11/4/23, 11:45 PM Course: AGRI 21(2022)-A (LEC***) PRINCIPLES OF GENETICS

Lecture Topic 12: DNA Recombination

Welcome to Lecture Topic 12!

Please read and study the discussion below taken from Chapter 11 of our reference book. Watch the two videos to
supplement your understanding of the discussion.

Don't forget to take the Quiz on Nov 2, 2023. Thank you.

Recombination is Essential for Genetic Exchange and DNA Repair

In a previous topic, we were introduced to genetic crossing over, a phenomenon that occurs in eukaryotes during
prophase I of meiosis. During crossing over, members of homologous chromosomes pair with each other and
exchange chromosome segments, resulting in chromosomes that are combinations of genetic information from
paternal and maternal sources. This genetic exchange is a major source of genetic diversity through sexual
reproduction. We have also discussed a previous topic the various ways in which genetic information can be
exchanged between DNA molecules from bacteria or bacteriophages. In these examples of recombination, the
process of exchange begins with either single-stranded or double-stranded DNA breaks.

Not only is recombination essential for the exchange of genetic information during meiosis, but it is also
important for the repair of damaged DNA. Various types of agents can create double-stranded breaks in DNA
molecules. These agents include ionizing radiation and free radicals. DNA recombination is also part of the ALT
mechanism used to maintain eukaryotic telomeres. When cells are defective in recombination mechanisms, they are
prone to accumulating mutations in genes and chromosomes, which in turn may lead to the development of tumors,
diseases, or even cell death.

Now that we have discussed the chemistry and replication of DNA, we can consider how recombination occurs at the
molecular level. In general, our discussion pertains to genetic exchange between any two homologous double-
stranded DNA molecules, whether they are viral or bacterial chromosomes or eukaryotic homologs during meiosis.

Models of Homologous Recombination

Genetic exchange at equivalent positions along two chromosomes with substantial DNA sequence homology
is referred to as general, or homologous recombination.

Several models attempt to explain homologous recombination, but they all have certain features in common. First, all
are based on proposals first put forth independently by Robin Holliday and Harold L. K. Whitehouse in 1964. Second,
they all depend on the complementarity between DNA strands to explain the precision of the exchange. Finally, each
model relies on a series of enzymatic processes in order to accomplish genetic recombination.

One such model, beginning from a single-stranded DNA break, is shown in Figure 10.1. It begins with two paired DNA
duplexes, or homologs [Step (a)], in each of which an endonuclease introduces a single-stranded nick at an
identical position [Step (b)]. The internal strand endings produced by these cuts are then displaced and
subsequently pair with their complements on the opposite duplex [Step (c)]. Next, a ligase seals the loose ends [Step
(d)], creating hybrid duplexes called heteroduplex DNA molecules, held together by a cross-bridge structure. The
position of this cross bridge can then move down the chromosome by a process referred to as branch migration
[Step (e)], which occurs as a result of a zipper-like action as hydrogen bonds are broken and then reformed between
complementary bases of the displaced strands of each duplex. This migration yields an increased length of
heteroduplex DNA on both homologs.

https://soul.su.edu.ph/course/view.php?id=30949#section-15 1/4
11/4/23, 11:45 PM Course: AGRI 21(2022)-A (LEC***) PRINCIPLES OF GENETICS

Figure 10.1 Model depicting how genetic recombination can occur as a result of the breakage and rejoining of
heterologous DNA strands. Each stage is described in the text. The electron micrograph shows DNA in a χ-form
structure similar to the diagram in (g); the DNA is an extended Holliday structure, derived from the ColE1 plasmid of
E. coli.

If the duplexes bend [Step {f}] and the bottom portion shown in the figure rotates 180° [Step (g)], an
intermediate planar structure called a χ (chi) form—or Holliday structure—is created. If the two strands on opposite
homologs previously uninvolved in the exchange are now nicked by an endonuclease [Step (h)] and ligation occurs as

https://soul.su.edu.ph/course/view.php?id=30949#section-15 2/4
11/4/23, 11:45 PM Course: AGRI 21(2022)-A (LEC***) PRINCIPLES OF GENETICS

in Step (i), two recombinant duplexes are created. Note that the arrangement of alleles is altered as a result of
this recombination.

Whereas the preceding model involves single-stranded breaks, similar recombination models have been
proposed that involve double-stranded breaks in one of the DNA double helices. In these models, endonucleases
remove nucleotides at the breakpoint, creating 3' overhangs on each strand. One of the broken strands invades the
intact double helix of the other homolog, and both strands line up with the intact homolog. DNA repair synthesis then
fills all gaps, and two Holliday junctions are formed. Endonuclease cleavages and ligations finalize the exchange. The
end result is the same as our original model: genetic exchange occurs between homologous DNA molecules.

QUIZ

Quiz for Lecture Topic 12

This is a short quiz to test if you have read and studied the discussion for this topic. You have 2
minutes to submit your answers, otherwise they will not be recorded. Good luck!
Not attempted Due 2 November 2023

Holliday Model of Homologous Recombination

DNA Recombination Model with Double-stranded Breaks

https://soul.su.edu.ph/course/view.php?id=30949#section-15 3/4
11/4/23, 11:45 PM Course: AGRI 21(2022)-A (LEC***) PRINCIPLES OF GENETICS

https://soul.su.edu.ph/course/view.php?id=30949#section-15 4/4