80

Chapter 19

Chapter 19

Variation and Selection in Populations

Synopsis:
This chapter involves the study of how genetic laws impact the genetic makeup of a population.
Mendelian principles are the basis for the Hardy-Weinberg law which allows one to calculate allele and
genotype frequencies from one generation to the next. The Hardy-Weinberg law can be used only if
other forces are not acting on the allele frequency. Those forces include selection, migration, mutation,
and population size.
Population geneticists try to determine the extent to which a trait is determined by genetic factors
and how much is determined by environmental factors. Knowing if a trait is largely determined by
genetic factors introduces the possibility for animal and plant breeders to select and maintain
populations with desired traits.

Significant Elements:
After reading the chapter and thinking about the concepts, you should be able to:

Determine allele frequencies in a population given the frequencies of genotypes.

Determine genotype frequencies in a population given the frequencies of alleles.

Determine genotype and/or allele frequencies in the next generation given the genotype or allele
frequencies in the present generation.

Determine if a population is in equilibrium.

Determine allele and genotype frequencies after migration has occurred.

Describe how heritability of a trait can be determined.

Problem solving tips:

p and q represent allele frequencies for a gene with two alleles; the sum of p
+ q must equal 100% of the alleles (or gametes) in the population. In other
words, p + q = 1.

p2, 2pq, q2 represent genotype frequencies; p2 corresponds to one

homozygous genotype, 2pq represents the heterozygous genotype and q 2 is
the other homozygous genotype.

Determining allele and genotype frequencies can be done two slightly

different ways. One method involves converting the initial numbers of each

81

Chapter 19

genotype to frequencies and then doing all calculations as frequencies. In this

case the frequency of the p allele = the frequency of the p/p homozygotes +
1/2 the frequency of the heterozygotes. The frequency of the q allele = the
frequency of the q/q homozygotes + 1/2 the frequency of the heterozygotes.
If the problem asks for numbers of individuals then the genotype frequencies
will be multiplied by the number of individuals in the population, as in
problem 19-2 solution #1. The problems in this chapter will be solved using
this method. Alternately the problems can be solved by calculating the
numbers of each allele present in the population. When doing this the total
must be divided by the total number of alleles, as in problem 19-2 solution
#2.

When any values of p and q that sum to 1 are binomially expanded (p 2 + 2pq
+ q2) the resulting genotype frequencies are those of a population in HardyWeinberg equilibrium.

The binomial expansion is the equivalent of a Punnett square. If p (or the A

allele) = 0.7 and q (or the a allele) = 0.3 then the binomial expansion gives
the genotype frequencies in the next generation as 0.49 AA + 0.42 Aa + 0.09
aa. You can use a Punnett square to generate the same genotype frequencies
in the F1 generation:
0.7 A
0.3 a

0.7 A
0.49 AA
0.21 Aa

0.3 a
0.21 Aa
0.09 aa

If the allele frequencies are calculated correctly then p + q MUST sum to 1. If

the genotype frequencies are calculated correctly than p 2 + 2pq + q2 MUST
sum to 1.

Once a population is at equilibrium allele and genotype frequencies do not

change.

If a genotype is selected against or if populations are combined (by

migration) or if there is significant mutation (usually together with selection)
allele frequencies will change.

Selection for one genotype and selection against another genotype are
balanced at a particular allele frequency (equilibrium frequency).

Chapter 19

Genetic drift is most often seen in small populations.

Polygenic traits are controlled solely by alleles of two or more gene;

82

multifactorial traits include polygenic traits and traits that are influenced by
both genes and environment.

Genetic and environmental contributions to a phenotype are sorted out by

setting up conditions in which the genetic background is consistent (to
analyze environmental contributions) or conditions in which the environment
is constant (to analyze genetic contributions).

83

Chapter 19

Solutions to Problems:
Vocabulary
19-1. a. 3; b. 5; c. 8; d. 7; e. 6; f. 1; g. 9; h. 2; i. 4.
Section 19.1 the Hardy-Weinberg Law
19-2. Solution #1:
a. Calculate genotype frequencies:
GGGG = 120 / 200 = 0.6; GGGB = 60 / 200 = 0.3; GBGB = 20 / 120 = 0.1.
b. Frequency of GG = 0.6 + 1/2 (0.3) = 0.75; frequency of GB = 0.1 + 1/2 (0.3) = 0.25.
c. Binomially expand p and q: (0.75 GG)2 + 2 (0.75 GG) (0.25 GB) + (0.25 GB)2 = 0.563 GGGG +
0.375 GGGB + 0.063 GBGB = 1.
Solution #2:
a. The genotype frequencies are calculated as in Solution #1 part a above.
b. The allele frequencies are determined by totaling all alleles within each genotype.
GGGG = 120 individuals with 2 GG alleles = 240 GG alleles
GGGB = 60 individuals with one GG allele = 60 GG alleles
GGGB = 60 individuals with one GB allele = 60 GB alleles
GBGB = 20 individuals with two GB alleles = 40 GB alleles
There are 300 GG alleles/400 total alleles so the frequency of GG (p) = 0.75. There are 100 GB
alleles/400 total alleles so the frequency of GB (q) = 0.25.
c. The expected frequencies can be calculated using the allele frequency and the terms of the HardyWeinberg law, p2 + 2pq + p2 = 1; 0.5625 GGGG + 0.375 GGGB + 0.0625 GBGB = 1.
19-3. For each population determine the allele frequency Use the genotype frequencies to calculate the
allele frequency. Binomially expand the allele frequencies to generate the genotype frequencies of the
next generation (the Hardy-Weinberg genotype frequencies).
a. The frequency of A = 0.25 + 1/2 (0.5) = 0.5; the frequency of a = 0.25 + 1.2 (0.5) = 0.25.
Binomially expanding A (p) and a (q) gives genotype frequencies of 0.25 AA + 0.5 Aa + 0.25 aa.
Because these genotype frequencies (the Hardy-Weinberg equilibrium) are the same as those of the
original population the original population is in equilibrium.

Chapter 19

84

b. The frequency of A = 0.1 + 1/2 (0.74) = 0.47; the frequency of a = 0.16 + 1/2 (0.74) = 0.53.
Binomially expanding A and a gives genotype frequencies of 0.221 AA + 0.498 Aa + 0.281 aa.
These are the Hardy-Weinberg genotype frequencies for p = 0.47 and q = 0.53. They are NOT the
same as the original genotype frequencies, so population b is not in equilibrium
c. Population c is not in equilibrium - the Hardy-Weinberg genotype frequencies are 0.61 AA +
0.34 Aa + 0.05 aa.
d. Population d is not in equilibrium the Hardy-Weinberg genotype frequencies are 0.50 AA +
0.41 Aa + 0.08 aa.
e. Population e is in equilibrium the Hardy-Weinberg genotype frequencies are 0.81 AA + 0.18 Aa
+ 0.01 aa.
19-4.
a. There are 60 D+D+ flies with normal wings and 90 DD+ flies with Delta wings. The frequencies
of these two genotypes are 60/150 = 0.4 D+D+ and 90/150 = 0.6 DD+. The allele frequency for D
= 0 DD homozygotes + 1.2 (0.6) = 0.3; allele frequency for D+ = 0.4 + 1/2 (0.6) = 0.7.
b. The following frequencies of F1 zygotes will be produced: 0.49 D+D+ + 0.42 DD+ + 0.09 DD =
1. The homozygous DD zygotes do not live, so the viable progeny are 0.49 + 0.42 = 0.91 (91%) of
the original zygotes. If there are 160 viable adults in the F 1 generation, then there must have been
160 / 0.91 = 176 F1 zygotes.
c. As noted in part b above, the viable progeny are only 91% of the F1 zygotes (0.49 D+D+ + 0.42
DD+ = 0.91. This equation must be normalized before you can calculate expected numbers of
progeny. Therefore, 0.49 D+D+ / 0.91 = 0.54 D+D+ = proportion of the viable offspring with the
D+D+ genotype and 0.42 D+D / 0.91 = 0.46 D+D = proportion of the viable offspring with the
D+D genotype. Thus, the numbers of viable individuals with these genotypes are: 0.54 D+D+ x
160 = 86 D+D+ and 0.46 D+D x 160 = 74 D+D.
d. No. The lethality of the DD genotype means that there are genotype-dependent differences,
violating one of the basic tenets of the Hardy-Weinberg equilibrium. Therefore this population
will never achieve equilibrium.
19-5.
a. The frequency of M = 0.5 + 1/2 (0.2) = 0.6; the frequency of N = 0.3 + 1/2 (0.2) = 0.4. The
expected genotype frequencies in the next generation are calculated using these allele frequencies in

85

Chapter 19

the binomial expansion p2 + 2pq + p2 = 1: 0.36 MM + 0.48 MN + 0.16 NN = 1. These HardyWeinberg genotype frequencies are not the same as the genotype frequencies of the initial
population, therefore the initial population is not in equilibrium.
b. The genotype frequencies in the F1 will be: 0.36 MM + 0.48 MN + 0.16 NN = 1. From this the
allele frequencies in the F1 generation can be calculated: M = 0.36 + 1/2 (0.48) = 0.6 and N =
0.16 + 1/2 (0.48) = 0.4.
c. The same as in part b.
19-6.
a. Convert the genotypes into frequencies by dividing each by the total number (= 1480).
QFQFRCRC

0.137

QFQGRCRC

0.068

QGQGRCRC

0.068

QFQFRCRD

0.251

QFQGRCRD

0.126

QGQGRCRD

0.126

QFQFRDRD

0.112

QFQGRDRD

0.056

QGQGRDRD

0.056

Calculate the allele frequencies for Q and R genes separately. In this population the frequencies of
the Q gene are 0.5 QFQF + 0.25 QFQG + 0.25 QGQG = 1. The allele frequencies are QF = 0.625
and QG = 0.375. The expected genotype frequencies in the next generation (Hardy-Weinberg
equilibrium) is therefore 0.39 QFQF + 0.47 QFQG + 0.14 QGQG = 1. The population is NOT in
equilibrium for the Q gene. The genotype frequencies for the R gene in this population are 0.27
RCRC + 0.5 RCRD + 0.22 RDRD = 1. The RC allele frequency = 0.52 and the RD allele frequency
= 0.48. The expected genotype frequencies in the next generation (Hardy-Weinberg equilibrium) is
0.27 RCRC + 0.5 RCRD + 0.23 RDRD =1. In this case the observed genotype frequencies are very
close to the Hardy-Weinberg genotype frequencies, so the population IS in equilibrium for the R
gene.

Chapter 19

86

b. the fraction that will be QFQF in the next generation is the expected genotype frequency in part a:
0.39 for the QFQF genotype.
c. The fraction that will be RCRC in the next generation will again be the expected frequency
calculated based on the allele frequency: 0.27.
d. This probability is not influenced by allele frequencies calculated in part a above. Instead this is a
standard probability question starting with parents of specific genotypes as discussed in Chapter 2
(see problem 2-4). This cross is QFQG RCRD and QFQF RCRD. The probability is the product of
the individual probabilities for each of the genes. There is a 1/2 chance that the female will
contribute the QG allele and a 1/1 chance that the male will contribute the QF allele = 1/2 chance
of a QFQG child. Both parents are heterozygous for the R gene, so there is a 1/4 chance the child
will be homozygous RDRD. There is a 1/2 chance the child will be male. The overall probability is
(1/2)(1/4)(1/2) = 1/16 that the child will be a QFQGRDRD male.
19-7. If a population is in Hardy-Weinberg equilibrium then the allele frequency p is squared to give
the genotype frequency p2. Thus, each different allele frequency of p has a different p2 value and a
different set of genotype frequencies at equilibrium.
19-8. The 3:1 ratio is seen when two heterozygous individuals are crossed. This ratio is not relevant for
a population where the crosses are of many different sorts some will be homozygous dominant x
homozygous dominant, others homozygous dominant x heterozygous, some homozygous dominant x
homozygous recessive, some heterozygous x homozygous recessive and others homozygous recessive x
homozygous recessive. The ratio of wild type : mutant progeny in the population will depend on
specific allele frequencies in that population.
19-9.
a. The frequencies of the genotypes in the sailor population are: MM = 324/400 = 0.81, MN = 72/400
= 0.18 and NN = 4/400 = 0.01. The frequency of the N allele = 0.01 + 1/2 (0.18) = 0.1.
b. In order to calculate the allele and genotype frequencies in the children you must calculate the allele
frequencies of each original population and then combine them in the correct proportions. From
part a the frequency of N = 0.1 so the frequency of M = 0.9. In the Polynesian population the allele
frequencies are N = 0.94 and M = 0.06. When these two populations mix randomly then 40% of
the M and N alleles will come from the sailor population (400/1000) and the remaining 60% will be
provided by the Polynesians. Therefore the frequency of N in the mixed population = 0.4 (0.1) +

87

Chapter 19

0.6 (0.94) = 0.04 + 0.564 = 0.604. The frequency of M in the mixed population = 0.4 (0.9) + 0.6
(0.06) = 0.36 + 0.036 = 0.396. The genotype frequencies in the next generation will be: (0.604)2
NN + 2 (0.604) (0.396) MN + (0.396)2 MM = 0.365 NN + 0.478 MN + 0.157 MM = 1. If there
are 1,000 children then there will be 478 children with the MN genotype.
c. The observed genotype frequencies in the children are 100/1000 NN + 850/1000 MN + 50/1000
MM = 0.1 NN + 0.85 MN + 0.05 MM. The allele frequencies among the children therefore are N =
0.1 + 1/2 (0.85) = 0.525 and M = 0.05 + 1/2 (0.85) = 0.475.
19-10.
a. If a population is in Hardy-Weinberg equilibrium then q2 = q. If 1/250,000 people are affected by
the autosomal recessive disorder alkaptonuria then the genotype frequency of the recessive
homozygote (q2) = 4 x 10-6. Therefore q = 4 x 10-6 = 0.002.
b. Remember that the population is in Hardy-Weinberg equilibrium, that q = 0.002 and p (the normal
allele) = 0.998. Therefore the frequency of carriers (heterozygotes) = 2 (0.998) (0.002) = 0.004
= 4 x 10-3. The ratio of carriers / affected individuals = 4 x 10-3 / 4 x 10-6 = 1,000:1.
c. If the unaffected woman has an affected child then she must be a carrier, as was the father of the
child. She has remarried. If her new husband is homozygous normal she can never have an affected
child. However if the new husband is a carrier or is affected himself then she can have an affected
child. The total probability of an affected child is the sum of each of these individual probabilities.
The probability that she will have an affected child by this marriage = [(0.004 probability that
her new husband is a carrier) (0.5 probability that he passes on his mutant allele) (0.5 probability
that she passes on her mutant allele)] + [(4 x 10-6 probability that her new husband is affected) (1
probability that he passes on a mutant allele) (0.5 probability that she passes on her mutant allele)]
= [0.001] + [2 x 10-6] = 0.001002 = 0.001. As you can see, the probability that her second
husband is affected is insignificant. Therefore the probability that this woman will have an affected
child with her second husband is only dependent on the frequency of heterozygotes in the
population.
d. No - if one of the genotypes is selected against then the frequency of p and q will change each
generation. The population will never reach equilibrium (see problem 19-4 and Figure 19.8).
19-11.
a. If a gene has three alleles then p + q + r = 1. The genotype frequencies of a population at HardyWeinberg equilibrium would therefore be represented by the binomial expansion of the allele

Chapter 19

88

frequencies: (p + q + r)2 = (1)2 = p2 + 2pq + q2 + 2pr + r2 + 2qr = 1. As discussed in the

Problem Solving Tips at the beginning of this chapter, a Punnett square will give the same genotype
frequencies:
p

p
p2

q
pq

r
pr

pq

qr

pr

q2
pr

r2
b. In the Armenian population the allele frequencies are IA = 0.36, IB = 0.104 and i = 0.536. If the
population is in Hardy-Weinberg equilibrium then the binomial expansion of these allele
frequencies gives: 0.13 IAIA + 0.075 IAIB + 0.011 IBIB + 0.111 IBi + 0.287 ii + 0.386 IAi = 1.
Therefore the frequencies of the four blood types in this population are 0.516 A, 0.122 B, 0.075
AB and 0.287 O.
19-12.
a. When considering an X linked gene, women have two alleles. Therefore the allele and genotype
frequencies in women are calculated exactly as we have been doing p + q =1 and p2 + 2pq + q2
=1. However men are hemizygous (have only one allele for genes on the X chromosome) so the
men's allele frequencies and genotype frequencies are the same p + q = 1.
b. If 1/10,000 males is a hemophiliac then the allele frequency of the mutant allele (q) = 1 x 10-4.
Therefore the frequency of affected females = q2 = 1 x 10-8. If there are 1 x 108 women in the
United States then only one of them should be afflicted with hemophilia.
19-13.
a. Remember that colorblindness is an X-linked recessive trait. Therefore boys are hemizygous the
allele frequency for C = 8324/9049 = 0.92 and for c = 725/9049 = 0.08.
b. If the population is in Hardy-Weinberg equilibrium then the girls should have the same allele
frequencies as the boys. The genotype frequency of colorblindness (cc) in the girls = 40/9072 =
0.0044 and if the girls are in equilibrium then the allele frequency of c = 0.0044 = 0.066. This
does not equal 0.08 (from part a).
Alternately if the population is in equilibrium then the allele frequency of c in the boys should
accurately predict the genotype frequencies in the girls. Thus c2 should equal the frequency of
colorblind girls - (0.08)2 = 0.0064 which does not equal the cc genotype frequency of 0.0044.
Therefore this sample does not demonstrate Hardy-Weinberg equilibrium.

89

Chapter 19

c. Based on this information the frequency of the cpcp genotype in girls = 3/9072 = 3.3 x 10-4 and
the frequency of cp = 3.3 x 10-4 = 0.018. The frequency of the cdcd genotype in the girls =
37/9072 = 0.0041 and the frequency of cd = 0.0041 = 0.064. The frequency of the C allele is 1
(0.018 + 0.064) = 0.918.
d. The genotype frequencies among the boys are the same as the allele frequencies in the girls
calculated in part c. Thus in boys C = 0.918 (normal vision), cd = 0.064 (colorblind) and cp =
0.018 (colorblind). In the girls the genotype frequencies are: CC = 0.843 (normal vision), Ccd
= 0.118 (normal vision), Ccp = 0.033 (normal vision), cpcp = 3.3 x 10-4 (colorblind), cdcd =
0.004 (colorblind) and cdcp = 0.002 (normal vision).
e. These results make it much more likely that the population is in equilibrium. As seen in part c,
the allele frequency of C is the same in boys and girls and the allele frequency of c in the boys
is the same as the total frequencies of cd + cp in girls. Likewise the frequencies of genotypes
with normal vision (0.918 predicted in boys : 0.92 observed in boys and 0.996 predicted in girls vs
0.996 observed in girls) vs colorblind vision (0.082 predicted in the boys vs 0.08 observed and
0.004 predicted in the girls vs 0.004 observed) are the same in boys and girls.
19-14. The genotype frequency of the recessive, non-eye rolling phenotype (ugh ugh) is 410/500 = 0.82
for the French population and 125/200 = 0.625 for the Kenyan population. If each of these groups is in
equilibrium then the allele frequency for ugh in the French group is 0.82 = 0.906 and in the Kenyan
group the ugh allele frequency = 0.625 = 0.791. If the two groups married and had children at
random then the French group would provide 500/700 = 71.4% of the total ugh alleles and the Kenyans
would provide 200/700 = 28.6% of these alleles (see problem 19.9). Therefore the frequency of the
ugh allele in the mixed group = 0.714 (0.906) + 0.286 (0.791) = 0.873. The Ugh allele frequency = 1
0.873 = 0.127. The genotype frequencies in the children will be 0.762 ugh ugh + 0.222 Ugh ugh +
0.016 Ugh Ugh = 1. If there are 1,000 children then there will be 762 ugh ugh children that will not
express the eye rolling trait and 238 Ugh - children that will roll their eyes.
19-15.
a. Diagram the cross:
vg+ vg+ x vg vg F1 vg+ vg F2 1/4 vg+ vg+ : 1/2 vg+ vg : 1/4 vg vg (3 wild type : 1
vestigial). If the vestigial F2 flies are selected against then the remaining F2 genotypes are 1/3 vg+
vg+ and 2/3 vg+ vg. Therefore the genotype frequencies in the F2 are 0.33 vg+ vg+ and 0.67

Chapter 19

90

vg+ vg; the allele frequencies in the F2 for vg+ = 0.33 + 1/2 (0.67) = 0.67 and for vg = 1/2
(0.67) = 0.33.
b. The expected genotype frequencies in the F3 progeny are 0.449 vg+ vg+ + 0.442 vg+ vg + 0.109
vg vg = 1, or 0.891 wild type and 0.109 vestigial.
c. If the F3 vestigial flies are selected against then the altered F 3 genotypic ratio becomes 0.449 vg+
vg+ + 0.442 vg+ vg = 0.891. In order to calculate allele frequencies this equation must be
normalized (see problem 19-4c), becoming 0.504 vg+ vg+ + 0.496 vg+ vg = 1. The F3 allele
frequencies: vg+ = 0.504 + 1/2 (0.496) = 0.752 and vg = 1/2 (0.496) = 0.248. The genotype
frequencies in the F4 generation will be: 0.566 vg+ vg+ + 0.373 vg+ vg + 0.062 vg vg = 1. The
genotype frequencies in the F4 generation will be: 0.566 vg+ vg+ + 0.373 vg+ vg + 0.062 vg vg =
1. Therefore the F4 allele frequencies are vg+ = 0.566 + 1/2 (0.373) = 0.753 and vg = 0.062 +
1/2 (0.373) = 0.247.
d. If all of the F4 flies are allowed to mate at random then there is no selection and the population
will be in Hardy-Weinberg equilibrium. Therefore the F5 genotype and allele frequencies will be
the same as those in the F4 generation in part c above: 0.566 vg+ vg+ + 0.373 vg+ vg + 0.062 vg
vg = 1; vg+ = 0.753 and vg = 0.247.
19-16.
a. Convert the genotypes to frequencies: 60/150 t+t+ = 0.4 t+t+ and 90/150 t+t = 0.6 t+t. The allele
frequencies are: t+ = 0.4 + 1/2 (0.6) = 0.7 and t = 1/2 (0.6) = 0.3.
b. First determine the frequencies of the three genotypes if all lived, then remove the inviable mice
from your calculations and normalize the genotype frequencies in order to calculate the allele
frequencies. The expected genotypes frequencies of the zygotes in the next generation are: 0.49
t+t+ + 0.42 t+t + 0.09 tt = 1. However the tt zygotes die, so the remaining genotype frequencies
are: 0.49 t+t+ + 0.42 t+t = 0.91. When this is normalized it becomes 0.538 t+t+ + 0.462 t+t = 1.
If 200 progeny mice are scored there will be 108 normal mice and 92 tailless mice.
c. The Dom 1 population has 64 members and the genotype frequencies are 0.25 t+t+ and 0.75 t+t.
Thus the allele frequencies are t+ = 0.625 and t = 0.375. The Dom 2 population has 84 members

91

Chapter 19

and the genotype frequencies are 0.571 t+t+ and 0.429 t+t. The allele frequencies here are t+ =
0.785 and t = 0.215. If the populations interbreed randomly then the Dom 1 parents will provide
64/148 = 0.432 of the alleles (gametes) found in the next generation and Dom 2 will provide 0.568
or 56.8% of the alleles. Therefore the combined gamete frequencies are: t+ = 0.432 (0.625) +
0.568 (0.785) = 0.716 and t = 0.432 (0.375) + 0.568 (0.215) = 0.284. The genotype frequencies in
the next generation will be (0.716)2 t+t+ + 2(0.716)(0.284) t+t + (0.284)2 tt = 1. Of course the tt
zygotes die (see part b above) so the normalized genotype frequencies of the two viable genotypes
are 0.558 t+t+ and 0.442 tt.
Section 19.2 Causes of Allele Frequency Changes
19-17. A fully recessive allele is not expressed in a heterozygous organism, so there is no selection
against the heterozygotes. Selection against the homozygous recessive genotype will decrease the
frequency of the recessive allele in the population, but it will never totally remove it, as the
recessive allele is hidden in the heterozygote (Figure 19.8). In addition, a recessive allele sometimes
confers an advantage when present in the heterozygote, as seen for the sickle cell allele in areas
where malaria is prevalent. Finally, mutation can produce new recessive alleles in the population.
19-18. The farther from equilibrium, the greater the q, so the population with an allele frequency
of 0.2 will have the larger q.
19-19.
a. The allele frequency of b = 0.25 = 0.5 so the allele frequency of B = 0.5.
b. To calculate q, first determine q in both generations. For tank 1 (and all three tanks), q = 0.5 (see
part a) and q in the next generation (q') = 0.16 = 0.4. Therefore q for tank 1 is q'- q = 0.4 - 0.5
= -0.1. The same calculations are carried out for the other two tanks: q for all tanks = 0.5; q' for
tank 2 = 0.5 so q = 0; q' for tank 3 = 0.55 and q = 0.05.
b. q

Tank 1
-0.1

Tank 2
0.0

Tank 3
0.05

c. wBb
1.0
1.0
1.0
d. wbb
<1.0
1.0
>1.0
c. If the fitness of the BB genotype = 1 (wBB = 1) and the b allele is totally recessive then the
fitness of the Bb genotype (wBb) = 1 also (see row c. in the table above).
d. In all three tanks the original frequency of the b allele (q) = 0.5. In tank 1 the frequency of b in the
progeny (q') = 0.4 so the frequency of the bb (small tail) males has decreased. Therefore the fitness

Chapter 19

92

of the small tailed males decreased, so wbb <1.0. In tank 2 q' = 0.5 (the b allele frequency
remained the same) so wbb = 1.0. In tank 3 q' = 0.55 (the b allele frequency increased) so wbb
>1.0.
19-20. The equilibrium frequency will be different for the two populations. Equilibrium frequency
is a balance between selection and mutation and the selection is very different in the two
populations.
19-21.
a. The affected genotype dies before reproductive age. Therefore the fitness value (w) = 0 and the
selection coefficient (s) = 1 for the affected genotype. There is no selection pressure against the
carrier or the homozygous normal genotypes, so for both of these w = 1 and s = 0.
b.

w = p2 wRR + 2pq Rr + q2 wrr = (0.96)2 x 1.0 + 2 (0.96) (0.04) x 1.0 + (0.04)2 x 0 = 0.9984
(See equation 21.4a).
q = -srr p q2 / w = -1 (0.96) (0.04)2 / 0.9984 = -1.54 x 10-3 (see equation 21.7).

c. If the mutation rate from CF+ to CF- is 1 x 10-6 then the expected evolutionary equilibrium
^
frequency ( q ) of the CF- allele is:
-6
-6
-3
-3
^
q = ( / s p) = [(10 ) / (1 x 0.96)] = [10 / 0.96] = 1.02 x 10 . This number (1.02 x 10 )
is smaller than the observed q which is 0.04. This implies that there must be some heterozygote
advantage to the mutant CF- allele.
d. Interestingly one recent study suggests the hypothesis that CF+/CF- heterozygotes may be better
able to survive outbreaks of cholera. This is possible because people with cholera have diarrhea
that pumps water and chloride ions out of the small intestine. The CFTR protein encoded by the
CF gene is a chloride ion channel. CF+/CF- heterozygotes thus lose less water than CF+/CF+
individuals when infected with cholera. Thus the heterozygotes are less likely to die of dehydration.
Section 19.3 Analyzing Quantitative Variation
19-22.
a. Using genetic clones, only environmental effects contribute to variation.
b. Monozygotic twins are genetically identical so they can be thought of as genetic clones whereas
dizygotic twins are genetically different. Comparison of MZ and DZ twins provides an
assessment of the effect of genes versus environment.

93

Chapter 19

c.

Cross-fostering is removing offspring from a mother and placing with several different mothers to
randomize the effects of different mothering environments. This is done to reduce environmental
effects when determining heritability of a trait.

19-23. High heritability indicates that the phenotypic differences observed are due in large part to
genetic differences. Choice b would be true.
19-24.
a. Calculations of heritability vary in different environments, so by controlling environmental
similarity we can get a better estimate of the effect of genetic factors in that population.
b. Concordance values for MZ twins would allow a more accurate estimation of heritability, as
MZ twins have a genetic similarity of 1 but an environmental similarity about equal to either nontwin siblings or DZ twins.
19-25.
a. The table shows the average differences for each category. If a trait has a high heritability the MZ
(monozygotic) twins would have small average differences compared with DZ (dizygotic) twins or
siblings. You also expect that MZ twins raised together would have about the same (small) average
differences as MZ twins raised apart since the environment would not contribute much. Thus, the
table shows that Height has the highest heritability because both categories of MZ twins have
about the same average differences, and these are much lower than the average differences for DZ
twins or siblings. The table also implies that Weight has the lowest heritability since the average
differences for MZ twins raised apart are much higher than for MZ twins raised together; in fact,
the average differences for MZ twins raised apart is almost the same as for DZ twins or siblings.
IQ is somewhere in between, since the average differences for MZ twins raised apart is apparently
significantly less than the differences for DZ twins or siblings. However, the data shows that
environment is a very strong influence on the phenotype of IQ scores since there is considerably
more variation among MZ twins raised apart than for MZ twins raised together.
b. The data from the CDC does not affect the conclusions from part a. During the 42-year period
of the study, there can have been very little change in the genetic composition of such a large group
of people (all 15 year-old boys in the United States), yet there have been very significant changes in
the height and weight of this population. The difference of roughly 15 pounds indicates that
environment (most probably, diet) plays a critical role in determining the phenotype of weight. The
data from the CDC does seem to show that the environment also plays more of a role in
determining the phenotype of height than would have been inferred from the data in part a.

Chapter 19

94

19-26. Heritability values depend on the frequency of the trait in the population, on the total
amount of phenotypic variation in the population and on environmental factors that could
influence the trait. Comparing frequency and total phenotypic variation for the trait between the two
populations could provide evidence that heritability values for the two populations would be expected
to be different.
19-27.
a. There is a founder effect of the descendants coming from a small number of individuals, so
whatever recessive alleles were present in the population are more likely to be combined. Therefore
the frequency of some alleles and genotypes can be higher in that population. Other alleles may not
have been included in that original gene pool.
b. An advantage to studying the Finnish population is that there is genetic homogeneity and
probably fewer genes (potential modifiers) that may affect the trait and therefore can be more
easily dissected. A disadvantage is that some mutations that are present in general population
may not be found in this small, inbred population and therefore will not be identified in studies
of Finns.
19-28. The second trait, because the greater number of loci increases the potential for the
accumulation of new mutations that will affect the trait. The first trait will reach a selective plateau
more quickly.
19-29.
a. The relationship can be expressed with the formula 2n +1 where n = number of genes.
b. For one gene one of the extreme phenotypes (either homozygous genotype) will be found with a
frequency of 1/4 in the F2 generation. When you know the ratio of one of the extreme phenotypes
in the progeny you can us (1/4)n to calculate the number of genes. In this case (1/4)n = 1/256, so
(1/4)4 = 1/256 and n=4.

95

Chapter 19

19-30.
a. Remember that A', B', C' and D' each add 2 cm to leaf length, while A, B, C and D add 4 cm each
to leaf length. Each allele is incompletely dominant and the alleles of all four genes have additive
effects. Thus the genotype A'A' B'B' C'C' D'D' will give the shortest leaves, 16cm long. The
phenotype will increase by an additional 2 cm with each non-prime allele that is added to the
genotype. Therefore there are 9 possible phenotypes (2n + 1 as in problem 19-19a). The probability
of each allele = (1/2) and there are a total of 8 alleles controlling the phenotype. Therefore the
probability of any single genotype is (1/2)8 = 1/256. The simplest, although quite time consuming,
way to calculate the frequency of each phenotype is to set up the Punnett square of all possible
genotypes, as shown in Figure 3.22. In this case there are 16 different genotypes of gametes:
A'B'C'D', AB'C'D', A'BC'D', A'B'CD', A'B'C'D, ABC'D', AB'CD', AB'C'D, A'BC'D, A'BC'D,
A'B'CD, A'BCD, AB'CD, ABC'D, ABCD', ABCD. The frequency of each phenotype is: 16 cm (8
prime alleles), 1/256; 18 cm (7 prime alleles), 8/256; 20 cm (6 prime alleles), 28/256; 22 cm (5
prime alleles), 56/256; 24 cm (4 prime alleles), 70/256; 26 cm (3 prime alleles), 56/256; 28 cm
(2 prime alleles), 28/256; 30 cm (1 prime allele), 8/256; and 32 cm (0 prime alleles or 8 nonprime alleles), 1/256.
b. If the allele frequencies for gene A are A = 0.9 and A' = 0.1 then the genotype frequencies are AA
(p2) = 0.81, AA' (2pq) = 0.18 and A'A' (q2) = 0.01. For gene B the genotype frequencies are
BB = 0.81, BB' = 0.18 and B'B' = 0.01. For gene C the genotype frequencies are CC = 0.01,
CC' = 0.18 and C'C' = 0.81. For gene D the genotype frequencies are DD = 0.25, DD' = 0.5
and D'D' = 0.25.
c. When the frequency of the alleles in the population is not uniform as it is in part a, you have to use
the allele or genotype frequencies to determine the probability of obtaining a particular genotype, in
this case AA BB CC DD. The probability of this genotype is the product of the appropriate
genotypes for the individual genes calculated in part b above. Therefore the probability of AA BB
CC DD = (0.81) (0.81) (0.01) (0.25) = 0.0016.