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The centromere and telomeres are speci c functional regions of a chromosome. Centromeres isolated from yeast
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15. The Eukaryotic Chromosome
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The Eukaryotic Chromosome
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chromosomes have three consensus areas. Telomeres are tandem repeats of a short ( ve-base-pair to eight-base-pair) segment. Telomeric sequences are added to the ends of chromosomes by the enzyme telomerase, which uses RNA as a template for adding DNA nucleotides. The number of telomeric repeats varies, declining as a cell ages. Telomeric repeat number may control the ability of a cell to replicate and may be implicated in cancerous growth. Substructuring in the eukaryotic chromosome is demonstrated by G-, C-, and R-banding techniques. C-bands (constitutive heterochromatin) appear to be around the centromeres. These bands consist primarily of satellite DNA, which seems to have a structural role in the chromosome. G-bands (Giemsa bands) presumably represent intercalary heterochromatin and, also presumably, do not have an active transcriptional role. R-bands (reverse bands) appear be-
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tween the G-bands and represent intercalary euchromatin, the site of transcribed, structural genes. STUDY OBJECTIVE 3: To analyze the nature of the DNA in
eukaryotic chromosomes
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Eukaryotes have very large genomes with huge differences in DNA content between organisms similar in complexity, leading to the C-value paradox. Eukaryotic chromosomes contain both unique and repetitive DNA. Highly repetitive DNA is structural (centromeres, telomeres). Junk DNA is mainly short and long interspersed elements.These SINEs and LINEs are often present in hundreds of thousands of copies and can account for 50% of an organism s DNA. They are retrotransposons, transposons that jump by way of an RNA intermediate. Some functional genes also occur in many copies, such as ribosomal RNA genes, histone genes, and globin genes.
S O L V E D
PROBLEM 1: Why is higher-order chromosomal structure expected in eukaryotes but not prokaryotes Answer: The simplest explanation is the difference in amount of the genetic material in prokaryotes and eukaryotes. Since the average human chromosome has several centimeters of DNA, that DNA must be contracted to a size in which it can be moved during mitosis and meiosis without tangling and breaking. Nucleosomes provide the rst order of coiling, and then several levels of coiling of the nucleosomed DNA bring it down to a manageable size for nuclear divisional processes. PROBLEM 2: Why might we expect to see chromosomal
P R O B L E M S
Answer: The various patterns of chromosomal puf ng are expected because puf ng indicates transcription, the activity of speci c genes. Thus, since various tissues are different because they have different proteins, each tissue is expected to have a unique suite of active genes and thus a unique suite of puffs. Similarly, different stages in an insect s development would require different genes to be active, and different puffs should therefore appear at different stages of development. Some genes are active all the time because they specify proteins, such as ribosomal protein genes, that are needed all the time. Finally, environmental insults such as heat shock are known to induce a group of genes that are needed to react to the speci c insult, resulting in a suite of puffs that respond consistently to an environmental insult.
puffs that are tissue- and stage-speci c, constitutive, and environmentally induced
E X E R C I S E S
THE EUKARYOTIC CELL
A N D
P R O B L E M S *
1. Summarize the major differences between eukaryotes and prokaryotes, including the structures of their DNAs.
THE EUKARYOTIC CHROMOSOME
2. Summarize the evidence that the eukaryotic chromosome is uninemic.
3. What results would you get in the experiment shown in gure 15.1 if the eukaryotic chromosome were not uninemic, but instead had some other number of complete DNA molecules (e.g., binemic) 4. What are the major protein components of the eukaryotic chromosome What are their functions 5. What evidence is used to determine the length of DNA associated with a nucleosome What is a nuclease-hypersensitive site What functions are associated with these sites
*Answers to selected exercises and problems are on page A-18.
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