Objectives Summarize the effects of the different types of natural selection on gene pools.

Relate changes in genetic equilibrium to mechanisms of speciation.

Explain the role of natural selection in convergent and divergent evolution.

Review Vocabulary gene: DNA segment that

controls protein production and the cell cycle (p.211)

New Vocabulary gene pool

allelic frequency genetic equilibrium genetic drift stabilizing selection directional selection disruptive selection speciation

geographic isolation reproductive isolation polyploid

gradualism

punctuated equilibrium adaptive radiation divergent evolution convergent evolution

404 THE THEORY OF EVOLUTIOi\'

Mechanisms of Evolution

Using Prior Knowledge You may recognize the birds shown here as meadowlarks. These birds range throughout much

of the United States. Meadowlarks look so similar that it's often difficult to tell them apart. Although they are closely related and occupy the same ranges in parts of the central United States, these different meadowlarks do not normally interbreed and are classified as distinct species.

Using your knowledge of birds and animal behavior, infer what prevents competition between two meadowlark species that occupy

the same area.

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I

Western meadowlarkSturnella neglecta

Eastern meadowlarkSturnella magna

Population Genetics and Evolution

When Charles Darwin developed his theory of natural selection in the 1800s, he did so without knowing about genes. Since Darwin's time, scientists have learned a great deal about genes and modified Darwin's ideas accordingly. At first, genetic information was used to explain the variation among individuals of a population. Then, studies of the complex behavior of genes in populations of plants and animals developed into the field of study called population genetics. The principles of today's modern theory of evolution are rooted in population genetics and other related fields of study and are expressed in genetic terms.

Populations, not individuals, evolve

Can individuals evolve? That is, can an organism respond to natural selection by acquiring or losing characteristics? Recall that genes determine most of an individual's features, such as tooth shape or flower color. If an organism has a feature-called a phenotype in genetic terms-that is poorly adapted to its environment, the organism may be unable to survive and reproduce. However, within its lifetime, it cannot evolve a new phenotype by natural selection in response to its environment.

First generation

Phenotype frequency

Allele frequency

RR RR RR' RR' RR RR' RR RR'
Second generation Phenotype Allele
frequency frequency
White = 0.125 R = 0.75
Pink = 0.25 R' = 0.25
Red = 0.625
RR RR' RR RR' RR R'R' RR RR White = 0 Pink = 0.5 Red = 0.5

R = 0.75 R' = 0.25

Rather, natural selection acts on the range of phenotypes in a population. Recall that a population consists of all the members of a species that live in an area. Each member has the genes that characterize the traits of the species, and these genes exist as pairs of alleles. Just as all of the individuals make up the population, all of the genes of the population's individuals make up the population's genes. Evolution occurs as a population's genes and their frequencies change over time.

How can a population's genes change over time? Picture all of the alleles of the population's genes as being together in a large pool called a gene pool. The percentage of any specific allele in the gene pool is called the allelic frequency. Scientists calculate the allelic frequency of an allele in the same way that a baseball player calculates a batting average. They refer to a population in which the frequency of alleles remains the same over generations as being in genetic equilibrium. In the Connection to Math at the end of the chapter, you can read about the

mathematical description of genetic equilibrium. You can study the effect of natural selection on allelic frequencies in the BioLab at the end of the chapter.

Look at the population of snapdragons shown in Figure 15.11. A pattern of heredity called incomplete dominance, which you learned about earlier, governs flower color in snapdragons. If you know the flower-color genotypes of the snapdragons in a population, you can calculate the allelic frequency for the flower-color alleles. The population of snapdragons is in genetic equilibrium when the frequency of its alleles for flower color is the same in all its generations.

Changes in genetic equilibrium

A population that is in genetic equilibrium is not evolving. Because allelic frequencies remain the same, phenotypes remain the same, too. Any factor that affects the genes in the gene pool can change allelic frequencies, disrupting a population's genetic equilibrium, which results in the process of evolution.

Figure 15.11 Incomplete dominance produces three phenotypes: red flowers (RR), white flowers (R'R'), and pink flowers (RR'). Although the phenotype frequencies of the generations vary, the allelic frequencies for the Rand R' alleles do not vary.

15.2 MECHANISMS OF EVOLUTION 405

Figure 15.12

Genetic drift can result in an increase of rare alleles in a small population. Notice the child has six fingers on each hand.

Figure 15.13

These swallowtail butterflies live in different areas of North America. Despite their slight variations, they can interbreed to produce fertile offspring.

You have learned that one mechanism for genetic change is mutation. Environmental factors, such as radiation or chemicals, cause many mutations, but other mutations occur by chance. Of the mutations that affect organisms, many are lethal, and the organisms do not survive. Thus, lethal mutations are quickly eliminated. However, occasionally, a mutation results in a useful variation, and the new gene becomes part of the population's gene pool by the process of natural selection.

Another mechanism that disrupts a population's genetic equilibrium is genetic drift-the alteration of allelic frequencies by chance events.

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406 TIIF THEORY OF FVOLVrIO;\,

Genetic drift can greatly affect small populations that include the descendants of a small number of organisms. This is because the genes of the original ancestors represent only a small fraction of the gene pool of the entire species and are the only genes available to pass on to offspring. The distinctive forms of life that Darwin found in the Galapagos Islands may have resulted from genetic drift.

Genetic drift has been observed in some small human populations that have become isolated due to reasons such as religious practices and belief systems. For example, in Lancaster County, Pennsylvania, there is an Amish population of about 12 000 people who have a unique lifestyle and marry other members of their community. By chance, at least one of the original 30 Amish settlers in this community carried a recessive allele that results in short arms and legs and extra fingers and toes in offspring, Figure 15.12. Because of the small gene pool, many individuals inherited the recessive allele over time. Today, the frequency of this allele among the Amish is high-1 in 14 rather than 1 in 1000 in the larger population of the United States.

Genetic equilibrium is also disrupted by the movement of individuals in and out of a population. The transport of genes by migrating individuals

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is called gene flow. When an individual leaves a population, its genes are lost from the gene pool. When individuals enter a population, their genes are added to the pool.

Mutation, genetic drift, and gene flow may significantly affect the evolution of small and isolated gene pools, such as those on islands. However, their effect is often insignificant in larger, less isolated gene pools. Natural selection is usually the most significant factor that causes changes in established gene pools-small or large.

Natural selection acts on variations

As you've learned, traits have variation, as shown in the butterflies pictured in Figure 15.13. Try measuring variations in MiniLab 15.2.

Recall that some variations increase or decrease an organism's chance of survival in an environment. These variations can be inherited and are controlled by alleles. Thus, the allelic frequencies in a population's gene pool will change over generations due to the natural selection of variations. There are three different types of natural selection that act on variation: stabilizing, directional, and disruptive.

are not evolving.

Collect Data

Detecting a Variation Pick almost any trait-height, eye color, leaf width, or seed size-and you can observe

how the trait varies in a population. Some variations are an advantage

to an organism and some are not.

Procedure

o Copy the data table shown here, but include

the lengths in millimeters (numbers 25 through 45) that are missing from this table.

Length in mm 20 21 22 23 24 - 46 47 48 49 50

Checks

My DataNumber of Shells

Class DataNumber of Shells

fJ Use a millimeter ruler to measure a peanut shell's length.

In the Checks row, check the length you measured.

D Repeat step 2 for 29 more shells.

D Count the checks under each length and enter the total in the row marked My Data.

D Use class totals to complete the row marked Class Data.

Analysis

1. Collect and Organize Data Was there variation among the lengths of peanut shells? Use class data to support your answer.

2. Draw Conclusions If larger peanut shells were a selective advantage, would this be stabilizing, directional, or disruptive selection? Explain your answer.

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15.2 MECH!\N)S,'>lS OF EVOLUTION 407

Figure 15.14

Different types of natural selection act over the range

of a trait's variation. The red, bell-shaped curve indicates a trait's variation in a population. The blue, bell-shaped curve indicates the effect of a natural selection.

Selection for -----# average size spiders

Normal variation

1l Stabilizing selection favors average individuals. This type of selection reduces variation in a population.

a Directional selection favors one of the extreme variations of a trait and can lead to the rapid evolution of a population.

Normal

Selection for

Selection for light limpets

t! Disruptive selection favors both extreme variations of a trait, resulting eventually in no intermediate forms of the trait and leading to the evolution of two new species.

408 THE THEORY OF EVOLUTION

Stabilizing selection is natural selection that favors average individuals in a population, as shown in Figure 15.14. Consider a population of spiders in which average size is a survival advantage. Predators in the area might easily see and capture spiders that are larger than average. However, small spiders may find it difficult to find food. Therefore, in this environment, average-sized spiders are more likely to survive-they have a selective advantage, or are "selected for."

Directional selection occurs when natural selection favors one of the extreme variations of a trait. For example, imagine a population of woodpeckers pecking holes in trees to feed on the insects living under the bark. Suppose that a species of insect that lives deep in tree tissues invades the trees in a woodpecker population's territory. Only woodpeckers with long beaks could feed on that insect. Therefore, the long-beaked woodpeckers in the population would have a selective advantage over woodpeckers with very short or average-sized beaks.

Finally, in disruptive selection, individuals with either extreme of a trait's variation are selected for. Consider, for example, a population of marine organisms called limpets. The shell color of limpets ranges from white, to tan, to dark brown. As adults, limpets live attached to rocks. On light-colored rocks, white-shelled limpets have an advantage because their bird predators cannot easily see them. On dark-colored rocks, darkcolored limpets have the advantage because they are camouflaged. On the other hand, birds easily see tancolored limpets on either the light or dark backgrounds. Disruptive selection tends to eliminate the intermediate phenotypes.

Natural selection can significantly alter the genetic equilibrium of a population's gene pool over time. Significant changes in the gene pool could lead to the evolution of a new species over time.

The Evolution of Species

You've just read about how natural processes such as mutation, genetic drift, gene flow, and natura] selection can change a population's gene pool over time. But how do the changes in the makeup of a gene pool result in the evolution of new species? Recall that a species is defined as a group of organisms that look alike and can interbreed to produce fertile offspring in nature. The evolution of new species, a process called speciation (spee shee AY shun), occurs when members of similar populations no longer interbreed to produce fertile offspring within their natural environment.

Physical barriers can prevent interbreeding

In nature, physical barriers can break large populations into smaller ones. Lava from volcanic eruptions can isolate populations. Sealevel changes along continental shelves can create islands. The water that surrounds an island isolates its populations. Geographic isolation occurs whenever a physical barrier divides a population.

A new species can evolve when a population has been geographically isolated. For example, imagine a population of tree frogs living in a rain forest, Pi gure 15.15. If small populations of tree frogs were geographically isolated, they would no longer be able to interbreed and exchange genes. Over time, each small population might adapt to its environment through natural selection and develop its own gene pool. Eventually, the gene pools of each population might become so different

Word Origin

speciation from the Latin word species, meaning "kind"; Speciation is a process that produces two species from one.

Figure 15.15

When geographic isolation divides a population of tree frogs, the individuals no longer mate across populations. Explain and Illustrate

t Tree frogs are a single population.

1l The formation of a river may divide the frogs into two populations. A new form may appear in one population.

ti' Over time, the divided populations may become two species that may no longer interbreed, even if reunited.

Parent plant (2n) Meiosis begins

Figure 15.16

Many flowering plants, such as this California tarweed, are polyploids-individuals that result from mistakes made during meiosis.

polyploidy from the Greek word polys, meaning "many"; Polyploid plants contain multiple sets of chromosomes.

Fertilization

Fertilization

Normal meiosis

Normal gametes (n)

that they could no longer interbreed with the other populations. In this way, natural selection results in new speCIes.

Reproductive isolation can result in speciation

As populations become increasingly distinct, reproductive isolation can arise. Reproductive isolation occurs when formerly interbreeding organisms can no longer mate and produce fertile offspring.

There are different types of reproductive isolation. Two examples are given here. One type occurs when the genetic material of the populations becomes so different that fertilization cannot occur. Some geographically separated populations of salamanders in California have this type of reproductive isolation. Another type of reproductive isolation is behavioral. For example, if one population of tree frogs mates in the fall, and another mates in the summer, these two populations will not mate with each other and are reproductively isolated.

410 THE THEORY OF EVOLUTI001

New .......... ~ polyploid species

Zygote (4n)

--_ •• _~ Sterile plant

Zygote (3n)

A change in chromosome numbers and speciation

Chromosomes can also playa role in speciation. Many new species of plants and some species of animals have evolved in the same geographic area as a result of polyploidy (PAR lih ploy dee), illustrated in Figure 15.16. Any individual or species with a multiple of the normal set of chromosomes is known as a polyploid.

Mistakes during mitosis or meiosis can result in polyploid individuals. For example, if chromosomes do not separate properly during the first meiotic division, diploid (2n) gametes can be produced instead of the normal haploid (n) gametes. Polyploidy may result in immediate reproductive isolation. When a polyploid mates with an individual of the normal species, the resulting zygotes may not develop normally because of the difference in chromosome numbers. In other cases, the zygotes develop into adults that probably cannot reproduce. However, polyploids within a population may interbreed and form a separate species.

Polyploids can arise from within a species or from hybridization between

;pecies. Many flowering plant species md many important crop plants, such IS wheat, cotton, and apples, originated by polyploidy.

Speciation rates

Although polyploid speciation takes only one generation, most other mechanisms of speciation do not occur as quickly. What is the usual rate of speciation?

Scientists once argued that evolution occurs at a slow, steady rate, with small, adaptive changes gradually accumulating over time in populations. Gradualism is the idea that species originate through a gradual change of adaptations. Some evidence from the fossil record supports gradualism. For example, fossil evidence shows that sea lilies evolved slowly and steadily over time.

In 1972, Niles Eldredge and Stephen J. Gould proposed a different hypothesis known as punctuated equilibrium. This hypothesis argues that speCIatIOn occurs relatively quickly, in rapid bursts, with long periods of genetic equilibrium in between. According to this hypothesis, environmental changes, such as higher temperatures or the introduction of a competitive species, lead to rapid changes in a small population's gene pool that is reproductively isolated from the main population. Speciation happens quickly-in about 10 000 years or less. Like gradualism, punctuated equilibrium is supported by fossil evidence as shown in Figure 15.17.

Biologists generally agree that both gradualism and punctuated equilibrium can result in speciation, depending on the circumstances. It shouldn't

Figure 15.17

The fossil record of elephant evolution supports the view of punctuated equilibrium. Several elephant species may have evolved from an ancestral population in a short time.

Elephas

0
1
2
primigenius
3
4

5
6 about 55 million years ago

15.2 MECHANISMS OF EVOLUTION 411

Figure 15.18 Evolutionary biologists have suggested that the ancestors of all Hawaiian Island honeycreepers migrated from North America about

5 million years ago. As this ancestral bird population settled in the diverse Hawaiian niches, adaptive radiation occurred.

Possible ancestral Laysan finch

surprise you to see scientists offer alternative hypotheses to explain observations. The nature of science is such that new evidence or new ideas can modify theories.

~m!!,!·J,;m3 Analyze and critique the strengths and weaknesses of gradualism and punctuated equilibrium.

Patterns of Evolution

Biologists have observed different patterns of evolution that occur throughout the world in different natural environments. These patterns support the idea that natural selection is an important agent for evolution.

Diversity in new environments An extraordinary diversity of unique plants and animals live or have lived on the Hawaiian Islands, among

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Extinct mamo

them a group of birds called Hawaiian honeycreepers. This group of birds is interesting because, although similar in body size and shape, they differ sharply in color and beak shape. Different species of honeycreepers evolved to occupy their own niches.

Despite their differences, scientists hypothesize that honeycreepers, as shown in Figure 15.18, evolved from a single ancestral species that lived on the Hawaiian Islands long ago. When an ancestral species evolves into an array of species to fit a number of diverse habitats, the result is called adaptive radiation.

Adaptive radiation in both plants and animals has occurred and continues to occur throughout the world and is common on islands. For example, the many species of finches that Darwin observed on the Galapagos Islands are a typical example of adaptive radiation.

Crested honeycreeper

... JKauai Aiihau

Akiapolaau

Molokai
Oahu
\ Maui
Lanai
Hawaii
Apapane
t
•
Maui
parrotbill
Grosbeak
finch
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Akikiki

412 TIlE TI-UDRY OF EVOLUTION

Figure 15.19

Unrelated species of plants such

as the Euphorbia (A) and this organ pipe cactus (8) share a similar fleshy body type and no leaves.

Adaptive radiation is a type of divergent evolution, the pattern of evolution in which species that once were similar to an ancestral species diverge, or become increasingly distinct. Divergent evolution occurs when populations change as they adapt to different environmental conditions, eventually resulting in new species.

Different species can look alike

A pattern of evolution in which distantly related organisms evolve similar traits is called convergent evolution. Convergent evolution occurs when unrelated species occupy similar environments in different parts of the world. Because they share similar

environmental pressures, they share similar pressures of natural selection.

For example, in Figure 15.19 you see an organ pipe cactus (family Cactaceae) that grows in the deserts of North and South America and a plant of the family Euphorbiaceae that looks similar and lives in African deserts. Although these plants are unrelated species, their environments are similar. You can see that they both have fleshy bodies and no leaves. That convergent evolution has apparently occurred in unrelated species, is further evidence for natural selection.

~im!l,!,";m3 Compare and contrast convergent and divergent evolution.

Understanding Main Ideas

1. Explain and illustrate why the evolution of resistance to antibiotics in bacteria is an example of directional natural selection.

2. How can geographic isolation change a population's gene pool?

3. Why is rapid evolutionary change more likely to occur in small populations?

4. How do gradualism and punctuated equilibrium differ? How are they similar? Include in your answer the patterns of extinction observed in both theories.

Thinking Critically

5. Hummingbird moths are night-flying insects whose behavior and appearance are similar to those of hummingbirds. Explain how these two organisms demonstrate convergent evolution.

------4. j iJ i!' j14'1 i W .-----

6. Experiment Biologists discovered two squirrel species living on opposite sides of the Grand Canyon. They hypothesize that the species evolved from a common ancestor. What observations or experiments could provide evidence for this hypothesis? For more help, refer to Experiment

in the Skill Handbook.

15.2 ;vlECHA]\;ISlvlS OF EVOLUTION 413

Mathematics and Evolution

In the early 1900s, G. H. Hardy, a British mathematician, and W Weinberg, a German doctor, independently discovered how the frequency of a trait's alleles in a population could be described mathematically.

Suppose that in a population of pea plants,

36 plants are homozygous dominant for the tall trait (TT), 48 plants are heterozygous tall (Tt), and 16 plants are short plants (tt). In the homozygous tall plants, there are (36) (2), or 72,

T alleles and in the heterozygous plants there are 48 T alleles, for a total of 120 T alleles in

the population. There are 48 t alleles in the h.eterozygous plants plus (16) (2), or 32, t alleles 111 the short plants, for a total of 80 t alleles in the population. The number of T and t alleles in ~he population is 200. The frequency of T alleles IS 1201200 or 0.6, and the frequency of t alleles is 801200, or 0.4.

The Hardy-Weinberg principle The Hardy-Weinberg principle states that the frequency of the alleles for a trait in ~ stable population will not vary. This statement IS expressed as the equation p + q = 1, where p is the frequency of one allele for the trait, and q is the frequency of the other allele. The sum of the frequencies of the alleles always includes 100 percent of the alleles, and is therefore stated as 1.

Squaring both sides of the equation produces. the equation p2 + 2pq + q2 = 1. You can use this equation to determine the frequency of ?'enotypes in a population: homozygous dominant individuals (p2), heterozygous individuals (2pq), and recessive individuals (q2). For example, in the pea plant population described above, the frequency of the genotypes would be

(0.6) (0.6) + 2(0.6) (0.4) + (0.4) (0.4) = 1

Ii Ii

II 416 TI-TE THEORY OF EVOLUTJO]\,'

Ii

A population of penquins

The frequency of the homozygous tall genotype is 0.36, the heterozygous genotype is 0.48, and the short genotype is 0.16.

In any sexually reproducing, large population, o'enotype frequencies will remain constant if no

b .

mutations occur, randommating occurs, no nat-

ural selection occurs, and no genes enter or leave the population.

Implications of the principle The HardyWeinberg principle is useful for sev~ral reasons. First, it explains that the genotypes 111 populations tend to remain the same. Second, because a recessive allele may be masked by its dominant allele, the equation is useful for determining the recessive allele's frequency in the population. Finally, the Hardy-Weinberg principle is useful in studying natural populations to determine how much ~atural selection may be occurring in the population.

Math in Biology

. - .. -~~~----------======~------

Draw Conclusions The general population of the United States is getting taller. Assuming that height is a genetic trait, does this observation viclate the Hardy-Weinberg principle? Explain your answer.

/1lJ To find out more about the l.;..-/ Hardy-Weinberg principle, visit