Genetics From Genes To Genomes 5th Edition Solutions Manual 1
Genetics From Genes To Genomes 5th Edition Solutions Manual 1
Genetics From Genes To Genomes 5th Edition Solutions Manual 1
chapter
5
Linkage, Recombination, and the
Mapping of Genes on Chromosomes
Synopsis
In Chapter 5, we learn the consequences of the fact that hundreds or even thousands of genes can
be found on one chromosome. If two genes lie close together on a chromosome, their alleles
do not assort independently; instead, the genes are linked and parental classes of gametes will be
formed more frequently than recombinant classes. The Recombination Frequency (RF) increases
to a maximum of 50% as the distance between the two genes increases. The formation of
recombinant classes depends on the occurrence of physical crossovers between nonsister
chromatids that take place during prophase of the first meiotic division, after the chromosomes
have replicated and homologous chromosomes have paired with each other. The greater the
distance between the genes, the more likely the chance of a crossover and the greater the RF; the
50% limit to RF occurs when genes are sufficiently far apart that at least one crossover occurs
between them in every meiosis. By measuring RF, geneticists can map the location of genes
along the chromosomes.
In most organisms, RF can be tracked only by looking at individual progeny of a cross
involving at least one parent who is a heterozygote for the two genes in question. But in the
fungi S. cerevisiae and N. crassa, all the products of a meiosis remain together in a sac called an
ascus. As a result, researchers can infer more details about the locations and kinds of crossovers
that took place during the meiosis that produced that ascus.
Finally, Chapter 5 discusses what happens when recombination takes place during mitosis -
rather than meiosis. In contrast with meiotic recombination, which is a programmed event that
must occur at least once on every chromosome during every meiosis, mitotic recombination
reflects rare mistakes that occur (usually in response to DNA damage). Mitotic recombination
provides a valuable tool to researchers because it allows them to create mosaic organisms in
which different cells have different genotypes.
Key terms
syntenic genes – genes located on the same chromosome
parental types/classes – gametes whose 5-1alleles are in the same combinations as in the
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gametes that gave rise to previous generation; or progeny generated by parental
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Education.
gametes
recombinant types/classes – gametes whose alleles are in different combinations
chapter 5
resulting from a meiosis in which a crossover took place between the two genes in
question.
linkage – when two genes are located close enough to each other on the same
chromosome so that a doubly heterozygous parent makes more parental type than
recombinant type gametes
recombination frequency (RF) – the proportion of the total number of gametes or
progeny that are recombinant types (recombinants/total)
map unit (mu)/centimorgan (cM) – a measure of genetic distance between linked
genes; 1 mu = 1 cM = 1% RF.
locus (singular)/loci (plural) – a specific location on a particular chromosome
interference – a phenomenon in which a crossover event on a chromosome prevents
or limits the occurrence of a second crossover event nearby on the same chromosome.
Interference is thought to take place as a way to help ensure that at least one crossover
occurs on each bivalent, so that homologous chromosomes segregate accurately
during the first meiotic division.
coefficient of coincidence (c.o.c.) – the ratio of the number of double crossovers
observed in a cross to the number of double crossovers expected if the two crossovers
were independent of each other. The interference = 1 – c.o.c. This makes sense because
the amount of interference reflects a decrease in the frequencies of double crossovers
(DCOs).
chi-square test – a statistical device used to measure the likelihood that a particular set
of observed experimental results could have been obtained as chance deviations
from the expectations from a particular hypothesis to be tested (the null
hypothesis)
significant result – experimental results that are highly unlikely to be explained by the
null hypothesis. A significant result allows you to reject the null hypothesis. An
insignificant result does not allow you to reject the null hypothesis. Note that
these definitions mean that the chi-square test can never prove a null hypothesis,
but can only reject the null hypothesis.
degrees of freedom (d.o.f) – In the chi-square test for the goodness of fit, the d.o.f. =
number of classes – 1. The d.o.f. is required for interpreting the chi-square test.
p value – the probability that the observed results could be obtained by chance deviations
from the expectations of the null hypothesis. If the p value is low (usually below 0.05
for genetic analysis), you can reject the null hypothesis.
ascus – in fungi, a sac that contains all four products of a single meiosis. These haploid
products are the ascospores (or simply spores). A tetrad is the collection of the 4
spores in one ascus. (In Neurospora, the ascus is an octad containing 8 spores due to
the occurrence of one round of mitosis after meiosis is complete.)
ordered versus unordered tetrad – In ordered tetrads (N. crassa), the geometry of the
spores in the ascus reflects the movement of chromatids during the two meiotic
divisions. In unordered tetrads (S. cerevisiae), the four ascospores are arranged at
random within the ascus.
5-2
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chapter 5
parental ditype (PD) – a tetrad in which all four spores are parental types (two spores
of each of the two parental types possible)
nonparental ditype (NPD) – a tetrad in which all four spores are recombinant types
(two spores of each of the reciprocal recombinant types)
tetratype (T) – a tetrad with four spores all of different genotypes (the two reciprocal
parental types and the two reciprocal recombinant types)
first-division segregation pattern (MI) and second-division segregation pattern
(MII) – arrangements of spores in ordered tetrads (octads in Neurospora). In an MI
pattern, if an imaginary line is drawn through the center of the ascus, all the spores
with one allele are on one side of the line and all those with the other allele are on
the other side of the line (4:4). In an MII pattern, pairs of spores with both alleles
are found on either side of the imaginary line at the ascus middle (2:2:2:2).
twin spots – adjacent patches of tissues that are phenotypically and genotypically
different from each other, and phenotypically and genotypically different from the
surrounding cells. Twin spots can result from mitotic crossing-over depending on
the arrangement of alleles and the sites of the mitotic crossovers.
mosaic – an organism harboring cells of different genotypes. For example, a fruit fly with
twin spots is a mosaic.
sector – in yeast, a region of a colony of cells in which the cells have a different phenotype
and genotype from the other cells in the same colony
Key Equations
• Recombination frequency (RF) = # recombinant progeny / total # progeny.
(Multiply × 100 to express the RF as %.)
• 1% RF = 1 mu or 1 cM.
• Coefficient of coincidence (c.o.c) = # DCOs observed / # DCOs expected. To
calculate the # DCOs expected, multiply the crossover frequency in interval 1 ×
crossover frequency in interval 2, and then multiply this value by the total number
of progeny.
• Interference = 1 – c.o.c.
• Χ2 = Σ [(# observed - # expected)2 / # expected].
(In other words, Χ2 is the sum over all classes of [(O-E)2 / E] calculated for each
class.)
• Degrees of freedom (d.o.f.) = # classes -1.
• In Χ2 analysis, the ultimate goal is to determine the probability p that the observed
results could have been obtained by chance if the null hypothesis is correct. To
determine p, you need to calculate Χ2 and d.o.f., and then you must consult a table
of critical values such as Table 5.2 on p. 147.
• See the equations in the box Three Easy Rules for Tetrad Analysis that follows.
5-3
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chapter 5
Problem
Solving
For many of the problems in this chapter, you will still need to start by answering the same
questions you dealt with in the first four chapters:
How many genes are involved in the cross?
For each of these genes, what is the dominance relationship between the alleles?
For each of these genes, is it X-linked or autosomal?
In the problems in this chapter, you now have to consider the additional possibility that genes may
be syntenic (on the same chromosome) and that they are genetically linked.
When thinking about genes that are linked, you must write the genotypes in a way that
represents the linkage. Remember that there is one allele per homolog, so aa becomes a / a. It is
usually advantageous to indicate the homologous chromosomes with horizontal, parallel lines so
that you can track which alleles of which genes were present on which chromosomes. Thus, a+
b+ c / a b c+ would be diagrammed as:
Although the two lines in the diagram above represent the two homologous chromosomes, you
should always keep in mind that recombination actually occurs at the four-strand stage, after
each chromosome has replicated into two sister chromatids. This fact plays an important role in
understanding why there is a 50% limit of the RF measured between two genes, and it also helps
you visualize what is occurring when you are analyzing tetrads in fungi.
5-4
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Education.
chapter 5
IV.
V.