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No Fear provides access to Shakespeare for students who normally couldn’t (or wouldn’t) read his plays. It’s also a very useful tool when trying to explain Shakespeare’s wordplay!
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I tutor high school students in a variety of subjects. Having access to the literature translations helps me to stay informed about the various assignments. Your summaries and translations are invaluable.
Kathy B.
Teaching Shakespeare to today's generation can be challenging. No Fear helps a ton with understanding the crux of the text.
Kay H.
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While the leading strand and lagging strand replicate differently, each
individual nucleotide added to each strand is attached through the same
mechanism. In this section, we will examine the mechanism of nucleotide
attachment. Note: classes such as AP Biology do not require you to know
the topics covered in this section.
The Building Blocks of DNA Replication are Deoxyribonucleotides
The building blocks added on to a growing daughter strand are individual
nucleotides. Remember, in DNA the -OH group at the 2' position of the ribose
ring is missing. As a result, the substrates for DNA synthesis are called 2'
deoxyribonucleotides.
Figure %: 2' Deoxyribonucleoside triphosphate
Attached to each deoxyribose ring is a base group (C, G, A, or T) and a
triphosphate group. The three phosphates are designated alpha, beta, and gamma
(alpha being the closest to the ribose ring). These phosphates play key roles in
the addition of subsequent nucleotides to the daughter strand.
Addition Occurs Via a Nucleophilic Attack
Deoxyribonucleoside triphosphates, as we just stated, are the building blocks
of DNA. Recall, furthermore, that a complete polynucleotide strand of DNA has only
one phosphate group and that through this phosphate group each nucleotide is attached
to the next. Why then is the substrate a triphosphate instead of just a monophosphate?
The answer to this question lies in the chemistry underlying the addition of nucleotides
to a growing daughter strand of DNA.
While each nucleotide added to a growing DNA chain lacks an -OH group at its 2'
position, it retains its 3' -OH. This hydroxyl group is used to attack the alpha
phosphate group of an incoming nucleoside triphosphate. In the attack, the
3' -OH replaces the beta and gamma phosphates that are ejected from the complex
as a pyrophosphate molecule. The result is the formation of the phosphodiester
bond between the growing daughter strand and the next nucleotide. The 3' -OH of
the newly added nucleotide is now exposed on the end of the growing chain and can
attack the next nucleotide in the same way.
Figure %: Addition of Nucleotides to a Growing Daughter Strand
The figure above presents a simplified schematic of a growing polynucleotide
chain. The lines represent the ribose sugar with one 3' -OH branching from
it. Each p represents a phosphate group. This figure illustrates a number of
key points of DNA replication. First, we see that the parent strand is oriented
in the 3' to 5' direction. Second, each new nucleotide added to the growing
daughter strand is complementary to the nucleotide on the parent strand that is
across from it and a bond forms between them. Finally, we see how the 3' -OH group
displaces the two outermost phosphate groups of an incoming nucleotide in order to
add it to the growing chain.
The Driving Force of the Addition Reaction
Each incoming nucleotide supplies the energy for its addition in the high-energy
bond between the beta and gamma phosphates that are ejected upon addition. It is
not the release of the pyrophosphate that drives the reaction, but rather the subsequent
hydrolysis that takes place. A much larger amount of energy is released when the two
phosphates are separated into individual phosphates through the hydrolysis reaction.