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Chapter 9: Evolution at Multiple Loci
MULTIPLE CHOICE
1. What two major concepts in biology were unified in the middle of the twentieth century?
a. Natural selection and drift
b. Discontinuous and continuous variation
c. Mutation and migration
d. Natural selection and Mendelian genetics
e. Population genetics and quantitative genetics
ANS: D DIF: Easy REF: 9.1 TOP: I
MSC: Factual
2. The rediscovery of Mendelian inheritance caused increased scrutiny of Darwin’s theory of natural
selection because of an apparent conflict in the two concepts. This conflict was based on the
observation that Darwin focused on continuous traits, whereas Mendelian genetics deals with
a. discrete characters.
b. alleles.
c. flowers.
d. genotypes.
e. All of the above
ANS: A DIF: Easy REF: 9.1 TOP: I.A
MSC: Applied
3. The observation of an offspring phenotype outside of the range of parental phenotypes is most likely
due to
a. additive genetic effects.
b. mutation.
c. selection.
d. latent variation.
e. C and D
ANS: E DIF: Easy REF: 9.1 TOP: I.C
MSC: Factual
5. A haplotype is
a. all of the alleles at a single locus.
b. a product of epistasis.
c. a set of alleles at different loci along a chromosome.
d. all of the alleles in a population.
e. None of the above
ANS: C DIF: Easy REF: 9.2 TOP: II.A
MSC: Factual
6. Consider a population with two loci (A, B), each with two alleles (A, a; B, b). What are the haplotype
frequencies in this population if the allele frequencies are A = 0.5, a = 0.5, B = 0.7, b = 0.3?
a. AB = 0.35, Ab = 0.15, aB = 0.35, ab = 0.15
b. AB = 0.45, Ab = 0.05, aB = 0.25, ab = 0.25
c. AB = 0.4, Ab = 0.1, aB = 0.3, ab = 0.2
d. AB = 0.125, Ab = 0.25, aB = 0.25, ab = 0.125
e. Cannot be determined based on the information provided
ANS: E DIF: Medium REF: 9.2 TOP: II.A
MSC: Applied
8.
10. What is the largest possible value that linkage disequilibrium can reach if all allele frequencies are 0.5?
a. 0.25
b. 0.5
c. 0.75
d. 1
e. None of the above
ANS: A DIF: Easy REF: 9.2 TOP: II.D
MSC: Applied
11. If the value of linkage disequilibrium (D) is equal to –0.25, one can infer that the alleles are
a. in repulsion.
b. in coupling.
c. in linkage disequilibrium.
d. A and C
e. None of the above
ANS: D DIF: Medium REF: 9.2 TOP: II.D
MSC: Applied
13. Which of the following processes that can create linkage disequilibrium is illustrated in the figure
below?
a. Drift
b. Selection
c. Mutation
d. Migration
e. All of the above
ANS: D DIF: Easy REF: 9.2 TOP: II.E
MSC: Factual
14. In the figure below, the grey boxes represent the offspring of parents that are heterozygous at locus 1
(Aa) and locus 2 (Bb). Which of the grey boxes represent repulsion double heterozygote offspring?
a. Light grey
b. Medium grey
c. Dark grey
d. B and C
e. All of the above
ANS: C DIF: Easy REF: 9.2 TOP: II.F
MSC: Factual
15. Assuming the value of linkage disequilibrium (D) is positive, recombination will generate
a. more repulsion pairs.
b. more coupling pairs.
c. no change, since recombination does not change expected haplotype frequencies.
d. an increase in D.
e. None of the above
ANS: A DIF: Medium REF: 9.2 TOP: II.F
MSC: Applied
16.
In the figure above, a new beneficial allele, B, arises on a chromosome containing the ordered loci A,
B, C, D, and E, depicted in the top panel. After reproduction, those chromosomes possessing the B
allele increase in frequency. Over time, recombination breaks up the association of the B allele with
the A, C, D, and E alleles. Assuming that the five loci are spaced evenly along the chromosome, the
first pair of allele combinations to be broken up will be
a. B and A.
b. B and C.
c. B and D.
d. B and E.
e. All alleles will break up at an equal rate
ANS: D DIF: Easy REF: 9.2 TOP: II.G
MSC: Applied
17.
In the figure above, each new line represents the frequency of a new beneficial mutation. The figure
describes the process of
a. selective sweeps.
b. periodic selection.
c. genetic hitchhiking.
d. background selection.
e. A and B
ANS: E DIF: Medium REF: 9.2 TOP: II.G
MSC: Factual
19.
20. The following figure demonstrates the genotypic adaptive landscape for bacteria growing in media
without antibiotics. The height of the bars associated with a genotype indicates the fitness of that
genotype. The R allele confers resistance to antibiotics, and the C allele compensates for the loss of
fitness that the R allele confers in the absence of antibiotics. The rc haplotype lacks both compensation
and resistance. Based on this landscape, why are resistance and compensation unlikely to be lost even
in the absence of antibiotics?
21. Genotypic adaptive landscapes may better represent true adaptive landscapes than phenotypic adaptive
landscapes when
a. single mutations can have a large effect on the phenotype.
b. phenotypic landscapes are hard to mathematically model.
c. phenotypic landscapes do not account for drift.
d. genotypic landscapes are easier to visualize.
e. genotypic landscapes are smooth.
ANS: A DIF: Easy REF: 9.3 TOP: III.B
MSC: Factual
24. In the figure below, what does the straight line indicate?
26. The figure below describes the percent oil content in corn from different generations. What is the
expected slope of a parent–offspring regression for corn oil content?
a. 0.69
b. 1.6
c. 2.1
d. 2.5
e. Cannot be determined based on the information provided
ANS: A DIF: Difficult REF: 9.4 TOP: IV.C
MSC: Applied
27. Texas longhorn cattle have been bred to increase the tip to tip width of their horns, and the heritability
of this trait has been well studied by Professor David Hillis at the University of Texas, Austin. In the
current generation, the mean horn length is 52 inches. If only a subset of this population, with a mean
horn length of 61 inches, are allowed to reproduce, and the narrow-sense heritability of this trait = 0.2,
what is the expected horn length in the offspring generation?
a. ~1.8 inches
b. ~9 inches
c. ~54 inches
d. ~61 inches
e. Cannot be determined based on the information provided
ANS: C DIF: Difficult REF: 9.4 TOP: IV.D
MSC: Applied
SHORT ANSWER
1. A species of lotus produces a nearly continuous spectrum of flower colors, ranging from white to deep
red. The color of the flower is determined by 10 loci, each with two codominant alleles. What concept
does this example illustrate?
ANS:
Polygenic phenotypic variation
2. In a population of dragonflies, wing structure is determined by two loci (A, B), each with two alleles
(Aa, Bb). Abnormal wings are produced in aa individuals unless they possess at least one B allele. This
is an example of
ANS:
epistasis.
3. Consider two microsatellite loci (A, B) each with two alleles determined by the number of repeats of
the microsatellite motif (A = 15 or 17; B = 27 or 31). You genotype an individual in this population
and determine that its haplotype is 15/27 and 15/27. Are these alleles in coupling and/or repulsion?
Explain your answer.
ANS:
These alleles could be in either coupling or repulsion depending on how the researcher decides to refer
to the four alleles.
4. Consider two loci (A, B) each with two alleles (Aa, Bb). The frequency (f) of A is known to be 0.3, the
frequency (f) of b is known to be 0.25, and the coefficient of linkage disequilibrium (D) between loci A
and B is known to be 0.15. What is the expected haplotype frequency of haB? Please show your work.
ANS:
Equation: D = fafB – haB
Rearrange to: haB = fafB – D
fa = 1 – 0.3 = 0.7
fB = 1 – 0.25 = 0.75
D = 0.15
Then substitute and solve:
haB = (0.7)(0.75) – 0.15
haB = 0.525 – 0.15
haB = 0.375
5. In Lake Malawi cichlid fish, one can observe two color variants, Black Barred (BB) and Orange Blotch
(OB). Sex in these fish is not determined by a sex chromosome but by a gene on an autosome, which
produces either males or females. Both of these phenotypes are controlled by one biallelic locus each.
OB females have a selective advantage in terms of their survivability: The OB phenotype allows
females to blend into their background. OB males have a selective disadvantage because their
reproductive coloration is disrupted and they have little mating success. How would this process
influence the association of alleles at the color locus and sex determining locus?
ANS:
Natural selection would generate linkage disequilibrium between the two loci such that the female
allele would become associated with the OB allele and the male allele would become associated with
the BB allele.
6. If recombination between locus A and locus B is 0.15, and the initial value of linkage disequilibrium is
0.20, what will the value of linkage disequilibrium be after one generation of recombination? Show
your work.
ANS:
D = –rD
D = –0.15 0.20
D = –0.03
D(t + 1) = 0.20 – 0.03 = 0.17
7. Multiple drug-resistant Staphylococcus aureus (MRSA) is a bacterium that has developed resistance to
a wide range of antibiotics. Its resistance to penicillin, methicillin, and oxacillin is a result of the serial
use of these antibiotics, which created a selective environment that favored resistance to the drug being
used at the time. This resistance provides the bacteria with the ability to thrive in an environment with
standard antibiotics, thereby making it a dangerous human health hazard. Why would one expect
MRSA to retain penicillin resistance even if this antibiotic is rarely used?
ANS:
The MRSA strain has experienced several rounds of periodic selection, which has generated a single
nonrecombining chromosome that conveys resistance to multiple drugs. In the absence of sexual
reproduction and recombination, these alleles are unlikely to be disrupted.
8. Explain why a population may never reach its most fit phenotype.
ANS:
If the adaptive landscapes have multiple peaks, the population will tend to climb the nearest peak, even
if it is not the highest peak. In the absence of drift, natural selection will keep the population on this
local maximum, thereby preventing it from reaching the global maximum.
ANS:
Single mutations can have large phenotypic effects.
10. A study of susceptibility to major depressive disorder in monozygotic twins separated at birth revealed
that the broad-sense heritability of this trait is 0.10, and its total phenotypic variance is 0.25. What is
the variance of the genotypic value of this disorder? Show your work.
ANS:
H2 = VG/(VG+VE) and VP = VG+VE
Therefore:
H2 = VG/VP
VP H2 = VG
0.25 0.1 = 0.025
VG = 0.025
11. Without using mathematical symbols, explain the difference between broad- and narrow-sense
heritability, including why one is more relevant to evolutionary studies.
ANS:
Broad-sense heritability describes the proportion of the total phenotypic variance described by the
genetic variance. Narrow-sense heritability describes the proportion of the total phenotypic variance
described by the additive genetic variance. The narrow-sense heritability is more valuable to
evolutionary studies as it focuses strictly on the additive genetic variation, and it is the additive genetic
variance on which natural selection operates. In contrast, the broad-sense heritability includes many
variance components whose impact on the phenotype are context dependent. Therefore, these variance
components may not be exposed to the action of natural selection.
12. Explain why one cannot compare the estimated heritability of intelligence tests from different
populations from the United States.
ANS:
All heritability estimates are within-population measures not between-population measures. This is
due partly to the different environmental variances found in different populations.
Now if you will look at a map of Alaska you will see that the
Porcupine River is like the letter U laid over on its side; that is to say,
its head waters are in Alaska and the stream then flows east over
the International boundary into the Yukon Territory, thence north by
northeast across the Arctic Circle and when it reaches latitude 137
degrees and longitude about 67-1/2 degrees, it makes a sharp bend
and flows back west by southwest for a couple of hundred miles,
when it empties into the Yukon River, between the towns of Beaver
and Fort Yukon.