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III. Molecular Genetics
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15. The Eukaryotic Chromosome
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The Eukaryotic Chromosome
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enerally, eukaryotic cells are large, and prokaryotic cells are small. For example, an average eukaryotic cell is about 50 m in diameter, whereas an average bacterium is about 5 m in length. The average virus is about 0.05 m in diameter. These size differences occur because eukaryotic cells have complex substructures and internal architecture that prokaryotic cells lack. Since we believe that prokaryotic cells depend on diffusion to exchange materials with the environment, they would have to be small. And viruses, intracellular parasites, would of necessity be very small. There are, of course, exceptions. In 1999, a team of scientists from Germany, Spain, and the United States isolated large sulfur bacteria off the Namibian coast of Africa and named them Thiomargarita namibiensis, the sulfur pearl of Namibia.These bacteria can be almost half a millimeter in diameter, the size of the period at the end of this sentence ( g. 1). Based on the sequence of 16S ribosomal DNA, these bacteria were shown to be close relatives of other marine sulfur bacteria. They are almost one hundred times the volume of the bacteria previously believed to be largest, Epuliscium shelsoni, known only from the intestine of the brown surgeon sh. The smallest prokaryotes are the Mycoplasmas, at about 0.2 m in diameter, rivaling the viruses in size. They are animal pathogens and decomposing organisms. The smallest eukaryote, Ostreococcus tauri, a green alga found in the plankton, was discovered in 1994 from a water sample in a French lagoon on the Mediterranean Sea by a group of French sci-
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entists. These organisms are less than 1 m in diameter. Scientists believe that the lower limit on the size of a cell (not counting viruses) is about 200 nm (0.2 m), based on the size of DNA and ribosomes that a cell must contain. With T. namibiensis as the largest prokaryote, we note that the largest eukaryotic cell with a single nucleus is most likely the ostrich egg. The largest organisms are the blue whale, Balaenoptera musculus, weighing in
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at 118,000 kilograms; giant redwood trees, Sequoiadendron giganteum, 100 meters tall and weighing 5.5 million kilograms; a quaking aspen clone, Populus tremuloides, weighing 6 million kilograms; and Armillaria bulbosa, a fungus. In 1992, three scientists from the University of Toronto and Michigan Technological University, using restriction fragment length polymorphisms (RFLPs) and polymerase chain reaction (PCR) techniques, showed that the huge hyphal mass of this tree-root colonizing fungus growing in a forest in northern Michigan was a single organism. It covered about eight hectares, probably weighed more than 10,000 kilograms, and probably has existed for more than 1,500 years. Although we don t want to get distracted by the oddities and extremes of nature, size differences are remarkable.
The bacterium Thiomargarita namibiensis shown with a fruit y (Drosophila virilis, 3 mm in length) for size comparison. The arrow points to a single bacterial cell, 0.5 mm wide, bright with sulfur inclusions. Above the cell are empty sheaths of dead bacteria. (From H.N. Schulz, et al., Dense populations of a
giant sulfur bacterium in Namibian Shelf Sediments in Science, Vol. 284, pp. 493 95, April 16, 1999. Reprinted by permission of the American Association for the Advancement of Science.)
stretched molecules relax. From other sources, primarily UV absorbance studies, it was estimated that the largest Drosophila chromosome had about 43 109 daltons of DNA. Results from the viscoelastic measurements indicated the presence of DNA molecules of between 38 and
44 109 daltons. Viscoelastic measurements of inversions, which changed the ratio of the arms but not the overall size of the chromosome, yielded similar results. However, a translocation that radically changed the size of the chromosome to 59 109 daltons resulted in an
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