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The Endosymbiont Hypothesis

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The Endosymbiont Hypothesis
An interesting question can produce some interesting answers. One of these is the question of what living things are made of. There are many possible answers, depending on who you ask. A young child asking his parents may simply be told: “Stuff”. A chemist may go into the detail of the basic common organic elements: Carbon, Hydrogen, Oxygen, Nitrogen and Phosphorous (Starr et al., 2009) and the different compounds that are formed. As biologists, we tend to look at the composition of living things at a relatively larger scale viz. from a scale of less than 0.1 nm to a scale of more than 1 nm (Starr et al., 2009a). I say relatively because, although this is still very small, it is much larger than the scale at which chemists work. This scale encompasses the realm of cells and cellular structures. The basic unit of life, as defined by the cell theory, pioneered by Schwann and Schleiden in 1838 (Mazzarello, 1999; Wayne, 2009), is the cell. Since that time, a lot of development has occurred in the quest to find the basis of life. It is now known that there are in fact two distinct kinds of cells, i.e. the prokaryotic and eukaryotic cells. The distinct differences between the two are easily discernible with modern technology such as electron microscopes. These differences include: the presence of a nucleus in the eukaryotes, and the absence of one in the prokaryotes; Eukaryotes also have distinct membrane-bound organelles, while prokaryotes do not (Stanier et al., 1970a; Mauseth, 2009). It is after discovery and examination of these organelles that first led to the Endosymbiont Hypothesis or Endosymbiont Theory, as it is sometimes called (both terms are used throughout). In this essay we shall look at the history of the theory, how it was formed, who the most influential collaborators were, and how it is applied to the evolution of certain organelles, in particular, the chloroplast and mitochondria.
The development of the Endosymbiont Theory occurred over a



References: DOUGLAS, S.E., 2004. The Molecular Biology of Cyanobacteria, in: Advances in Photosynthesis and Respiration. Springer Netherlands, pp. 91–118. ENCYCLOPEDIA BRITANNICA 2009, 2009. Heinrich Anton de Bary. Encyclopedia Britannica 2009 Ultimate Reference Suite. FENCHEL, T., FINLAY, B.J., 1991. Endosymbiotic Methanogenic Bacteria in Anaerobic Ciliates: Significance For the Growth Efficiency of the Host. Journal of Eukaryotic Microbiology 38, 18–22. HARTMANN-PETERSEN, P., GERRANS, G.C., Hartmann-Petersen, R., 2007. Encyclopaedia of Science & Technology, 2nd ed. New Africa Books (Pty) Ltd., Claremont, South Africa. MARGULIS, L., 1971. The Origin of Plant and Animal Cells. American Scientist 59, 230 – 235. MARRS, K.A., 2004. The Endosymbiotic Theory [WWW Document]. The Endosymbiotic Theory. URL http://www.biology.iupui.edu/biocourses/N100/2k4endosymb.html MAUSETH, J.D., 2009 MAZZARELLO, P., 1999. A unifying concept: the history of cell theory. Nature Cell Biology 1, E13. MOORE, R., CLARK, W.D., VODOPICH, D.S., 1998. Chapter 3: Structure and Function of Plant Cells, in: Botany. WCB/McGraw-Hill, United States of America. MORDEN, C.W., SHERWOOD, A.R., 2002. Continued evolutionary surprises among dinoflagellates. Proc Natl Acad Sci USA 99, 11558 –11560. NASS, S., NASS, M.M.K., 1963. INTRAMITOCHONDRIAL FIBERS WITH DNA CHARACTERISTICS. The Journal of Cell Biology 19, 613 –629. RIS, H., PLAUT, W., 1962. Ultrastructure of DNA-containing areas in the chloroplast of chlamydomonas. The Journal of Cell Biology 13, 383 –391. SAPP, J., CARRAPIÇO, F., ZOLOTONOSOV, M., 2002. Symbiogenesis: The Hidden Face of Constantin Merezhkowsky. History & Philosophy of the Life Sciences 24, 413 – 440. STANIER, R.Y., DOUDOROFF, M., ADELBERG, E.A., 1970a. Chapter 2: The nature of the microbial world, in: The Microbial World. Prentice-Hall, Inc., Englewood Cliffs, New Jersey, p. 52. STANIER, R.Y., DOUDOROFF, M., ADELBERG, E.A., 1970b. Chapter 25: Mutualistic Endosymbioses, in: The Microbial World. Prentice-Hall, Inc., Englewood Cliffs, New Jersey, p. 741. STARR, C., TAGGART, R., EVERS, C., STARR, L., 2009a. Molecules of Life, in: Biology: The Unity and Diversity of Life. Brooks/Cole, Cengage Learning, Belmont, CA. STARR, C., TAGGART, R., EVERS, C., STARR, L., 2009b. Time Line for Life’s Origin and Evolution, in: Biology: The Unity and Diversity of Life. Brooks/Cole, Cengage Learning, Belmont, CA. STARR, C., TAGGART, R., EVERS, C., STARR, L., 2009c. Where did organelles come from, in: Biology: The Unity and Diversity of Life. Brooks/Cole, Cengage Learning, Belmont, CA. TORDAY, J.S., REHAN, V.K., 2012. Prokaryotes versus Eukaryotes, in: Evolutionary Biology: Cell-Cell Communication and Complex Disease. Wiley - Blackwell, Hoboken, New Jersey. WALLIN, I.E., 1923. Symbionticism and prototaxis, two fundamental biological principles. Anat. Rec. 26, 65–73. WALLIN, I.E., 1925. On the nature of Mitochondria: IX. Demonstration of the bacterial nature of mitochondria. American Journal of Anatomy 36, 131–149. WAYNE, R., 2009. Plant Cell Biology: From Astronomy to Zoology. Elsevier Inc., United States of America. WELLS, W.A., 2005. There’s DNA in those organelles. The Journal of Cell Biology 168, 853.

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