Variation and Population

Genetics
Variation and Population
Genetics

Christopher J. Paradise, PhD

A. Malcolm Campbell, PhD
Variation and Population Genetics
Copyright Christopher J. Paradise and A. Malcolm Campbell. 2016.

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 Abstract
This book describes and analyzes genetic and environmental factors that
cause variation in individuals and populations. Data will be used to evalu-
ate the processes by which variation is generated in organisms and how
variation affects natural selection. Genetic factors include mutation, in-
dependent assortment, crossing over, and recombination. Environmental
factors include gradients and differences in abiotic conditions. Genotype
frequencies can be used to determine allele frequencies and this infor-
mation can be used to determine whether a population is evolving at
a genetic locus. The Hardy-Weinberg equilibrium will be applied as a
null model to make this determination. Non-Mendelian genetics can
affect the evolution of viruses and reassortment in viruses will be used to
illustrate another mechanism that generates variation in organisms and
how this mechanism relates to rapid evolution of viruses and the need
for annual flu vaccines.

Keywords
populations, phenotypes, genotype, traits, natural selection, inheritance,
homozygous, heterozygous, alleles, mutation, independent assortment,
meiosis, recombination, crossing over, environmental gradient, law of
segregation, law of independent assortment, genotype frequencies, allele
frequencies, Hardy-Weinberg equilibrium, Hardy-Weinberg theorem,
genetic drift, gene flow
 Contents
Preface...................................................................................................ix
Acknowledgments....................................................................................xi
Introduction.........................................................................................xiii
Chapter 1 Individual Organisms Exhibit Variation Caused
by Genetics and theEnvironment......................................1
Heritable Variation, Mutations and Independent
Assortment.....................................................................1
Variation Caused by the Environment................................8
Chapter 2 Population Genetic Information can be Used
toPredict Evolution.........................................................15
Chapter 3 Annual Flu Vaccines are Needed Because of
Non-Mendelian Genetics.................................................25
Ethical, Legal, and Social Implications:
Some People Refuse the Flu Vaccine for
Themselves and Their Children.....................................30
Conclusion............................................................................................35
Glossary................................................................................................37
Index....................................................................................................39
 Preface
This book about variation in populations and population genetics is part
of a thirty book series that collectively surveys all of the major themes
in biology. Rather than just present information as a collection of facts,
the reader is treated more like a scientist, which means the data behind the
major themes are presented. Reading any of the thirty books by P aradise
and Campbell provides readers with biological context and comprehen-
sive perspective so that readers can learn important information from a
single book with the potential to see how the major themes span all size
scales: molecular, cellular, organismal, population and ecologic systems.
The major themes of biology encapsulate the entire discipline: informa-
tion, evolution, cells, homeostasis and emergent properties.
In the twentieth century, biology was taught with a heavy emphasis
on long lists of terms and many specific details. All of these details were
presented in a way that obscured a more comprehensive understanding.
In this book, you will learn about biological variation, population ge-
netics and the Hardy-Weinberg theorem, and some of the supporting
evidence behind our understanding. The historic and more recent experi-
ments and data will be explored. Instead of believing or simply accepting
information, readers of this book will learn about the science behind
variation and population genetics, the way professional scientists do
with experimentation and data analysis. In short, data are put back into
the teaching of biological sciences.
Readers of this book who wish to see the textbook version of this
content can go to www.bio.davidson.edu/icb where the pedagogically-
designed and interactive Integrating Concepts in Biology for introduc-
tory biology college courses or a high school AP Biology course can be
accessed.
 Acknowledgments
Publishing this book would not have been possible without the generous
gift of Dr. David Botstein who shared some of his Breakthrough Prize
with co-author AMC. Davids gift allowed us to hire talented artists (Tom
Webster and his staff at Lineworks, Inc.) and copyeditor Laura Loveall.
Thanks go to Kristen Mandava of Mandava Editorial Services for project
management and guidance. In particular, we are indebted to Katie Noble
and Melissa Hayban for their many hours and attention to detail.
Kristen Eshleman, Paul Brantley, Bill Hatfield and Olivia Booker
helped us with technology at Davidson College. We are grateful to ad-
ministrators Tom Ross, Clark Ross, Carol Quillen, Wendy Raymond,
Verna Case, and Barbara Lom who had confidence in us and encouraged
us to persist despite setbacks along the way.
Thanks to my wife Amy Brooks for her constant support during the
development of this textbook, and my daughter Evelyn for her endless
energy. Thanks to Malcolm Campbell for his steadfast resolve and opti-
mism. Without him, this book would not exist. Thanks to collaborator
Laurie Heyer for taking my sometimes half-baked math ideas and turn-
ing them into powerful and elegant Bio-Math Explorations. I learned
a lot from both of them. While the math is largely absent from this
book, our collaboration with her made this a better book. Nancy Stamp
at Binghamton University, and Bill Dunson and Richard Cyr at The
Pennsylvania State University influenced me greatly in how I think as
a scientist and approach my teaching. Finally, I thank my students
in Integrated Concepts in Biology II, who enthusiastically participated
in our experiment to redesign introductory biology, starting with the
text and ending with a new approach to teaching biology.
 Introduction
Look around any classroom. How many people in the room look the
same? What is the makeup of the class in terms of hair, eye, or skin color?
What about height? There is some variation in all of these traits. Much
of that variation has a genetic component and all of the variation relates
to information. In this chapter, information at the level of the individual
will be consideredfirst by investigating the causes of variation among
individuals and then by examining how genetic information within in-
dividuals plays out at the population level. Genetic and environmental
changes lead to variation within species.
 CHAPTER 1

Individual Organisms
Exhibit Variation
Caused by Genetics and
theEnvironment

This book focuses on information in individual organisms, but readers

should keep in mind that individuals make up populations, which are
typically defined as groups of individuals of the same species living in the
same place at the same time. Within most populations, individuals vary
in any number of phenotypes, or the set of observable characteristics
they possess, resulting from the interaction of genotype (their genetic
composition) with the environment. Only very rarely do two individuals
look exactly alike and that there is often much variation among individu-
als. Variation in this context is the extent to which phenotypes differ from
individual to individual. In most populations, each individual is different.
Consider just two examples: eye color and height; both vary quite a bit in
almost any human population.

Heritable Variation, Mutations and

Independent Assortment
In one of the first scientific studies of variation of characteristics, Sir Francis
Galton examined the relationship between the average height of a couple
and the height of their adult offspring (Figure 1). Galton was a British
scientist, explorer, and inventor. Galton used statistics and mathematics
and the slope of a best-fit line to quantify the influence of parents heights
2 VARIATION AND POPULATION GENETICS

74
y = 0.65x + 23.94
72
height of child (inches)

70

68

66

64

62

60
62 64 66 68 70 72 74
mean height of parents (inches)

Figure 1 Data showing the relationship between height of parents

and offspring. The steeper line indicates a slope of one, and the
shallower line indicates the best-fit line indicated by equation. The
size of the circles is proportional to the number of comparisons. Total
sample size = 928 offspring and 205 sets of parents.
Source: Redrawn with original data from Galton 1889.

on the height of their offspring. Sir Francis Galton invented the concept
of linear regression to help quantify this relationship.
There are many phenotypic traits that vary among humans. Some
obvious ones that can be investigated, besides eye color and height, are
weight, hair color and skin color. Every characteristic has some compo-
nent of heritability; that is, there are one or more genes that control the
phenotype for each individual. Eye color is an inherited trait influenced
by more than one gene. Height is thought to be controlled by multiple
genes, but there is also an environmental component of height, including
factors such as nutrition and stress level during development. Malnutri-
tion or exposure to toxins may negatively affect growth, whereas exposure
to hormones may enhance growth. Galton discovered that the heights of
parents correlate with their offspring, but not perfectly. Galton found that
if the average height of parents was low, a high percentage of offspring
were taller than their parents average height, as indicated by the points
above the dashed line for averages below 67 inches (Figure 1). Likewise,
tall parents tended to have more offspring below their average height.
 Individual Organisms Exhibit Variation 3

These two trends yielded a best-fit line that had a slope of 0.65, much
less than 1.0. In addition, he found a high degree of variability around
the average for any set of parents, as indicated by the scattering of points
around the best-fit line. Less variation would lead to more or all points
falling on the best-fit line.
Individuals within populations that vary in characteristics also have
varying degrees of success. To illustrate the importance of variation to his
thesis, Darwin defined natural selection as ...preservation of favour-
able individual differences and variations, and the destruction of those
which are injurious... Although Darwin used variation as a central
concept in his Theory of Evolution by Means of Natural Selection, how
variation among individuals in a population was generated or maintained
was not known in Darwins time. There are several different mechanisms
that produce variation among individuals, and they will be explored them
in this chapter.
As Galton observed, a characteristic may be heritable to some extent
and yet highly variable. Therefore, the genetic code is one source of vari-
ability. Charles Darwin wrote in The Origin of Species that if variations in
phenotypes, however slight, are in any way advantageous to an individual,
those variations tend to increase the ability of the individual to survive and
reproduce. If the phenotype is inherited by offspring, then genes playing
a role in determining that phenotype will spread in the population. Al-
though how inheritance of characteristics worked was not understood in
Darwin or Galtons time, their insights and data were important founda-
tions in the study of how variation among individuals affected evolution.
Much of the variation among individuals in a population is caused
by genetic differences; that is, variation in deoxyribonucleic acid (DNA)
sequences. An example that illustrates this source of variation is a study
of blood pressure in rats (Rattus norvegicus) made by Giuseppe Bianchi
and his colleagues. As in humans, some rats have higher blood pressure
than others. The scientists developed two colonies of rats by breeding rats
that had low blood pressure with each other in one colony, and rats that
had higher blood pressure with each other in the other colony. After 85
generations of doing this, the blood pressures were very similar among
individuals within each colony, but very different between individuals in
different colonies (Figure 2).
4 VARIATION AND POPULATION GENETICS

180

160 = systolic pressure

blood presure (mm Hg 1 s.e.)

= diastolic pressure
140

120

100

80

60

40

20

0
high BP low BP
rat colony

Figure 2 Mean blood pressures for rats in the two colonies, measured
in millimeters of mercury. All high blood pressure rats were
homozygous for the adducin genes (Y and R). Low blood pressure
rats were all homozygous for the F gene only.
Source: Data from Bianchi et al., 1994.

The DNA of several genes was tested to determine whether the rats
were homozygous (possessing two copies of the same version of a gene) or
heterozygous (possessing two different versions of a gene) for these genes.
When Bianchi and colleagues tested twenty rats from each colony, they
found that all rats tested were homozygous for each gene within each strain,
with one exception. The exception was a protein called adducin, which is
composed of two subunits (called and ) whose genes are located on two
different chromosomes. The different versions, or alleles, of each protein
subunit differed by only one amino acid. They named the alleles of the
proteins Y, F, R, and Q, depending on the subunit and the amino acid
that was present at the variable position. For the subunit, the scientists
found that a tyrosine (with the one letter code of Y) substituted for a phe-
nylalanine (F) at position 316. Thus Y stands for the mutated subunit in
the high blood pressure colony and F stands for the normal, wild-type
subunit. For the subunit, an arginine (R) substitutes for glutamine (Q)
at position 529. Thus R stands for the mutated subunit in the high blood
pressure colony and Q stands for the normal, wild-type subunit.
 Individual Organisms Exhibit Variation 5

138

136
blood pressure (mm Hg 1 s.e.) 134

132

130

128

126

124

122

120
Q/Q Q/R R/R
type of beta adducin in low blood pressure rats

Figure 3 Mean systolic blood pressures of the three combinations

of two versions of the adducin gene in rats from the low blood
pressure colony. Q and R refer to the amino acid present at position
529 on the protein subunit of adducin. Error bars = 1 standard
error (SE).
Source: Data from Bianchi et al., 1994, in text.

The high blood pressure rats were found to be homozygous for the
and R versions of the adducin genes. The low blood pressure rats
Y

were homozygous for the F adducin gene, and the colony had all three
possible genotypes for the subunit: homozygous R, homozygous Q,
and heterozygous with both R and Q. When Bianchi and his colleagues
compared the blood pressures of these rats, they found that, although
lower than the high blood pressure rats, there were differences among the
three genotypes in the low blood pressure rats (Figure 3).
The scientists crossbred rats from the two colonies. Of course, rats
from the high blood pressure colony were homozygous for both adducin
genes. They chose rats from the low blood pressure colony that were also
homozygous for the Q gene. The resulting offspring in the first genera-
tion were rats heterozygous for both genes; that is, they carried one copy
of Y and R from their high blood pressure parent and one copy of F
and Q from their low blood pressure parent. All the first generation rats
were then crossbred to produce rats of every possible combination of the
6 VARIATION AND POPULATION GENETICS

two versions of the two genes. Blood pressure of these rats was measured
to determine the impact of having any combination of these genes.
The resulting average systolic blood pressures of rats in the second
generation were all in between the systolic blood pressures measured in
rats from the two colonies (the original parental generation). After two
generations of crossbreeding rats from the two colonies, the scientists ob-
tained rats with intermediate blood pressures. However, there were still
some differences. Rats that were homozygous for Y and R, same as rats
from the high blood pressure colony, and rats that were homozygous for
Y and heterozygous QR both still had higher blood pressure than rats
that were FF and either RR or QR. Rats that were heterozygous at
the subunit typically had intermediate blood pressures, as did rats that
were homozygous for F and Q.
This study illustrates several processes that increase variation among
individuals. Bianchi and his colleagues concluded that a point mutation,
a change in a single nucleotide in a DNA sequence, was responsible for
producing the two alleles of the and subunits of adducin. These muta-
tions increased the variation among rats, despite the fact that the scientists
separated them out by selecting for rats that had either high or low blood
pressure in the two colonies. The variation increased because after the
mutations there were two alleles of each gene, leading to two versions of
each adducin subunit. That variation produced several combinations of
the adducin protein.
Generations of inbreeding produced individuals that were mostly ho-
mozygous, at least for all but one of the genes tested. Because the scien-
tists were selecting for individuals with a certain phenotype, individuals
with other phenotypes were subsequently lost from the population because
those rats were removed by the researchers. Eventually most of those alleles
that cause undesirable traits in a population were removed from the popu-
lation. In the case of the adducin gene in the low blood pressure colony,
the heterozygous condition produced rats that had the lowest systolic
blood pressure. Although this outcome may be surprising, it illustrates that
individual variation sometimes produces a phenotype that is selected for.
Rats with high blood pressure have associated health problems, such
as red blood cell and kidney dysfunctions. This led the scientists to con-
clude that slightly harmful mutations had occurred over the generations
 Individual Organisms Exhibit Variation 7

of inbreeding. The harmful mutations were able to remain in the popula-

tion because the rats were bred in the laboratory. In nature, these harm-
ful mutations are eliminated by natural selection. Although the scientists
identified the two adducin genes as playing a role in high blood pressure,
there are other genes involved, as must be the case in order to obtain
rats in the second generation of crossbreeding that possessed intermediate
blood pressures. The two alleles of the two subunits of adducin interact
when in different combinations to cause wide variation in high blood
pressure, as indicated by the mean blood pressures of rats in the second
generation of crossbreeding. However, none of these rats has blood pres-
sure as low or high as their grandparents, and that variation is caused
by other genes also controlling blood pressure. After two generations of
crossbreeding, the different alleles of these other genes, from the homo-
zygous grandparents, had been reshuffled independently of the adducin
genes, increasing the variation of responses.
The observation of blood pressure variation and the wide range of
combinations of the two versions of the two adducin genes leads to an-
other conclusion about how variation increases and is maintained in indi-
viduals. Independent assortment, when non-homologous chromosomes
migrate without regard to each other during meiosis, leads to recombi-
nation. Recombination leads to new combinations of genes in offspring
that did not occur in the parents, by independent assortment and cross-
ing over, which occurs when paired chromosomes exchange portions of
their DNA during meiosis.
Bianchi and his colleagues knew that the genes coding for the two
subunits of adducin were located on different chromosomes; so they
could predict that when two heterozygous individuals mated, offspring
of those individuals would exhibit one of nine different combinations of
the two genes (Table 1A).
The heterozygotes were obtained from mating of rats from each col-
ony that were homozygous for different versions of the two genes. A het-
erozygote possesses two alleles for a particular gene, resulting in variation
within an individual. However, if the genes were on the same chromo-
some, the variation of outcomes would be limited (Table 1B). This latter
scenario assumes no crossing over. If there were crossing over, the extent
of variation would also increase.
8 VARIATION AND POPULATION GENETICS

Table 1 Variation of combinations of the two adducin genes. All

parents in the matings are heterozygous for both the adducin subunits
and can contribute one of two alleles for each gene to their offspring.
F and Y are alleles of the subunit, and R and q are alleles of the
subunit. A, Variation when the genes are located on two different
chromosomes, showing independent assortment. B, Variation when
the genes are on the same chromosome, with no crossing over.
A range of alleles
contributed from male parent
F/Q F/R Y/Q Y/R
F/Q FF/QQ FF/QR FY/QQ FY/QR
Range of allelles
F/R FF/QR FF/RR FY/QR FY/RR
contributed from
Y/Q FY/QQ FY/QR YY/QQ YY/QR
female parent
Y/R FY/QR FY/RR YY/QR YY/RR
B range of gene versions
contributed from male parent
F/Q F/Q

Range of allelles F/Q FF/QQ FY/QR

contributed from
female parent Y/R FY/QR YY/RR

Source: C. Paradise.

Variation Caused by the Environment

While genetic mechanisms can increase variation among individuals,
variation might also be caused by the environment. For instance, if ex-
pression of various genes is affected by the environment, variation among
individuals in a population may be affected. The effect of the environ-
ment on individual variation can be examined across an environmen-
tal gradient, the gradual change in an environmental factor from one
place to another. For instance, levels of metal contamination in the soil
surrounding metal ore smelting operations decrease as distance from the
smelter increases. Very few plants live in soils near smelters that have been
operating for many decades. Nicholas Caiazza and James Quinn studied
two species of plants near a smelting operation in eastern Pennsylvania,
slender sandwort (Arenaria patula) and Japanese honeysuckle (Lonicera
japonica). They collected plants from sample sites that varied in their dis-
tance from the smelter, along the pollution gradient. In addition, they
documented the level of metal pollutants at each site where plants were
collected (Figure 4).
 Individual Organisms Exhibit Variation 9

8000

zinc concentration in soils (ppm)

7000
= sandwort
6000
= honeysuckle
5000

4000

3000

2000

1000

0
near medium far

15
copper concentration in soils (ppm)

12

0
near medium far
proximity to smelting operation

Figure 4 Contamination in soils surrounding a smelting operation in

Pennsylvania. Sandwort plants were collected at three sites (near,
medium, far) and honeysuckle plants were collected at two sites
(near and far). A, Zinc concentrations in parts per million (ppm).
B, Copper concentrations in parts per million (ppm).
Source: From Caiazza and Quinn, 1980, Table 1.

Caiazza and Quinn counted stomata, the gas exchange pores on

leaves, and hairs on the leaves that they collected from the various
sites over a period of 2 years (Figure 5). They also took seeds from the
plants and grew them under standard conditions in a nearby courtyard;
this procedure would help the researchers determine whether differences
they might find near the smelter were due to the environmental gradient
or genetic differences that had evolved among the various populations.
After a period of time, they collected leaves from these courtyard grown
plants and performed the same analyses (see Figure 5).
10 VARIATION AND POPULATION GENETICS

honeysuckle stomata density honeysuckle hair density

lower leaf stomata density (#/mm2)

upper leaf hair density (#/mm2)

900 3.0
= near
800 = medium 2.5
= far
700 2.0

600 1.5

500 1.0

400 0.5

300 0.0
field 77 field 79 courtyard field 77 field 79 courtyard

sandwort stomata density sandwort hair density

upper leaf stomata density (#/mm2)

220 5

lower leaf hair density (#/mm2)

210 4
200
3
190
2
180

170 1

160 0
field 77 field 79 courtyard field 77 field 79 courtyard

Figure 5 Stomata and hair densities of two plants collected at two

times and grown in controlled conditions in a courtyard. Sandwort
plants from the intermediate site were not grown in the courtyard.
Source: Data from Caiazza & Quinn 1980, Tables 2 and 3.

Both species of plants exhibit variation across the gradient. The sand-
wort shows more consistent variation with distance from the smelter, and
honeysuckle still has variation when plants from different sites are grown
under controlled conditions. However, the variation across years is evi-
dence that some environmental factor affected the density of stomata and
hairs. Caiazza and Quinn speculated that heavy metals in the air affect
plants. Air pollution from the smelter affected the soil conditions, but
air quality varies more from year to year than soil conditions, and this
could have led to annual changes in variation among the sites. There is
also a genetic component, which is more evident with honeysuckle, but
there is also an environmental component that increases variation among
individuals. The number of stomata tends to be lower and hairs are denser
closer to the smelter, and that may be in response to polluted air. The
fewer stomata, the fewer pollutants will enter the plants when they open
for gas exchange. A high density of hairs on leaves may trap pollutants,
acting as a filter. The drawback to these changes is that plants will not be
 Individual Organisms Exhibit Variation 11

able to obtain as much carbon dioxide as other plants with more stomata,
but this may still be a phenotype that is advantageous and selected for in
a polluted environment.
In another study, Curtis Lively studied an acorn barnacle (Chthamalus
anisopoma) that has variation in shell shapes along a rocky intertidal habi-
tat off the Gulf of California. Barnacles are non-motile marine animals
that cement themselves to rocks and other substrates in tidal zones, in
order to keep from being washed away by the waves.
Lively knew that a certain type of snail, Acanthina angelica, specialized
on feeding on barnacles. This snail has a special spine on its shell that it
uses to pry open barnacles from the top and gain access to the animal in
its shell. Lively knew from the observations of other naturalists that few
barnacles live near where a snail had a refuge, which the snail uses for
protection from desiccation. At intermediate distances from snail refuges,
barnacles with the bent shell type occur sparsely, but more frequently
than barnacles with the cone shell type. The bent shell type has the
opening of the barnacle on the side rather than at the top like the cone
shell type has. Barnacles emerge from the opening, either on the side or
the top, to filter feed in the water. Far from snail refuges, barnacles with
the cone shell type occur at high density and more frequently than bent-
shell barnacles.
To determine whether the variation in shell type was due to an envi-
ronmental factor that affected the development of barnacle shell shape,
Lively needed to distinguish between correlation and causation; the in-
creased presence of bent-shell barnacles could be correlated with the pres-
ence of snails, or it could be caused by the presence of some other factor.
The scientist set up an exclusion experiment with plots where the bar-
nacles were allowed to develop with the snails present and with the snails
excluded. An exclusion experiment is where one or more species are ex-
cluded from experimental plots or habitats. Barnacles with the bent shell
type developed only in the presence of snails (Figure 6).
He then set up other plots where he placed barnacles of each shell
shape, and half of those plots with the snail predator and other half with-
out, and tracked the survival of barnacles with each shell shape over time.
In the absence of the snail predator, barnacles with either shell type sur-
vived equally well (both about 90% after 5 days) and always with higher
12 VARIATION AND POPULATION GENETICS

= cone
120
= bent

number of barnacles ( 1 s.e.)

100

80

60

40

20

0
snail present snail absent
predator treatment

Figure 6 Number of acorn barnacles (Chthamalus anisopoma) in

plots with and without the snail predator (Acanthina) in the predator
exclusion experiment.
Source: From Lively, 1986, Table 1.

survival than in the presence of the snail predator. However, in the pres-
ence of the snail predator barnacles with the bent shell type had much
higher survival (about 80%) than barnacles with the cone shell type
(about 40%).
The barnacle study demonstrates that a predator can cause develop-
mental changes. The bent shell variety is less common overall than the
cone and never developed in the absence of the predator, but a significant
number of them developed in the presence of the predator. However, the
cone shell phenotype was present in both the presence and absence of the
snail. Even in the presence of the predator, there was a large number of
cone-shaped barnacles, indicating that these individuals either did not re-
ceive the predator cue during development or that they were genetically
incapable of growing into the bent shape. As was the case with honey-
suckle and sandwort, environmental factors can alter the physical ap-
pearance of animals but only within the scope of their genetic potential.
The bent shape may be rare in the absence of the predator because of a
decrease in feeding efficiency, but the benefit of having this shell shape in
the presence of the predator is that it increases the probability of survival.
Careful experimentation with proper controls can determine the
causes and extent of natural variation in populations and how this
 Individual Organisms Exhibit Variation 13

variation relates to information in populations. Variation among indi-

viduals can lead to the evolution of populations, just as natural selection
acts to alter the relative abundance of individuals with particular charac-
teristics. This chapter demonstrates that variation among individuals and
information in populations increases through mutation, recombination,
independent assortment, and environmental factors. Those variations can
have implications to survival and reproduction, as was observed with the
barnacles, and may be acted upon by the mechanisms of evolution. In
the next chapter, one of the major mechanisms and concepts that scien-
tists study with regard to variation in populations, the Hardy-Weinberg
theorem, is examined.

Bibliography
Bianchi G, Tripodi G, Casari G, et al.: Two point mutations within the
adducin genes are involved in blood pressure variation, Proc Natl
Acad Sci USA 91(9):39994003, 1994.
Caiazza NA Jr, Quinn JA: Leaf morphology in Arenaria patula and
Lonicera japonica along a pollution gradient, Bull Torrey Bot Club
107(1):918, 1980.
Darwin C: On the origin of species by means of natural selection, 1859,
ZETACRAFT Publishing 2010.
Galton F: Natural inheritance, 1889, MacMillan.
Lively CM: Predator-induced shell dimorphism in the acorn barnacle
Chthamalus anisopoma, Evolution 40(2):232242, 1986.
 Index
Acanthina angelica, 11 Flu vaccines, 25
Acorn barnacle (Chthamalus concerns about long-term health
anisopoma), 1112 risks of, 32
Acquired immune deficiency refusal of, 3033
syndrome (AIDS), 27 Fontanilla, Ian, 15
Adducin, 4, 6 Food and Drug Administration
Advisory Committee on (FDA), 32
Immunization Practices
(ACIP), 33 Galton, Francis, 12
Air pollution, 10 Gametes, 15
Allele frequencies, 16, 17 Gene flow, 21
Alleles, 4 Genetic code, 3
Alzheimer disease, 32 Genetic drift, 21
American Academy of Pediatrics Genotype, 1
(AAP), 33 Genotype frequencies, 16, 17
Arenaria patula. See Slender Glutamine, 4
sandwort Glycoprotein, 15
Arginine, 4 Guillain-Barr syndrome (GBS), 33
Avian influenza viruses, 28, 30

Barnacle. See Acorn barnacle H1N1, 27, 29

Bianchi, Giuseppe, 37
Bird flu, 29
Blood pressure variations,
in rats, 37
Caiazza, Nicholas, 810
Centers for Disease Control and
Prevention (CDC), 29, 32
Chthamalus anisopoma. See Acorn
barnacle
Crossing over, 7
Darwin, Charles, 3
Environmental gradient, 8
Epidemic, 25
Evolutionary processes
population genetic information
used to predicting, 1523
Exclusion experiment, 11
H5N1, 2930
Hardy-Weinberg (HW) genotype
frequencies, of MN genetic
locus, 16
Hardy-Weinberg equilibrium, concept
of, 17, 18, 2022
Hardy-Weinberg theorem, 17
Hemagglutinin, 27
Heterozygous, 4
Homozygous, 4
Human immunodeficiency virus
(HIV), 27
Independent assortment, 7
Influenza, 2532
Influenza A H1N1, 29, 31
Influenza A H3N2, 29, 31
Influenza A virus, 2629
Influenza B viruses, 29, 31
Inheritance, 3
40 INDEX

Japanese honeysuckle (Lonicera Quinn, James, 810

japonica), 810
Law of independent assortment, 15
Law of segregation, 15
Linear regression, 2
Live-attenuated virus, 32
Lively, Curtis, 11
Lonicera japonica. See Japanese
honeysuckle
Meiosis, 7
Mendels laws of inheritance, 15
Messenger RNA (mRNA), 26
Mutation, 6
Nasal spray vaccine, 32
National Academy of Sciences
Institute of Medicine, 32
National Institutes of Health
(NIH), 32
Natural selection, 3
Neuraminidase, 27
1918 pandemic, 2526
Origin of Species, The, (book), 3
Pandemics, 25, 29, 33
Peoples refusing flu vaccines, reasons
for, 3033
Phenotypes, 1, 3
Phenylalanine, 4
Population genetic information, used
to predicting evolutionary
processes, 1523
Populations, 1
Rattus norvegicus, 38
Recombination, 7
Refusal of vaccines, reasons
for, 3033
Ribonucleic acid (RNA) genome, 25
Single-stranded RNA (ssRNA) virus,
26, 28
Slender sandwort (Arenaria
patula), 810
Snails, 1112
Swine influenza viruses, 28
Thimerosal, 3233
Traits, 2
Tyrosine, 4
Vaccines, 25
reasons for refusing, 3033
Variation
caused by the environment, 813
of characteristics, 1
of combinations of two adducin
genes, 78
genetic differences, 3
and genetic information, 15
importance of, 3
leading to evolution of
populations, 13
among rats blood pressure, 37
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