Romil Shah

Period 1
AP Biology

Chapter 18: Evolutionary Changes in Populations

I. Introduction
a. A population consists of all individuals of the same species that live in a particular place
at the same time. Individuals with a population vary in many recognizable characters.
b. Some of this variation is due to the environment, some is due to heredity.
c. Biologists study variation in a particular character by taking measurements of that
character in a population. By comparing the character and parents and offspring, it is
possible to estimate the amount of observed variation that is genetic, as represented by the
number of frequency, and kinds of alleles in a population.
d. An allele is one of two or more alternate forms of a gene. Alleles occupy corresponding
positions, or loci, or homologous chromosomes.
e. Population genetics is the study of genetic variability within a population and of the
forces that act on it.
f. There are 5 factors responsible for the evolutionary change: nonrandom mating,
mutation, genetic drift, gene flow, and natural selection.
II. Genotype, phenotype, and allele frequencies
a. Each population possesses a gene pool, which includes all the alleles are all loci present
in the population. Because diploid organisms have a maximum of two different alleles at
each genetic locus, a single individual typically has only a small fraction of the alleles
present in a population's gene pool.
b. The evolution of populations is best understood in terms of genotype, phenotype, and
allele frequencies.
c. Each genotype frequency is a portion of a particular genotype in the population.
d. Phenotype frequency is the portion of a particular phenotype in population. If each
genotype corresponds to a specific phenotype and genotype frequencies are the same.
e. And allele frequency is the proportion of a specific allele (that is, of A or a) in a
particular population. A mentioned earlier, each individual, being diploid, just 2 alleles at
each genetic locus.
III. The Hardy-Weinberg principle
Romil Shah
Period 1
AP Biology

a. The frequencies of alleles and genotypes do not change from generation to generation
unless enclosed by outside factors.
b. A population whose allele and genotype frequencies do not change from generation to
generation is said to be at genetic equilibrium.
c. However, if allele frequencies change over successive generations, evolution is a
occurring.
d. The explanation for the stability of successive generations in populations at genetic
equilibrium was provided independently by Godfrey Hardy, and English mathematician,
and Wilhelm Weinberg, a German physician, in 1908.
e. The resulting Hardy-Weinberg principle shows that if the population is large, the process
of inheritance does not by itself cause changes in allele frequencies.
f. It also explains what dominant alleles are not necessarily more common than recessive
ones.
g. When alleles are dominant and recessive, it is usually impossible to visually distinguish
heterozygous individuals from homozygous dominant individuals. The Hardy-Weinberg
principle lets us use the frequencies to calculate the expected genotype frequencies and
allele frequencies.
h. These values provide a basis of comparison with a population's allele frequencies in
succeeding generations.

A. Genetic equilibrium occurs if certain conditions are met

a. The Hardy-Weinberg principle of genetic equilibrium tells us what to expect when a

sexually reproducing population is not evolving. Their relative proportions of alleles
and genotypes in successive generations will always be the same, provided the
following conditions are met:
i. Radom mating. In unrestricted random mating, each individual in the
population has an equal chance of mating with any individual of the
opposite sex.
ii. No debt mutations. There must be no mutations that convert A to “a” or
vice versa.
Romil Shah
Period 1
AP Biology

iii. Large population size. Allele frequencies in a small population are more
likely to change by random fluctuations (that is, by, genetic drift) then are
allele frequencies in a large populations.
iv. No migration. There can be no exchange of alleles with other populations
that might have different allele frequencies.
v. No natural selection. If natural selection is occurring, certain phenotypes
(and their corresponding genotypes) are favored over others.
Consequently, the allele frequencies will change, the population will
evolve.

B. Human MN blood groups are a valuable illustration of the Hardy- Weinberg principle

a. Humans have dozens of antigens on the surfaces of their blood cells. (An antigen is a
molecule, usually a protein or a carbohydrate, that is recognized as a foreign by cells or
another organism’s immune system.)
b. One group of antigens, designated the MN blood group, stimulates the production of
antibodies when injected into rabbits or guinea pigs.
c. However, humans do not produce antibodies for M and N, so the MN blood group is not
medically important, for example, when giving blood transfusions.
d. The MN blood group is of interest to population geneticists because alleles for the MN
blood group, usually designated M and N, are codominant (genotype MN produces
antigen M only, genotype NM produces antigen N only, and the heterozygous
genotype MN produces both antigens).
e. The MN characteristic is not subject to natural selection and that it does not produce a
visible phenotype that may affect random mating (Genetic Equilibrium)
IV. Microevolution
a. The degree of departure between the observed allele or genotype frequencies and those
expected by the Hardy- Weinberg principle indicates the amount of evolutionary change.
This type of evolution−generation-to-generation changes in allele or genotype frequencies
within a population−is sometimes referred to as microevolution, because it often involves
relatively small or minor changes, usually over two generations. Changes in the allele
Romil Shah
Period 1
AP Biology

frequencies of a population result from five micro-evolutionary processes: nonrandom

mating, mutation, genetic drift, gene flow, and natural selection. These micro-evolutionary
processes are the opposite of the conditions that must be met if a population is in genetic
equilibrium. When one or more of these processes act on a population, allele or genotype
frequencies change from one generation to the next.

A. Nonrandom mating changes genotype frequencies

a. When individuals select mates on the basis of phenotype (thereby selecting the
corresponding genotype), they bring about evolutionary change in the population. Two
examples of nonrandom mating are interbreeding and assertive mating.
b. The mating of genetically similar individuals that are more closely related than if they
had been chosen at random from the entire population is known as inbreeding.
Although inbreeding does not change the overall allele frequency, the frequency of
homozygous genotypes increases with each successive generation of inbreeding. The
most extreme example of inbreeding is self-fertilization, which is particularly common
in certain plants.
c. Inbreeding does not appear to be detrimental in some populations, but in others it causes
inbreeding depression, in which inbred individuals have lower fitness than those not
inbred
d. Fitness is the relative ability of a given genotype to make a genetic contribution to
subsequent generations; fitness is usually measured as the average number of surviving
offspring of one genotype compared to the average number of surviving offspring of
competing genotypes. Inbreeding depression, as evidenced by fertility declines and
high juvenile mortality, is thought to be caused by the expression of a full recessive
alleles as homozygosity increases with inbreeding
e. Assortative mating, in which individuals select mates based on their phenotype, is
another example of nonrandom mating.
f. Such selection of mates with the same phenotype is known as positive assortative mating
(as opposed to less common phenomenon, negative assortative mating, in which mates
with similar phenotypes are selected)
Romil Shah
Period 1
AP Biology

g. Positive assortative mating is practiced by many human societies, in which men and
women tend to marry individuals like themselves in such characteristics such as height
or intelligence. Like inbreeding, assortative mating usually increases homozygosity at
the expense of heterozygosity in the population and is not change the overall allele
frequencies in the population however, Assortative mating changes genotype
frequencies only at the loci involved in mate choice, whereas inbreeding affects
genotype frequencies in the entire genome

B. Mutation increases variation within a population

a. Variation is introduced into a population through mutation, which is the unpredictable

change in the deoxyribonucleic acid. Mutations, which of the source of all new alleles,
result from (1) a change in the nucleotide base pairs of gene, (2) a rearrangement of
genes within chromosomes so that their interactions produce different effects, or (3) a
change in the chromosome structure. Mutations occur unpredictably and
spontaneously. The rate of mutation appears relatively constant for a particular of
locus, but may vary several orders of magnitude among genes within a certain species
and among different species.
b. Not all mutations passed from one generation to the next. Those occurring in somatic
(body) cells are not inherited. When an individual with such a mutation dies, mutation
is lost. So mutations, however, occur in reproductive cells. These mutations may or
may not overtly affected offspring, because most of the DNA cell is "silent" and is not
code for specific polypeptides or proteins that are responsible for physical
characteristics. Even if the mutation occurs in the DNA that codes for a polypeptide, it
may still have little effect on the structure or function of that polypeptide.
c. However, when a polypeptide is sufficiently altered to change its function, the mutation
is usually harmful. By acting against seriously abnormal phenotypes, natural selection
eliminates or reduces to low frequencies the most harmful mutations.
d. Mutations do not determine the direction of evolutionary change.
e. The production of new mutations simply increases the genetic variability that is acting
on natural selection and, therefore, increases the potential for new adaptations.
Romil Shah
Period 1
AP Biology

f. Mutation itself causes small deviations allele frequencies from those predicted by the
Hardy-Weinberg principle. Although allele frequencies may be changed by mutation,
these changes are typically several orders of magnitude smaller than changes caused by
other evolutionary forces, such as genetic drift.

C. In genetic drift, random events change allele frequencies

a. The size of the population has important effects on allele frequencies because random
events, or chance, tend to cause the changes of the relatively greater magnitude in a
small population. If the population consists of only a few individuals, an allele present
at a low frequency in the population could be completely lost by chance.
b. The production of rant illusionary changes in a small breeding populations is known as
genetic drift. Genetic drift results in changes in allele frequencies in a population from
one generation to another. One allele may be eliminated from the population purely by
chance, regardless of whether the allele is beneficial, harmful, or of no particular
advantage or disadvantage. Thus, genetic drift decreases genetic variation within a
population, although it tends to increase genetic differences among different
populations.

C. When bottlenecks occur, genetic drift becomes a major evolutionary force

a. Because of fluctuations in the environment, such as depletion in food supply or an

outbreak of disease, a population may rapidly and markedly decreased from time to
time. The population is said to go through a bottleneck during which genetic drift can
occur in the small population of survivors. As the population again increases in size,
many allele frequencies may be quite different from those in the population preceding
the decline.

D. The founder in effect occurs when a few "founders" establish a new colony

b. When one or a few individuals from a large population establish, or how found, a
colony, they bring with them only a small fraction of the genetic variation present in
the original population.
Romil Shah
Period 1
AP Biology

c. Typically, the allele frequencies in the newly founded population are quite different
from those of the parent population. The genetic drift that results when a small number
of individuals from a large population found a new colony is called the founder effect.

E. Gene flows generally increases variation within a population

d. Individuals of a species tend to be distributed in local populations that are genetically

isolated to some degree from another population.
e. The migration of breeding individuals between populations causes a corresponding
movement of alleles, or gene flow, that has significant evolutionary consequences. As
alleles flow from one population to another, they usually increase the amount of
genetic variability within the recipient population. If sufficient gene flow occurs
between two populations, they become more similar genetically. Because gene flow
reduces the amount of variation between two populations, it tends to counteract the
defects of natural selection and genetic drift, both of which often cause populations to
become increasingly distinct.
f. If migration by members of the population is considerable, and if populations differ in
their allele frequencies, then significant genetic changes occur in local populations.

F. Natural selection changes allele frequencies in a way that increases adaptation

a. Natural selection is the mechanism of evolution first proposed by Darwin in which

members of a population that are more successfully adapted to the environment are
more likely to survive and reproduce. Over successive generations, the proportion of
favorable alleles increases in the population in contrast with other micro-evolutionary
processes, natural selection leads to add active evolutionary change natural selection
not only explains why organisms are well adapted to the environments in which they
live with but also helps account for the remarkable diversity in life.
b. Natural selection is the differential reproduction of individuals with different traits, or
phenotypes, in response to the environment. Natural selection preserves individuals
with favorable phenotypes and eliminates the loose with unfavorable phenotypes.
Individuals that survive and to produce fertile offspring have a selective advantage.
Romil Shah
Period 1
AP Biology

c. The mechanism of natural selection does not develop a "perfect" organism. Rather, it
weeds out those individuals whose phenotypes are less adapted to environmental
challenges, while allowing better adapted individuals to survive and passed their alleles
to their offspring.
V. Natural selection operates on an organism’s phenotype
a. Natural selection does not act directly on an organism's genotype.
b. The phenotype represents an interaction between the environment and all the alleles in
the organism’s genotype.
c. Much more common is the interaction of alleles of several different loci for the
expression of a single phenotype.
d. When characters (characteristics) are under polygenic control (as in human height), a
range of phenotypes occurs, with most of the population located in the median range and
fewer at either extreme.
e. Three kinds of selection cause changes in the normal distribution of phenotypes in a
population: stabilizing, directional, and disruptive selection.
f. Stabilizing selection (360)
i. The process of natural selection associated with a population well adapted
to its environment is known as stabilizing selection.
ii. Stabilizing selection selects against phenotype extremes.
iii. Although stabilizing selection decreases the amount of variation in a
population, variation is rarely eliminated by this process, because other
micro-evolutionary processes against a decrease in variation.
g. Directional selection
i. In the environment changes over time, directional selection may favor
phenotypes and one of the extremes of the normal distribution. Over
successive generations, one phenotype gradually replaces another.
ii. Directional selection only occurs, if alleles favored under the new
circumstances are already present in the population.
h. Disruptive selection
Romil Shah
Period 1
AP Biology

i. Sometimes extreme changes in the environment may favor two or more

different phenotypes at the expense of the mean. More than one
phenotype may be favored in the new environment.
ii. Disruptive selection is a special type of directional selection in which
there is a trend in several directions rather than just one.
iii. It results in a divergence, or splitting apart, of distant groups of individuals
within a population.
iv. Natural selection induces changes in the types in the frequencies of alleles
populations only if there is pre-existing inherited variation. Genetic
variation is the raw material for evolution change, because it provides the
diversity on which natural selection acts.
VI. Genetic variation in populations
a. Populations contain abundant genetic variation that was originally introduced by
mutation. Sexual reproduction, with associated crossing over, independent assortment of
chromosomes during meiosis, and random union of gametes, also contributes to genetic
variation.
b. Genetic polymorphism exists among alleles and the proteins for which they code
c. one way of evaluating genetic variation and a population is to examine genetic
polymorphism, which is the presence in a population of two or more alleles with a given
locus. Genetic polymorphism is extensive populations, although many of the alleles are
present at low frequencies.
d. One way biologists estimate the total amount of genetic polymorphism is in populations
is by comparing the different forms of a particular protein. Each for consists of a slightly
different amino acid sequence that is coded for by a different allele.
e. Slight variations in amino acid sequences in the different forms of a particular enzyme
causes each migrate at a different rate, which can be detected using special stains or
radioactive labels. Determining the sequence of nucleotides in DNA from individuals in a
population provides a direct estimate of genetic polymorphism.
i. Balance polymorphism exists for long periods
Romil Shah
Period 1
AP Biology

f. Balanced polymorphism is a special type of genetic polymorphism in which two or more

alleles present a population over many generations as a result of natural selection.
Heterozygote advantage and frequency-dependent selection are mechanisms that preserve
balanced polymorphism.
i. Genetic variation may be maintained by heterozygote advantage
g. Natural selection sometimes helps maintain genetic diversity in a population, including
alleles that are unfavorable in the homozygous state.
h. This phenomenon, known as heterozygote advantage, is demonstrated in humans by the
selective advantage of heterozygous carriers of the sickle cell allele.
i. Genetic variation may be maintained by frequency-dependent selection.
i. In cases of frequency, dependent selection in the fitness of a particular phenotype
depends on how frequently it appears in the population. Often a phenotype has a greater
selective value when rare than when common in the population.
j. Such phenotypes lose their selective advantage as they become more common.
k. Frequency-dependent selection often acts to maintain genetic variation in populations of
prey species. In this case, the creditor catches and consumes the more common phenotype
but may ignore the rarer phenotypes. Consequently, the less common phenotype produces
more offspring and therefore makes it greater relative contribution to the next generation.
i. Neutral variation may give no selective advantage or disadvantage
l. Some of the genetic variation observed in a population make for no apparent selective
advantage or disadvantage in a particular environment.
m. Variation that does not alter the ability of an individual to survive and reproduce and is,
therefore, not adaptive is called neutral variation.
n. Not extent of neutral variation in organisms is difficult to determine. It is relatively easy
to demonstrate that an allele is beneficial or harmful it its effect is observable.
o. But the variation in alleles that involves only slight differences in the proteins they code
for may or may not be neutral. These alleles may be influencing the organism in subtle
ways that are difficult to measure or assess. Also, an allele that is neutral in one
environment may be beneficial or harmful in another.
Romil Shah
Period 1
AP Biology

i. Populations in different geographic areas often exhibited genetic

adaptations to local environment.
p. In addition to the genetic variation among individuals within a population, genetic
differences often exists among different populations within the same species, a
phenomenon known as geographic variation. One type of geographic variation is a cline,
which is a gradual change in a species’ phenotype and genotype frequencies through a
series of geographically separate populations as a result of an environmental gradient. A
cline exhibits in variation in the expression of such attributes as color, size, shape,
physiology, or behavior. Clines are common among species with continuous ranges over
larger geographic areas.

q. The genetic basis of these clinical differences was experimentally demonstrated in a set
of classical experiments in which series of population from different geographic areas were
grown in the same environment.