492 Chapter 19 | The Evolution of Populations

19.1 | Population Evolution

By the end of this section, you will be able to:
• Define population genetics and describe how population genetics is used in the study of the evolution of
populations
• Define the Hardy-Weinberg principle and discuss its importance

The mechanisms of inheritance, or genetics, were not understood at the time Charles Darwin and Alfred Russel Wallace
were developing their idea of natural selection. This lack of understanding was a stumbling block to understanding many
aspects of evolution. In fact, the predominant (and incorrect) genetic theory of the time, blending inheritance, made it
difficult to understand how natural selection might operate. Darwin and Wallace were unaware of the genetics work by
Austrian monk Gregor Mendel, which was published in 1866, not long after publication of Darwin's book, On the Origin
of Species. Mendel’s work was rediscovered in the early twentieth century at which time geneticists were rapidly coming
to an understanding of the basics of inheritance. Initially, the newly discovered particulate nature of genes made it difficult
for biologists to understand how gradual evolution could occur. But over the next few decades genetics and evolution were
integrated in what became known as the modern synthesis—the coherent understanding of the relationship between natural
selection and genetics that took shape by the 1940s and is generally accepted today. In sum, the modern synthesis describes
how evolutionary processes, such as natural selection, can affect a population’s genetic makeup, and, in turn, how this can
result in the gradual evolution of populations and species. The theory also connects this change of a population over time,
called microevolution, with the processes that gave rise to new species and higher taxonomic groups with widely divergent
characters, called macroevolution.

Evolution and Flu Vaccines

Every fall, the media starts reporting on flu vaccinations and potential outbreaks. Scientists, health experts,
and institutions determine recommendations for different parts of the population, predict optimal production
and inoculation schedules, create vaccines, and set up clinics to provide inoculations. You may think of the
annual flu shot as a lot of media hype, an important health protection, or just a briefly uncomfortable prick in
your arm. But do you think of it in terms of evolution?
The media hype of annual flu shots is scientifically grounded in our understanding of evolution. Each year,
scientists across the globe strive to predict the flu strains that they anticipate being most widespread and
harmful in the coming year. This knowledge is based in how flu strains have evolved over time and over
the past few flu seasons. Scientists then work to create the most effective vaccine to combat those selected
strains. Hundreds of millions of doses are produced in a short period in order to provide vaccinations to key
populations at the optimal time.
Because viruses, like the flu, evolve very quickly (especially in evolutionary time), this poses quite a
challenge. Viruses mutate and replicate at a fast rate, so the vaccine developed to protect against last year’s
flu strain may not provide the protection needed against the coming year’s strain. Evolution of these viruses
means continued adaptions to ensure survival, including adaptations to survive previous vaccines.

Population Genetics
Recall that a gene for a particular character may have several alleles, or variants, that code for different traits associated
with that character. For example, in the ABO blood type system in humans, three alleles determine the particular blood-type
protein on the surface of red blood cells. Each individual in a population of diploid organisms can only carry two alleles
for a particular gene, but more than two may be present in the individuals that make up the population. Mendel followed
alleles as they were inherited from parent to offspring. In the early twentieth century, biologists in a field of study known
as population genetics began to study how selective forces change a population through changes in allele and genotypic
frequencies.

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Chapter 19 | The Evolution of Populations 493

The allele frequency (or gene frequency) is the rate at which a specific allele appears within a population. Until now
we have discussed evolution as a change in the characteristics of a population of organisms, but behind that phenotypic
change is genetic change. In population genetics, the term evolution is defined as a change in the frequency of an allele in
a population. Using the ABO blood type system as an example, the frequency of one of the alleles, IA, is the number of
[1]
copies of that allele divided by all the copies of the ABO gene in the population. For example, a study in Jordan found a
frequency of IA to be 26.1 percent. The IB and I0 alleles made up 13.4 percent and 60.5 percent of the alleles respectively,
and all of the frequencies added up to 100 percent. A change in this frequency over time would constitute evolution in the
population.
The allele frequency within a given population can change depending on environmental factors; therefore, certain alleles
become more widespread than others during the process of natural selection. Natural selection can alter the population’s
genetic makeup; for example, if a given allele confers a phenotype that allows an individual to better survive or have more
offspring. Because many of those offspring will also carry the beneficial allele, and often the corresponding phenotype,
they will have more offspring of their own that also carry the allele, thus, perpetuating the cycle. Over time, the allele will
spread throughout the population. Some alleles will quickly become fixed in this way, meaning that every individual of the
population will carry the allele, while detrimental mutations may be swiftly eliminated if derived from a dominant allele
from the gene pool. The gene pool is the sum of all the alleles in a population.
Sometimes, allele frequencies within a population change randomly with no advantage to the population over existing allele
frequencies. This phenomenon is called genetic drift. Natural selection and genetic drift usually occur simultaneously in
populations and are not isolated events. It is hard to determine which process dominates because it is often nearly impossible
to determine the cause of change in allele frequencies at each occurrence. An event that initiates an allele frequency change
in an isolated part of the population, which is not typical of the original population, is called the founder effect. Natural
selection, random drift, and founder effects can lead to significant changes in the genome of a population.

Hardy-Weinberg Principle of Equilibrium

In the early twentieth century, English mathematician Godfrey Hardy and German physician Wilhelm Weinberg stated the
principle of equilibrium to describe the genetic makeup of a population. The theory, which later became known as the
Hardy-Weinberg principle of equilibrium, states that a population’s allele and genotype frequencies are inherently stable—
unless some kind of evolutionary force is acting upon the population, neither the allele nor the genotypic frequencies would
change. The Hardy-Weinberg principle assumes conditions with no mutations, migration, emigration, or selective pressure
for or against genotype, plus an infinite population; while no population can satisfy those conditions, the principle offers a
useful model against which to compare real population changes.
Working under this theory, population geneticists represent different alleles as different variables in their mathematical
models. The variable p, for example, often represents the frequency of a particular allele, say Y for the trait of yellow in
Mendel’s peas, while the variable q represents the frequency of y alleles that confer the color green. If these are the only
two possible alleles for a given locus in the population, p + q = 1. In other words, all the p alleles and all the q alleles make
up all of the alleles for that locus that are found in the population.
But what ultimately interests most biologists is not the frequencies of different alleles, but the frequencies of the resulting
genotypes, known as the population’s genetic structure, from which scientists can surmise the distribution of phenotypes.
If the phenotype is observed, only the genotype of the homozygous recessive alleles can be known; the calculations provide
an estimate of the remaining genotypes. Since each individual carries two alleles per gene, if the allele frequencies (p and q)
are known, predicting the frequencies of these genotypes is a simple mathematical calculation to determine the probability
of getting these genotypes if two alleles are drawn at random from the gene pool. So in the above scenario, an individual
pea plant could be pp (YY), and thus produce yellow peas; pq (Yy), also yellow; or qq (yy), and thus producing green peas
(Figure 19.2). In other words, the frequency of pp individuals is simply p2; the frequency of pq individuals is 2pq; and the
frequency of qq individuals is q2. And, again, if p and q are the only two possible alleles for a given trait in the population,
these genotypes frequencies will sum to one: p2 + 2pq + q2 = 1.

1. Sahar S. Hanania, Dhia S. Hassawi, and Nidal M. Irshaid, “Allele Frequency and Molecular Genotypes of ABO Blood Group System in a Jordanian
Population,” Journal of Medical Sciences 7 (2007): 51-58, doi:10.3923/jms.2007.51.58.
494 Chapter 19 | The Evolution of Populations

Figure 19.2 When populations are in the Hardy-Weinberg equilibrium, the allelic frequency is stable from
generation to generation and the distribution of alleles can be determined from the Hardy-Weinberg equation. If
the allelic frequency measured in the field differs from the predicted value, scientists can make inferences about
what evolutionary forces are at play.

In plants, violet flower color (V) is dominant over white (v). If p = 0.8 and q = 0.2 in a population of 500
plants, how many individuals would you expect to be homozygous dominant (VV), heterozygous (Vv), and
homozygous recessive (vv)? How many plants would you expect to have violet flowers, and how many
would have white flowers?

In theory, if a population is at equilibrium—that is, there are no evolutionary forces acting upon it—generation after
generation would have the same gene pool and genetic structure, and these equations would all hold true all of the time.
Of course, even Hardy and Weinberg recognized that no natural population is immune to evolution. Populations in nature
are constantly changing in genetic makeup due to drift, mutation, possibly migration, and selection. As a result, the only
way to determine the exact distribution of phenotypes in a population is to go out and count them. But the Hardy-Weinberg
principle gives scientists a mathematical baseline of a non-evolving population to which they can compare evolving
populations and thereby infer what evolutionary forces might be at play. If the frequencies of alleles or genotypes deviate
from the value expected from the Hardy-Weinberg equation, then the population is evolving.

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Chapter 19 | The Evolution of Populations 495

Use this online calculator (http://openstaxcollege.org/l/hardy-weinberg) to determine the genetic structure of a

population.

19.2 | Population Genetics

By the end of this section, you will be able to:
• Describe the different types of variation in a population
• Explain why only heritable variation can be acted upon by natural selection
• Describe genetic drift and the bottleneck effect
• Explain how each evolutionary force can influence the allele frequencies of a population

Individuals of a population often display different phenotypes, or express different alleles of a particular gene, referred
to as polymorphisms. Populations with two or more variations of particular characteristics are called polymorphic. The
distribution of phenotypes among individuals, known as the population variation, is influenced by a number of factors,
including the population’s genetic structure and the environment (Figure 19.3). Understanding the sources of a phenotypic
variation in a population is important for determining how a population will evolve in response to different evolutionary
pressures.

Figure 19.3 The distribution of phenotypes in this litter of kittens illustrates population variation. (credit: Pieter Lanser)

Genetic Variance
Natural selection and some of the other evolutionary forces can only act on heritable traits, namely an organism’s genetic
code. Because alleles are passed from parent to offspring, those that confer beneficial traits or behaviors may be selected
for, while deleterious alleles may be selected against. Acquired traits, for the most part, are not heritable. For example, if an
athlete works out in the gym every day, building up muscle strength, the athlete’s offspring will not necessarily grow up to
be a body builder. If there is a genetic basis for the ability to run fast, on the other hand, this may be passed to a child.