Natural Selection and Mechanisms of Evolution (II

)
Descent with Modification: A Darwinian View of Life The Origin of Species The Evolution of Populations

Prepared by Raajeswari Rajendran

The Smallest Unit of Evolution

One common misconception about evolution is that individual organisms evolve, in the Darwinian sense, during their lifetimes Natural selection acts on individuals, but populations evolve. Genetic variations in populations ◦ Contribute to evolution

The Modern Synthesis
 

Population genetics provides a foundation for studying evolution Microevolution ◦ Is change in the genetic makeup of a population from generation to generation Population genetics ◦ Is the study of how populations change genetically over time ◦ Reconciled Darwin’s and Mendel’s ideas The modern synthesis ◦ Integrates Mendelian genetics with the Darwinian theory of evolution by natural selection ◦ Focuses on populations as units of evolution

◦ ◦

Gene Pools and Allele Frequencies

A population
◦ Is a localized group of naturally occurring individuals that are capable of interbreeding and producing fertile offspring.

The gene pool
◦ Is the total aggregate of genes in a population of species at any one time ◦ Consists of all gene loci in all individuals of the population

The Hardy-Weinberg Theorem

The Hardy-Weinberg theorem ◦ Describes a population that is not evolving ◦ States that the frequencies of alleles and genotypes in a population’s gene pool remain constant from generation to generation provided that only Mendelian segregation and recombination of alleles are at work. Mendelian inheritance ◦ Preserves genetic variation in a population ◦

Preservation of Allele Frequencies - Hardy-Weinberg Equilibrium  In a given population where gametes contribute to
the next generation randomly, allele frequencies will not change.

Hardy-Weinberg equilibrium
◦ Describes a population in which random mating occurs ◦ Describes a population where allele frequencies do not change

Gametes for each generation are drawn at random from the gene pool of the previous generation:

A population in HardyWeinberg equilibrium

80% CR (p = 0.8)

20% CW (q = 0.2)

Sperm
CR (80%) CW (20%)

p2
CR (80%)

pq

Eggs

p2 64% CRCR 16% CRCW

CW (20%)

qp

16% CRCW

4% CWCW

q2 If the gametes come together at random, the genotype frequencies of this generation are in Hardy-Weinberg equilibrium: 64% CRCR, 32% CRCW, and 4% CWCW

Gametes of the next generation: 64% CR from CRCR homozygotes 4% CW from CWCW homozygotes

+ +

16% CR from CRCW homozygotes 16% CW from CRCW heterozygotes

= =

80% CR = 0.8 = p 20% CW = 0.2 = q

With random mating, these gametes will result in the same mix of plants in the next generation:

64% CRCR, 32% CRCW and 4% CWCW plants

If p and q represent the relative frequencies of the only two possible alleles in a population at a particular locus, then
◦ p2 + 2pq + q2 = 1 for the next generation ◦ And p2 and q2 represent the frequencies of the homozygous genotypes and 2pq represents the frequency of the heterozygous genotype

A population in Hardy-Weinberg equilibrium 

Frequency of different alleles of a gene in a population can be estimated by examining the proportion of individuals that have particular phenotypes and genotypes. E.g. human MN blood group – controlled by 2 codominant alleles, LM and LN. The heterozygote is blood type MN and the homozygotes are blood types M and N. A population of Aboriginal people from Elcho Island in Australia’s Northern Territory has 28 individuals with blood type M, 129 individuals with blood type MN and 195 individuals with blood type N. (Refer to handout (Knox) – pg 838839).

 

   
  

Current generation: Frequency of LM allele in the population – 185/704 = 0.26 = p. Frequency of LN allele in the population – 519/704 = 0.74 = q. p+q=1 Next generation: p2 + 2pq + q2 = 1 Blood group M, (p2) = (0.26)2 = 0.07 Blood group N, (q2) = (0.74)2 = 0.54 Blood group MN, (2pq) = (2 x 0.26 x 0.74) = 0.39

If

the genotype frequencies for the next generation is similar to the frequency predicted by the Hardy-Weinberg theorem, then evolution has not taken place.

Conditions for Hardy-Weinberg Equilibrium

The Hardy-Weinberg theorem
◦ Describes a hypothetical population

In real populations
◦ Allele and genotype frequencies do change over time

Five assumptions/conditions to apply the Hardy-Weinberg theorem

The five conditions for non-evolving populations are rarely met in nature
◦ ◦ ◦ ◦ ◦ Extremely large population size No gene flow No mutations Random mating No natural selection

1.Population size
The first assumption is population size is large.  If the population is very small, or if it goes through periodic ‘bottlenecks’ of low numbers, alleles in the population may drift to high or low frequencies purely by chance.  These random changes in allele frequencies in small isolated populations is termed genetic drift and can lead to rapid fluctuations in allele frequencies.

Genetic Drift
 

Statistically, the smaller a sample ◦ The greater the chance of deviation from a predicted result Genetic drift ◦ Describes how allele frequencies can fluctuate unpredictably from one generation to the next ◦ Tends to reduce genetic variation


CR CR CRCW CWCW CRCW CR CR CR CR CRCW CRCW CRCW CR CR CR CR Only 5 of 10 plants leave offspring CRCR

CWCW CRCW

CR CR Only 2 of 10 plants leave offspring CRCR CRCR CR CR CRCR CRCW CR CR

CR CR CRCR CR CR CRCR CRCR CRCR

CWCW CRCW CWCW

Generation 1 p (frequency of CR) = 0.7 q (frequency of CW) = 0.3

Generation 2 p = 0.5 q = 0.5

Generation 3 p = 1.0 q = 0.0

The Bottleneck Effect

In the bottleneck effect
◦ A sudden change in the environment may drastically reduce the size of a population ◦ The gene pool may no longer be reflective of the original population’s gene pool

(a)

Shaking just a few marbles through the narrow neck of a bottle is analogous to a drastic reduction in the size of a population after some environmental disaster. By chance, blue marbles are over-represented in the new population and gold marbles are absent. Original population Bottlenecking event Surviving population

Bottleneck effect

Understanding the bottleneck effect ◦ Can increase understanding of how human activity affects other species bringing about a fall in the population of species making it vulnerable to a genetic drift.

(b) Similarly, bottlenecking a population of organisms tends to reduce genetic variation, as in these northern elephant seals in California that were once hunted nearly to extinction.

The Founder Effect

The founder effect
◦ Occurs when a few individuals become isolated from a larger population ◦ Can affect allele frequencies in a population ◦ This too leads to a genetic drift.

2. No gene flow / migration
The second assumption is that individuals with particular alleles do not migrate at different rates.  Migration of individuals with particular alleles at different rates can lead to changes in allele frequencies.

3. Mutation rate
The third assumption is that different alleles mutate at the same rate and therefore unlikely to lead to rapid changes in the population.  If alleles mutate at different rates, they may lead to changes in the allele frequencies of the next generation.

4. Random mating
The fourth assumption is that random mating occurs.  Non-random mating will distort HardyWeinberg frequencies.

5. Darwinian fitness/no natural selection

Constant Hardy-Weinberg frequencies will occur only if there are no differences in the Darwinian fitness of individual genotypes. The assumption made here is that the environment is constant and has no changes. Fitness is measured as the relative contribution of offspring to subsequent generations due to differential survival and or reproduction. If environment changes and natural selection is in progress, then the allele frequencies will change.

Given the above assumptions, the genotype frequencies are expected to remain constant from generation to generation.  If one or more of these assumptions is not met, allele frequencies in a population may change and the population may evolve.

Population Genetics and Human Health

We can use the Hardy-Weinberg equation
◦ To estimate the percentage of the human population carrying the allele for an inherited disease

Genetic variation
When Darwin proposed natural selection as the primary mechanism for evolution, he emphasized the importance of heritable differences among individuals.  He knew that natural selection could not cause evolutionary change unless individuals differed in their inherited characteristics.  What are the sources of genetic variation?

Sources of genetic variation
1. Mutation – source of new alleles in a population  2. Crossing-over and recombination  3. Gene flow – immigration and emigration  4. Random segregation and independent assortment  5. Random fertilization  6. Random mating
 

Genetic variation ◦ Occurs in individuals in populations of all species ◦ Is not always heritable Mutation and sexual recombination produce the variation that makes evolution possible The two processes, mutation and sexual recombination ◦ Produce the variation in gene pools that contributes to differences among individuals

Mutation Rates

Mutation rates
◦ Tend to be low in animals and plants ◦ Average about one mutation in every 100,000 genes per generation ◦ Are more rapid in microorganisms

Sexual Recombination

In sexually reproducing populations, sexual recombination
◦ Is far more important than mutation in producing the genetic differences that make adaptation possible

Three major factors alter allele frequencies and bring about most evolutionary change
◦ Natural selection ◦ Genetic drift ◦ Gene flow

Natural Selection

Differential success in reproduction ◦ Results in certain alleles being passed to the next generation in greater proportions Natural selection is the primary mechanism of adaptive evolution Natural selection ◦ Accumulates and maintains favorable genotypes in a population ◦

Gene Flow

Gene flow
◦ Causes a population to gain or lose alleles ◦ Results from the movement of fertile individuals or gametes ◦ Tends to reduce differences between populations over time

Variation Within a Population
 

Both discrete/quantitative and quantitative characters ◦ Contribute to variation within a population Discrete characters ◦ Can be classified on an either-or basis Quantitative characters ◦ Vary along a continuum within a population ◦

Variation Between Populations

Most species exhibit geographic variation
◦ Differences between gene pools of separate populations or population subgroups
1 2.4 3.14 5.18 6

7.15

8.11

9.12

10.16

13.17

19

XX

1 9.10

2.19 11.12

3.8 13.17

4.16 15.18

5.14

6.7 XX

Some examples of geographic variation occur as a cline, which is a graded change in a trait along a geographic axis
Heights of yarrow plants grown in common garden

RESULTS The average plant sizes in the common garden were inversely correlated with the altitudes at which the seeds were collected, although the height differences were less than in the plants’ natural environments.

Atitude (m)

Mean height (cm)

EXPERIMENT Researchers observed that the average size of yarrow plants (Achillea) growing on the slopes of the Sierra Nevada mountains gradually decreases with increasing elevation. To eliminate the effect of environmental differences at different elevations, researchers collected seeds from various altitudes and planted them in a common garden. They then measured the heights of the resulting plants.

Sierra Nevada Range

Great Basin Plateau

CONCLUSION The lesser but still measurable clinal variation in yarrow plants grown at a common elevation demonstrates the role of genetic as well as environmental differences.

Seed collection sites

A Closer Look at Natural Selection

From the range of variations available in a population
◦ Natural selection increases the frequencies of certain genotypes, fitting organisms to their environment over generations

Evolutionary Fitness

The phrases “struggle for existence” and “survival of the fittest”
◦ Are commonly used to describe natural selection ◦ Can be misleading

 

Reproductive success
◦ Is generally more subtle and depends on many factors

Fitness
◦ Is the contribution an individual makes to the gene pool of the next generation, relative to the contributions of other individuals

Relative fitness
◦ Is the contribution of a genotype to the next generation as compared to the contributions of alternative genotypes for the same locus

Neutral Variation

Neutral variation
◦ Is genetic variation that appears to confer no selective advantage

Darwin explored the Galápagos Islands
◦ And discovered plants and animals found nowhere else on Earth

The origin of new species, or speciation
◦ Is at the focal point of evolutionary theory, because the appearance of new species is the source of biological diversity

Evolutionary theory
◦ Must explain how new species originate in addition to how populations evolve

Macroevolution
◦ Refers to evolutionary change above the species level

The biological species concept emphasizes reproductive isolation  Species

◦ Is a Latin word meaning “kind” or “appearance”

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 in nature and produce viable, fertile offspring but are unable to produce viable fertile offspring with members of other populations

(a) Similarity between different species. The eastern meadowlark (Sturnella magna, left) and the western meadowlark (Sturnella neglecta, right) have similar body shapes and colorations. Nevertheless, they are distinct biological species because their songs and other behaviors are different enough to prevent interbreeding should they meet in the wild. (b) Diversity within a species. As diverse as we may be in appearance, all humans belong to a single biological species (Homo sapiens), defined by our capacity to interbreed.

Reproductive Isolation

Reproductive isolation
◦ Is the existence of biological factors that impede members of two species from producing viable, fertile hybrids ◦ Is a combination of various reproductive barriers

Prezygotic barriers
◦ Impede mating between species or hinder the fertilization of ova if members of different species attempt to mate

Postzygotic barriers
◦ Often prevent the hybrid zygote from developing into a viable, fertile adult

Prezygotic and postzygotic barriers
Prezygotic barriers impede mating or hinder fertilization if mating does occur Habitat isolation Individuals of different species Temporal isolation Behavioral isolation
Mating attempt

Mechanical isolation

HABITAT / ECOLOGICAL ISOLATION

TEMPORAL ISOLATION BEHAVIORAL / ETHOLOGICAL ISOLATION MECHANICAL ISOLATION

(b)

(g)

(d) (e) (a) (f) (c)

Post-zygotic barriers Gametic isolation Reduce hybrid viability
Fertilization

Reduce hybrid fertility

Hybrid breakdown
Viable fertile offspring

GAMETIC ISOLATION

REDUCED HYBRID VIABILITY

REDUCED HYBRID FERTILITY HYBRID BREAKDOWN (k)

(j) (m) (l) (h) (i)

Limitations of the Biological Species Concept

The biological species concept cannot be applied to
◦ Asexual organisms ◦ Fossils ◦ Organisms about which little is known regarding their reproduction

Other Definitions of Species

The morphological species concept
◦ Characterizes a species in terms of its body shape, size, and other structural features

The paleontological species concept
◦ Focuses on morphologically discrete species known only from the fossil record

The ecological species concept
◦ Views a species in terms of its ecological niche

The phylogenetic species concept
◦ Defines a species as a set of organisms with a unique genetic history

Speciation – Formation of New Species
Speciation can take place with or without geographic separation Speciation can occur in two ways ◦ Allopatric speciation ◦ Sympatric speciation and parapatric (a) speciation

Allopatric speciation. A population forms a new species while geographically isolated from its parent population.

(b) Sympatric speciation. A small

population becomes a new species without geographic separation.

Allopatric (“Other Country”) Speciation

In allopatric speciation
◦ Gene flow is interrupted or reduced when a population is divided into two or more geographically isolated subpopulations

Once geographic separation has occurred
◦ One or both populations may undergo evolutionary change during the period of separation

A. harrisi

A. leucurus

Sympatric (“Same Country”) Speciation

In sympatric speciation and parapatric speciation
◦ Speciation takes place in geographically overlapping populations

Causes of sympatric speciation -Polyploidy

Polyploidy
◦ Is the presence of extra sets of chromosomes in cells due to accidents during cell division ◦ Has caused the evolution of some plant species – grass and wheat ◦ There are two types of polyploidy – autopolyploidy and allopolyploidy

An autopolyploid
◦ Is an individual that has more than two chromosome sets, all derived from a single species
Gametes produced by flowers on this branch will be diploid. Offspring with tetraploid karyotypes may be viable and fertile—a new biological species.

Failure of cell division in a cell of a growing diploid plant after chromosome duplication gives rise to a tetraploid branch or other tissue.

2n 2n = 6 4n = 12 4n

An allopolyploid
◦ Is a species with multiple sets of chromosomes derived from different species

Unreduced gamete with 4 chromosomes Meiotic error; chromosome number not reduced from 2n to n

Hybrid with 7 chromosomes

Unreduced gamete with 7 chromosomes

Viable fertile hybrid (allopolyploid)

Species A 2n = 4

2n = 10 Normal gamete n=3 Species B 2n = 6 Normal gamete n=3

Habitat Differentiation – Parapatric speciation

Parapatric speciation is a mode of speciation in which divergence occurs among populations that have contiguous distributions and hence are incompletely geographically separated. ◦ As a result of divergent selection, gene flow between the populations becomes prgressively reduced and eventually the differentiated populations become reproductively isolated. ◦ Sympatric and parapatric modes of speciation assume that speciation takes place without complete geographical isolation.

Allopatric and Sympatric Speciation: A Summary

In allopatric speciation
◦ A new species forms while geographically isolated from its parent population

In sympatric speciation
◦ The emergence of a reproductive barrier isolates a subset of a population without geographic separation from the parent species

Adaptive Radiation

Adaptive radiation
◦ Is the evolution of diversely adapted species from a common ancestor upon introduction to new environmental opportunities

The Hawaiian archipelago
◦ Is one of the world’s great showcases of adaptive radiation

N

Dubautia laxa

1.3 million years MOLOKA'I KAUA'I MAUI 5.1 million years O'AHU LANAI 3.7 million years

Argyroxiphium sandwicense HAWAI'I 0.4 million years

Dubautia waialealae

Dubautia scabra

Dubautia linearis

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