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Tamarin: Principles of Genetics, Seventh Edition
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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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DNA replication, including formation of final Okazaki fragment
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Removal of nal primers after the replication of linear DNA creates single-stranded ends.
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the chromosomes. First, the guanine-rich DNA can form complex structures. Biochemists have discovered that four guanines can form a planar G-tetraplex, with the four bases hydrogen bonded to each other ( g. 15.29). Several structures have been hypothesized to explain the novel ends of these chromosomes ( g. 15.30). Second, proteins have been discovered that bind to the 3 ends of telomeres. In the ciliate Oxytricha nova, a protein called the telomere end-binding protein (TEBP) attaches to the 3 ends of telomeres and protects them ( g. 15.31). Finally, a novel structure called the t-loop has been discovered at the end of mammalian telomeres. This loop forms at the ends of chromosomes under the direction of a protein called TRF2 (telomere repeat-binding factor), which causes the 3 end of the chromosome to loop around and interdigitate into the double helix, forming the loop ( g. 15.32). How do cells keep track of the number of their telomeric repeats Proteins have been isolated that bind to telomeres (Rap1 in Saccharomyces cerevisiae, TRF1 in human beings). By mutating these proteins or the telomeric sequences, scientists have changed the equilibrium number of telomeric repeats.This led to the current model that the cell counts the number of these proteins bound to the telomeres, not the number of telomeres directly, to know whether telomeres should be added. This is a very active area of research. In yeast, protozoa, and other single-celled organisms, telomerase is active, keeping the ends of the chromo-
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Telomerase extends telomeres using telomerase RNA (red) as a template. Gap lling (green) by DNA polymerase and ligase complete the double helix. (Source: Data
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from Shippen-Lentz and Blackburn, Science, 247:550, 1990.)
somes at the appropriate lengths. These cells can divide potentially forever. However, in most cells of higher organisms, telomerase is not active, and the ends of the chromosomes get shorter with each cell division. At a certain telomeric length, the cells no longer divide. However, if telomerase becomes active, and the ends of the chromosomes lengthen, a signal is conveyed to keep cells dividing, which can lead to cancerous growth. In fact, human telomerase was isolated from an immortal cell line (HeLa) derived from cervical cancer cells. Thus,
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
15. The Eukaryotic Chromosome
The McGraw Hill Companies, 2001
The Eukaryotic Chromosome
A G-tetraplex can form from four guanines in a plane, hydrogen bonded with each other. (Source: Data from
Yong Wang and Dinshaw J. Patel, Solution structure of the human telomeric repeat d[AG3 (T2AG3)3] G-tetraplex, Structure, 1:263 82, December 15, 1993.)
Figure 15.31 Telomere of the ciliate Oxytricha nova shown bound by the dimeric protein called telomere end-binding protein (TEBP). The and subunits of the protein form a deep cleft in which the 3 end of the telomere lies. The folding of the protein into its nal form around the DNA may only occur after the DNA has bound, explaining how the DNA could be recognized and placed into such a deep cleft. (Reprinted
courtesy of Dr. Martin Horvath.)
The C-Value Paradox
Why do eukaryotes have so much DNA, and why is there huge variation in the DNA content between species of comparable complexity These questions de ne the C-value paradox, in which C refers to the quantity of DNA in a cell. For an example of the paradox, although human beings have 3.3 billion base pairs in the haploid genome, an amoeba has more than 200 billion base pairs. And although an average bony sh has over 300 billion base pairs of DNA in its haploid genome, the Japanese puffer sh has less than half a billion base pairs. If the basic bony sh pattern can be created with less than half a billion base pairs, why does the average bony sh have over 600 times that much DNA What is this excess DNA doing To explain the C-value paradox, researchers examined the repetitiveness of DNA, and more recently, probed and sequenced DNA to understand its properties.
Based on G-tetraplexes ( g. 15.29), the illustrated structure can form at the very tip of a telomere. The sequence d(GGTTGGTGTGGTTGG) is shown forming a four-stranded structure. (Reproduced, with permission, from the
Annual Review of Biophysics and Biomolecular Structure, Volume 23, 1994 by Annual Reviews, Inc.)
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