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In the next several chapters, we examine the details of protein synthesis. We will see that DNA does possess the complexity required to direct protein synthesis. Although complementarity restricts the base opposite a given base in a double helix, there are no restrictions on the sequence of bases on a given strand. Later, we will show that each sequence of three bases in DNA speci es a particular amino acid during protein synthesis. The ge-
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Hydrogen bond N HC N To deoxyribose C N C C N CH O Adenine H N C N H N H O C C CH3
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H H N C N C O N
5 O Base Base
3 O
CH CH
OH 3
To deoxyribose
Guanine Figure 9.14
Cytosine
Hydrogen bonding between the nitrogenous bases in DNA.
Figure 9.15 Polarity of the DNA strands. Polarity is established by the 3 and 5 carbons of a given sugar. For example, moving down the left strand, the polarity is 5 3 (read as ve-prime to three-prime). Moving down the right strand, the polarity is 3 5.
C P P C P 5 5
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
9. Chemistry of the Gene1
The McGraw Hill Companies, 2001
Chemistry of Nucleic Acids
Alternative Forms of DNA
Melting (denaturation) temperature ( C)
Relationship of the number of hydrogen bonds (G-C content) and the thermal stability of DNA from different sources. (From J. Marmur and P. Doty, Jr., Relationship of the Number
of Hydrogen Bonds and the Thermal Bonds and the Thermal Stability of DNA from Different Sources, Journal of Molecular Biology, 5:109 112. Copyright 1962 Academic Press LTD.)
netic code gives the relationship of DNA bases to the amino acids in proteins.
The form of DNA we have described so far is called B DNA. It is a right-handed helix: it turns in a clockwise manner when viewed down its axis. The bases are stacked almost exactly perpendicular to the main axis, with about ten base pairs per turn (34 ; see g. 9.13c). However, DNA can exist in other forms. If the water content increases to about 75%, the A form of DNA (A DNA) occurs. In this form, the bases tilt in regard to the axis, and there are more base pairs per turn. However, this and other known forms of DNA are relatively minor variations on the right-handed B form. In 1979, Alexander Rich and his colleagues at MIT discovered a left-handed helix that they called Z DNA because its backbone formed a zigzag structure (fig. 9.18). Z DNA was found by X-ray crystallographic analysis of very small DNA molecules composed of repeating G-C sequences on one strand with the complementary C-G sequences on the other (alternating purines and pyrimidines). Z DNA looks like B DNA with each base rotated 180 degrees, resulting in a zigzag, left-handed structure ( g. 9.19). (The original con guration of the bases is referred to as the anti con guration; the rotated con guration is called the syn con guration.) Originally, it was thought that Z DNA would not prove of interest to biologists because it required very high salt concentrations to become stable. However, it was found that Z DNA can be stabilized in physiologically normal conditions if methyl groups are added to the cytosines. Z DNA may be involved in regulating gene expression in eukaryotes. We return to this topic in chapter 16 (box 9.3).
Replication
Percent G + C
Watson and Crick hinted in their 1953 paper how DNA might replicate. Their observation stemmed from the property of complementarity. Since the base sequence on one strand is complementary to the base sequence on the opposite strand, each strand could act as a template for a new double helix if the molecule simply unzipped, allowing each strand to specify the sequence of bases on a new strand by complementarity ( g. 9.17). Mutability would occur due to mispairings, other errors in replication, or damage to the DNA.
T T C C G A C G C T A A G A T T T
A A G G C T A G C C T A A G A T T Replication fork (Y-junction)
New strands
Location
DNA must reside in the nucleus of eukaryotes, where the genes occur on chromosomes, or in the chromosomes of prokaryotes and viruses. In both prokaryotes and eukaryotes, the majority of the cell s DNA is in the chromosomes. And all viruses contain either DNA or RNA. Thus, DNA ful lls all the requirements of a genetic material. RNA can ful ll the same requirements in RNA viruses and viroids.
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