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AP Biology
(Text from Biology, 6th Edition, by Campbell and Reece)

Terms and Concepts for Mechanisms of Evolution

In On the Origin of Species by Means
of Natural Selection, Charles
Darwin argued that the organisms now inhabiting the Earth descended
from ancestral species. He proposed natural selection as a mechanism
for evolution, stating that a population of organisms can chang
change over
generations if individuals with certain heritable traits leave more
offspring than others. Natural selection caused evolutionary
characteristi that enhance
adaptation, a prevalence of inherited characteristics
organisms survival and reproduction.
Darwinss field research took place on the voyage of the HMS Beagle
around South America. He was able to observe many different species
of plants and animals, diverse environments, and study South
Sout American
fossils. An especially important area was the Galpagos
Galpagos Islands, where Darwin learned that animal
species on the islands resembled those living on the South American mainland.
Descent with modification,
Darwinss idea that all organisms were
modification, later known as evolution, was Darwin
related through descent from an unknown ancestor. As the descendants came to live in different
habitats over the years, they accumulated adaptations that fit them to specific ways of life. His main
ideas can be summarized:
Natural selection is differential success in reproduction.
Natural selection
lection occurs thr
through an interaction between the environment and the variability
inherent among the individual organisms making up a population.
The product of natural selection is the adaptation of population of organisms to their
Darwin knew that organisms population size would increase exponentially unless certain factors
limited survival. In each generation, environmental factors selected some traits over others. These
favored traits allowed organisms to produce more offspring than those without such traits, resulting in
the frequency of favored traits in a population, otherwise known as
a evolution.
In contrast, artificial selection is the breeding of domesticated plants and animals. Humans have
modified other species over many generations by selecting individuals with the desired traits as
breeding stock. For example, all dogs are th
thee same species, but artificial selection has resulted in the
numerous breeds of dogs present today.
It is important to remember that natural selection can only amplify or diminish heritable variations. An
organism may become modified during its lifetime
lifetime,, but such characteristics will not be passed on to
the next generation. Additionally, individuals do not evolve a population is the smallest unit that
can evolve.
AP Biology
(Text from Biology, 6th Edition, by Campbell and Reece)

Terms and Concepts for Mechanisms of Evolution


Examples of Natural Selection

A prevalent example of natural selection is the evolution of insecticide resistance in hundreds of insect
species. Widespread use of pesticides has resulted in populations that are unaffected. Initially, a
certain pesticide may kill 99% of the insects, with subsequent sprayings being less and less effective.
The survivors will pass their resistance gene onto future generations, resulting in a population that has
adapted to change in its environment.
Extensive use of antibiotics has had a similar effect on certain diseases caused by bacteria.

Fossils are relics or impressions of organisms from the past, preserved in rock. Most fossils are found
in sedimentary rocks formed from the sand and mud that settle to the bottom of seas, lakes, and
marshes. Fossils within layers show that a succession of organisms has populated Earth throughout
time. Georges Cuvier developed the study of fossils, known as paleontology.
paleontology He recognized that
extinction occurred often throughout the history of life and that each stratum of fossils was
characterized by fossil species, with older stratum containing fossils increasingly dissimilar to modern
life. However, he was an opponent of evolution and advocated catastrophism,
catastrophism the idea that each
boundary corresponded in time to a catastrophe that destroyed many of the species living there at that
Later, towards the end of the 18th century, Jean Lamarck placed fossils in an evolutionary context.
Lamarck compared current species with fossil forms and saw several lines of descent. However, he
thought that acquired characteristics could be inherited for example, the long neck of the giraffe was
obtained through successive generations as giraffes stretching their neck in several generations
resulted in the final length. There is no evidence that acquired characteristics can be inherited. Even
so, his theory deserves credit for its claim that evolution is the best explanation for the fossil record
and diversity of life, its recognition of Earths age, and its emphasis of adaptation to the environment
as a primary product of evolution.

Similarity in characteristics resulting from common ancestry is known as
homology Anatomical homologies are anatomical similarities between
species grouped in the same taxonomic category. For example, mammals
have similar forelimb structure, despite differences in function. The
forelegs, flippers, wings, and arms of different mammals are variations on a
common structural theme. Such signs of evolution are called homologous
structures These structures include vestigial organs,
organs structures of
AP Biology
(Text from Biology, 6th Edition, by Campbell and Reece)

Terms and Concepts for Mechanisms of Evolution

marginal importance to the organism. Vestigial organs are remnants of structures that had important
functions in ancestors. For example, whales have small pelvic bones that are no longer in use.
Humans have a remnant of the tailbone (coccyx).
Embryological homologies homologies can be seen during embryonic development. All vertebrate
embryos have structures called pharyngeal pouches in their throat regions at some stage in
Molecular homologies all species of life use the same genetic machinery of DNA and RNA, with an
essentially universal genetic code.
Some homologies are shared by all life because they date back to the ancestral past. Homologies that
evolved more recently are shared by smaller groups. For example, the vertebrate branch contains the
same basic five-digit limb structure illustrated for mammals. Evolutionary relationships among species
are also documented in similarities between DNA and proteins.
In contrast, analogous structures are similar due to their function, but not their structure. These
structures evolved independently, such as the wings of birds, bats, and insects. They are a result of
convergent evolution.
evolution Homologies are a result of divergent evolution.

The geographic distribution of species, biogeography,
biogeography first suggested evolution to Darwin. Species
tend to be more closely related to other species from the same area than to other species with the
same way of life but located in different areas. Islands generally have endemic populations, meaning
plants and animals found nowhere else in the world. However, these species are closely related to
those from the nearest mainland or neighboring island.
The HardyHardy-Weinberg theorem states that the frequency of alleles and genotypes in a populations
gene pool remain constant over the generations unless acted upon by agents other than Mendelian
segregation and recombination of alleles. This is analogous to shuffling a deck of cards: no matter how
many times the deck is reshuffled, the deck itself remains the same. A populations gene pool will not
change over time if the only factors influencing it are meiosis and random fertilization. The HardyHardyWeinberg equation,
equation p + 2pq + q = 1, is used to find frequencies of alleles and genotypes. This
mechanism of inheritance preserves the genetic variation that is the substrate for natural selection.
A population in Hardy-Weinberg equilibrium must satisfy five conditions:
1. It must be very large. Small populations can be influenced by genetic drift, which is chance
fluctuation in the gene pool.
AP Biology
(Text from Biology, 6th Edition, by Campbell and Reece)

Terms and Concepts for Mechanisms of Evolution

2. There can be no migration. Gene flow, the transfer of alleles between populations due to
movement of individuals or gametes, can increase or decrease genotypic frequencies.
3. No net mutations can take place. This will alter the gene pool.
4. There must be random mating. If individuals pick mates with certain genotypes, then random
mixing of gametes does not occur.
5. There can be no natural selection. Differential survival and reproductive success of genotypes
will alter their frequencies.
Deviation from Hardy-Weinberg equilibrium usually results in evolution.
Microevolution is a generation-to-generation change in a populations frequencies of alleles. Such
changes can be tracked by using the Hardy-Weinberg equation. Microevolution occurs even if the
frequencies of alleles are changing for only one genetic locus. The causes of microevolution are genetic
drift, natural selection, gene flow, and mutation. The two major factors are genetic drift and natural

Genetic Drift
Genetic drift is change in a populations allele frequencies due to chance. Populations can be shrunk
down to a small size, in which genetic drift may have significant impacts.
In the bottleneck effect,
effect disasters drastically reduce the size of a population. The
small surviving population may not provide the same representation of alleles as the
original population did some alleles will be overrepresented, others
underrepresented, and others may even be eliminated. Genetic drift may continue
to change the gene pool until the population is large enough for sampling errors to
be less significant. Bottlenecking usually reduces the overall genetic variability in a
population. For example, cheetahs, due to overhunting, have had their numbers
reduced to only three small populations in the wild. Their genetic variability is close
to that of inbred laboratory mice.
Genetic drift can also occur when a few individuals from a larger population colonize an isolated area.
The smaller the sample size, the less the genetic makeup of the colonists will represent the gene pool
of the larger population they left. Genetic drift in a new colony is known as the founder effect.
effect An
example is the relatively high frequency of inherited disorders in human populations begun by a small
number of colonists.

Natural Selection
AP Biology
(Text from Biology, 6th Edition, by Campbell and Reece)

Terms and Concepts for Mechanisms of Evolution

Natural selection causes alleles being passed to the next generation in numbers disproportion to the
relative frequencies in the present generation. Because of differences between individuals in a
population, some may have a better chance of survival and ability to reproduce than others. For
example, a light-colored rabbit in a snowy environment may have a better chance of survival than a
dark-colored rabbit. Out of the various agents that cause microevolution, selection is the one likely to
adapt a population to its environment. It accumulates and maintains favorable genotypes in a

Gene Flow
Gene flow is genetic exchange due to the migration of fertile individuals or gametes between
populations. This reduces differences between populations and can combine neighboring populations
into a single population with a common gene pool.

Variation within Populations

Quantitative characters are those that vary along a continuum, such as height. Discrete characters
can be classified on an either-or basis, usually because they are determined by a single gene locus with
different alleles that affect distinct phenotypes.
When two or more forms of a discrete character are represented in a population, the different forms are
called morphs.. A population is polymorphic for a character if two or more distinct morphs are each
represented in high enough frequencies to be readily noticeable. Polymorphism does not apply to
quantitative characters.
Genetic variation can be measured at the level of whole genes and at the molecular level of DNA.
Gene diversity is the average percent of loci that are heterozygous. For example, the genome of a fruit
fly has 13,000 loci, and the flies, on average, are heterozygous at about 14% of their loci. Population
geneticists measure nucleotide diversity by comparing the nucleotide sequences of DNA samples
from two individuals and then pooling the data from other comparisons of two individuals. For
example, nucleotide diversity in humans is 0.1%, meaning that two humans from the same population
will have the same nucleotide at 999 out of every 1,000 nucleotide sites in their DNA.

Variation between Populations

Most species show geographic variation,
variation differences in gene pools between populations or subgroups
of populations. Since environmental factors are likely to differ from one place to another, natural
selection can contribute to geographic variation. One type of geographic variation is a cline,
cline a graded
change in some trait along a geographic axis. A gradation in some environmental variable may produce
AP Biology
(Text from Biology, 6th Edition, by Campbell and Reece)

Terms and Concepts for Mechanisms of Evolution

a cline. For example, average body size of many North American species of birds and mammals
increase gradually with increasing latitude.

New alleles originate only by mutation, or change in the nucleotide sequence of DNA. A mutation
affecting any gene locus is a rare accident. Most mutations occur in somatic cells and are not passed
on to additional generations. Random chance determines whether a mutation will be beneficial to an
organism, how it will alter a gene, and where it will strike. It is only on rare occasions that mutant
allele will enhance reproductive success of an individual and assist in survival. This is more likely in a
changing environment. In organisms with very short generation spans, such as bacteria, mutations
generate genetic variation at a rapid pace. On the other hand, plants and animals, with a longer
generation-to-generation time scale, sexual recombination is the main factor behind genetic variation.

Sexual Recombination
Through meiosis, homologous chromosomes from each parent will trade genes by crossing over, and
then the homologous chromosomes will be segregated into separate gametes. Each zygote will have a
unique assortment of alleles due to the random union of a sperm and an ovum.

Recessive alleles that are less favorable than their dominant counterparts, or even harmful in the
present environment, can persist in a population through heterozygous individuals. This variation is
exposed to selection only when both parents carry the same recessive allele and combine two copies in
one zygote. Heterozygote protection maintains a huge pool of alleles that may not be suitable for
present conditions, but could bring new benefits when the environment changes.

Balanced Polymorphism
The ability of natural selection to maintain stable frequencies of two or more phenotypic forms in a
population is called balanced polymorphism
olymorphism Heterozygote advantage is one mechanism by which
natural selection preserves variation. If individuals who are heterozygous at a particular locus have a
greater chance of survival and reproductive success than homozygotes, then two or more alleles will be
maintained at that locus by natural selection. An example of heterozygote advantage is seen in those
who are heterozygous for sickle-cell anemia while they do not have the disease, they are resistant to
malaria. The environment favors them over those that are homozygous dominant, who are susceptible
to malaria, and those who are homozygous recessive, who have sickle-cell disease.
AP Biology
(Text from Biology, 6th Edition, by Campbell and Reece)

Terms and Concepts for Mechanisms of Evolution

Another mechanism promoting balanced polymorphism is frequencyfrequency- dependent selection,
selection in which
survival and reproduction of any one morph declines if that phenotypic form becomes too common in
the parasite. Relationships between parasites and hosts may involve frequency-dependent selection.

Neutral Variation
Some genetic variations observed in populations seem to have little, if any, effect on reproductive
success. Diversity of human fingerprints is an example of neutral variation,
variation which seems to confer no
selective advantage for some individuals for other. There is no consensus as to how much genetic
variation is neutral or if any variation can be considered truly neutral. Variation that appears neutral
may influence survival and reproductive success in ways that are difficult to measure.
Darwinian fitness is the contribution an individual makes to the gene pool of the next generation
relative to the contributions of other individuals. Population geneticists define relative fitness as the
contribution of a genotype to the next generation compared to the contributions of alternative
genotypes for the same locus. Survival alone does not guarantee reproductive success. Sterile plants
or animals have a relative fitness of 0. Of course, survival is a prerequisite for reproducing, and
longevity increases fitness if it results in certain individuals leaving more descendants than other
individuals have. Also, an individual that matures quickly and becomes fertile at an early age may have
greater reproductive potential than individuals that live longer but mature late. Many factors that
affect survival and fertility determine an individuals evolutionary fitness.
It is important to remember that a whole organism is subject to natural selection. Relative fitness of
an allele depends on the entire genetic context in which it works. Alleles that contribute nothing to an
organisms success, or even those that may be slightly maladaptive, may be perpetuated because they
are present in individuals whose overall fitness is high.
Directional selection is common during periods of environmental change. It shifts the frequency
curve in one direction or the other by favoring initially relatively rare individuals. Diversifying
selection occurs when environmental conditions favor individuals on both extremes of a phenotypic
range over intermediate phenotypes it results in distinctly different phenotypes. Stabilizing
selection favors more common intermediate variants, reducing variation and stabilizes for a particular
phenotypic character.
AP Biology
(Text from Biology, 6th Edition, by Campbell and Reece)

Terms and Concepts for Mechanisms of Evolution

Secondary sexual characters, those not directly associated with reproduction, cause distinctions in
appearance called sexual dimorphism.
dimorphism This can be manifested in a sixe difference or in features such
as colorful plumage of male birds, a male lions mane, or a male deers antlers. Sexual dimorphism is a
product of sexual selection. Intrasexual selection is a direct competition among individuals of one
sex (usually males in vertebrates) for mates of the opposite sex. Males can use secondary sexual
equipment to battle competitors. In intersexual selection,
selection also called mate choice, individuals of one
sex (usually females) are choosy in selecting mates of the other sex. Males with the most impressive
features are the most attractive to females. Secondary sexual characteristics that help a male gain a
mate increase reproductive success are propagated because every time a female chooses a mate based
on a certain appearance or behavior; she perpetuates the alleles that caused her to make that choice
and allows a male with a particular phenotype to perpetuate his alleles.

Macroevolution involves evolutionary change on a large scale, encompassing the origin of new
taxonomic groups, evolutionary trends, adaptive radiation, and mass extinction. Speciation is the key
process because any genus, family, or higher taxon originates with a new species that may be the first
member of a higher taxon.

The Biological Species Concept

The biological species concept defines a species as a population or group of populations whose
members have the potential to interbreed with one another in nature to produce viable, fertile
offspring, but who cannot produce viable, fertile offspring with members of other species. In other
words, a biological species is the largest set of populations in which genetic exchange is possible and
that is genetically isolated from other such populations. Reproductive isolation prevents populations
belonging to different species from interbreeding, even if their ranges overlap.

Prezygotic Barriers
Prezygotic barriers impede mating between species or hinder the fertilization of ova if members of
different species attempt to mate.
Habitat I solation Two species that live in different habitats within the same area may
encounter each other rarely, if at all.
Behavioral Isolation Special signals that attract mates or elaborate courtship behavior are
reproductive barriers among closely related animals. For example, eastern and western
meadowlarks remain distinct biological species because of differences in their songs.
AP Biology
(Text from Biology, 6th Edition, by Campbell and Reece)

Terms and Concepts for Mechanisms of Evolution

Temporal Isolation Two species that breed during different times of day, different seasons,
or different years cannot mix their gametes.
Mechanical Isolation Closely related species may fail to mate because they are
anatomically incompatible.
Gametic Isolation Even if gametes of different species meet, they rarely fuse to form a
zygote. For animals whose eggs are fertilized in the female reproductive tract, sperm of one
species may not survive in the environment of the female reproductive species. In cases of
external fertilization, sperm may not be able to penetrate eggs of another species.

Postzygotic Barriers
If a sperm cell from one species does fertilize an ovum of another species, then postzygotic barriers
usually prevent the hybrid zygote from developing into a viable, fertile adult.
Reduced Hybrid Vitality Genetic incompatibility between the two species may abort
development of the hybrid at some embryonic stage.
Reduced Hybrid Fertility Even if two species mate and produce viable hybrid offspring,
reproductive isolation is intact if the hybrids are completely or largely sterile.
Hybrid Breakdown In some cases, first-generation hybrids are viable and fertile, but these
hybrids cannot produce viable offspring if they mate with other hybrids or either parental
In allopatric speciation
ciation speciation takes place in populations with geographically separate ranges.
Gene flow is initially interrupted between two populations because they are separated in space. In
sympatric speciation,
speciation speciation takes place in geographically overlapping populations.

Allopatric Speciation
Several geologic processes can fragment a population into two or more isolated populations. For
example, a mountain range could emerge, a land bridge could form, or a large lake could separate into
several smaller ones. Geographic isolation and allopatric speciation can also occur if individuals
colonize a new, geographically remote area and become isolated from the parent population.
Likelihood of allopatric speciation increases with a small and isolated population.

Adaptive Radiation
Evolution of diversely adapted species from a common ancestor is called adaptive radiation.
This type of evolution occurs on island chains far enough apart for populations to develop in
isolation, but close enough for occasional dispersion events. For example, the many finch
AP Biology
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Terms and Concepts for Mechanisms of Evolution

species on the Galpagos Islands may have developed from one species of finch originating on
the mainland.

Evolution of Reproductive Barriers

It is important to understand that geographic isolation does not qualify as reproductive
isolation in the biological sense. Reproductive barriers relevant in the biological species
concept prevent interbreeding even in the absence of geographic isolation. Additionally,
speciation is not because of a drive to construct reproductive barriers around a population. In
most cases, such reproductive barriers are probably coincidental as a result of changes in gene

Sympatric Speciation
In a sympatric speciation, new species arise within the range of parent populations.

Polyploid Speciation in Plants

Some plant species have their origins in accidents during cell division that resulted in extra sets
of chromosomes a condition called polyploidy.
polyploidy An autopolyploid is an individual with more
than two chromosome sets, all derived from a single species. This tetraploid can now mate
with itself or with other tetraploids. However, such mutants cannot interbreed with diploid
plants of the original population.
Another type of polyploidy species, more common than autopolyploids, is an allopolyploid.
This is a polyploid species originating from the contribution of two different species to a
polyploid hybrid. Origin of an allopolyploid begins when two different species interbreed and
combine chromosomes. Allopolyploids are usually sterile because the haploid set of
chromosomes from one species cannot pair during meiosis with the haploid set from the other
species. While infertile, a hybrid can still propagate itself asexually. Through various
mechanisms, a sterile hybrid can be changed into a fertile polyploid.

Sympatric Speciation in Animals

Polyploid speciation is much less common in animals than in plants. However, other
mechanisms can lead to sympatric speciation in animals. For example, animals can become
reproductively isolated within the geographic range of a parent population if genetic factors
cause them to become dependent on resources not used by the parent population. For
example, different species of wasps pollinate a particular fig species. A genetic change that
caused wasps to select a different fig species would segregate mating individuals of this new
phenotype from the parent population.
AP Biology
(Text from Biology, 6th Edition, by Campbell and Reece)

Terms and Concepts for Mechanisms of Evolution

Another example is in the cichlid population of Lake Victoria there are nearly 200 species of
closely related fish belonging to the cichlid family. This subdivision of populations into groups
specialized for exploiting different food sources and other resources in the lake is one process
that probably contributed to adaptive radiation. In some cases, nonrandom mating, in which
females select males with a certain appearance, was probably a key factor in sympatric
speciation of the cichlids.


Advocates of punctuated equilibrium argue that species diverge in spurts of relatively rapid change,
instead of slowly and gradually. Most of the morphological modification takes place as they first bud
from parent species. They will then change only a little, even as they give rise to additional species. In
contrast, gradualism supports less abrupt changes.
Newer, more advanced structures can evolve from older, more simple structures that fulfilled the same
purpose. For example, the human eye is a refined optical organ constructed from multiple parts that
probably originated from more basic versions, such as clusters of photoreceptor cells. Evolution can
also arise by gradual refinement of existing structures for new purposes. Such structures that evolve in
one context but become co-opted for another function are called exaptations.
exaptations For example, the
hollow, lightweight bones of birds may have originated due to increased agility.
The shape of an organism depends partly on the relative growth rates of different parts during
development. This proportioning is called allometric growth.
growth Evolution of morphology that arises by a
modification in allometric growth is an example of heterochrony,
heterochrony which is evolutionary change in the
rate or timing of developmental events. Heterochrony can also change the timing of reproductive
development relative to development of somatic organs. If rate of reproductive development
accelerates compared to somatic development, the sexually mature stage of a species may retain body
features that were juvenile structures in an ancestral species. This is called paedomorphosis.
Heterochrony affects the evolution of morphology by altering the rates of development of various body
parts or by changing the timing of onset or completion of a particular parts development.
Macroevolution can also result from changes in genes that control the placement and special
organization of body parts. For example, genes called homeotic genes determine where basic features
will develop or how various parts will be arranged. Once class of homeotic genes called Hox genes
provide positional information in an animal embryo.
AP Biology
(Text from Biology, 6th Edition, by Campbell and Reece)

Terms and Concepts for Mechanisms of Evolution

Phylogeny is the evolutionary history of a species or group of related species. This has a basis in
continental drift, in which plate movements rearrange geography. About 250 million years ago, plate
movements brought land masses together into a supercontinent named Pangaea.
Pangaea Continental drift
also explains the current distribution of organisms.


The fossil record is the ordered array in which fossils appear within layers, or strata, of sedimentary
rocks that mark the passing of geologic time. Relative dating of fossils uses index fossils, fossils of
creatures that are widespread but present only in a specific time period, to determine the age of other
fossils. Absolute dating allows paleontologists to give an age in years. Radiometric dating,
dating the
measurement of certain radioactive isotopes in fossils or rocks, is the method most often used to
determine the ages of rocks and fossils on a scale of absolute time. Each radioactive isotope has a
fixed rate of decay. An isotopes halfhalf -life,
life the number of years it takes for half of the original sample
to decay, is unaffected by environmental variables. Thus, measuring the amount of an isotope left can
be used to date fossils.
Tracing phylogeny is one of the main goals of systematics
systematics, the study of biological diversity in an
evolutionary context.

Taxonomists assign to each species a two-part Latinized name, or binomial.
binomial The first part of a
binomial is the genus,
genus while the second part is the specific epithet,
epithet referring to one species within
the genus. A major objective of systematics is to group species into broader taxonomic categories.
Hieararchical classification goes from domains, to kingdoms, to phlyums, to classes, to orders, to
families, to genuses, and then to a species. The phylogenetic trees that systematists construct
reflect the hierarchical classification of taxonomic groups nested within more inclusive groups.

A phylogenetic diagram based on cladistics is called a cladogram.
cladogram It is a tree constructed from a series
of dichotomies, or two-way branch points. Each evolutionary branch in a cladogram is called a clade.
A clade consists of an ancestral species and all of its descendants such a group is said to be