Bibliography And Biography Differences Between Prokaryotes

Lynn Margulis, (born March 5, 1938, Chicago, Illinois, U.S.—died November 22, 2011, Amherst, Massachusetts), American biologist whose serial endosymbiotic theory of eukaryoticcell development revolutionized the modern concept of how life arose on Earth.

Margulis was raised in Chicago. Intellectually precocious, she graduated with a bachelor’s degree from the University of Chicago in 1957. Soon after, she married American astronomer Carl Sagan, with whom she had two children; one, Dorion, would become her frequent collaborator. The couple divorced in 1964. Margulis earned a master’s degree in zoology and genetics from the University of Wisconsin at Madison in 1960 and a Ph.D. in genetics from the University of California, Berkeley, in 1965. She joined the biology department of Boston University in 1966 and taught there until 1988, when she was named distinguished university professor in the department of botany at the University of Massachusetts at Amherst. She retained that title when her affiliation at the university changed to the department of biology in 1993 and then to the department of geosciences in 1997.

Throughout most of her career, Margulis was considered a radical by peers who pursued traditional Darwinian “survival of the fittest” approaches to biology. Her ideas, which focused on symbiosis—a living arrangement of two different organisms in an association that can be either beneficial or unfavourable—were frequently greeted with skepticism and even hostility. Among her most important work was the development of the serial endosymbiotic theory (SET) of the origin of cells, which posits that eukaryotic cells (cells with nuclei) evolved from the symbiotic merger of nonnucleated bacteria that had previously existed independently. In this theory, mitochondria and chloroplasts, two major organelles of eukaryotic cells, are descendants of once free-living bacterial species. She explained the concept in her first book, Origin of Eukaryotic Cells (1970). At the time, her theory was regarded as far-fetched, but it has since been widely accepted. She elaborated in her 1981 classic, Symbiosis in Cell Evolution, proposing that another symbiotic merger of cells with bacteria—this time spirochetes, a type of bacterium that undulates rapidly—developed into the internal transportation system of the nucleated cell. Margulis further postulated that eukaryotic cilia were also originally spirochetes and that cytoplasm evolved from a symbiotic relationship between eubacteria and archaebacteria (seearchaea).

Her 1982 book Five Kingdoms, written with American biologist Karlene V. Schwartz, articulates a five-kingdom system of classifying life on Earth—animals, plants, bacteria (prokaryotes), fungi, and protoctists. The protist kingdom, which comprises most unicellular organisms (and multicellular algae) in other systems, is rejected as too general. Many of the organisms usually categorized as protists are placed in one of the other four kingdoms; protoctists make up the remaining organisms, which are all aquatic, and include algae and slime molds. Margulis edited portions of the compendium Handbook of Protoctista (1990).

Another area of interest for Margulis was her long collaboration with British scientist James Lovelock on the controversial Gaia hypothesis. This proposes that the Earth can be viewed as a single self-regulating organism—that is, a complex entity whose living and inorganic elements are interdependent and whose life-forms actively modify the environment to maintain hospitable conditions.

In addition to Margulis’s scholarly publications, she wrote numerous books interpreting scientific concepts and quandaries for a popular audience. Among them were Mystery Dance: On the Evolution of Human Sexuality (1991), What Is Life? (1995), What Is Sex? (1997), and Dazzle Gradually: Reflections on Nature in Nature (2007), all cowritten with her son. She also wrote a book of stories, Luminous Fish (2007). Her later efforts were published under the Sciencewriters Books imprint of Chelsea Green Publishing, which she cofounded with Dorion in 2006.

Margulis was elected to the National Academy of Sciences in 1983 and was one of three American members of the Russian Academy of Natural Sciences. She was awarded the William Procter Prize of Sigma Xi, an international research society, and the U.S. National Medal of Science in 1999. In 2008 she received the Darwin-Wallace Medal of the Linnean Society of London. She was a coauthor, with Dorion, of Encyclopædia Britannica’s article on life.


Eukaryotes have long been thought to have arisen by evolving a nucleus, endomembrane, and cytoskeleton. In contrast, it was recently proposed that the first complex cells, which were actually proto–eukaryotes, arose simultaneously with the acquisition of mitochondria. This so–called symbiotic association hypothesis states that eukaryotes emerged when some ancient anaerobic archaebacteria (hosts) engulfed respiring αproteobacteria (symbionts), which evolved into the first energy–producing organelles. Therefore, the intracellular compartmentalization of the energy-converting metabolism that was bound originally to the plasma membrane appears to be the key innovation towards eukaryotic genome and cellular organization. The novel energy metabolism made it possible for the nucleotide synthetic apparatus of cells to be no longer limited by subsaturation with substrates and catalytic components. As a consequence, a considerable increase has occurred in the size and complexity of eukaryotic genomes, providing the genetic basis for most of the further evolutionary changes in cellular complexity. On the other hand, the active uptake of exogenous DNA, which is general in bacteria, was no longer essential in the genome organization of eukaryotes. The mitochondrion–driven scenario for the first eukaryotes explains the chimera–like composition of eukaryotic genomes as well as the metabolic and cellular organization of eukaryotes.

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