Crossing Over, Recombination, and Linkage

The fi st step in gene mapping is to establish linkage with a known polymorphic

marker (one with at least two alleles in the population). Th s can be done by
recombination
mapping to determine whether the gene is near a particular marker.
Multiple markers on diferent chromosomes are used to establish linkage (or the
lack of it).
Recombination mapping is based on crossing over during meiosis, the type of cell
division that produces haploid ova and sperm. During prophase I of meiosis,
homologous
chromosomes line up and occasionally exchange portions of their DNA.
Th s process (shown in Figure II-5-2) is termed crossover.
Figure II-5-2. The Process of Crossing Over Between
Homologous Chromosomes
When a crossover event occurs between two loci, G and M, the resulting
chromosomes
may contain a new combination of alleles at loci G and M. When a new
combination occurs, the crossover has produced a recombination. Because
crossover
events occur more or less randomly across chromosomes, loci that are located
farther apart are more likely to experience an intervening crossover and thus a
recombination
of alleles. Recombination frequency provides a means of assessing the
distance between loci on chromosomes, a key goal of gene mapping.
This process is illustrated in Figure II-5-3.
If the gene of interest (with alleles G1 and G2) and the marker (with
alleles M1 and M2) are on diferent chromosomes, the alleles will remain
together in an egg or a sperm only about 50% of the time.
They are unlinked.
If the gene and the marker are on the same chromosome but are far
apart, the alleles will remain together about 50% of the time. The larger
distance between the gene and the marker allows multiple crossovers to
occur between the alleles during prophase I of meiosis. An odd number
of crossovers separates G1 from M1 (recombination), whereas an even
number of crossovers places the alleles together on the same chromosome
(no recombination). The gene and marker are again defined as
unlinked.
If the gene and the marker are close together on the same chromosome,
a crossover between the two alleles is much less likely to occur. Therefore,
G1 and M1 are likely to remain on the same chromosome more
than 50% of the time. In other words, they show less than 50%
recombination.
The gene and the marker are now defined as linked.