Biol. 2320 - Fall 2000 - In-Class Exercise (due today)
and Thought Question #3 Due 10/2/00

In-Class Exercise or "If you find a replication fork in the road, leave it where you found it"

The problem (solution due at the end of class today): The replication fork of an actively replicating DNA duplex moves in a single direction, with daughter strands synthesized on the separated parental strands. Both daughter strands, like all DNA, are synthesized in a 5'®3' orientation. One of the two daughter strands is polymerized into the replication fork (the leading or continuous strand) while the other is read out of the replication fork (the lagging or discontinuous strand). The figure from your text shown on the following page (fig 13.8c) is a misrepresentation. It shows one DNA polymerase enzyme directing continuous strand synthesis and another separate DNA polymerase directing discontinuous strand synthesis. In fact, DNA polymerase is a dimer (i.e. two enzymes hooked together) made up of one monomer that directs continuous strand synthesis and a slightly different monomer that directs discontinuous strand synthesis. The dimer can only move in the direction of the replication fork. How can one enzyme dimer, moving in the direction of the replication fork, synthesize both the continuous and discontinuous strands? The enzyme cannot simply stretch. Use the model replication kit provided to come up with a mechanism for how discontinuous strand synthesis might work (we know how continuous strand synthesis works).

Replication Kit:

Double sphere:                 DNA polymerase asymmetric dimer
Clipped Rubber Hoses:     Segment of double stranded DNA template (with 5' end marked by pipette tip)
Pipe Cleaners                   Okazaki fragments
Small clips                         Hydrogen bonds for Okazaki fragments

Rules:

Synthesize the new discontinuous strand from one end of the strand to the other using the Okazaki fragments (again, ignore leading strand synthesis for now).

You can only synthesize in a 5’®3' direction; therefore the parent strand must be read in a 3'®5' (plugs define 5' ends of parental strands).

Position the enzyme dimer right at the replication fork (unclip the strands as you go).

You can only add an Okazaki fragment when the appropriate portion of the strand is in contact with the enzyme dimer at the replication fork.

Hints:

DNA is not a rigid molecule, particularly when in single stranded form. It is much more flexible, like the model tubing we have used. Do not let your preconceived notions of DNA structure dictate what you consider to be possible.

[5 pts] Turn in one copy of this today:

Group members:

____________________________

____________________________

 

Describe the solution you came up with in no more than two sentences - you can also use this space to draw what you did:

Thought question:
Each individual group member is to write a separate on a separate sheet
 and hand in on 10/2/00
Typed if at all possible

1.      [2 points] We just had to go through contortions to generate a workable model that describes the three dimensional structure of the replication fork. This is all because the DNA polymerase is a dimer and not two types of free monomer. Provide a biologically relevant reason why cells might need to keep the two sites of polymerization (the enzymes) together. Explain in no more than 2 sentences.

2.      [3 points] You just clipped together a whole series of Okazaki fragments, discontinuously synthesized from the lagging strand. When Meselson and Stahl performed their experiment that confirned that DNA was replicated semi-conservatively, they used density gradient centrifugation to separate different types of DNA. Why didn't they observe Okazaki fragments?