zoology conference in Boston. The audience, including myself, was fascinated by her display of knowledge in the fields of biology and evolution. As Professor Gould tried to refute her, his voice trembled with frustration. It was impossible; Lynn was always ready with some biological example or data to refute his, or anybody’s, arguments. As Geoffrey Cowley said: As the results accumulate, Margulis keeps spinning bigger ideas ... If [her] pronouncements embarrass her more solemn colleagues, Margulis couldn’t care less. Her ideas have always caused discomfort, but as the Yale zoologist has observed, they’re so rich that if one in 10 turns out to be right, that alone will change the agenda for biological research. The enthusiasm that she inspired in those around her with her tireless curiosity was unbounded. Some ten years later, when I was the head of the Scientific Programs Department at CosmoCaixa, the Science Museum of Madrid, we invited her to a symposium on evolution. At the end, people crowded together around her for an autograph, a short chat, a photo, or just a chance to touch her! She conveyed a passion for knowledge, for thinking, and for rebelliousness against long-held dogmas. She made us step outside of our human dimension in order to understand the fundamental role of microbes in evolution and in the environment. Many of her ideas have been confirmed and have found their place in school textbooks in Spain; her books are on the shelves of every science library. Her voice resonates in the classrooms of our universities. And, inspired by our first meeting, there is a microscope in my home together with test tubes full of mud and microbes. When she happened to be in Madrid, she always had [ 85 ]

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a room in my home; now she has a place in my heart, filled with emotion and gratitude. Today, after working for many years in basic research in microbiology, as well as in the dissemination and social application of science, I have set off on a new career path, one that strongly focuses on the environment, specifically on the production, breeding and genetic line selection of queen bees. Unfortunately, Lynn barely had time to learn the details of my new interest. Iâ&#x20AC;&#x2122;m sure she would have encouraged me in my efforts. They allow me to spend much of my time close to natureâ&#x20AC;&#x201D;a joy that I know Lynn would have gladly shared with me. 3

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teaching in the rain Martí Domínguez University of Valencia

Al meu pais la pluja no sap ploure: o plou poc o plou massa; si plou poc és la sequera

si plou massa és la catàstrofe.

Qui portarà la pluja a escola?

Qui li dirà com s’ha de ploure? In my country rain doesn’t know how to rain: either it rains too little or too much; if too little, there is drought,

if too much, there is disaster.

Who will send the rain to school? Who will teach it how to rain?

Raimon (singer-songwriter)

Someone like me, a dedicated biologist with a strong interest in entomology and botany, could not have remained indifferent to the contributions Lynn Margulis made to these branches of science. In addition to her research activity, scientific contributions to cell biology and evolution, and her revolutionary ideas, I admired Lynn’s natural ability to communicate, easily conveying her enthusiasm and thrilled to share her knowledge, not from a magisterial position, but person to person. [ 87 ]

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We shared several interests, including the history of science, literature, philosophy, and science dissemination. I started my career writing popular-science articles for a weekly magazine, El Temps. Later I became Editor-in-Chief of Mètode, a popular science magazine published quarterly by the University of Valencia. Thanks to the efforts of the journal’s team and the quality of the authors’ contributions, Mètode has become a reference in science communication and the recipient of several awards. My goal when I took over the journal was to promote knowledge of and critical reflection on recent scientific developments and events, expanding the content beyond a summary of the research carried out at the University of Valencia. We are proud to have published several contributions by Lynn. With her insights, we recognized the effect of using color to enhance scientific illustrations, whether photos or drawings. Lynn’s lectures and talks were always accompanied by extraordinary visual materials. She felt that if those resources exist, why not use them to show the wonderful world to which we have dedicated our professional lives? Articles on science are granted very little space in the press and other media. In newspapers and magazines most of the attention is paid to discoveries with clinical implications and to remedies for diseases. In many cases, what passes as news is no more than very preliminary information on new drugs or treatments under investigation and whose translation into clinical practice will take years, if they ever become available at all. Science is culture and Lynn understood it this way. She wrote for Mètode on at least three occasions. In issue no. 38 (2003), in her article “Symbiosis and termites” she wrote “Termites belong to a family of wood-eating insects that live in symbiosis with bacteria and protists that live in their digestive tract.” Two more articles by [ 88 ]

teaching in the rain | martí domínguez

Lynn Margulis and Ernst Mayr, whose obituary she published in the journal Mètode, no. 45, 2005. Photo courtesy of Mètode.

her were published in 2005: an obituary of Ernst Mayr (see photo), who had died early that year, and an article entitled “Syphilis and Nietzsche’s madness,” in which she wrote that, after careful study and documentation, “there is detailed evidence to suggest that the philosopher should have passed through each of the three stages of syphilis: the chancre of primary syphilis, immediately after infection; the terrible pox, fever and pain in secondary syphilis, which emerges months or years later; and the dreaded third phase: paresis.” The philosopher’s disease was the topic that she used to describe her explorations of disease mechanisms but above all, of the origin, morphology, and function of a group of microbes for which she always showed a preference, spirochetes. [ 89 ]

once upon a time lynn margulis

In 2001, when Lynn Margulis was awarded an Honoris Causa Doctorate degree by the University of Valencia, two researchers at the university, Francesc Mezquita and Antonio Camacho, interviewed her for Mètode. In that interview, published in issue no. 31, Lynn described both her personal journey as a scientist, from her early interest in science to her relationships with the many scientists with whom she collaborated throughout her long career, and the ideas that led to the development of her theories. Importantly, she also highlighted the names of the forgotten scientists whose work she would later expand upon in some of her most highly recognized scientific achievements. I remember back to 1990, when I attended a lecture by Lynn that had been organized by the Catalan Society for Biology at the University of Valencia. From the beginning, I was seduced by her words, her spontaneous smile, and the twinkle in her eyes. I was impressed by her innate intelligence and her ability to deeply relate to people. She understood human nature as well as she understood the microorganisms she studied. And she was determined. I remember a field trip we made to the Prat de Cabanes Marsh, in Castelló, with Juli Peretó. It began to rain cats and dogs but that didn’t matter to Lynn. She was too excited about what she might find in the soil. Indeed, she was absolutely oblivious to the rain, although our clothing was quickly soaked. Juli, at that time Vice-Rector of the University of Valencia, was in the car answering phone calls and did not have a moment of rest. Suddenly Lynn realized that it was certainly raining very hard and decided that she had collected enough samples. We ran, dripping wet, to the car, laughing and shouting. I had to turn on the car heater to avoid us getting chilled and to defog the window so that [ 90 ]

teaching in the rain | martĂ domĂnguez

we could drive home. If we ever meet again we will remember that day. And I remember you, Lynn, not just as a visionary biologist, but as a delightful human being as well. 3

[ 91 ]

swimming with bacteria Núria Gaju

Autonomous University of Barcelona, Bellaterra

Potser el secret és que no hi ha secret i aquest camí l' hem fet tantes vegades que ja ningú no se'n sorprèn; potser caldria que trenquéssim la rutina fent algun gest desmesurat, alguna sublimitat que capgirés la història. Potser, també, del poc que tenim ara no sabem fer-ne l'us que cal; qui sap. Maybe the secret is that there is no secret and we have taken this road so many times that no one is surprised; perhaps we should break the routine, make an excessive gesture, something sublime that overthrows history. Perhaps, too, of the little we now have we do not know its proper use, who knows. Miquel Marti i Pol (1929-2003), L' àmbit de tots els ámbits

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Microbial ecology is a field of biology that studies the many interactions that occur in nature. Interactions among microorganisms have long been recognized but initially they could be interpreted only from observation and deduction. Similar interactions probably led to partnerships at the beginning of life on Earth, some of which stably persisted. In the fossil record, stromatolites, which have their current counterparts in microbial mats, have provided an approach to understanding the first steps in the evolution of life and the establishment of the first ecosystems on our planet. The endosymbiotic theory proposes that the ancestors of at least two types of eukaryotic organelles—mitochondria and plastids— were independent prokaryotic organisms that became permanent endosymbionts of other cells. This is what, in 1986, before defending my thesis, I read with great interest in the book Early Life by Lynn Margulis, published in Spanish by Editorial Reverté as El origen de la célula. For two decades, Lynn had been defending this theory, which thanks to advances in microscopy and molecular biology is now well established but still subject to modification—as is all scientific knowledge—pending newer findings. The endosymbiotic theory explained the origin of mitochondria in an association between aerobic and anaerobic bacteria, which at the beginning might have been causal but became stable thereafter. That first relationship between these two types of microorganisms may have started as one between predator and prey. The study of predation among prokaryotes was one of the topics of my doctoral thesis. In the mid 1980s, talking about it with Lynn, she expressed intense interest in the work and encouraged my research. I was very grateful for her support especially since when one is at the initial stages of one’s own research it is unusual to receive this kind of attention from [ 94 ]

Swimming with bacteria | nĂşria gaju

admired scientists. She showed the same enthusiasm towards the focus of our labâ&#x20AC;&#x2122;s work, the predatory bacterium Daptobacter. We were studying prokaryotic predation in anoxygenic lakes and the control exerted by predatory bacteria on phototrophic bacterial populations

Lytic plates of predatory bacteria on a Chromatiaceae lawn filter. Photo courtesy of N. Gaju.

in these habitats. At that time, our goal was the taxonomic and functional characterization of natural microbial communities (in Lake Banyoles, Lake Estanya and in the Ebro Delta, Catalonia, Spain) in order to better understand the factors that regulate their behavior. [ 95 ]

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More recently, our research has expanded to the applied aspects of this knowledge of natural ecosystems, both planktonic and benthic; for example, the impact of contaminants such as oil and pesticides on natural communities. In all their many different aspects, whether basic research or potential industrial applications, microbial relationships have been of fundamental importance in our studies.

Light microscopy image of field samples showing predator cell attached to the surface of Chromatium sp. Photo courtesy of N. Gaju.

Transmission Electron Micrographs of thin sections of Daptobacter from samples taken in Lake Estanya. Photo courtesy of I. Esteve.

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Swimming with bacteria | núria gaju

After Lynn’s first visit to the Autonomous University of Barcelona, around 1982, we met often in what, at least for me, were very rewarding encounters. It is very difficult to fully describe her incredible career, as a researcher and as a teacher, and it is equally difficult to describe her generosity and human qualities. What can I say about her contribution to science? In changing our picture of the evolution of life on our planet? Her work, her expansion of our knowledge, and the details of her discoveries can be read about in the books and articles that show her wide range of interests and motivation. But her activities as a teacher and a disseminator of science around the world were no less important. My enthusiasm for her work can be traced back to 1985, when I attended her course “Origin and Evolution of Cells” held at our university. In the summer of 1988 I was awarded an internship to work with Dr. Alan Liss at the State University of New York at Binghamton in the NASA-PBI Program. We were searching for the presence of a calmodulin-like protein in Spirochaeta, as evidence of the endosymbiotic origin of undulipodia. During that time, I also had the opportunity to visit Lynn, both at the University of Boston and at UMass Amherst, where she moved that year. Lynn helped me in the development of my research and I have continued to benefit from her advice, which also deepened my appreciation of the microbial world. She was always a welcoming host, opening the doors of her house to me. I cannot remember any encounter with her that was not interesting. Lynn was fortunate to have a very large expanded family, one whose members included colleagues and friends from around the world, and she was very devoted to us. I will never forget her, not only for her contributions to science and our own field of work, but also for the woman she was. 3 [ 97 ]

bitten by the science bug Laura Villanueva

Royal NIOZ, Den Burg-Texel, The Netherlands

As a horse after a race, and a hunting dog when he hath hunted, and a bee when she hath made her honey, look not for applause and commendation; so neither doth that man that rightly doth understand his own nature when he hath done a good turn: but from one doth proceed to do

another, even as the vine after she hath once borne fruit in ther own proper season, is ready for another time.

Marcus Aurelius (121-180), book V of Meditations

In 1999, I attended a meeting organized in the old building of the Institute of Marine Sciences in Barcelona. At the time, I was a sophomore at the University of Barcelona and it was my first encounter with Lynn. Seeing her in person was quite a big deal for me, as her ideas, and the curiosity and enthusiasm that she conveyed in her books and articles had led me to change my decision from entering medical school to studying biology. I remember I was running late for her talk; I ended up taking a taxi and then sprinting through the streets of Barcelona... and although I had never followed any actor/musicianâ&#x20AC;&#x2122;s career I was now unashamedly Lynn Margulisâ&#x20AC;&#x2122; groupie! During that talk (and in the many others of hers that I heard over the years), I could not help but feel lucky to be a scientist, even one still in training. She explained things in a fascinating and even humorous way. I was surprised to hear her speaking Spanish, [ 99 ]

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frequently stopping to ask for the audienceâ&#x20AC;&#x2122;s help in remembering a word she had forgotten. That was typical of her unique personality. After that talk I decided I would do whatever it took to study those wonderful ecosystems that had been involved in early life and the evolution of microbial strategies. After asking here and there and reading everything that fell into my hands related to the topic I was sent to talk to Prof. Ricardo Guerrero (Lynn and Ricardo had initiated a scientific collaboration years ago) and it was in his lab that I started my scientific career, receiving my PhD under his supervision. Lynn often came to Barcelona, invited to participate in different activities (scientific and publishing meetings, workshops, field work, conferences, etc.). During those visits she often lectured at the University of Barcelona. The students crowded the Aula Magna, overfilling its capacity for 360 people, just to hear her. She was our source of inspiration. At the end of those lectures, you could see people leaving with bright eyes and full of research energy. It was on one of those occasions that we were introduced officially, becoming better acquainted over dinner. From that moment onwards we became not just colleagues but also friends. I was thrilled in that although I was just a student she took my opinions seriously. Over the years, as I worked on my thesis, we met several times. Almost coincidentally, I ended up doing a post-doc in the Department of Microbiology of the University of Massachusetts-Amherst (U-Mass), which meant having Lynn as a work neighbor. That was a truly great but also difficult time for me. I was not very happy about the course of either my life or my scientific career and my chats with Lynn over coffee in her wonderful office were often a â&#x20AC;&#x153;lifesaver.â&#x20AC;? We used to talk about spirochetes, our favorite microbes, which she had studied and described so well in many of her [ 100 ]

bitten by the science bug | laura villanueva

books. Those microorganisms that â&#x20AC;&#x153;look like tightly coiled snakes,â&#x20AC;? as she described them. They can be found, and she did, in marine and fresh waters, deep muddy sediments, and in the gastrointestinal tracts of several different animals. Spirochetes include the genera Spirochaeta, Treponema, Borrelia, and Leptospira. They are a group of motile spiral-shaped bacteria. Some of them are causative agents of diseases, such as syphilis (Treponema pallidum) and Lyme disease (Borrelia burgdorferi). Lynn, like many other microbiologists, abhorred the study of microorganisms as pathogens. Firstly, because within this great diversity of microorganisms there are only a few pathogens, and, secondly, because microorganisms are of extraordinary interest in the study of life and evolution. We also talked a lot about the microorganisms living in microbial mats, a keen area of interest that we shared. For my thesis, I had studied the ecophysiological and mo-

Microbial mat from the Ebro Delta. Photo by R. Guerrero.

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lecular characteristics of estuarine microbial mats, in an approach that revealed many of the microbial strategies for survival in a broad range of environments. I understood why microbiologists and other scientists from related disciplines were fascinated by microbial mats: these prokaryotic communities are modern-day representatives of what might have been the first ecosystems on Earth. To understand the function of a microbial mat as a community, the physical and chemical microenvironments in which the microorganisms live must be well known and in detail. The community just below the mat surface experiences steep vertical gradients of light intensity and redox conditions that change markedly during the diel cycle. Indeed, motile photosynthetic organisms optimize their position with respect to the resultant light gradient. One can enter into the study of a microbial mat and never stop because of its seemingly endless number of extraordinary features. But in Lynn’s office we also talked about life. As women and scientists, we struggled with similar challenges: Can I be a good, devoted scientist and have a family? Can I fulfill all of my personal and professional desires? When you are in your mid-twenties you are not terribly worried about those things but then you grow older and have to make choices. You fall in love, have your heart broken, and all the while have to keep on going with your research as if nothing had happened. Certainly, Lynn had endured similar experiences and was generous in sharing them. When I read Lynn’s book Luminous Fish I was relieved to learn that I was not alone in my feelings. I was grateful that she wrote that wonderful guide for all of us, especially women who have been bitten by the science bug. In Luminous Fish she said: “You can’t be a good wife, good mother and good scientist. One of them has to go.” [ 102 ]

bitten by the science bug | laura villanueva

Scanning electron micrograph of the microbial diversity of a spirochete bloom obtained from Ebro Delta microbial mats, filtered on a 3-µm-filter. Photo by L. Villanueva.

I remember with great joy one of Lynn’s birthdays—I was still at the U-Mass—when I gave her a scanning electron micrograph of a bloom of microbial mat spirochetes (Spirosymplokos) from the Ebro Delta (see photo). Apparently, presents were not to be given on Lynn’s birthday but spirochetes always melted her heart. After her death, someone from her lab contacted me to ask me if I wanted to keep that photo, which had been hanging in her office ever since. I could not think of anything better to remember the time we spent together. As I type these words I look at that photograph, now hanging in my office. Her energy and inspiration underlie my decision to continue in science. Lynn, you have left us, but so much of you remains with us. 3 [ 103 ]

a woman who left her mark Rubén López and Concepción Ronda

Center for Biological Research, CSIC, Madrid

Una luciérnaga entre el musgo brilla Y un astro en las alturas centellea,

Abismo arriba, y en el fondo abismo,

¿qué es al fin lo que acaba y lo que queda? A firefly glows among the moss And a star sparkles on the sky, Abyss above and below abyss,

What is it finally that finishes and what is it that remains?

Rosalía de Castro (1837-1885)

The death of Lynn Margulis, on November 22, 2011 was a tragic and unexpected loss. Among her many friends, associates, and colleagues, the desire arose to express and to share, if only briefly, our stories of our professional and personal experiences with Lynn. It would be a way to acknowledge her influence, on us and so many other people, and to take note of the recognition she received in Spain, specifically in Catalonia where she spent long and frequent research stays. For a microbiology enthusiast like me (Rubén López), who in 1967 was preparing his PhD thesis, the news that Lynn Margulis had published an article in which she stated that the eukaryotic cell was the result of successive symbiogenetic inputs of different pro[ 105 ]

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karyotic cells was truly shocking. This was a brilliant demonstration of the seminal role of microorganisms in major evolutionary events. Lynn soon became a scientist with cult status. Maybe this consideration explained why it was that in 1987 I hesitated to send her, as an editor of the Proceedings of the National Academy of Sciences, the manuscript that I had co-authored with other members of my research group at the Institute for Immunology and Microbial Biology. My closest friends warned me of her strictness and thoroughness when reviewing manuscripts for a journal so coveted by scientists. These expectations were fulfilled. Lynn took many months to accept my group’s work, since in the two first evaluations the manuscript did not meet her requirements of quality. Indeed, she decided to call in a third referee, who finally recommended publication. Shortly afterwards, Concha (my wife’s short name), also a scientist, and myself met Lynn in Madrid, at a congress in which I presented research with data that expanded on the PNAS paper. Lynn was in the audience and I had the satisfaction of telling her that the journal’s acceptance of our manuscript had launched our careers, as it brought the recognition of the scientific community. That day, we shared lunch with her, Ricardo Guerrero—one of our dearest friends— and Antonio Lazcano, two of Lynn’s close friends. The four of us will never forget that lunch with her, during which we could appreciate her enthusiasm, wit, and humor. Since then, seeing Lynn during her stays in Madrid was always a pleasure for Concha and for me. She visited our house on several occasions, once because she had to change her dress after having spilled something on it. We share this anecdote because it exemplifies her behavior, her affectionate spontaneity with so many people, who loved and admired her. Her bonhomie alternated with the fi[ 106 ]

a woman who left her mark | ruben lópez & concepción ronda

ery passion with which she sharply defended her professional contributions and her ideas. She was truly a woman who left her mark on the world. About the origin of the eukaryotic cell, she proposed that both the chloroplasts, that provide us with oxygen, and the oxygen-breathing mitochondria evolved from independent former bacteria as a consequence of symbiosis. Later, molecular biology fully confirmed her daring proposals, proving them scientifically sound enough to explain the appearance of the eukaryotic cell. Currently, school textbooks include those concepts, a measure of the scientific impact of her ideas. After Lynn’s visits, Concha and I frequently commented on her extraordinary humility. We knew that, in her frequent visits to Barcelona and also to Madrid, she missed being able to get around by bike as easily as she did in Amherst. We knew that, besides walking, the bicycle was her favorite means of transportation for short (and not so short) distances, regardless of the weather. Concha, besides her scientific research, was interested in the application of new information technologies to research. In this field, she coordinated projects such as the development of science and technology indicators and quality assessment measures. These assessment tools would reflect the social and economic impact of scientific activity, something of particular interest and importance in our country. They would also illustrate the paradigmatic shift in scientific communication processes that followed the development of the Internet. The information they provided would allow the development of a web portal on science and technology in Europe. Lynn showed her appreciation for this work. She was an inveterate user of any technology that would allow her to store in an organized and accessible manner the enormous amount of scientific informa[ 107 ]

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tion that she was continuously gathering and generating. In 2004, the director of our Center for Biological Research asked me to suggest a prestigious scientist to participate in the events surrounding the inauguration of our new institute, located

Lynn Margulis and Rub茅n L贸pez during her visit to the Center for Biological Research in Madrid, part of an invitation to deliver a lecture celebrating the 50th anniversary of the Center. Photo courtesy of R. L贸pez.

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a woman who left her mark | ruben lópez & concepción ronda

on the campus of Complutense University. I had the honor to invite Lynn Margulis. During her lecture she defended the relevant role of the “uptake of genomes” through symbiogenesis, as a large mobile element that contributed to driving evolution. Certainly, a controversial idea. But to Lynn, proposing new ideas based on her observations was always her banner. Less than a year before her death, we attended a lecture she delivered at an international meeting in Madrid. Although clearly suffering from a bad cold, once on stage and responding to the incisive questions from the audience, there emerged the extraordinary scientist, who seemed to have miraculously recovered her strength in order to staunchly defend her most recent insights. It was the last opportunity we had to admire this singular woman, who once had said: “Life is bacterial, and those organisms that are not bacteria have evolved from bacterial organisms.” As a scientist and human being, Lynn shared our concerns about the problems of our times and the society in which we live, including the emergence of new diseases and the reemergence of several infectious diseases. Today, infection has to be studied and treated as an ecological process, as a strategy to control pathogenic microorganisms. Lynn agreed with our emphasis on the importance of establishing strong interdisciplinary links among researchers. She understood the need to look for long-term measures based on accurate basic research into the evolution of resistance and molecular analyses of the microorganisms involved in infectious diseases. As a scientist, Lynn was imaginative, brilliant, and combative. As a friend, empathic, literate, and warm-hearted. 3

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not gone with the wind Bego単a Vendrell

Institute of Marine Sciences, CSIC, Barcelona

Nuestras horas son minutos cuando esperamos saber, y siglos cuando sabemos

lo que se puede aprender Our hours are like minutes when we wish to learn,

and like centuries when we know what can be learned

Antonio Machado (1875-1939)

I remember the first time I saw Lynn. I was a high school student very interested in ecology and my wish was to go to university to study biology. In high school, thanks to my biology teacher, I was fascinated by the endosymbiotic theory, as a creative process, and by the Gaia theory. I even carried my fascination home with me, where I spoke enthusiastically to my parents about these amazing ideas so important to understanding life and its evolution on our planet. I must have talked so much about these subjects because my parents, knowing that Lynn Margulis was going to deliver a talk at CosmoCaixa in Barcelona, insisted that I attend. I returned ecstatic; that bright-eyed woman conveyed her ideas with such a passion but yet so simply that any doubts were immediately banished. I especially re[ 111 ]

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call that she referred to Charles Darwin as papรก Carlos and I have to confess that since then I have sometimes made use of that expression. I want to emphasize the importance of that first encounter for a young student like me, who early on had to decide on the course of her future. The importance lies in the enthusiasm and sincerity that I perceived in Lynn, who clearly loved what she did and truly believed in it. Today, as a researcher, I look for those same didactic qualities in high-level scientists and especially from our own teachers. Those are the qualities that drive students to pursue their careers with the same dedication. In 2004 I attended her course on Gaia at Schumacher College in the south of Devon, UK. Lynn turned out to be a very warm-

Lynn and the author during a walk around the fields of Schumacher College. Photo courtesy of B. Vendrell

[ 112 ]

not gone with the wind | begoña vendrell

hearted, very affable person, an excellent communicator, still passionate about her work and multifaceted. It struck me how she was able to transmit this passion to us, and how she always surprised the audience with comments on other subjects —not always related to biology— telling jokes or reciting poems by Emily Dickinson, whose poetry I came to know thanks to Lynn. Shortly after the fall of that year, Lynn visited the University of Barcelona to participate in a symposium on the origin and evolution of life. The meeting was being held at the Faculty of Biology and had been organized by members of Omnis cellula, including myself. I was shocked to see the crowded lecture hall: people had even crowded the windowsills and stairs! Above all, I remember that many young people had come also from other faculties (for example, my brother; although he was a mathematician, he was fascinated by Lynn’s ideas and eager to attend her lecture). I am indebted to Lynn because she strengthened in me a creative and holistic way of thinking. She made me intellectually open-minded and helped me to develop a perspective of a closely connected world. Inspired by her work, I consider ecology and the relationships between organisms to be at the basis of the evolutionary process. The idea that life has the capacity to modify and create new habitats and resources is closely linked with the concept of Gaia, and for me it cannot be separated from Lynn. She claimed that bacteria and other microorganisms were essential actors in the evolution of life on Earth. This idea marked a very subtle boundary between the organic and the inorganic, and acknowledged that not all interactions among organisms and between them and the environment are known. To understand Gaia in its globality can be difficult. Gaia, as Lynn said, was symbiosis seen from space; life as a product of the interaction be[ 113 ]

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tween cells is the poetry of autopoiesis. Some of these ideas, for example, that over time the environment has become more specialized and organized, can, —from my perspective as an ecology student— be related to those of Margalef's regarding ecological succession and the theory of boundaries. In recognition of this link, I decided to incorporate this line of thinking into my doctoral dissertation. For me, Lynn was one of the few people able to think creatively, seeing the world from a global perspective, and relating reductionists schemes in science to this globality. She was also a source of courage and inspiration: for her tenacity, her strength when confronted with adversity or heated criticism, and for ever being amazed by Life. I must also thank her for “introducing” me to the work of several Russian scientists, whom I never would have read had it not been for her. Three key words to define you, Lynn are warmth, creativity, freedom. 3

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sharing knowledge , sharing life Jordi Mas-Castellà

International Microbiology, Barcelona

…“It is your garden now, little children”, said the Giant,

and he took a great axe and knocked down the wall. And when people were going to market at twelve o’clock they found the Giant playing with the children in the most beautiful garden they had ever seen.”

Oscar Wilde (1854-1900), The selfish giant

I first met Lynn, at the Autonomous University of Barcelona (UAB) in the late 1980s, during one of her frequent visits and long stays. Our group in the Microbiology Department was studying microbial mats and the physiology of phototrophic bacteria. By then Lynn was already an internationally renowned scientist, based on her endosymbiotic theory to explain the evolution of the eukaryotic cell. I should mention that, at the time, science in its purest state could be found in the laboratory of Ricardo Guerrero and his group at the UAB, which I had joined as a graduate student. There, in addition to Lynn, I met James Lovelock, Julian Davies, Alex J. B. Zehnder, Hans Trüper, Rita Colwell, Thomas D. Brock, Elio Schaechter, and many other great microbiologists, invited by Ricardo. Microbial mats had been one of the main research topics in our laboratory for many years. Several members of that original group [ 115 ]

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now occupy important positions in different research centers and have continued working on various aspects of microbial mats, taking new approaches and using more advanced techniques. Microbial mats form at sediment-water interfaces, have a thickness ranging from several millimeters to a few centimeters, and consist of vertically stratified communities of benthic microorganisms, mainly distinct photosynthetic bacteria. The upper layers of the mat are responsible for oxygenic photosynthesis and are thus usually composed of oxygenic photoautotrophs such as cyanobacteria and diatoms. Below are layers of anoxygenic photosynthetic bacteria, mainly species of purple sulfur and green sulfur bacteria. Phototrophic organisms place themselves at different positions in these structures according to the vertical distribution of environmental factors. The stability of this system would not be possible without the action of sulfate-reducing bacteria.

Lynn Margulis collecting samples from microbial mats of the Ebro Delta. Photo courtesy of R. Guerrero.

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sharing knowledge, sharing life | jordi mas-castellĂ

The fact that microbial mat communities are strictly prokaryotic makes them excellent models for studying the origin and early evolution of life, a subject that fascinated both Lynn and the group led by Ricardo Guerrero. In natural environments such as freshwater stratified lakes, phototrophic prokaryotes also form multilayered communities, reproducing the typical structure of a microbial mat. The nature of this physical structure determines the stability and the persistence of the layering, as well as the scale of the lamination. I did much of the experimental work for my thesis at the University of Massachusetts, Amherst. There, in the Department of Geoscience and in the Department of Biochemistry, I received generous help from Lynn and the other members of her group. Some of them, including Steve Goodwin (currently Dean of the College of Natural Sciences), Greg Hinkle, Lorraine Olendzenski, and Betsey Dexter Dyer, spent research stays at the Autonomous University of Barcelona and we shared many adventures, standing in muddy sludge up to our knees to sample microbial mats. Those years were the beginning of long scientific relationships and lasting friendships. The subject of my thesis, the biosynthesis and accumulation of biopolymers by bacteria inhabiting microbial mats, had become a hot topic in the field of microbial ecology. Lynn was the most passionate scientist I have ever met. She reminds me in some ways of Erin Brockovich, as portrayed on screen by Julia Roberts. She was so enthusiastic about what she was defending that it was impossible not to join her. Why would such a woman spend so much time and personal effort to convince the world that many millions of years ago different cells merged to give rise to a new kind of organism? She was science personified; defending her view of the truth among the ever-present critical [ 117 ]

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An evening meeting with members of the Cercle d’Amics de la Ciència (Friends of Ciencia Club) of the Catalan Foundation for Research in Barcelona in 2003. R. Guerrero, L. Margulis, and J. Mas. Photo by M. Piqueras.

voices. She taught me that a good scientist must also be a good communicator. If you cannot tell the world what you have discovered, what your contribution to human knowledge is, then you are not 100 % a scientist. Scientific discoveries are news worth spreading and are a part of the activities that define a scientist’s career. Lynn spent a good part of her life informing the world about the implications of her greatest contribution to science: the endosymbiotic theory of the origin of the eukaryotic cell. But Lynn was a very special person in other ways as well. She influenced everyone she met—the time you spent with her was not neutral, was not quiet. Given her readily conveyed abundance of energy and enthusiastic spirit, I was amazed to realize how close she was to her children and family. When she talked about them, she spoke with the voice of a loving mother and grandmother. She [ 118 ]

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loved every minute of life. She was the model combination of a brilliant mind and a very warm heart. In the early 1990s, she was my host during my stay in Amherst, when I worked with Clint Fuller at the Biochemistry Department at U-Mass, experimenting with the polyhydroxyalkanoates accumulated by phototrophic bacteria. She was a tremendous help and I am grateful for all that she did for me. Lynn and I co-authored, together with other Spanish scientists, a paper published in the journal Microbiologia SEM (vol. 11: 397-399), about the survival mechanisms of bacteria in microbial ecosystems. Her spirit, her willingness to defend what she thought was right will always be a strong part of my memories of her. She represents for me the survival of truth, with a voice louder than the voices of those who believe in the immutability of life. 3

[ 119 ]

a groundbreaking Balbina Nogales

University of the Balearic Islands, Palma de Mallorca

Mais les brav’s gens n’aiment pas que L’on suive une autre route qu’eux,

Non les brav's gens n'aiment pas que L’on suive une autre route qu'eux, Tout le monde se rue sur moi,

Sauf les culs-de-jatte, ça va de soi But the good folks don’t like it

If you take a different road than they do No, the good folks don’t like it

If you take a different road than they do Everyone pounces on me

Except those with no legs: that goes without saying.

George Brassens (1921-1981), La mauvaise réputation

The area around Lake Banyoles comprises several ecosystems of great interest. Not only from the point of view of science but also socially and economically because it offers many leisure and sports opportunities. A large number of international scientific publications have originated from work carried out at Lake Banyoles. Among the renowned authors of those papers is Ramon Margalef (1919–2004), who carried out ecological studies there. By the time I started to [ 121 ]

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collaborate with members of the Microbiology Department of the Autonomous University of Barcelona (UAB), during the second half of the 1980s, the work of our group and that of many others researchers had already caught the attention of Lynn Margulis. Within the karstic Banyoles area, the small Lake Cisó, with its unique characteristics, was the subject of ongoing studies by the microbial ecology team in which I completed both my undergraduate studies and my doctoral thesis. In aquatic systems, as in most natural environments, microbial populations form communities that establish multiple interactions with abiotic and biotic components and, either plants, animals, or other microorganisms. The relations between populations in communities are determined, among other factors, by the available resources, in addition to the physicochemical conditions of the environment. For Lynn, the interactions that were essential to evolution were mutualism and cooperation. Such interactions result in a more efficient use of resources and are indicative of successful evolutionary practice. These were also essential to her own work in which she traced several elements and functions of eukaryotic organisms to their prokaryotic origins. In Lake Cisó, important populations of the ciliated Coleps hirtus were detected in stratified waters. The cytoplasm of these ciliates contained algal endosymbionts, which we observed and subsequently identified as Chlorella vulgaris. Other endosymbionts of the genus Chlorella were also observed in the ciliate Prorodon. Betsey Dyer, who had been a student of Lynn’s, was actively involved in identifying these organisms during her stay at the UAB. Their algal endosymbionts provide Coleps hirtus and Prorodon with organic matter and oxygen from oxygenic photosynthesis, [ 122 ]

a groundbreaking | balbina nogales

Cover of the book Environmental Evolution, edited in 1992 by L. Margulis and Lorraine Olendzsenski. A second edition of the book appeared in 2000. The picture on the front cover shows blooms of Chromatium, a phototrophic purple-sulfur bacteria that periodically turns the surface of Lake Cis贸 red.

thereby affording them trophic versatility, as these ciliates feed on detritus, bacteria, algae, and other ciliates. Interactions between these microorganisms can be considered mutualistic because the algae also benefit from them. [ 123 ]

once upon a time lynn margulis

On the contrary, my thesis, was dedicated to examining antagonistic relationships involving photosynthetic anaerobic bacteria in Lake Cis贸, main primary producers in the lake. Certainly, under natural conditions negative interactions, including competition and parasitism, are also important for example as factors influencing populations size and dynamics. My research demonstrated the presence in Lace Cis贸 of heterotrophic bacteria able to cause the lysis of the photosynthetic bacteria of the genus Chlorobium and capable of growing at expenses of these photosynthetic cells. Colocalization of antagonistic heterotrophic bacteria with maximum population densities of Chlorobium in the stratified lake provided good evidence of the relevance of this type of interactions in this peculiar ecosystem. That was a productive time for me as a young scientist, and the experience that I gained there has led me, over the years, to highlight the importance of microbial ecology as a field of basic research. Our work and that of others contributes to safeguarding the environment and to effective environmental remediation subsequent to the impact of human activity. Today, in our group at the University of the Balearic Islands, our focus is on the role of microbial communities in the recovery of marine ecosystems impacted by acute or chronic hydrocarbon pollution, which is the major source of pollution in marine ecosystems. In the summer of 1991, I attended the 7th International Symposium on Photosynthetic Prokaryotes, which was held at the University of Massachusetts in Amherst. Lynn invited me, a young and quite shy doctoral student, to the dinner she had prepared in the garden of her home for a large group of scientists. I could never have imagined being among those important people but I enjoyed it immensely! [ 124 ]

a groundbreaking | balbina nogales

My personal remembrances of Lynn can be summarized mainly by her: 1. Curiosity. This was easily discovered simply by looking into her eyes and by her spirited approach to all that stimulated it. 2. Determination and dynamism. She would start her lectures by asking the audience questions, and she would not stop asking until she got an answer. For Spanish students, who at least during my student days were almost unanimously reluctant to speak in public and almost always tried to hide somewhere in the crowd, this was often a quite uncomfortable experience. 3. Proximity to students. Despite being a highly renowned scientist, Lynn was always keen on having contact with young students. 4. Generosity. For example, although she was always very busy, she took extreme care in editing the first scientific paper that I wrote. Her efforts greatly improved the manuscript in many ways and taught me a lot about how to write a paper. In fact, more than 90 % of the paper was red-inked. But her comments and corrections were so constructively formulated that I was not completely depressed after receiving the corrected version. It taught me a lot and I was extremely grateful to her for that. I will always remember Lynn as someone very special and I am glad that I had the opportunity to meet her personally. If I have to define her in keywordsâ&#x20AC;&#x201D;an easy task!â&#x20AC;&#x201D;I would say she was a free spirit. Free enough to be able to break through dogmas and free enough to recognize what she ignored. 3 [ 125 ]

a woman for all seasons Carles Pedrós-Alió

Institute of Marine Sciences, CSIC, Barcelona

L’únic important és convertir-nos en protagonistes i deixar de ser espectadors. No sé si canviaré res en la meva vida però la immensitat blanca del desert d’aigua restarà sempre a prop, només a un petit vol de la imaginació. Sempre a punt per a quan calgui.

The only important thing is to become involved and to stop being spectators. I do not know whether I will change

anything in my life, but the white vastness of the desert of water will always remain close to me, only a short flight of my imagination. Always ready whenever it is needed.

Carles Pedrós-Alió, Desert d’aigua

Many years have passed since I first met Lynn Margulis, and many changes have occurred since that time when few things appeared clear to the student I was then. Today I appreciate and recognize the circumstances that led me down the path of science. I am a microbiologist by training, but I became very interested in ecology, and microbial ecology is one of those fields that fall in between disciplines. This is a challenge but a very stimulating one because microbial ecology provides us with a view of the ecology of microorganisms that finally reveals the principles underlying their distribution and behavior. Over the last few decades I have focused my [ 127 ]

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research on marine microbiology. My team and I have carried out most of our work in the sea, with marine plankton (phytoplankton, bacteria, archaea, flagellates and ciliates) and their genomes as the subject of our work. The sea is the largest ecosystem on the planet and, at the same time, the least known. Each milliliter of seawater is inhabited by a thousand living protists, about one million bacteria and archaea, and close to ten million viruses. These planktonic organisms carry out the bulk of photosynthesis and respiration in the sea and therefore are responsible for the balance of CO2 between the ocean and the atmosphere. About half of the oxygen we respire comes from marine photosynthetic organisms, and more than 95 % of respiration in the oceans is due to microorganisms. Some members of the microbial plankton, such as diatoms and dinoflagellates, are well known. But most members, especially the smallest ones, have yet to be recognized or described. Microorganisms less than 2 Âľm in diameter make up picoplankton. We already know that the picoplankton is responsible for most of the photosynthesis in the open sea, whereas we are just beginning to understand their diversity and physiology. This is because of their morphology and the difficulty of isolating them in pure culture. However, in the last decade the application of a number of genomics techniques in oceanography has helped to identify many of these microorganisms and their functions without the need to isolate them. This is part of the work we are currently developing at the Institute of Marine Sciences (CSIC). Our work, nevertheless, and fortunately, is carried out in many locations: during marine oceanographic cruises in the Mediterranean, Atlantic, the Arctic, and Antarctica, as well as in the Atacama Desert (Chile) and the thermal springs of Costa Rica. [ 128 ]

a woman for all seasons | carlos pedr贸s-ali贸

Cover of the Spanish translation of Early Life, Ed. Revert茅, 1988, Barcelona

In June 1973, as an undergraduate, I attended the first meeting of the International Society for the Study of the Origin of Life, which was being held in Barcelona. I was too ignorant and too young not to be awed by the proximity of Nobel laureates such as Severo Ochoa and Melvin Calvin. Thus, it was not surprising that I did not pay much attention to the presentation by Lynn Margulis. I did notice, however, that she was one of the extremely few female scientists [ 129 ]

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invited to the meeting. In retrospect, the fact that she was invited to a major meeting six years after the publication of her momentous paper in the Journal of Theoretical Biology must have symbolized the acceptance by her peers of her theory of the symbiotic origins of mitochondria and chloroplasts. For the next few decades, I enjoyed her lively and passionate talks and conversations. But my appreciation of her work became serious when I read her 1981 book Symbiosis in Cell Evolution. By that time, I had just finished my PhD, at the University of Wisconsin, and was slightly more mature, scientifically speaking. I was overwhelmed by the amount of facts that Lynn had patiently collected from many different sources, some lost in scientific literature over a century old, and by the precise way in which the information was put together and integrated into a coherent theory. It was therefore with enthusiasm that, together with Cristina EnrĂquez de Salamanca, I accepted the job of translating her book Early Life in 1985. I remember that it took us several days just to translate the first sentences of the Preface, as Lynn had used a combination of poetic and scientific language to describe what the living world might have looked like a few billion years ago. Fortunately, the rest of the book was a lot easier. Together with Carl Woeseâ&#x20AC;&#x2122;s proposal of the three-branched tree of life, the symbiotic origin of eukaryotes put forward by Lynn Margulis has established the foundations of modern biology. It has been a privilege to live through these two revolutionary changes in our understanding of life. Lynn Margulis was the most provocative, innovative, cross-disciplinary scientist I have ever known. 3

[ 130 ]

the living experience

of natural history museums Al铆cia Dur贸 i Sans

Institute of Marine Sciences, CSIC, Barcelona

But nature is a stranger yet; The ones that cite her most

Have never passed her haunted house, Nor simplified her ghost.

To pity those that know her not Is helped by the regret

That those who know her, know her less The nearer her they get.

Emily Dickinson (1830-1886)

In writing about Lynn, the first thing that I feel I have to do is to thank her for the almost ten years of knowledge she shared with me. In this short space, I will try to describe some of the many things I keep in my memory and mostly in my heart. During these months after her death, I have been sharing unique moments and common feelings about her with others who knew her. It seems that although Lynn is no longer with us her presence endures. And I remember not only lived moments but ideas and projects. This is how I have continued talking to Lynn: thinking, creating, building, doing. Those are verbs that we used frequently, ever since our first meeting. [ 131 ]

once upon a time lynn margulis

Starting in 1989, biology became a reality at the Science Museum of “la Caixa” Foundation in Barcelona, in the space devoted to the Living Planet. Those of us responsible for the project wanted a museum of the future and the opportunity to introduce the most innovative approaches. I met Lynn in 1998, when I was working as a biologist and museologist. An essential aspect of the exhibit was that in presenting the many facets of life it should be interactive, addressing the origin, appearance, and diversification of organisms and the nature of the first ecosystems. Lynn’s first two bibliographic suggestions were the books she had co-authored with her son Dorion: Microcosmos and Garden of microbial delights. Our first talks turned into a brainstorm of ideas, scribbles on paper, bibliography, more squiggles on paper, references, ... I shrank under an avalanche of concepts, possibilities, optimism, but I was eager to share her passion for science and for transmitting knowledge. How can I forget our glances of mutual understanding in our fascination for learning and in trying to describe “What is Life?” Over the years, we continued talking and I frequently returned to her descriptions of the origin of life, the evolution of microorganisms, and microbial ecosystems. I loved her idea of showing science through art and art through science: Perhaps it was this intuitive feel for interdisciplinarity that gave rise to her theory of symbiogenesis. And it is one of the concepts underlying the new museum, which was our joint project, Biology and Museology in a modern conception. For example, to display the differences between unicellular and multicellular organisms we used both living microorganisms and scientific drawings, which allowed us to show many details. That is how I discovered the work of Christie Lyons, the artist who contributed those drawings. In fact, Lynn [ 132 ]

the living experience of natural history museums | alícia duró i sans

A Winogradsky column as developed by Lynn and the author with the help of Dr. Jordi Urmeneta (microbiologist, University of Barcelona) and Marcel·lí Antúnez (artist). Photo courtesy of A. Duró.

[ 133 ]

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donated part of her own collection of Christie’s drawings to the museum. It was also in this period that I discovered the poetry of Emily Dickinson. One of her poems stands at the beginning of this text. One question was how to show a concept as important and revolutionary as symbiogenesis. What museologist elements would we use? Talking and thinking we found common ground. She summarized her definition by saying: “symbiogénesis is 1 + 1 = 1.” “What does this mean?” I asked. Her answer was that 1 species + 1 species = 1 species. “Wow! Now we’ve got it.” We arrived at the key point, at the essence of the phenomenon, but it was not mathematical language; rather it was museum language. And this explanation has served to open the eyes of everyone who comes across it. At first you think it is a mistake, but we are talking the language of symbiogenesis. It allowed us to develop the script for an audio-visual to illustrate symbiogenesis and its accompanying elements, all of which can be seen in the museum. And so we arrived at “creating” the first ecosystem that at the same time was also meant as a tribute to Professor Ramon Margalef. We constructed 2-m high Winogradsky columns made of transparent glass that evoked dynamic sculptures but clearly revealed the diversity of microbial communities. An exhibit in an interactive museum is like a specimen in a natural history collection: it includes all the information associated with its story. In this case, the exhibit reflected the interdisciplinary efforts of scientists, artists, and museologists: again, a fusion. Lynn enjoyed and considered it as an innovative project that had become reality, where “microbes” began to have a worthy place. It was time to change the public’s view of those “bad guys” by showing their fundamental role in the origin, maintenance, and evolution of life. [ 134 ]

the living experience of natural history museums | alícia duró i sans

One of the several drawings of the Tree of Life created for the exhibition “What is life” at the Science Museum, “la Caixa” Foundation, Barcelona, Spain. Image courtesy of A. Duró.

[ 135 ]

once upon a time lynn margulis

Throughout this process Lynn asked questions and provoked new ideas. She transmitted her immense desire to learn. We shared the vision of museums “with content,” where everyone can experience hands-on science and the joy in understanding and discovering even simple things. With Lynn, every moment was crucial and all who worked with her were captivated by her vocation. A very special remembrance is that of having worked on the project the Tree of Life: The Five Kingdoms. We started this project in 2002, once more scribbling as we brainstormed about how to convey scientific knowledge and enjoying the intense satisfaction in finding the best solution. We worked hard and time flowed but we remained engaged in the task, never feeling tired. We created a large drawing that specified each phylum (including Chaetognatha, which is often left out or misspelled). Drawing is the first taxonomic tool and we used it to communicate with the museum’s designers, who were able to convert the Tree into a-three dimensional visualization. It was a sculpture that presided over the 2006 temporary exhibition on diversity. Lynn’s creativity motivated me. She always encouraged me to continue with the project even though there were days when it seemed impossible! In 2004, the new Museum of Science, CosmoCaixa, was inaugurated. Now, people of all ages can experience the fascination of science and discovery, and, perhaps for the first time, become similarly captivated by Lynn’s ideas. I will never forget Lynn: a woman scientist always ready to fight for and defend her beliefs. 3

[ 136 ]

gaia and more Héctor Ruiz

Editor-in-chief, Omnis cellula, Barcelona

L’entitat autopoètica més petita coneguda i simple és una cel·lula bacteriana. La més gran és probablement Gaia —la vida i el seu entorn planetari (Lovelock 1988)… La constitució d’una

filosofia de la vida basada en la vida mateixa és una tasca de les noves generacions d’estudiants de ciència.

The smallest and simplest known autopoietic entity

is a bacterial cell. The biggest is probably Gaia —life and its planetary surrounding (Lovelock 1988)… The

establishment of a philosophy based on life itself is a job for the new generations of science students.

Lynn Margulis, Omnis cellulla

In thinking about Lynn, the qualities I would choose to emphasize are perseverance, when it came to defending her ideas, her great ability to communicate, and her kindness and generosity towards our efforts to popularize science. I will always be grateful to her for her many selfless collaborations with Omnis cellula, in the form of articles, lectures, and above all, the rights to publish the Catalan edition of Microcosmos. My own path in science communication began with my biology studies. I did research in genetics and spent several research stays in the United States: at the University of Washington, the Univer[ 137 ]

once upon a time lynn margulis

sity of Southern California, and the Jet Propulsion Laboratory. I also taught biology here in Spain. Eventually, however, I became interested in developing other initiatives, which moved me away from a research and teaching career, yet without leaving the world of science. This was due, in part, to an awareness of the difficulty of pursuing a scientific career in Spain. I admire those who have not given up and who have continued to move forward, achieving highquality scientific work, training young researchers, and affirming their faith in the professional paths they have chosen. The difficulties for Spanish scientists are mainly â&#x20AC;&#x201D;but not exclusivelyâ&#x20AC;&#x201D; economic. For our government, whatever its political tendencies, scarce resources are allotted for research even in the best of times. Today, given the current economic difficulties, there have been many deep cuts in research budgets. But there are other problems as well, especially at the universities, in the form of deeply vested interests, the sclerotic organization of departments, and the limited access to the system, all of which are real challenges and the responses to them are not always objective ones. I had long been interested in the use of information technology in the field of education, especially at the elementary and high school levels. Clearly, digital materials can enrich the school curriculum and increase studentsâ&#x20AC;&#x2122; learning capacity and motivation. It is a paradox that the computer, as the ultimate tool of the 21st century, is hardly available in classrooms. I thus became involved in creating quality products to support the teaching of science, aimed at both teachers and students. These initiatives required effort, not only on my part but also from the people who believed in the project. And we were successful, founding Digital-Text, a company directing the production of multimedia school textbooks, which became a leader [ 138 ]

gaia and more | héctor ruiz

Issue 9 (2005) of Omnis cellula, with Lynn Margulis' contribution of the article (translated into Catalan) “Simbiogènesi: Motor de l’evolució.” Image courtesy of the magazine.

in its field. The establishment of the company grew out of our experience in the area of scientific communication, with our magazine Omnis Cellula, which is currently a publication of the Catalan Society for Biology, of the Institute for Catalan Studies. In a joint project with the University of Barcelona, we published the Catalan edition of Lynn Margulis’ book Microcosmos, in 2008. Our long-held wish to translate the book into our language had become reality, and we had the opportunity to thank Lynn personally [ 139 ]

once upon a time lynn margulis

for helping us to obtain the rights. Microcosmos is a jewel of popular science. More than 25 years have passed since it was first published in English, but its relevance has not become outdated. The book immerses us in the origins of our universe. Lewis Thomas (1913â&#x20AC;&#x201C;1993), a friend of Lynnâ&#x20AC;&#x2122;s and a highly respected researcher, physician, and author of another jewel in the popular science literature, The Youngest Science: Notes of a Medicine Watcher, wrote the foreword for Microcosmos, which was originally published by Summit Books, New York, in 1986. In that book, Lynn Margulis and Dorion Sagan provide a history of life from the perspective of microbes, which were the first inhabitants of early Earth, beginning more than 3500 million years ago. Indeed, they were the only life form until the appearance of nucleated cells, about 2000 million years later. I first met Lynn when I was just a sophomore in Biology with a great interest in the origin of life. It was during a course in Microbial Ecology, with excursions to the Ebro Delta, to which I was invited by Prof. Ricard Guerrero. I was thrilled to find out that Lynn knew who I was before I had even introduced myselfâ&#x20AC;&#x201D;presumably Dr. Guerrero had talked to her about me. After that first meeting, I had the opportunity of sharing many wonderful moments with her almost every time she visited Catalonia. In fact, she was once invited by Omnis cellula to take part in a symposium on evolution. I had asked her to talk to students about the origin of the eukaryotic cell as I myself never grew tired of listening to her ideas on symbiogenesis. However, the greatest pleasure was being able to pursue those ideas more deeply over lunch or dinner with her and other scientists. She made a deep impact on my life both as a scientist and as a person. Lynn once told me that studying life by narrowing it down to equations and physical laws was just like analyzing a poem's me[ 140 ]

gaia and more | héctor ruiz

Héctor Ruiz and Begoña Vendrell with Ricardo Guerrero and Lynn Margulis at a symposium in Barcelona in 2004. Photo courtesy of H. Ruiz.

ter: it missed out on its true essence. I am deeply indebted to her for having shaped my way of seeing the world, away from anthropocentrism towards an appreciation of the role played by the microcosmos. I am also indebted to her for a great part of my love for science communication. Her perseverance is still a great source of inspiration in my everyday life. As a tribute to Lynn, I will always be proud of having added the word ‘symbiogenesis’ to the Gran Diccionari de l’Enciclopèdia Catalana, and especially of having published Microcosmos in Catalan. If I had to define Lynn in just three words, I would say: persistent, intelligent, and passionate. 3 [ 141 ]

my dear lynn.

a humble and late eulogy Xavier Maym贸-Gatell

CEO, Fimarge Wealth Management, Andorra

Farewell to Thee! But not farewell

To all my fondest thoughts of Thee:

Within my heart they still shall dwell; And they shall cheer and comfort me. Nothing is lost that Thou didst give,

Nothing destroyed that Thou hast done.

Anne Bront毛 (1820-1849)

It is not the critic that counts: not the one who points out

how the strong [woman] stumbles or where the doer of deeds could have done them better. The credit belongs to the [woman] who is actually in the arena, whose face is

marred by dust and sweat and blood, who strives valiantly, who errs and comes up short again and again, because there is no effort without error or shortcoming, but who

knows the great enthusiasms, the great devotions, who spends [herself] for a worthy cause; who at best, knows, in

the end, the triumph of high achievement, and who, at the worst, if [she] fails, at least [she] fails while daring greatly,

so that [her] place shall never be with those cold and timid souls who knew neither victory nor defeat.

(Adapted from) Theodore Roosevelt (1858-1919)

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I learned of my dear Lynn’s passing away more than a year after it happened. Immersed in the physical and emotional strenuousness of a career in the financial world, I had lost contact with a life —and the people— I deeply loved so long ago. Now approaching my 50s, I feel that the scientific and university life that I left at the age of 32 would be, again, a great life for me. I miss research, I miss teaching and, most of all, I miss people like my dear Lynn. The business world is, as I often say, wide and shallow. No depth. It is very interesting in many ways, true, but there is no depth to be found. Of course, citing Ralph Waldo Emerson, “it is not length of life, but depth of life”. My dear Lynn lived deeply. I have no doubts about that. Up until the early 2000s, I had met my dear Lynn numerous times, both in the USA and in Catalonia, always in relation to science and scientific meetings. Many times, I found her devoting her spare time helping popularize science. It worked marvelously for me. Few people, like my friend and mentor Prof. Ricard Guerrero, like my dear Lynn, have two qualities that should be required but are seldom found in scientists: outstanding generosity and capacity to communicate the excitement of the unknown. In the 1980s I discovered the world of microorganisms and the scientific theories on the origin of life from Ricard Guerrero, while I was a university student in Barcelona. A hidden world waiting to be understood! What a challenge! I couldn’t wait to be immersed in it. Then, in the summer of 1990, Prof. Guerrero invited me to spend a few months at the University of Massachusetts at Amherst (U-Mass), learning the lab techniques needed to study anaerobic [ 144 ]

my dear lynn. a humble and late eulogy | xavier maymó-gatell

Lynn at her Amherst home with her father in the summer of 1990. Photo by X. Maymó-Gatell.

bacteria. During my stay, my dear Lynn generously offered me a place to live, in her own home. We never worked together, but the time that I spent with my dear Lynn at home and with her son, Dorian Sagan —later I would meet his father at Cornell University—, made for an unforgettable summer. Many good friends who have contributed to this book are more qualified to speak about my dear Lynn’s professional accomplishments, so I won’t even try. Let me offer a different perspective instead. Let me speak about the things that transformed me that summer, thanks to my dear Lynn. I was very young —23 years of age was considered “very young” [ 145 ]

once upon a time lynn margulis

in Catalonia. Although at the same age Alexander the Great of Macedon had conquered half the World, in my time and country we could barely leave home for more than a couple of days without our mothers crying inconsolably for their loss. I knew almost nothing about life; real life I mean. I had read a lot, sure, but as an outstanding Catalan publisher —Carmen Balcells— once said: “Reading is a good substitute for life. But have no doubt: it is much better to live than to read. In fact, I only read when I cannot live.” That was an apt description of my personal situation back then. Also, I knew little about the greatness of great scientists, so I went to U-Mass without inhibitions or fears. On the one hand, it made me more impressionable regarding basic, fundamental things; the important ones. On the other hand, it allowed me to learn more, to absorb more personal knowledge and experiences that, in regard to some of them, I only really brought to conscience years later, when I found them there, patiently waiting within me. Let me tell you about a few of the things my dear Lynn enlightened me about: PRO-ACTIVITY needs a good dose of bravery to be effected. Of course, you cannot be brave without fear (then you would be thoughtless). That makes it a much more interesting personal quality. My dear Lynn had it. For someone born in the USA, it may not seem an important quality; one could say, “that’s the way things work.” Not true. At least not in many other parts of the world. Southern European culture, for example, is reactive. People wait for things to happen; then act. I am not that way. I never was, but I didn’t know that until I met her. She continuously took intuitively firm stances before anyone could demonstrate them, and that required the assumption of risk, and it took courage. What an example she was. We all have fears, but few of us choose not to be governed by them. [ 146 ]

my dear lynn. a humble and late eulogy | xavier maymó-gatell

To her friends, to her students, to her research, to her values. My dear Lynn was determined and persistent in everything she did. That gave her consequence of thought and action and, in turn, gave it to others. Nothing worthy in the advancement of human nature may be achieved without being thoroughly bound to it by personal commitment. Abraham Lincoln, in a speech at New Haven (March 6, 1860) said: “Let us have faith that right makes might; and in that faith, let us, to the end, dare to do our duty as we understand it,” regardless of the difficulties. GENEROSITY. Lincoln again: “When you make it to the top, turn and reach down for the person behind you.” Of course I was not behind her; in fact, I was many feet below her talent. Nevertheless, she opened her world to me completely. I had access to anything I needed, both materially and intellectually. Coming from a country where people tend to be more generous with themselves than with others, it was a wonderful shock. There I was, young and unknown. There she was, always busy, so renowned, with so many other important things to do than help me… but she did. NO CEILING. Being in the USA is like having a home with no roof. You feel the cold when it comes and you get wet when it rains, but on a sunny day, you can jump, for as long as you want, as high as your possibilities will let you. There’s no ceiling over your head to stop you. The opportunities to see and carry yourself far and beyond are limitless. In the USA, your life and what you do with it rests, as Ricard Guerrero would say, in your hands, your head and your heart. I love this feeling. Europe, as a whole, has lost it. Southern Europe, as a whole, never had it. Together with Prof. Guerrero—an outstanding exception in Spain—, my dear Lynn made that great feeling grow even larger. That summer I had a strong sense of liberation from COMMITMENT.

[ 147 ]

once upon a time lynn margulis

Lynn and Xavier Maymó, on a hiking trip in Massachusetts in the summer of 1990. Photo by X. Maymó-Gatell.

the oppressive ambience of Spain, and all the while, as I gained this mental and emotional freedom, my dear Lynn was nearby, constantly sharing her encouragement, enthusiasm and unlimited passion. As Mark Twain put it: “Keep away from people who try to belittle your ambitions. Small people always do that, but the really great make you feel that you, too, can become great.” Last, but no less important… CHICKEN. Yes, chicken. I had no idea one could eat chicken every day and be reasonably healthy. After a few weeks in my dear Lynn’s home, I had the strong feeling I would start crowing spasmodically more and more often, to be finally trans[ 148 ]

my dear lynn. a humble and late eulogy | xavier maymĂł-gatell

formed into my own Cock-a-Doodle Dandy 1. Roasted, fried, boiled, grilled, marinated, in soup, in salads, boneless, skinless, shredded, you name it. The ability of my dear Lynn to cook chicken in a seemingly infinite number of ways left me flabbergasted. In those years, it was reported that chickens were given hormones, i.e., estrogens and such, to attain faster growth. At night I used to dream that big breasts would come out of my chest and, contrary to what some may think, it was not a pleasant dream... Since then, I have never eaten as much chicken as I did that summer. Also, rarely have I ever been treated again as my dear Lynn treated me: with intellectual respect, with patience and understanding, with care and motherly love. After my Ph.D. at Cornell University went well, publishing in Science and all, I decided to make a 180 degree turn in my professional life. It was more than 15 years ago. Since then, I have managed several companies in the financial sector, and still do today. But, in my own defense, I have never worked as a banker. I guess I have retained my set of values and a bit of common sense. My dear Lynn, I miss you; your daring propositions, your generosity, your science, you. As Lincoln said once: â&#x20AC;&#x153;towering genius disdains a beaten path. It seeks regions hitherto unexplored.â&#x20AC;? I am sure your vital energy, still part of our world, will help us in our efforts to imitate the towering genius you always showed freely to anyone ready to see it. The pain derived from the loss can never be greater than the happiness to have shared a bit of this life with you. 3 1

In the play, written in 1949 by Sean O'Casey and set in the mid-century Irish countryside, the struggle between repression and liberty, which I clearly felt at the time, is brought to life as a fantasy in which a magic cockerel forces the characters to make choices about how they live their lives.

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To laugh often and love much;

to win the respect of intelligent persons and the affection of children;

to earn the approbation of honest critics and to endure the betrayal of false friends; to appreciate beauty;

to find the best in others; to give of oneâ&#x20AC;&#x2122;s self;

to leave the world a little better,

whether by a healthy child, a garden patch or a redeemed social condition;

to have played and laughed with enthusiasm and sung with exultation;

to know that even one life has breathed easier because you have lived.

This is to have succeeded.

Bessie Anderson Stanley (1879-1952)

[ 150 ]

let s go to ithaca* Gemma Reguera

Michigan State University, East Lansing, MI, USA

As you set out on the way to Ithaca hope that the road is a long one,

filled with adventures, filled with understanding. The Laestrygonians and the Cyclopes,

Poseidon in his anger: do not fear them,

you’ll never come across them on your way

as long as your mind stays aloft, and a choice emotion touches your spirit and your body. The Laestrygonians and the Cyclopes,

savage Poseidon; you’ll not encounter them unless you carry them within your soul,

unless your soul sets them up before you.

Constantine Petrou Cavafy (1863-1933)

There are journeys in life that go beyond covering a distance. Those that cause you to make sharp turns that bring you into a different landscape, filled with different people and mindsets. I can now look back and clearly identify those defining moments. I can also appreciate, even after many years, just how important they were, how much *Some parts of this text are based on “Cartas al presidente” (Letters to the President) section of SEM@foro, magazine of the Spanish Society for Microbiology, December 2012, pp. 80-81.

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they inspired me, and how much they influenced who I am and what I do today. Let me outline that route for you, not because it is particularly important but because it provides the context to evoke with emotion one single person who crossed my path and by doing so profoundly influenced me. This is not about me, this is about Lynn. I was born and raised in Spain, yet here I am in the United States, working as an associate professor in the Department of Microbiology and Molecular Genetics at Michigan State University. My university is located in the city of East Lansing. Ever since its funding, in 1855, Michigan State has remained loyal to the true values of the land-grant mission: to promote the well-being of society through the advancement of technology. It was, after all, the first land grant institution in the country and its educational model was based on agriculture, science, and engineering and their applications towards social advancement. Faithful to its origins, it is one of the very few universities that stands out for its programs in agriculture and education. The Department of Microbiology, where I work, also has the merit of being widely recognized for its research in microbial ecology and evolution. I could not have found a better setting for my research interests: understanding microbial processes and how they impact the environment. I am fascinated by microbes because they adapt to our disruptive influence and are capable of bringing balance to our sometimes chaotic actions. Thus, it is not surprising that through research in microbiology we also seek to harness those very same microbial processes for their biotechnological applications. My group, for example, is engaged in studying the electrical activity of some bacteria and the ability of some species to produce electrically conductive nanocables or nanowires, which are used to reach out to the environment and to [ 152 ]

let’s go to ithaca

| gemma reguera

interact with and transfer respiratory electrons to toxic metals and radionuclides. In doing so, they mineralize these compounds and decontaminate the environment. I am just one of many Spanish investigators who made the decision to pursue a career abroad. My story is similar to those of others, although our various reasons for doing so may differ. I came to the USA because I wanted to do research in environmental microbiology and because I was longing for an educational system of intellectual freedom that promoted the individual rather than the masses. I don’t come from a wealthy family; I studied in Spanish state schools and at a Spanish university from beginning to end. My journey started after I had graduated in biology at the University of Oviedo, when I was awarded a fellowship to be an exchange student at the University of Massachusetts at Amherst. Fortunately, my knowledge of English allowed me to compete for a fellowship normally awarded to philologists. The before and after of this fellowship that took me to the USA are reference points in my professional life. The decision to leave Spain was a difficult one. As a microbiologist I couldn’t obtain enough financial support to stay in Spain to do a doctorate or to go abroad. The environment in Spain, poor in resources and perspectives for young scientists, only increased my feelings of suffocation and intellectual needs in what I always considered a rigid mass educational and scientific system. And even worse was the university system itself, which perpetuated a hierarchy not always based on quality. It pains me to say this, because I don’t want these negative impressions to mar the respect I have for the exceptional scientific and didactic work of a great many of my Spanish mentors and colleagues, who have always inspired and unconditionally supported me. The educational and scientific train[ 153 ]

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ing I received from them, not from the system, provided the intellectual and moral basis of my profession. I owe them a great deal of what I have achieved professionally. During those years in Amherst, I worked very hard. I was a teaching assistant in Spanish and also in Microbiology. These jobs not only allowed me to earn money but they also allowed me to take courses in the microbiology graduate program, where I was awarded first a master’s degree and then a doctorate. Perhaps more important than these degrees, Amherst exposed me to a new culture of education and science, and to scientists whose influence continue to inspire me. There are moments from my Amherst experience that seem to be frozen in time in my mind, perhaps because of how much they meant to me both scientifically and personally. One of them is, without a doubt, the day I met Lynn Margulis. I was first introduced to her soon after I arrived at the University of Massachusetts, by a friend of mine who was doing his Ph.D. in her lab. We were walking on campus one day, speaking in Spanish, and we heard this voice behind us: ¿No me vas a presentar a tu amiga? (“Aren’t you going to introduce me to your friend?”) Yes, that was Lynn. We spoke in Spanish for a while. I simply could not believe that this friendly, down to Earth woman was the great Lynn Margulis. After that, I remember going to her lab to admire the enormous cockroaches that her group raised there for their symbiotic studies. We also coincidentally met at the public swimming pool on many mornings, where we often chatted. Lynn was one of the greatest educators on campus. She often referred to the importance of creative thinking and scientific inquiry in her own education and never forgot to put them into practice herself in her efforts to foster the scientific development of her students. Her use of educational videos and books were pioneering. She knew [ 154 ]

let’s go to ithaca

| gemma reguera

how to reach students’ minds and how to excite them. As a graduate student, I found her style liberating: Her classes for graduate students were not based on memorization but on scientific reasoning. In lectures, we discussed the experiments underlying important discoveries and dissected the logic that gave rise to the experimental plan. For the first time I learned to reason while I studied and thus I stopped being bored in lectures. That, along with the theoretical training I brought with me from Spain, helped me to advance quickly and to stand out academically. Another great surprise and joy was the grading system. Students received the grade he or she deserved. In Spain, the number of top grades a teacher is able to give is determined by the number of students enrolled. I sadly remember my virology class, which I enjoyed so much and in which I earned the second-highest grade, a minimal 0.1 difference from the top score. But as only one top grade (an A or 4.0, the Spanish matrícula de honor) was allowed based on the number of students registered for the class, I received only a B (3.0, or sobresaliente). In the North American system, as I learned, grades could be “curved,” i.e., distributed according to a statistical system that normalizes grades based on the performance of all the students in the course. Thus, the best students get the top grade (“A”) and the grade becomes a point of reference. I felt this to be a fair system and it freed me to enjoy the courses simply because they nourished my need for knowledge. I was so motivated that I signed up for additional courses. I also took advantage of the academic flexibility of the programs and registered for courses in other departments. This multidisciplinary training is something I continue to develop and try to pass on both to my students and to the people in my research group. [ 155 ]

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The most overwhelming contrast between Spanish and North American university life was the resources available to me. For example, I was shocked to go from sharing a tiny desk with another student to having my own desk, work table, and bench space. The instruments available to me in my research group allowed me to carry out all kinds of experiments. And this directly affected the quantity and quality of the research I did. Scientists in the USA put not only their vocational goals but also their personal efforts into their scientific endeavors, and merit a salary according to their knowledge and work. Productivity and quality in research require economic investment in material resources and in personal compensation. Certainly, scientific research in Spain will suffer enormously from reductions in research allocations that are already tragically low. Research brings progress and capital and is therefore even more important in times of crisis. With my newly awarded Ph.D. in hand, I left for my first post-doc in Boston, returning 2 years later to the University of Massachusetts for a second post-doc. And thatâ&#x20AC;&#x2122;s when Lynn and I met for a second time, an unforgettable one. It was in 2005 and seemingly out of the blue I received a phone call from her (Lynn did not e-mail!). She wanted to talk about the discovery of bacterial nanowires, which we had just published in Nature. Just like that, unpredictable and direct as she was, she asked about the research, the bacteria, the nanowires,â&#x20AC;Ś and invited me to give a lecture in her class. It was her love for science that fascinated me the most. During my lecture, she was like an eager child, asking questions non-stop, intrigued by the discovery, by what the microorganisms were telling, going beyond the dry results, always looking at the big (big!!!!) picture. Her intellect, curiosity, and genuine enthusiasm were wondrous. She was so refreshingâ&#x20AC;&#x201D;every[ 156 ]

let’s go to ithaca

| gemma reguera

thing that I admire in a scientist: intelligence, encyclopedic knowledge, stubborn, naughty, passionate and very, very human. At the time of my lecture, I was pregnant. And with her usual frankness, she grilled me about the pregnancy, the baby, plans for the future, how I was going to handle a career and motherhood… she was like a hurricane of concern. She was also the first to call me at home as the due date approached, leaving a message in Spanish telling me that she had not seen me for a while and was worried. I was touched. In a culture of individualism, where so many people just don’t care, her genuine affection and loyalty made her shine. She was one of the first to meet my son, Darío, the one with the name of the Persian king and the poet, Ruben Darío (her own words!). She melted looking at the baby. Always the evolutionary biologist, she admired his health and strong complexion: Es un buen macho, Gemma (‘He is a good male, Gemma’). And she kept asking questions about my transition to motherhood (Was I nursing? “Yes, Lynn, I am and plan to continue for a few more months at least”) and about so many other things that all career women struggle with in our professional and personal lives. No other woman has inspired me more, both professionally and personally. I treasure the moments I spent with her. And here I am, eight years after Lynn’s phone call. In the end, science, like life in general, is about confronting challenges. My challenge involved leaving my country and culture behind, seeking a career elsewhere. If I had not accepted that challenge, I would not have experienced what I have shared with you, what has possibly been the most rewarding adventure of my life. I have learned how to live in two cultures, my own and my adopted one, and I am proud of having citizenship in both. Throughout the years I have kept in contact with old and new Spanish friends and mentors. Like many [ 157 ]

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Cover of the Spanish and English edition of Luminous fish.

of the young people that took on the adventure of a career abroad, I donâ&#x20AC;&#x2122;t plan on returning to the country where I was born. Now, I am in a place where I can do the science I have always enjoyed and where I have started a family. I am happy working at a university embedded in a system that provides me with the stability and resources to do what I love and to make it compatible with family life. But I must credit Lynn for reconnecting me with my Spanish academic roots. She was deeply involved with Spanish science and scientists. She was the bridge and catalyst between the two countries, just for the love of science. She introduced me to many Spanish researchers and promoted my research and my persona back in my own country. She always spoke well of meâ&#x20AC;Ś she was a terrific mentor. [ 158 ]

let’s go to ithaca

| gemma reguera

Lynn was an advocate of science. She was passionate about it. She worked tirelessly to disseminate science and to stimulate scientific inquiry. I am proud to say that her dedication to science dissemination inspired me to follow similar steps. The fruit of that influence is my contribution as associate blogger and editor to the science blog of the American Society for Microbiology “Small Things Considered” by my close friend and mentor Moselio (Elio) Schaechter, another key figure in the field and history of microbiology. Lynn was also unique because she loved to challenge and to be challenged. She was not afraid of anything or anybody. And she liked to provoke. I loved the answer she once gave to a journalist criticizing the Gaia hypothesis: “Don’t worry, Gaia is a tough bitch.” She was unique, a force of a nature that few can understand unless they had known her. If you want to know her most personal side, read her first book of fiction Luminous Fish: Tales of Science and Love. I leave you with her own words, the ones she wrote for me in that very same book one memorable day of January of 2007 in Barcelona, words that sum up my own personal journey: “With love of life and its study.” 3

[ 159 ]

members of all the kingdoms Juli Peretó

University of Valencia

Quin privilegi, haver pogut parlar

tants anys davant d'uns ulls que renovaven contínuament la seva joventut,

haver-me sentit ple de savieses més altes que la meva

i des d'elles haver pogut parlar, argumentar, sorprendre, rebatre, demostrar,

oblidar-me dels meus límits, de la meva poca gràcia, de la meva veu monòtona, de mi mateix i tot,

i en un cel de pissarres i de guix esdevenir tan sols un portador -indigne, ho sé, i obscur- de tantes meravelles! What a privilege having been able to talk

for so many years before eyes that continuously renewed their youth,

having felt myself full of wisdoms higher than mine

and from those wisdoms, having been able to talk, to argue, to surprise, to refute, to demonstrate, to forget my limits, my lack of grace,

my monotonous voice, and even myself,

and in a sky of blackboards and chalk becoming only a carrier -unworthy, I know, and dark— of so many wonders!

David Jou (poet and physicist), El professor

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Without any intention of making comparisons, like many others dedicated to science, research, education and science popularization, I have admired Lynn Margulis for her passionate devotion to the exploration of nature, both in the laboratory and in the field. Her observations, under an audacious scientific perspective, led her to propose the endosymbiotic theory of the origin of the eukaryotic cell. And she did it without disregarding previous contributions of other authors, especially Russian botanists, whose work had not been recognized by the official history of biology and that she contributed to disseminate. We had known Lynn’s fighting spirit long before meeting her, and we were also aware of the many controversies arisen as a result of her collaboration with James Lovelock about the Gaia theory. Although it is true that Gaia has not been yet completely accepted, its contribution to the understanding of the close relationship between biology and geology throughout the history of our planet cannot be denied. Lynn has been controversial; for example for her criticism of neo-Darwinism and, more recently, for having supported authors proposing extremely eccentric or erroneous ideas. In the mid 1960s, hardly anybody, perhaps not even Lynn herself, could foresee that her article On the origin of mitosing cells–finally published in the Journal of Theoretical Biology after having been rejected many times—would be the starting point of a recognition—slow but steady—of the role of symbiosis in the evolution of life. Today recognizing that nucleated complex cells are the result of the merger of lineages, and that symbiosis is one of the fundamental evolutionary mechanisms is accepted by canonical evolutionary biology. Darwinian orthodoxy of the divergent branches of the tree of life is completed by the observation of numerous convergences generating completely new lineages. [ 162 ]

members of all the kingdoms

| juli peretó

What is a cockroach today but the result of the incorporation of an ancient flavobacterium within a lineage of arthropods, about 150 million years ago, in a permanent, inseparable way? And so, countless examples of microorganisms that have shaped the metabolism, development and behavior of complex eukaryotes, either unicellular or multicellular. Nobody can understand the evolution of animals and plants without knowing the role of microorganisms that inhabit our internal and external folds and that, on given occasions, penetrate our cells. Symbiogenesis is a central concept in contemporary biology largely thanks to Lynn. So why should be strange that living beings who arrived just about 600 million years ago have had to evolve on a planet full of tiny creatures that had already explored—for more than 3000 million years—the biochemical opportunities offered by the planet? They could not avoid them. Lynn maintained a very close relationship with the University of Valencia, which awarded her with an honorary doctorate in 2001 and published several of her books, including Early Life (Bromera-PUV, translated into Catalan and Spanish). Her intellectual and personal generosity allowed us to have her in courses, seminars, lectures and outreach activities. Her passion to know nature and make others to know it and whatever she had learned about it made her a master in the art of popularizing science. Rarely have we seen students so attentive and captivated as they were by her words, nor have we seen so crowded halls. Lynn also wanted to participate in our outreach efforts and was an exceptional collaborator of Mètode, the magazine edited by biologist, journalist and writer Martí Dominguez. I first met Lynn in Valencia, along with Martí, when we organized a lecture for the students of the School of Biological Sciences and there [ 163 ]

once upon a time lynn margulis

Joan Or贸, Antonio Lazcano, Lynn Margulis and Stanley Miller. Valencia 1994 during a course to commemorate the Oparin centenary. Photo courtesy of J. Peret贸.

[ 164 ]

members of all the kingdoms

| juli peret贸

we interviewed her for the the weekly magazine El Temps, in 1992. The University of Valencia, my university, was founded in 1499 under the name of Estudi General. In this university I started my career as a professor of biochemistry and dedicated myself to this discipline because I was attracted by whatever had to do with an explanation of life and its functioning, origins and evolution. I preferred to do it in a field that provides an approach to the biological world that allows to unraveling it at a molecular scale. The tradition of evolutionary biochemistry was interrupted during the years of development of molecular biology in the second half of the twentieth century. However, it is impossible to understand life, at any scale, without the evolutionary approach. Practicing evolutionary biochemistry requires the contributions of other scientific disciplines and a broad perspective outside the narrow boundaries of university departments. It requires a bit of rebellion and courage that Lynn would poure. Although fortune and chance played a role on it, embarking myself on research on endosymbiont bacteria of insects related me intellectually to Lynn. Her visits to our laboratory, her talking with us, with Amparo Latorre and Andr茅s Moya, my mentors in that scientific field, were always stimulating. Lynn also gave away her generous enthusiasm impossible to forget. Life is a peculiar and somehow aberrant chemistry of organic compounds in aqueous solution in contact with the atmosphere that has changed over geological eras. There is no corner on the planet that has not been infected by life. Lynn taught us much about this adaptive metabolic virtuosity. She taught us the promiscuity of microorganisms and their ability to establish biochemical alliances that end up having a planetary impact. Microorganisms have, therefore, [ 165 ]

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a key role both in the functioning of terrestrial life at a global scale and in the evolution of more complex life forms, and this is something that Lynn always claimed. Her most categorical statement was that life on Earth depends on the microbial world. Lynn accompanied me in the journey from the chemistry of the cell to the planet as a whole and I learned that in no way is biochemistry confined within a membranous bag. In addition to teaching, doing research and organizing outreach scientific activities, I served as Vice-Rector of the University of Valencia and I am proud to mention the sensitivity of my institution towards the recognition with honors and awards of prominent scientists, among whom evolutionary biologists Francisco J. Ayala, Richard Dawkins, François Jacob and Lynn Margulis. In 1993, on the occasion of the celebration of 25 years of studies in biology, the University of Valencia included François Jacob in the faculty of doctors and years later published El desván de la evolución (The attic of evolution), a book with a selection of his texts, in addition to the speech he given at that time. Lynn Margulis was awarded an Honoris Causa Doctorate in 2001, during the celebrations of the fifth centenary of the university, an occasion that also led to the publication of the miscellany Una revolución en la evolución (A revolution in evolution). To François Jacob we owe the idea of molecular tinkering, the recycling to perform new functions, a metaphor that Darwin himself used to illustrate how orchids adapt to their insect pollinators. Margulis bequeathed us her sensitivity towards collaboration and interaction in nature: living beings are not isolated and pure entities as they are often presented in the childish schemes of some textbook. From my experience of doing research and teaching me[ 166 ]

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| juli peret贸

tabolism, I can state that symbiosis and tinkering are two engines for understanding life. And I have concluded that if evolutionary biology needs the information provided by biochemistry to understand the evolution at the molecular level, the opposite is also true: nothing makes sense in biochemistry unless we look at it under an evolutionary perspective. And the influence of Lynn is no stranger to this way of thinking. Lynn Margulis left us in November 2011, still young and active until the last moment. In December 2012 Rita Levi Montalcini undertook that trip, not so young, but also a great lady of science. More recently, and a few weeks apart, two other leading biologists, Fran莽ois Jacob and Christian de Duve also left us. All of them were the kind of people that do not leave you indifferent, either when reading or listening to them. The four of them had in their personality a concern for transmitting knowledge and being understood by the public so that, as de Duve said, the light of science should enlighten not only initiates. The works they left are lasting testimonies of their ideas, which we can all approach. In addition, if one has had also the opportunity of having listen to their words, having talked to them, it is an inner experience that we have an obligation to convey. Surely these scientists were exceptional people, but it is even more important to recognize their commitment to what they believed and their joy of sharing it. And that is something that before, now and always has to be assessed for the strength and character that all this imprints on life. 3

[ 167 ]

the art of storytelling Nicole Skinner

Institute for Catalan Studies, Barcelona, Catalonia Academia Europaea | Barcelona Knowledge Hub, Barcelona

Tell all the Truth but tell it slant Success in Circuit lies,

Too bright for our infirm Delight The Truth â&#x20AC;&#x2122;s superb surprise;

As Lightning to the Children eased With explanation kind

The Truth must dazzle gradually Or every man be blind -

Emily Dickinson (1830-1886)

There are numerous reasons why it is important to communicate science to society. We live in a world that has been shaped by and depends profoundly on science and technology. Scientific literacy, the knowledge and understanding of scientific concepts and processes, promotes lifelong learning skills that can be used in everyday life for better and more informed decision-making. It is also important to bring to light the potential economic benefits of investing in science, and how this investment will have high returns for society. Curiosity-driven, or basic, research provides new knowledge with the potential to create many opportunities. It can lead to fundamental discoveries but also to unforeseen developments, including some [ 169 ]

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of the most important technologies we use today, such as the World Wide Web, conceived and developed at the European Organization for Nuclear Research (CERN); the GPS, as an example of the commercial application of a military system, or even Google. To engage people with science we must show them what science is, what it means, and why we need it. Science satisfies our intellectual curiosity, humanity’s drive and desire to understand the natural and physical world in which we live in. But what exactly is science? According to the UK’s Science Council, it is “the pursuit and application of knowledge and understanding of the natural and social world following a systematic methodology based on evidence.” And yet science is a collective enterprise carried out by people, it is not perfect. Its uniqueness, however, lies in the fact that it is self-regulating. It has an inherent integrity that society should be able to trust. In Lynn’s words, “there is no Truth, but science is the best way of knowing.” Anyone who met Lynn Margulis personally would immediately sense that she was as passionate an individual as they come. In the words of Pam Pelletier, Science Program Director for Boston Public Schools, “Lynn was a bright fire in the world of science—and if you were lucky enough to get close, you could also feel the warmth of her humanity and her commitment to making teachers, students, everyone, better understand our biosphere and how it came to be.” Lynn’s most important contribution as a scientist was her theory of serial endosymbiosis, which represents a milestone in the history of science. But Lynn was also a teacher in the deepest sense of the word. Throughout her life, she took an interest in creating and developing teaching materials, producing a wonderful collection of curriculum units, educational videos, and books. In doing so, she [ 170 ]

the art of storytelling

| nicole skinner

Visit to the Government of the Balearic Islands during the celebration of the International Symposium, “Darwin: 150 years after the theory of evolution.” Palma de Mallorca, June 11–12 2009. From left to right: Wendy Ran, Nicole Skinner, Jacqueline Fortey, Richard Fortey, Bàrbara Galmés, Anna Omedes, Francesc Antich, Lynn Margulis, Sheldon Glashow, Antoni Riera, Bartomeu Llinàs, Misericòrdia Ramon, Ricard Guerrero. Photo courtesy of the Government of the Balearic Islands.

helped scientists to recognize that they also needed to be teachers, and teachers that they too had to be scientists. Besides her teaching abilities, Lynn had a zest for communicating science to the public. She was entertaining and stimulating; she had a vivacious energy and would discuss with great enthusiasm any topic at hand. But more importantly, perhaps, she retained a childish sense of wonder at what life is and had an innate ability to pass that excitement on to others—she made our inherent curiosity come alive. For Lynn, science was not a solitary practice. She was all about connecting ideas, thoughts and people, and she strongly believed [ 171 ]

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that the diffusion of knowledge and the promotion of scientific culture were equally important to research. “It’s not science to me unless it’s integrated,” Lynn said, “unless you have a story of something you can tell other people, and a story that makes sense.” And Lynn’s stories were the key to her success as a science communicator. It has been argued that 95 % of our scientific literacy comes from informal learning, outside of classroom walls, in the form of educational television and radio shows, movies and documentaries, science museums, zoos, aquariums, and national parks, and increasingly, through the countless digital games and resources available online. In both formal and informal science education, it is necessary to simplify concepts to meet the needs of the learners. But making science accessible does not mean dumbing it down. In this respect, Lynn Margulis was particularly skilled. Even when she was addressing an audience made up of specialists her lectures were easy to understand. Lynn visited most of our museums in Spain. In fact, as explained in other contributions in this book, she actively participated in the preparation of exhibitions related to the origin of life, contributing her ideas about how the information to be conveyed could be presented in the most broadly comprehensible way possible. Good science communicators will employ clear and direct language, avoiding the use of technical jargon. Even when explaining highly specialized concepts, they will translate complex science into everyday language in order to make it comprehensible to and accessible by a wider general audience. They value the importance of storytelling and are able to craft accounts that explain science with flare and with heart, in a way that relates directly to the audience. Passion, imaginative verbal skills, and creative writing are vital to effective science communication. [ 172 ]

the art of storytelling

| nicole skinner

“Life on earth is such a good story you cannot afford to miss the beginning. [...] Beneath our superficial differences we are all of us walking communities of bacteria. The world shimmers, a pointillist landscape made of tiny living beings.” Lynn narrates in her insightful and beautifully written chronicle of evolution, Microcosmos. She’s got our attention. Her words, her enthusiasm, have lit up our brain. And now, all we want to do is to continue reading, with close attention, the rest of story she is about to tell. 3

[ 173 ]

the ins and outs of life Carmen Chica

International Microbiology, Barcelona

Si yo volviera a nacer,

quisiera en la mar vivir. Velas quisiera zurcir, redes quisiera tejer.

barca quisiera tener,

remos para ir y venirâ&#x20AC;Ś Were I to be born again

I would like to live in the sea. Sails I would like to darn,

Networks I would like to weave. A boat I would like to have, and rows to come and goâ&#x20AC;Ś

Mirta Aguirre (1912-1980)

once upon a time

We could never have imagined writing about your absence. Yes, all of us are mortal, in spite of the potential immortality of our cells, but it is hard to internalize that you left with the twilight of that 22nd of November 2011, after barely five days of silence and uncertainty. At least two people, Diane Bylan and Hannah Greene, were with you when the stroke would cloud your vision and your [ 175 ]

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body felt the laziness against which your vitality had to surrender. But not immediately. In fact, when the emergency medical team arrived and carried you down the stairs you were energetically trying to take command, telling them what to do. “The old Lynn to the end,” this is what Barbara Patter, a close friend of Lynn’s in Amherst wrote to me. Using the words of a Spanish poet, Miguel Hernández, we can say, un manotazo duro, un golpe helado, un hachazo invisible y homicida, un empujón brutal te ha derribado (“a hard slap, a frozen blow, an invisible and murderous stroke of the axe, a brutal shove has brought you down”). Our grief and that of so many colleagues and friends here in Spain and across the world could not be measured. The life of a person can be seen as an bottomless well. The deeper it is, the more resources it contains to develop life to its fullest, and to enjoy with sensitivity what nature and human ingenuity have deployed. But sensitivity is a double-edged sword. It provides us with joy but also forces us to grieve over the catastrophes of nature and our limited ability to amend the gross abuses of human activity. The vast resources of Lynn’s well left her no time for rest, especially after she had immersed herself in studying the origins of life and of the Earth itself. Of the many treasures contained in Lynn’s well, the most intensely loved were music –Brahms, Schubert, Beethoven floated through the air while she worked or studied, wherever she was; literature –fiction, romance, fictionalized history, history of science, history in its purest form, and poetry, especially that of Emily Dickinson, whose house was next to that of Lynn. In fact Lynn’s house once was part of the Dickinson Homestead. The solitude that the poet chose and her free spirit fill the air of that shared space, against a background of flowing water and the sounds of the birds sheltered in the trees. [ 176 ]

the ins and outs of life | carmen chica

Emily Dickinsonâ&#x20AC;&#x2122;s Homestead (top), now a part of her museum, is next to Lynnâ&#x20AC;&#x2122;s house (bottom). Photos by M.Piqueras and C. Puche.

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welcome: lynn in spain

During her sabbatical at the Autonomous University of Barcelona (UAB) in 1985, Lynn and I drove out to the Serra de Collserola, which she wanted to explore on horseback. The manager of the riding center quickly noticed the skill of this â&#x20AC;&#x153;amazonâ&#x20AC;? (I, however, was desperately clinging to the horseâ&#x20AC;&#x2122;s mane) and trusted us to fend for ourselves. The horses were very familiar with the paths and the time they needed to return to the barn. I think this was one of the rare occasions that Lynn was forced to concede to a schedule other than her own. The horses paused, as was their habit, to rest in an esplanade at roughly the half-way point in the forest. Lynn did not share this need to stop and urged them to continue. No way! The horses did not budge until they were ready, returning at the scheduled time to the stable.

A game during the International Symposium on Photosynthetic Prokaryotes (ISPP2000) in Barcelona. Photo by E. Castillo.

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Lynn sent many people, young students then, from her lab for short research stays at our university, the UAB. These were memorable times of intensive field and lab work, as well as mutual learning. The students became acquainted with Spanish and Catalan culture (history, architecture, Gaudi, gastronomy, language). It is difficult to forget Greg Hinkle, nearly 2 m tall, drinking from the porr贸n and wearing the barretina that we gave him at the farewell party we organized when he returned to the States. And in Amherst, at her home, with its surrounding woods filled with squirrels, doves, cardinals, mockingbirds, and so many other kinds of suburban wildlife. Roosevelt, her dog, followed her everywhere, even accompanying her to her lectures, which he endured stoically from his place in the front row. He showed the patience that Lynn rarely had. Sometimes she interrupted her talk to reassure

Visit to the Alexandria Library, December 2003 Photo courtesy of M. Vallmitjana.

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him. Unfortunately, the owners of the magnificent building (usually rented for celebrations) in front of Lynnâ&#x20AC;&#x2122;s house did not always share her love for that dog, especially during wedding celebrations held in the large garden when, during the ceremony Roosevelt would help himself to a sampling of the buffet set up for the reception. In Amherst, the sky is as beautiful as the ground, especially in autumn, when the light filters through the yellow maples, illuminating the ground below. The soil is a lush multicolored carpet, where your feet sink deeply in the leaf cover, which rustles as you tread on it. Several places close to Lynnâ&#x20AC;&#x2122;s house are of great interest, both ecologically and simply as a beautiful landscape, criss-crossed with trails and natural preserved spaces, like the Hawley Bog Preserve, one of the few remaining examples of a New England bog in its natural state. And who can forget Lynnâ&#x20AC;&#x2122;s many eccentricities. A favorite example is a two-day excursion to Harvard Forest, with over 40 students in tow for fieldwork and workshops. Days before, Lynn had bought an enormous quantity of food at the supermarket that we needed to prepare for cooking. I chopped what felt like a mountain of pumpkins, peppers, potatoes, tomatoes, and all kinds of vegetables, which we then cooked in pots big enough for us to have hidden in. On no account would she have agreed about her students packing their own sandwiches or a lunch box. No, like a quartermaster in the army, provisions from A to Z and their transportation to the venue, a hotel-retreat with facilities for large meetings (kitchen, comfortable bedrooms, lecture halls), were organized. But Lynn had other plans for her overnight accommodations and set up a tent outside, welcoming the opportunity to sleep under the stars. Although eager to help her set up her tent, finding the right place for it and finally doing so was, at least for me, and I discovered that also for her, a challenge worthy of an expedition [ 180 ]

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Lynn and Carmen, U-Mass campus, 1999. Photo courtesy of M. Dolan.

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to Mt. Everest. But we reached it!!!! and we set off back to the hotel to have dinner and the first meeting. To go back however, posed an even more serious challenge because despite the star-lit sky we could barely find our way. We were relieved when, finally, we could see the lights of the hotel, where we arrived full of scratches from low-lying branches but also with mud and debris stuck to our clothing: we had inadvertently traipsed into the composting area. Ah, nature and its beauty! Our activities in Harvard Forest were centered around a historical geology itinerary. The surrounding fields were where the first Europeans had settled, built farms, and developed agriculture and animal husbandry. The settlement was gradually abandoned, a consequence of late 19th century industrialization. In 1938, an earthquake destroyed the trees and buildings. Since then, the property has been restored as a model for studying forest growth and succession and the dominance of plant species in a previously agricultural soil. At U-Mass Amherst, Tuesdays were Lynnâ&#x20AC;&#x2122;s busiest day. After coming home and preparing dinner, to which students were almost

A walk around Hawley Bogs. Barbara, Lynn, Carmen, Volodia, and Roosevelt. Photo courtesy of B. Duval.

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always invited, she would go back to the university to hold a seminar. With the help of Mike and Andrew, she prepared videos and reviewed material for discussion with her students, always in the company of Roosevelt. Working together at her department and her home, we put together the final version of the viewer’s guide to the video series Microcosmos. There was so much information to enter and her handwritten notes were extremely difficult to read (“Can you, Lynn, even read them?,” I asked her). At home, she was constantly in motion, surrounded not only by her sons, who came to visit, but also by students, colleagues, and friends. And as the door to Lynn’s house was always open, whether or not Lynn was at home, people who knew her would simply come in, call out to see if she was there, go into the kitchen, help themselves to whatever was in the fridge, and, if they had missed seeing Lynn, would leave behind a note to say that they had stopped by. This is not a conventional text, but then Lynn was not a conventional person. Not at all. She was a fighter, as her many colleagues would quickly learn, and she was a revolutionary. Her in-depth knowledge of biology and science in general were essential to her unique understanding of evolutionary biology. She was enthusiastic, charismatic, and inspiring. But she was also contradictory, that most human of features, and controversial until the end. She was unique, she was Lynn Margulis. From here I send out my love and admiration, out into the micro and macro cosmos with which she acquainted us, wishing to reach her in symbiosis with all the diversity that shapes them. If we had to define her in three keywords, we would say: Lynn Margulis•Lynn Margulis•Lynn Margulis. 3 [ 183 ]

I died for Beauty - but was scarce Adjusted in the Tomb,

When One who died for Truth was lain In an adjoining room -

He questioned softly “Why I failed”? “For Beauty”, I replied -

“And I - for Truth - Themself are one We Brethren, are”, He said.

And so, as Kinsmen, met a Night We talked between the Rooms -

Until the moss had reached our lips And covered up - our names.

Emily Dickinson (1830-1886)

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[Previous page] Emily Dickinsonâ&#x20AC;&#x2122;s gravestone decorated with people giftâ&#x20AC;&#x2122;s. Photo by C. Puche.

about the authors and collaborators authors

Ricardo Amils

Full Professor of Microbiology, Autonomous University of Madrid.

Senior Scientist, Astrobiology Center (INTA-CSIC). Interdisciplinary Scientist of the Mars Express Mission. Member of the Solar System Exploration Working Group of the European Space Agency

(2011-2013). Member of the Editorial Boards of International Microbiology, Astrobiology, Extremophiles, and Microbial Cell Factories.

Eva Barreno

Full Professor of Botany. University of Valencia. Director of the Plant Biodiversity and Ecophysiology group of the Cavanilles Institute

of Biodiversity and Evolutionary Biology. Interdisciplinary studies in lichens symbiotic microalgae and plants. PROMETEO award “Excellence in Research” (Generalitat Valenciana). Associate member

of the European project Keys to Nature. Academic coordinator of the UNESCO Chair of the University of Valencia.

Carmen Chica

Psychologist. Autonomous University of Barcelona, Master in Science Communication. Translator of books and articles on history and philosophy, and science. Managing coordinator of International

Microbiology. Member of the team of the Catalan Foundation for Scientific Research and Innovation for activities focused on science dissemination and promotion of scientific and technical studies (2001–2011). Animal lover.

[ 187 ]

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Martí Domínguez

Biologist, writer, and journalist. Professor of journalism and litera-

ture, University of Valencia. Editor-in-chief of Mètode, a journal on research dissemination. Winner of the Spanish journalism award.

His novels, based on relevant characters of art and science, such as Buffon, Goethe, Voltaire, Cézanne, have received numerous awards. Domínguez writes for the newspapers El País and La Vanguardia.

Some of his articles have been compiled in Peiximinuti (1993) and Bestiari (2000).

Alícia Duró i Sans

Biologist. Researcher and curator at the Institute of Marine Sciences,

CSIC. Museologist for twenty years at the Science Museum in Barcelona (CosmoCaixa), which introduces biology through hands-on

experiments, workshops, etc. Participant in biological and oceanographic cruises and research projects on marine zooplankton ecology,

biogeography, taxonomy (area of expertise: Chaetognatha), phylogeny, and natural history collections.

Marta Estrada

Biologist. Previous director of the Institute of Marine Sciences,

CSIC. She has conducted oceanographic exploration projects in the

Mediterranean and all the major oceans of the world. Her studies

focus on the ecology of phytoplankton and on populations of harmful algae, with the aim of identifying the mechanisms underlying

their proliferations. Member of the Royal Academy of Sciences and Arts of Barcelona and of the Institute for Catalan Studies. Recipient of the Creu of St. Jordi of the Catalan Government.

[ 188 ]

about the authors and collaborators

Isabel Esteve

Biologist. Full Professor of Microbiology, Autonomous University of Barcelona. Field of expertise: phototrophic microorganisms and high-

resolution microscopy techniques. Research: microbial mats of the Banyoles Lake area and Ebro Delta; bioremediation of heavy metals

by phototrophic microorganisms in natural environments; bacterial diversity in extreme environments; vertical migration of phototrophic bacterial populations in hypersaline microbial mats.

Ramon Folch

Biologist and ecosociologist. Director (1975–1982) of the Environmen-

tal Services of Autonomous Government of Catalonia. Environmental consultant in several countries, mainly in the Mediterranean, Africa, Americas, and Southeast Asia. Previous consultant in Environmental

Management to UNESCO and member of the UNESCO monitoring committee of the Rio Conference (Paris). Since 1994, director of his own company (ERF-Gestió i Comunicació Ambiental, SL), on

environmental project consultancy in the fields of territorial communication and management, as well as in project research and scientific publishing. Member of the Institute for Catalan Studies.

Núria Gaju

Biologist. Professor of Microbiology, Autonomous University of Barcelona (UAB). Research: anoxygenic phototrophic sulfur bacteria;

plancktonic and benthic natural ecosystems; predation and mutually beneficial associations as a model of bacterial symbioses; microbiology

of the phyllosphere; assessment of the heterogeneity of the community structure in response to environmental changes. Vice-Dean of the UAB–Finance, Services, and Infrastructure. [ 189 ]

once upon a time lynn margulis

Ricardo Guerrero

Full Professor of Microbiology at the University of Barcelona. Adjunct Professor at the University of Massachusetts-Amherst. PhD

in Sc., University of Barcelona (1970). Research Fellow at the University of California-Davis (1972-1973). Member of the Institute

for Catalan Studies, and its Scientific Secretary. Fellow of the Linnaean Society, the American Academy of Microbiology, and the Aca-

demia Europaea. President of the Spanish Society for Microbiology (2006-present). Founder, with Lynn Margulis (1998) of the journal

International Microbiology. Curator of the “Planet Life” exhibit of the Natural History Museum of Barcelona (2009-present). Professional and personal partner of Lynn Margulis from 1983 until her death.

Rubén López

Biologist. Research Professor at the Center for Biological Research of the CSIC. Postdoctoral fellow in international centers. He has collaborated with outstanding scientists, including J.H. Backing and A.

Tomasz, in the genetics, physiology, and metabolism of pneumococci, his field of expertise. Research: characterization of Streptococcus pneu-

moniae genes; sequencing mutations responsible for the unencapsulated phenotype; transcriptional analysis of DNA regions involved in the capsular polysaccharide and polysaccharide biosynthesis by different serotypes of Escherichia coli and by pneumococcal strains.

Jordi Mas-Castellà

Biologist, University of Barcelona. MBA from ESADE, Barcelona.

Research laboratory manager. Research in microbial ecology studying the accumulation of bacterial polymers in nature and its applications

in biotechnology. Postdoctoral fellow at the the University of Massa[ 190 ]

about the authors and collaborators

chusetts-Amherst, and at the University of Ghent, Belgium. Former

joint director and director of the Catalan Foundation for Research

and Innovation. Promoter and organizer of Science Week in Catalonia. Scientific adviser in public and private organizations.

Xavier Maymó-Gatell

Biologist, business executive and writer. “La Caixa” fellow and a Cornell University PhD. Research in bacterial reductive dechlorination of

chloroethenes; isolation and phylogenetic characterization of microorganisms. Assistant Professor of Microbiology. Has published in Science as a first author. In 1997, he left his professional life as a scientist, and

began a new career in business management. Has been a senior associate at McKinsey & Co., held a top management position at Crèdit

Andorrà and managed the wealth of an ultra-high-net-worth single family office. At present, Xavier is the CEO -and shareholder- of Fimarge, a wealth management co. in Andorra.

Balbina Nogales

Biologist. Associate professor at the University of the Balearic Islands. Research: Microbiology of marine hydrocarbon pollution and deg-

radation; microbial communities and ecologically relevant organisms (Pseudomonas, Roseobacter). Member of the Environmental Biotechnology Group of the EU-US Task Force on Biotechnology Research.

Carles Pedrós-Alió

Biologist, University of Wisconsin, Madison (USA). Research Profes-

sor at the Institute of Marine Sciences, CSIC. Previous director of the Department of Marine Biology and Oceanography. Editor for the

journals Marine Scientia and International Microbiology. Research: mi[ 191 ]

once upon a time lynn margulis

crobial ecology of freshwater and hypersaline systems of the Atacama Desert. Oceanographic cruises in Antarctica, the Mediterranean, and the Arctic. Winner of the Barcelona scientific research award.

Juli Peretó

Chemist-Biochemist. Associate Professor of Biochemistry and Molecular Biology, University of Valencia (UV). Research: origin of life, evolution of metabolism in bacteria endosymbionts of insects. Research stays

at the Universities of Essen, of Pennsylvania and of Paris XI in Orsay.

He has been Vice-Rector of the UV, Secretary and Vice-President of ISSOL (International Society for the Study of the Origin of Life). Director of the Erasmus Intensive Programme “Origin, Evolution and Future of the Biosphere”. Collaborator with the National Research Council in the creation of new research centers. Co-author, with Andrés Moya, of

the book Simbiosis. Seres que evolucionan juntos (Síntesis, 2011). Member

of the Institute for Catalan Studies.

Mercè Piqueras

Biologist, science writer, science editor, and translator. Associate

Editor of International Microbiology. Member of the Editorial Board of Treballs de la Societat Catalana de Biologia and Panacea. Freelance

contributor to various magazines (print and online) and newspapers. Scientific coordinator of the Museu Blau project of the Barcelona

Natural History Museum (2009–2011). Past President of the Cata-

lan Association for Scientific Communication (ACCC) and Past Vice-President of the Catalan Society for the History of Science

and Technology. ACCC representative at the European Union of

Science Journalist’s Associations (EUSJA). Author of the blog La lectora corrent (The common reader). [ 192 ]

about the authors and collaborators

Eduard Punset

Law degree from Madrid University and Master in economics at the

London School of Economics. He worked for ten years for the BBC

and The Economist in London and in the International Monetary Fund in Washington. In Spain he participated in the political transition to democracy and in the negotiation for Spanish entry to the

European Union as Minister of European Relations. He has been chairman of the Bull Technological Institute, Professor of innovation and technology of the Business Institute in Madrid and professor of

Science, Technology and Society at the Chemical Institute of Sarria (Ramon Llull University). He is best known as director and producer

of Redes (“Networks”) broadcasted in the Spanish Public Television channel La2; he has hosted this weekly television program on the

public understanding of science to a worldwide Spanish-speaking au-

dience since 1996. He has several books published in Spain , EEUU, Portugal and other countries.

Gemma Reguera

Biologist. Associate Professor in the Department of Microbiology

and Molecular Genetics at Michigan State University. Postdoctoral Fellow at Harvard Medical School. Research: mechanisms governing

microbial processes in time and space, microbial adaptive responses to the environment and their biotechnological applications; bacterial

interactions with surfaces and microbial processes within biofilms; applications of Geobacter spp.; search for biogeochemical reactions electrically coupled to the reduction of oxygen in sediments. Science

dissemination: associate blogger of M. Schaechter’s American Society for Microbiology blog Small things considered.

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Concepción Ronda

Biologist. Investigator at the Center for Biological Research, CSIC, Madrid. Research: lytic enzymes of the pneumococcal system; genetic

characterization of pathogenic microorganisms; morphogenesis of Sindbis viruses; mechanisms of antibiotic resistance. Participant in surveys of

European microbiology journals; development of science and technology quality assessment indicators and of criteria for accessing Spanish

microbiological and medical journals in the main international directories and databases.

Héctor Ruiz

Biologist. Founder and general director of Digital-Text, a company that specializes in the creation and commercialization of digital textbooks, interactive multimedia, educational contents, and didactic exercises. Visiting lecturer University of Southern California, University

of Washington, and University of Melbourne. Director of the popu-

lar science journals Eureka and Omnis cellula. Former president of the Edutech Clúster: an association of companies that promote educational technologies. He was awarded the Young Entrepreneur prize of the International Association of Young Enterprising 2011, Catalonia.

Moselio Schaechter

Distinguished Professor Emeritus, Tufts University, Boston (USA). He chaired the Department of Molecular Biology and Microbiology

at Tufts for 23 years. Currently, Adjunct Professor, San Diego State

University, and University of California at San Diego. President, American Society for Microbiology (1985–1986). He has devoted

himself to the width and breadth of the microbial world through a physiological approach, mainly regarding the growth of bacteria. In [ 194 ]

about the authors and collaborators

addition to a large body of scientific work, he has authored or edited

books on microbial physiology and pathogenesis, including Mechanisms of Microbial Diseases, Escherichia coli and Salmonella, and the Encyclopedia of Microbiology. He loves wild mushrooms, a fondness

readily apparent in his book In the Company of Mushrooms. Founder

of the blog Small Things Considered [http://schaechter.asmblog.org], where he shares the width and depth of microbial activities on this planet and his fascination for them.

Nicole Skinner

Biologist. Science communicator, editor, and translator. Scientific

Advisor at the Institute for Catalan Studies and Hub Officer for the Academia Europaea’s Barcelona Knowledge Hub, she specialises in the

organisation of science outreach activities and in the international pro-

motion of research and scientific publications. She is Managing Editor of the journal Contributions to Science and Adjunct Secretary and Webmaster of International Microbiology.

Mónica Solé Rojo

Biologist. Master in Scientific Communication and Ph.D. in Molecular Biology. Research in basic microbiology, genetics, and microbial

ecology. Former head of the Scientific Programs Department at Cos-

moCaixa, the Science Museum of Madrid. She has worked for many years in the fields of basic microbiology research, science dissemination, and projects related to the social application of science. Current

interest: Queen Bee breeding and selection, to obtain lines with interesting characters for honey production and other hive products, as

well as to counter the severe problem posed by pesticides and other environmental threats.

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Begoña Vendrell

Biologist and cultural anthropologist. Coordinator of the project El

mar a fondo (“The depth of the sea”) at the Institute of Marine Science (CSIC), an educational programme on oceans and seas developed in a

virtual interactive platform. Organizer of science courses for teachers and Education and outreach activities at international level. She has worked on benthic-pelagic coupling processes on the continental shelves of the

Antarctic, and work on traditional ecological knowledge of fishers in MPAs and surrounding areas. Associate Editor of Omnis cellula, a popular science magazine. Founder and member of APECS-Spain.

Montse Vallmitjana

Biologist. High school teacher. Organizer of scientific activities, con-

ferences, and workshops for students and teachers. Member of the Catalan Society for Biology. Coordinator of a NGO working for and in Burkina Faso. She carried out the first interview with Lynn Margulis in Catalonia, for the Catalan science magazine Ciència, in 1983.

Laura Villanueva

Biologist. Research scientist in Geomicrobiology, Department of Marine Organic Biogeochemistry, Royal Netherlands Institute for Sea

Research, NIOZ. Postdoctoral fellowships: Harvard Center for Sys-

tems Biology (MA, USA); University of Massachusetts, Department of Microbiology (MA, USA). PhD thesis, Department of Microbiol-

ogy, University of Barcelona, promotor: Prof. R. Guerrero. Research interests: molecular biology and genetics in microbiology and micro-

bial ecology (expression of functional genes in marine environments, combined methodological approaches in microbial ecology: signature lipid biomarkers and DNA/RNA methods, microbial mats). [ 196 ]

about the authors and collaborators

collaborators

Mercedes Berlanga

Biologist. Professor of food safety and hygiene, University of Barcelo-

na. Associate editor of the journal International Microbiology. Research: microbial ecology, antibiotic resistance, and nosocomial infections. Science disseminator.

Carme Puche

Filmmaker, writer, and journalist. Director of the short film Camille, about the artist Camille Claudel, which was awarded Best Screenplay at the International Sitges Film Festival (section: New Authors).

Carles Puche

Natural science illustrator. Teacher. Exhibition curator and photog-

rapher; collaborations with, among others, National Geographic, Cosmocaixa, Barcelona Botanical Gardens.

Jordi Rabascall

Designer and musician. Emotional and conceptual design. Art director, project manager, publisher.

Wendy Ran

Cell biologist. PhD, Boston University Medical School. Associate edi-

tor of International Microbiology journal. Freelance English language copy editor for several science and medical journals, investigators, publishers, and websites.

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Sandra Young

English teacher, translator of texts for books and journals (English into Spanish and Catalan, and viceversa). English proofreader.

[ 198 ]

This book was printed October 30, 2013 in Vanguard Gràfic (Sant Adrià del Besòs), after months of work of the whole team. We hope you enjoy it.