Professional Documents
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Variation and
Bundlethe more ment, crossing over, and recombination. Environmental factors
books you buy, include gradients and differences in abiotic conditions. Gen-
the greater your otype frequencies can be used to determine allele frequen-
discount! cies and this information can be used to determine whether a
population is evolving at a genetic locus. The Hardy-Weinberg Population
Genetics
equilibrium will be applied as a null model to make this deter-
THE CONTENT
mination. Non-Mendelian genetics can affect the evolution of
Energy Physics viruses and reassortment in viruses will be used to illustrate
Engineering another mechanism that generates variation in organisms and
Biotechnology how this mechanism relates to rapid evolution of viruses and
Biology the need for annual flu vaccines.
Mathematics
Christopher J. Paradiseis professor of biology and environ-
Chemistry
mental studies at Davidson College. He teaches introductory
biology, ecology, entomology, and topical seminars on ecotoxi-
THE TERMS cology and renewable natural resources. He also occasionally
Perpetual access leads a study abroad program in India. His research evaluates
for a one time fee anthropogenic factors that influence insect biodiversity at a
No subscriptions or variety of scales. His current research interests include effects of
access fees land use patterns on pollinator communities in parks.
Unlimited
A. Malcolm Campbellteaches biology at Davidson College,
concurrent usage
NC. He received national and international education awards:
Downloadable PDFs Genetics Society of America (2013); American Association for the
Free MARC records Advancement of Science (2012); and American Society for Cell
Biology (2006). He was the founding co-editor in chief of CBE Life
For further information,
Sciences Education; founding director of Genome Consortium
a free trial, or to order,
contact: for Active Teaching (GCAT); and member of the American Soci- Christopher J. Paradise
sales@momentumpress.net ety for Cell Biology governing council (20122014).
A. Malcolm Campbell
Variation and Population
Genetics
Variation and Population
Genetics
10 9 8 7 6 5 4 3 2 1
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
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)
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
= 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
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
Source: C. Paradise.
8000
4000
3000
2000
1000
0
near medium far
15
copper concentration in soils (ppm)
12
0
near medium far
proximity to smelting operation
600 1.5
500 1.0
400 0.5
300 0.0
field 77 field 79 courtyard field 77 field 79 courtyard
220 5
170 1
160 0
field 77 field 79 courtyard field 77 field 79 courtyard
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
80
60
40
20
0
snail present snail absent
predator treatment
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
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.
CHAPTER 2
Population Genetic
Information can be Used
toPredict Evolution
Mendels two laws of inheritance are the law of segregation and the law
of independent assortment. The law of segregation states that paired
chromosomes move to opposite nuclei during formation of gametes, and
the law of independent assortment states that non-homologous chro-
mosomes migrate independently of each other. In Chapter 1 the law
of independent assortment was used to describe how non-homologous
chromosomes migrate without regard to each other during meiosis. This
then affects the variation and genetic information in a population of or-
ganisms. In this chapter, the information content in the genes of a popu-
lation will be examined to determine whether the population is evolving
at a genetic locus.
The MN blood group in humans is determined by two alleles at one
locus on chromosome 4. Expression of the gene leads to production of a
glycoprotein that presents on the surface of red blood cells. These glyco-
proteins are used by the immune system to distinguish self from non-self.
MM individuals present one glycoprotein, NN present another, and MN
individuals present both, so a simple blood test can determine pheno-
type and genotype simultaneously. A team of scientists led by Ian Fon-
tanilla collected data on the MN locus in several populations of humans
in the Philippines. The populations that they selected were distributed
along a north-south gradient and across different islands. They collected
blood from over 500 individuals, each of which was asked to fill out a
questionnaire to assess information on place of birth, ethnic identity,
16 VARIATION AND POPULATION GENETICS
= MM
0.8 = HW MM
= MN
= HW MN
= NN
= HW NN
0.6
frequency
0.4
0.2
0.0
Isabela Panga Manilla Cam Sur Cebu Palawan Butuan Dav Sur
population
point is that the information content in genetic loci can be used, as well
as other information about populations, such as isolation and migration,
to speculate on evolutionary mechanisms that may or may not be acting
on such populations.
Bibliography
Arcellana AES, Guzman RMS, Fontanilla IKC: Distribution of MN blood
group types in local populations in Philippines, J Genet 90:e90e93,
2011. Online only: http://www.ias.ac.in/jgenet/OnlineResources/90/
e90.pdf
Hardy GH: Mendelian proportions in a mixed population, Science 28:
4150, 1908.
Hastings A: Hardy-Weinberg theorem, In: Encyclopedia of life sciences,
2001, Macmillan Publishers Ltd., Nature Publishing Group. Avail-
able online: www.els.net.
Weinberg W: On the demonstration of heredity in man, In: Boyer SH
IV, translator: Papers on human genetics, Englewood Cliffs, NJ, 1963,
Prentice-Hall, pp. 415.
CHAPTER 3
got its name. The 1918 pandemic was caused by the tiny virus called
influenza A (Figure 8). Like any organism, viruses contain genetic infor-
mation that determines their phenotype. Influenza is a single-stranded
RNA (ssRNA) virus, which means its chromosomes are composed of
only one strand of a RNA molecule. Unlike DNA, single-stranded RNA
(ssRNA) is structurally similar to messenger RNA (mRNA) because it is
composed of only one RNA polymer.
Each individual influenza virus contains eight ssRNA chromosomes,
and the virus encodes only eight proteins. Each protein can differ, as mul-
tiple alleles exist for each gene. Therefore, the right combination of these
eight RNA genes, each with multiple alleles and thus multiple proteins,
is what distinguishes a typical flu season from a pandemic, and the varia-
tions in those combinations, as will be seen, is what requires people to get
a vaccine each year.
The influenza virus reproduces by making new copies of the eight
ssRNA chromosomes and packaging them into groups. Other viral ge-
nomes come in many forms and sizes. Their genomes can be composed
FLU VACCINES NEEDED BECAUSE OF NON-MENDELIAN GENETICS 27
the rapid rate of human travel, it would be easy for this particular virus to
reach every corner of the world in a matter of days. However, just as rapid
is the response of world health organizations, like the CDC.
Science knows more about influenza now than it did in 1918. Mil-
lions of birds have been killed to prevent the spread of H5N1, but thou-
sands of subsistence farmers lost their life savings and their ability to feed
themselves. How does humanity weigh the tradeoffs of two bad out-
comes? Eliminating influenza is impossible, because the virus can hide
in many different animals, including whales, which some people still eat.
Therefore, annual vaccinations for susceptible populations and healthcare
workers are a good preventive measure, and the only solution we have
given the lack of a permanent one. Avian flu is many times more virulent
than the pandemics of H1N1, so a devastating pandemic of an avian flu
could happen.
Viral reproduction provides for the continuity of the species by trans-
mitting genetic information to its progeny. However, the RNA genome is
susceptible to errors, which increases the rate of variation due to imperfect
replication and high rates of recombination. In addition to responding to
a new strain of virus each year, a persons immune system remembers pre-
vious strains as a form of biological memory. Furthermore, physicians use
vaccines to transmit immunological memory to their patients to prevent
serious consequences from new infections. With each new flu season and
each new animal infection, humanity runs the risk of a new combination
of chromosomes resulting in a highly contagious and lethal virus. The
ability to randomly generate a new strain of virus is advantageous to the
virus and is a good example of how genetic information is not always
passed on in a predictable pattern, a wrinkle of complexity within idea of
transmission of information at the organism level.
revealed that 61% of Americans planning to refuse flu vaccine dont think
theyre at risk of a serious case of flu. They argue that the flu is not a life-
threatening disease. It will get a person sick, but the probability of dying
is very low. Yes, people die from the flu they say, but not the average,
healthy individual. Even among the elderly, the risk of death from the flu
is 1 in 1000.
Influenza, however, is not a mild illness; it can make a person very sick
for 3 to 5 days. This takes a toll on student learning, worker productivity,
and the economy. It can also lead to more hospital visits, taking up beds
even in intensive care units. It can be very serious, and in fact hundreds
of Americans do die each year of the flu. Not only can a person help
themselves avoid that fate, they can help reduce the spread of infections to
other individuals, including those that are immunocompromised and are
not able to get an annual vaccine, if the person remains flu-free.
Others argue that the vaccine may not be available until a human
population is in the middle of an epidemic, and then it is too late. How-
ever, it turns out that not everyone gets the flu at the same time. Even in
areas where there is widespread flu, studies suggest that only about 5% to
10% of the population has been infected, and there is the potential for
further infections and lengthening the flu season if a substantial propor-
tion of the population is unvaccinated.
Opponents of the flu vaccine say that it is ineffective. A person may
still get the flu even if they get the vaccine. Some annual flu vaccines have
been particularly ineffective. However, scientists know that just because two
events happen in sequence (a person gets the shot and then he or she gets the
flu) does not mean that one caused the other. The vaccine protects against
the three genotypes of the virus that scientists predict will be most common
or virulent that year. Scientists take an educated guess as to what three of
several hundred genotypes of flu virus they expect will have the greatest
virulence. As mentioned previously, early data collected by the CDC allow
for the guess to be more educated. In early 2015, influenza A H1N1,
influenza A H3N2, and influenza B viruses were the three strains found.
These can be used to develop and ramp up production of a vaccine, which
will be protective against these three strains but not others.
When a person gets the flu vaccine, their body produces antibodies to
those three specific strains. If a fourth strain comes to dominate later in
32 VARIATION AND POPULATION GENETICS
the flu season, the vaccine will be ineffective. Other strains can infect that
person, and there are lots of other non-influenza viruses going around
during flu season. In fact, influenza accounts for less than one-third of
flu-like illnesses during flu season. Because people who are vaccinated do
not get the most virulent flu strain, they actually suffer less flu-like illness
than the people who do not get vaccinated.
Then there are opponents who argue that if a person gets the nasal
spray vaccine they will shed the flu virus and likely infect others. How-
ever, this version of the virus is called live-attenuated virus, and even if it
is true that vaccinated individuals will shed viral particles, attenuation is
a process that takes an infectious agent and alters it so that it becomes
harmless or less virulent. Again, not getting vaccinated at all might in-
crease a persons susceptibility to influenza, and if they get the flu, they
will definitely shed the live virus for many days.
Some people take measures to strengthen their immune systems
against any invading pathogens. For instance, they use Chinese herbs,
elderberry syrup, and vitamin D, claiming that they get sick less intensely
and less frequently than their friends and neighbors. There is some clini-
cal evidence that vitamin D reduces the likelihood of contracting the flu,
but less or no evidence that other such remedies strengthen the immune
system, which is a nebulous concept to begin with.
There are concerns about the long-term health risks of flu vaccines.
These concerns revolve around the inclusion of thimerosal, which is a
mercury-based preservative, and other heavy metals in vaccines. Some
research suggests a link between annual flu shots and development of
Alzheimer disease, for instance. However, it turns out this is not well
supported, and parts of this argument are not even true. For instance,
single-dose units are made without thimerosal, because they are opened
and used only once, negating the potential for contamination. The
live-attenuated version of the vaccine (the nasal spray vaccine) is also a
single-dose unit without thimerosal. No new vaccines approved by the
Food and Drug Administration (FDA) for children contain thimerosal.
Although multi-dose preparations do contain thimerosal, the scientific
research does not support the argument that thimerosal-containing vac-
cines are harmful. In fact, the CDC, the FDA, the National Institutes of
Health (NIH), the National Academy of Sciences Institute of Medicine,
FLU VACCINES NEEDED BECAUSE OF NON-MENDELIAN GENETICS 33
Bibliography
Aufderheide JJ: 8 damn good reasons not to get the flu shot, Vactruth.com
(website): http://vactruth.com/2013/02/01/8-damn-good-reasons/.
Accessed March 5, 2014.
Bartlett E: Should I get a flu shot? 5 reasons why not, Holistic Squid
(website): http://holisticsquid.com/should-i-get-a-flu-shot/. Accessed
March 5, 2014.
CDC: Seasonal flu (website): http://www.cdc.gov/flu/. Accessed
December 17, 2008.
34 VARIATION AND POPULATION GENETICS
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EBOOKS Variation and Population Genetics
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This book describes and analyzes genetic and environmental
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factors that cause variation in individuals and populations. Data
LIBRARY will be used to evaluate the processes by which variation is
Create your own generated in organisms and how variation affects natural selec-
Customized Content tion. Genetic factors include mutation, independent assort-
Variation and
Bundlethe more ment, crossing over, and recombination. Environmental factors
books you buy, include gradients and differences in abiotic conditions. Gen-
the greater your otype frequencies can be used to determine allele frequen-
discount! cies and this information can be used to determine whether a
population is evolving at a genetic locus. The Hardy-Weinberg Population
Genetics
equilibrium will be applied as a null model to make this deter-
THE CONTENT
mination. Non-Mendelian genetics can affect the evolution of
Energy Physics viruses and reassortment in viruses will be used to illustrate
Engineering another mechanism that generates variation in organisms and
Biotechnology how this mechanism relates to rapid evolution of viruses and
Biology the need for annual flu vaccines.
Mathematics
Christopher J. Paradiseis professor of biology and environ-
Chemistry
mental studies at Davidson College. He teaches introductory
biology, ecology, entomology, and topical seminars on ecotoxi-
THE TERMS cology and renewable natural resources. He also occasionally
Perpetual access leads a study abroad program in India. His research evaluates
for a one time fee anthropogenic factors that influence insect biodiversity at a
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A. Malcolm Campbellteaches biology at Davidson College,
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NC. He received national and international education awards:
Downloadable PDFs Genetics Society of America (2013); American Association for the
Free MARC records Advancement of Science (2012); and American Society for Cell
Biology (2006). He was the founding co-editor in chief of CBE Life
For further information,
Sciences Education; founding director of Genome Consortium
a free trial, or to order,
contact: for Active Teaching (GCAT); and member of the American Soci- Christopher J. Paradise
sales@momentumpress.net ety for Cell Biology governing council (20122014).
A. Malcolm Campbell