BSC 1011C
General Biology II
Dr. Graeme Lindbeck
glindbeck@valenciacollege.edu

The Evolution of Populations

Outline

Introduction

One obstacle to understanding evolution is the common misconception that organisms evolve, in a Darwinian sense, in their lifetimes.

Natural selection does act on individuals by impacting their chances of survival and their reproductive success.
However, the evolutionary impact of natural selection is only apparent in tracking how a population of organisms changes over time.

It is the population, not its individual, that evolve.

Evolution on the scale of populations, called microevolution, is defined as a change in the allele frequencies in a population.

While many seeds land on the mine tailings each year, the only plants that germinate, grow, and reproduce are those that had already inherited genes enabling them to tolerate metallic soils.
Individual plants do not evolve to become more metal-tolerant during their lifetimes.

The Origin of the Species convinced most biologists that species are the products of evolution, but acceptance of natural selection as the main mechanism of natural selection was more difficult.

What was missing in Darwin's explanation was an understanding of inheritance that could explain how chance variations arise in a population while also accounting for the precise transmission of these variations from parents to offspring.
Although Gregor Mendel and Charles Darwin were contemporaries, Mendel's discoveries were unappreciated at the time, even though his principles of heredity would have given credibility to natural selection.

A. Population Genetics

1. The modern evolutionary synthesis integrated Darwinian selection and Mendelian inheritance

When Mendel's research was rediscovered in the early twentieth century, many geneticists believed that the laws of inheritance conflicted with Darwin's theory of natural selection.

Darwin emphasized quantitative characters, those that vary along a continuum.
- These characters are influenced by multiple loci.
Mendel and later geneticists investigated discrete "either-or" traits.

An important turning point for evolutionary theory was the birth of population genetics, which emphasizes the extensive genetic variation within populations and recognizes the importance of quantitative characters.

Advances in population genetics in the 1930s allowed the perspectives of Mendelism and Darwinism to be reconciled.
This provided a genetic basis for variation and natural selection.

A comprehensive theory of evolution, the modern synthesis, took form in the early 1940's.

It integrated discoveries and ideas from paleontology, taxonomy, biogeography, and population genetics.

The architects of the modern synthesis included geneticists Theodosius Dobzhansky and Sewall Wright, biogeographer and taxonomist Ernst Mayr, paleontologist George Gaylord Simpson, and botanist G. Ledyard Stebbins.

The modern synthesis emphasizes:

the importance of populations as the units of evolution,
the central role of natural selection as the most important mechanism of evolution, and
the idea of gradualism to explain how large changes can evolve as an accumulation of small changes over long periods of time.

While many evolutionary biologists are now challenging some of the assumptions of the modern synthesis, it shaped most of our ideas about how populations evolve.

2. A population's gene pool is defined by its allele frequencies

A population is a localized group of individuals that belong to the same species.

One definition of a species (among others) is a group of populations whose individuals have the potential to interbreed and produce fertile offspring in a nature.

Populations of a species may be isolated from each other, such that they exchange genetic material rarely, or they may intergrade with low densities in an intermediate region.

Members of a population are far more likely to breed with members of the same population than with members of other populations.

Individuals near the populations center are, on average, more closely related to one another than to members of other populations.

The total aggregate of genes in a population at any one time is called the population's gene pool.

It consists of all alleles at all gene loci in all individuals of a population.
Each locus is represented twice in the genome of a diploid individual.
- Individuals can be homozygous or heterozygous for these homologous loci.
If all members of a population are homozygous for the same allele, that allele is said to be fixed.
Often, there are two or more alleles for a gene, each contributing a relative frequency in the gene pool.

For example, imagine a wildflower population with two flower colors.

The allele for red flower color (R) is completely dominant to the allele for white flowers (r).

Suppose that in an imaginary population of 500 plants, 20 have white flowers (homozygous recessive - rr).

The other 480 plants have red flowers.
- Some are heterozygotes (Rr), others are homozygous (RR).
Suppose that 320 are RR and 160 are Rr.

Because these plants are diploid, in our population of 500 plants there are 1,000 copies of the gene for flower color.

The dominant allele (R) accounts for 800 copies (320 x 2 for RR + 160 x 1 for Rr).
The frequency of the R allele in the gene pool of this population is 800/1000 = 0.8, or 80%.
The r allele must have a frequency of 1 - 0.8 = 0.2, or 20%.

3. The Hardy-Weinberg Theorem describes a nonevolving population

The Hardy-Weinberg theorem describes the gene pool of a nonevolving population.

This theorem states that the frequencies of alleles and genotypes in a population's gene pool will remain constant over generations unless acted upon by agents other than Mendelian segregation and recombination of alleles.

The shuffling of alleles after meiosis and random fertilization should have no effect on the overall gene pool of a population.

In our imaginary wildflower population of 500 plants, 80% (0.8) of the flower color alleles are R and 20% (0.2) are r.

How will meiosis and sexual reproduction affect the frequencies of the two alleles in the next generation?

We assume that fertilization is completely random and all male-female mating combinations are equally likely.

Because each gamete has only one allele for flower color, we expect that a gamete drawn from the gene pool at random has a 0.8 chance of bearing an R allele and a 0.2 chance of bearing an r allele.

Using the rule of multiplication, we can determine the frequencies of the three possible genotypes in the next generation.

For the RR genotype, the probability of picking two R alleles is 0.64 (0.8 x 0.8 = 0.64 or 64%).
For the rr genotype, the probability of picking two r alleles is 0.04 (0.2 x 0.2 = 0.04 or 4%).
Heterozygous individuals are either Rr or rR, depending on whether the R allele arrived via sperm or egg.
- The probability of ending up with both alleles is 0.32 (0.8 x 0.2 = 0.16 for Rr, 0.2 x 0.8 = 0.16 for rR, and 0.16 + 0.16 = 0.32 or 32% for Rr + rR).

As you can see, the processes of meiosis and random fertilization have maintained the same allele and genotype frequencies that existed in the previous generation.
For the flower-color locus, the population's genetic structure is in a state of equilibrium, Hardy-Weinberg equilibrium.

Theoretically, the allele frequencies should remain at 0.8 for R and 0.2 for r forever.

The Hardy-Weinberg theorem states that the processes involved in a Mendelian system have no tendency to alter allele frequencies from one generation to another.

The repeated shuffling of a population's gene pool over generations cannot increase the frequency of one allele over another.

The Hardy-Weinberg theorem also applies to situations in which there are three or more alleles and with other interactions among alleles than complete dominance.

Generalizing the Hardy-Weinberg theorem, population geneticists use p to represent the frequency of one allele and q to represent the frequency of the other allele.

The combined frequencies must add to 100%; therefore p + q = 1.
If p + q = 1, then p = 1 - q and q = 1 - p.

In the wildflower example p is the frequency of red alleles (R) and q of white alleles (r).

The probability of generating an RR offspring is p² (an application of the rule of multiplication).
- In our example, p = 0.8 and p² = 0.64.
The probability of generating an rr offspring is q².
- In our example, q = 0.2 and q² = 0.04.
The probability of generating Rr offspring is 2pq.
- In our example, 2 x 0.8 x 0.2 = 0.32.

The genotype frequencies should add to 1:

p² + 2pq + q² = 1

For the wildflowers, 0.64 + 0.32 + 0.04 = 1.

This general formula is the Hardy-Weinberg equation.

Using this formula, we can calculate frequencies of alleles in a gene pool if we know the frequency of genotypes or the frequency of genotypes if we know the frequencies of alleles.

We can use the Hardy-Weinberg theorem to estimate the percentage of the human population that carries the allele for a particular inherited disease, phenyketonuria (PKU) in this case.

About 1 in 10,000 babies born in the United States is born with PKU, which results in mental retardation and other problems if left untreated.
The disease is caused by a recessive allele.

From the epidemiological data, we know that frequency of homozygous recessive individuals (q² in the Hardy-Weinberg theorem) = 1 in 10,000 or 0.0001.

The frequency of the recessive allele (q) is the square root of 0.0001 = 0.01.
The frequency of the dominant allele (p) is p = 1 - q or 1 - 0.01 = 0.99.
The frequency of carriers (heterozygous individuals) is 2pq = 2 x 0.99 x 0.01 = 0.0198 or about 2%.

Thus, about 2% of the U.S. population carries the PKU allele.

The Hardy-Weinberg theorom shows how Mendel's theory of inheritance plugs a hole in Darwin's theory of natural selection, the requirement for genetic variation.

Under older models of inheritance ("blending" theories), hereditary factors in an offspring were thought to be a blend of the factors inherited from its two parents.
This process tends to reduce genetic variation from generation to generation, leading to uniformity.
In Mendelian inheritance, there is no tendency to reduce genetic variation from one generation to the next as demonstrated by the Hardy-Weinberg theorem.

Populations at Hardy-Weinberg equilibrium must satisfy five conditions.

Very large population size. In small populations, chance fluctuations in the gene pool, genetic drift, can cause genotype frequencies to change over time.
No migrations. Gene flow, the transfer of alleles due to the movement of individuals or gametes into or out of our target population can change the proportions of alleles.
No net mutations. If one allele can mutate into another, the gene pool will be altered.
Random mating. If individuals pick mates with certain genotypes, then the mixing of gametes will not be random and the Hardy-Weinberg equilibrium does not occur.
No natural selection. If there is differential survival or mating success among genotypes, then the frequencies of alleles in the next variation will deviate from the frequencies predicted by the Hardy-Weinberg equation.

Evolution usually results when any of these five conditions are not met - when a population experiences deviations from the stability predicted by the Hardy-Weinberg theory.

B. Causes of Microevolution

1. Microevolution is a generation-to-generation change in a population's allele frequencies

The Hardy-Weinberg theory provides a baseline against which we can compare the allele and genotype frequencies of an evolving population.

We can define microevolution as generation-to-generation change in a population's frequencies of alleles.

Microevolution occurs even if the frequencies of alleles are changing for only a single genetic locus in a population while the others are at equilibrium.

2. The two main causes of microevolution are drift and natural selection

Four factors can alter the allele frequencies in a population:

genetic drift
natural selection
gene flow
mutation

All represent departures from the conditions required for the Hardy-Weinberg equilibrium.
Natural selection is the only factor that generally adapts a population to its environment.
- Selection always favors the disporportionate propagation of favorable traits.

The other three may effect populations in positive, negative, or neutral ways.

Genetic drift occurs when changes in gene frequencies from one generation to another occur because of chance events (sampling errors) that occur when populations are finite in size.

For example, one would not be too surprised if a coin produced seven heads and three tails in ten tosses, but you would be surprised if you saw 700 heads and 300 tails in 1000 tosses - you expect 500 of each.
The smaller the sample, the greater the chance of deviation from an idealized result.
Genetic drift at small population sizes often occurs as a result of two situations: the bottleneck effect or the founder effect.

Applied to a population's gene pool, we expect that the gene pool of the next generation will be the same as the present generation in the absence of sampling errors.

This requirement of the Hardy-Weinberg equilibrium is more likely to be met if the size of the population is large (theoretically, infinite).
The gene pool of a small population may not be accurately represented in the next generation because of sampling errors.
This is analogous to the erratic outcome from a small sample of coin tosses.

For example, in a small wildflower population with a stable size of only ten plants, genetic drift can completely eliminate some alleles.

The bottleneck effect occurs when the numbers of individuals in a larger population are drastically reduced by a disaster.

By chance, some alleles may be overrepresented and others underrepresented among the survivors.
Some alleles may be eliminated altogether.
Genetic drift will continue to impact the gene pool until the population is large enough to minimize the impact of sampling errors.

Bottlenecking is an important concept in conservation biology of endangered species.

Populations that have suffered bottleneck incidents have lost at least some alleles from the gene pool.
This reduces individual variation and adaptability.
For example, the genetic variation in the three small surviving wild populations of cheetahs is very low when compared to other mammals.
Their genetic variation is similar to highly inbred lab mice!

The founder effect occurs when a new population is started by only a few individuals that do not represent the gene pool of the larger source population.

At an extreme, a population could be started by single pregnant female or single seed with only a tiny fraction of the genetic variation of the source population.

Genetic drift would continue from generation to generation until the population grew large enough for sampling errors to be minimal.

Founder effects have been demonstrated in human populations that started from a small group of colonists.

Natural selection is clearly a violation of the conditions necessary for the Hardy-Weinberg equilibrium.

The later expects that all individuals in a population have equal ability to survive and produce viable, fertile offspring.
However, in a population with variable individuals, natural selection will lead some individuals to leave more offspring than others.
Selection results in some alleles being passed along to the next generation in numbers disproportionate to their frequencies in the present generation.
In our wildflower example, if herbivorous insects are more likely to locate and eat white flowers than red flowers, then plants with red flowers (either RR or Rr) are more likely to leave offspring than those with white flowers (rr).
If pollinators were more attracted by red flowers than white flowers, red flowers would also be more likely to leave more offspring.
Either mechanism, differential survival or differential reproduction, will increase the frequency of the R allele in the population and decrease that of the r allele.

Natural selection accumulates and maintains favorable genotypes in a population.

Gene flow is genetic exchange due to migration of fertile individuals or gametes between populations.

For example, if a nearby wildflower population consisted entirely of white flowers, its pollen (r alleles only) could be carried into our target population.
This would increase the frequency of r alleles in the target population in the next generation.

Gene flow tends to reduce differences between populations.

If extensive enough, gene flow can amalgamate neighboring populations into a single population with a common genetic structure.

Gene flow tends to reduce differences between populations.

If extensive enough, gene flow can amalgamate neighboring populations into a single population with a common genetic structure.
The migration of people throughout the world is transferring alleles between populations that were once isolated, increasing gene flow.

A mutation is a change in an organism's DNA.

A new mutation that is transmitted in gametes can immediately change the gene pool of a population by substituting the mutated allele for the older allele.

For any single locus, mutation alone does not have much quantitative effect on a large population in a single generation.
An individual mutant allele may have greater impacts later through increases in its relative frequencies as a result of natural selection or genetic drift.

While mutations at an individual locus is a rare event, the cumulative impact of mutations at all loci can be significant.

Each individuals has thousands of genes, any one of which could experience a mutation.
Populations are composed of thousands or millions of individuals that may have experienced mutations.

Over the long term, mutation is a very important to evolution because it is the original source of genetic variation that serves as the raw material for natural selection.

C. Genetic Variation, the Substrate for Natural Selection

1. Genetic variation occurs within and between populations

The variation among individuals in a population is a combination of inheritable and non-heritable traits.

Phenotype, the observable characteristics of an organism, is the cumulative product of an inherited genotype and a multitude of environmental influences.

For example, these butterflies are genetically identical at the loci for coloration, but they emerge at different seasons.

Only the genetic component of variation can have evolutionary consequences as a result of natural selection.

This is because only inheritable traits pass from generation to generation.

Both quantitative and discrete characters contribute to variation within a population.

Quantitative characters are those that vary along a continuum within a population.

For example, plant height in our wildflower population includes short and tall plants and everything in between.
Quantitative variation is usually due to polygenic inheritance in which the additive effects of two or more genes influence a single phenotypic character.

Discrete characters, such as flower color, are usually determined by a single locus with different alleles with distinct impacts on the phenotype.

Polymorphism occurs when two or more discrete characters are present and noticeable in a population.

The contrasting forms are called morphs, as in the red-flowered and white-flowered morphs in our wildflower population or the butterflies in the previous slide.
Human populations are polymorphic for a variety of physical (e.g., freckles) and biochemical (e.g., blood types) characters.

Polymorphism applies only to discrete characters, not quantitative characters, such as human height, which varies among people in a continuum.

Population geneticists measure genetic variation both at the level of whole genes and at the molecular level of DNA.

Gene diversity measures the average percent of gene loci that are heterozygous.

In the fruit fly (Drosophila), about 86% of their 13,000 gene loci are homozygous (fixed).
About 14% (1,800 genes) are heterozygous.

Nucleotide diversity measures the level of difference in nucleotide sequences (base pair differences) among individuals in a population.

In fruit flies, about 1% of the bases are different between two individuals.
Two individuals would differ at 1.8 million of the 180 million nucleotides in the fruit fly genome.

Humans have relatively little genetic variation.

Gene diversity is about 14% in humans.
Nucleotide diversity is only 0.1%.
- You and your neighbor have the same nucleotide at 999 out of every 1,000 nucleotide sites in your DNA.

Geographic variation results from differences in genetic structure either between populations or between subgroups of a single population that inhabit different areas.

Often geographic variation results from natural selection that modifies gene frequencies in response to differences in local environmental factors.
Alternatively, genetic drift can lead to chance variations among populations.
Geographic variation can occur on a local scale, within a population, if the environment is patchy or if dispersal of individuals is limited, producing subpopulations.

Geographic variation in the form of graded change in a trait along a geographic axis is called a cline.

Clines may represent intergrade zones where individuals from neighboring, genetically different, populations interbreed.
Alternatively, clines may reflect the influence of natural selection on gradation in some environmental variable.
- For example, the average body size of many North American species of birds and mammals increases gradually with increasing latitude, perhaps conserving heat by decreasing the ratio of surface area to volume.

Clines may reflect direct environmental effects on phenotype, but also genetic differences along the cline.

For example, average size of yarrow plants (Anchillea), gradually decreases with increasing variation.
Although the environment affects growth rate directly to some extent with altitude, common garden experiments have demonstrated that some of the variation has a genetic basis.

In contrast to clines, isolated populations typically demonstrate discrete differences.

For example, populations of house mice were first introduced to the island of Madiera in the 15^th century, but isolated populations developed that were separated by mountains.
Some isolated populations have evolved differences in karyotypes probably through genetic drift.

2. Mutation and sexual recombination generate genetic variation

New alleles originate only by mutation.

Mutations are changes in the nucleotide sequence of DNA.
Mutations of individual genes are rare and random.
Mutations in somatic cells are lost when the individual dies.
Only mutations in cell lines that produce gametes can be passed along to offspring.

Most point mutations, those affecting a single base of DNA, are probably harmless.

Most eukaryotic DNA does not code for proteins and mutations in these areas are likely to have little impact on phenotype.
Even mutations in genes that code for proteins may lead to little effect because of redundancy in the genetic code.
However, some single point mutations can have a significant impact on phenotype.
- Sickle-cell disease is caused by a single point mutation.

Mutations that alter the structure of a protein enough to impact its function are more likely to be harmful than beneficial.

A random change is unlikely to improve a genome that is the product of thousands of generations of selection.
Rarely, a mutant allele may enable an organism to fit its environment better and increase reproductive success.
This is especially likely if the environment is changing
These mutations may be beneficial now.
- For example, mutations that enable HIV to resist antiviral drugs are selected against under normal conditions, but are favorable under drug treatment.

Chromosomal mutations, including rearrangements of chromosomes, affect many genes and are likely to disrupt proper development of an organism.

However, occasionally, these dislocations link genes together such that the phenotype is improved.

Duplications of chromosome segments, whole chromosomes, or sets of chromosomes are nearly always harmful.

However, when they are not harmful, the duplicates provide an expanded genome.
These extra genes can now mutate to take on new functions.

Because microorganisms have very short generation times, mutation generates genetic variation rapidly.

In an AIDS patient, HIV generates 10¹⁰ new viruses per day.
With its RNA genome, mutation rate is higher than DNA genomes.
This combination of mutation and replication rate will generate mutations in the HIV population at every site in the HIV genome every day.
- In the face of this high mutation rate, single-drug treatments are unlikely to be effective for very long and the most effective treatments are multiple drug "cocktails."
  - It is far less probable that mutations against all the drugs will appear in individual viruses in a short time.

In organisms with sexual reproduction, most of the genetic differences among individuals are due to unique recombinations of the existing alleles from the population gene pool.

The ultimate origin of allelic variation is past mutations.

Random segregation of homologous chromosomes and random union of gametes creates a unique assortment of alleles in each individual.

Sexual reproduction recombines old alleles into fresh assortments every generation.

3. Diploidy and balanced polymorphism preserve variation

The tendency for natural selection to reduce variation is countered by mechanisms that preserve or restore variation, including diploidy and balanced polymorphisms.

Diploidy in eukaryotes prevents the elimination of recessive alleles via selection because they do not impact the phenotype in heterozygotes.

Even recessive alleles that are unfavorable can persist in a population through their propagation by heterozygous individuals.
Recessive alleles are only exposed to selection when two parents carry the same recessive allele and these are combined in one zygote.
This happens only rarely when the frequency of the recessive allele is very low.
Heterozygote protection maintains a huge pool of alleles that may not be suitable under the present conditions but that could be beneficial when the environment changes.

Balanced polymorphism maintains genetic diversity in a population via natural selection.

One mechanism in balance polymorphism is heterozygote advantage.

In some situations individuals that are heterozygous at a particular locus have greater survivorship and reproductive success than homozygotes.
In these cases, multiple alleles will be maintained at that locus by natural selection.

Heterozygous advantage maintains genetic diversity at the human locus for one chain of hemoglobin.

A recessive allele causes sickle-cell disease in homozygous individuals.
Homozygous dominant individuals are very vulnerable to malaria.
Heterozygous individuals are resistant to malaria.

The frequency of the sickle-cell allele is highest in areas where the malarial parasite is common.

The advantages of heterozygotes over homozygous recessive individuals who suffer sickle-cell disease and homozygous dominant individuals who suffer malaria are greatest here.
The sickle-cell allele may reach 20% of the gene pool, with 32% heterozygotes resistant to malaria and 4% with sickle-cell disease.

A second mechanism promoting balanced polymorphisms is frequency-dependent selection.

Frequency-dependent selection occurs when the reproductive success of any one morph declines if that phenotype becomes too common in the population.

The relationships between parasites and their hosts often demonstrate this type of relationship.

Hosts often vary in their defense against parasites and parasites in their ability to infect hosts.

Those parasites that are capable of infecting the most common host type will increase in abundance.
The rarer host types will increase as the genetic frequencies in the parasite population shifts.
These shifts in genetic frequencies among hosts and among parasites maintain variation in both populations.

Aspects of this teeter-totter of frequency-dependent selection can be seen in the host-parasite between clones of aquatic snails and a parasitic worm.

In these snails which reproduce asexually, the most common snail clones suffer the higher infection rates than the least common clone, suggesting frequency-dependent selection.

Some genetic variations, neutral variation, have negligible impact on reproductive success.

For example, the diversity of human fingerprints seems to confer no selective advantage to some individuals over others.
Much of the protein and DNA variation detectable by methods like electrophoresis may be neutral in their adaptive qualities.

The relative frequencies of neural variations will not be affected by natural selection.

Some neutral alleles will increase and others will decrease by the chance effects of genetic drift.

There is no consensus on how much genetic variation can be classified as neutral or even if any variation can be considered truly neutral.

It is almost impossible to demonstrate that an allele brings no benefit at all to an organism.
Also, variation may be neutral in one environment but not in another.
Even if only a fraction of the extensive variation in a gene pool significantly affects an organism, there is still an enormous reservoir of raw material for natural selection and adaptive evolution.

D. A Closer Look at Natural Selection as the Mechanism of Adaptive Evolution

Adaptive evolution is a blend of chance and sorting.

Chance creates new genetic variations by mutation and sexual recombination.
Sorting by natural selection favors the propagation of some variations over others.
This produces organisms that are better fit to their environments.

1. Evolutionary fitness is the relative contribution that an individual makes to the gene pool of the next generation

The common phrases "struggle for existence" and "survival of the fittest" are misleading if they are taken to mean direct competitive contests among individuals.

While some animals do engage in head-to-head contests, most reproductive success is the product of more subtle and passive factors.

Reproductive success may depend on a variety of factors.

For example, one barnacle may produce more offspring because it is more efficient in collecting food.
In a population of moths, some color variants may provide better camouflage from predators, increasing survival and the likelihood of reproduction.
Slight differences in flower shape, color, or fragrance may lead to differences in reproductive success.

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 use a more quantitative approach to natural selection, defining relative fitness as the contribution of one genotype to the next generation compared to the contributions of alternative genotypes for the same locus.

In a hypothetical population of wildflowers, RR and Rr plants have red flowers and rr plants have white flowers.
If red flowers produce more offspring than white flowers, we would set their relative fitness at 1 and set the relative fitness of the white flowers relative to the red.
If white flowers produce only 80% as many offspring as the red flowers, their relative fitness would be 0.8.

Survival alone does not guarantee reproductive success.

Relative fitness is zero for a sterile plant or animal, no matter how much longer it lives than others.
Similarly, organisms must survive to reproduce.

The many factors that affect both survival and fertility determine the evolutionary fitness of an individual.

It is the phenotype - physical traits, metabolism, physiology and behavior - not the genotype that interacts with the environment.

Selection acts on phenotypes.

Through the differential survival and reproductive success of phenotypes, natural selection adapts a population to its environment by increase or by maintaining favorable genotypes that produce the better phenotypes in the gene pool.

Natural selection works on the whole organism, the integrated composite of its many phenotypic features, not on a collage of parts.

The relative fitness of an allele depends on its entire genetic context, how it interacts with other genes and alleles.

The apparent advantages provided by one allele may be detrimental if not supported by other genes and alleles.
Plus, even alleles that are neutral or slightly maladaptive may be perpetuated if they occur in organisms with a high overall fitness.

2. The effect of selection on a varying characteristic can be directional, diversifying, or stabilizing

Natural selection can affect the frequency of a heritable trait in a population, leading to directional selection, diversifying selection, or stabilizing selection.

Directional selection is most common during periods of environmental change or when members of a population migrate to a new habitat with different environmental conditions.

Directional selection shifts the frequency curve for a phenotypic character in one direction by favoring what had been rare individuals.

Peter and Rosemary Grant documented directional evolution in beak size for the medium ground finch in the Galapagos Islands.

During wet years when seeds are abundant, all individuals consume relatively few large seeds.
However, during dry years when seeds are scarce, the small seeds are quickly depletedand birds with larger, stronger beaks that can crack large seeds are at an advantage, and their genes increase in the population.

Diversifying selection occurs when environmental conditions favor individuals at both extremes of the phenotypic range over intermediate phenotypes.

Diversifying selection can result in balanced polymorphism.

For example, two distinct bill types are present in black-bellied seedcrackers in which larger-billed birds are more efficient when feeding on hard seeds and smaller-billed birds are more efficient when feeding on soft seeds.

Stabilizing selection favors intermediate variants and acts against extreme phenotypes.

Stabilizing selection reduces variation and maintains the predominant phenotypes.

Human birth weight is subject to stabilizing selection.
Babies much larger or smaller than 3-4 kg have higher infant mortality.

3. Natural selection maintains sexual reproduction

Sex is an evolutionary enigma.

It is far inferior to asexual reproduction as measured by reproductive output.

If a population consisted of half sexual females and half asexual females, the asexual condition would increase.
- All of offspring of asexual females would be reproductive daughters.
- Only half of the offspring of sexual females would be daughters; the other half would be necessary males.

Theoretically, sex has a "two-fold disadvantage."

A female producing two offspring per generation would generate a population of eight females after four generations if reproducing asexually, but only one female if reproducing sexually.

Sex must confer some selective advantage to compensate for the costs of diminished reproductive output.

Otherwise, a migration of asexual individuals or mutation permitting asexual reproduction would outcompete sexual individuals and the alleles favoring sex.

In fact, most eukaryotes maintain sex, even in those species that can also reproduce asexually.

The "textbook" explanation for the maintenance of sex is that the process of meiosis and fertilization generate genetic variation on which natural selection can act.

However, this hypothesis that sex is maintained in spite of its disadvantages because it produces future adaptation in a variable world is difficult to defend.
Natural selection acts on the present, favoring individuals here and now that best fit the current, local environment.

A stronger hypothesis would present advantages to sex that place a value on genetic variation on a generation-to-generation time scale.

Resistance to disease may be one current value of variability overcoming the disadvantages of sex.

Parasites recognize and infect their hosts by attaching to receptor molecules on the host's cells.
Among the hosts, individuals are likely to have different alleles for these receptor molecules and, therefore, vary in their vulnerability to parasites.
At the same time, parasites evolve very rapidly in their ability to use specific host receptors.
Sex provides a mechanism for changing the distribution of alleles and varying them among offspring.
This coevolution in a host-parasite relationship has been called the "Red Queen effect."

4. Sexual selection may lead to pronounced secondary differences between the sexes

Males and females of a species differ not only in their reproductive organs, but often also in secondary sexual characteristics that are not directly associated with reproduction.

These differences, termed sexual dimorphism, may include size differences, coloration differences, enlarged or exaggerated features, or other adornments.
Males are usually the larger and showier sex, at least among vertebrates.

Sexual dimorphism is a product of sexual selection.

Intrasexual selection is direct competition among individuals of one sex (usually males) for mates of the opposite sex.

Competition may take the form of direct physical battles between individuals.
- The stronger individuals gain status.
More commonly ritualized displays discourage lesser competitors and determine dominance.

Intersexual selection or mate choice occurs when members of one sex (usually females) are choosy in selecting among individuals of the other sex.

Males with the most masculine features are the most attractive to females.
Interestingly, these features may not be adaptive in other ways and expose these individuals to extra risks.

However, even if these extravagant features have some costs, individuals that possess them will have enhanced reproductive success if they help an individual gain a mate.

Every time a females chooses a mate based on appearance or behavior, she perpetuates the alleles that caused her to make that choice
She also allows a male with that particular phenotype to perpetuate his alleles.

The underlying bases of female choice is probably not aesthetic.

Recent research is investigating the hypothesis that females use these sexual advertisements to measure the general health of a male.
Individuals with infections or other problems are likely to have a relatively dull, disheveled plumage.
These individuals are unlikely to win many females.
For the female that chooses a healthy mate, even if his inclination is just a prewired response to visual signals, the benefit is a greater probability of having healthy offspring.

5. Natural selection cannot fashion perfect organisms

There are at least four reasons why natural selection cannot produce perfection.

Evolution is limited by historical constraints.
- Evolution does not scrap ancestral anatomy and build from scratch.
- Evolution co-opts existing structures and adapts them to new situations.
Adaptations are often compromises.
- Organisms are often faced with conflicting situations that prevent an organism from perfecting any one feature for a particular situation.
  - For example, because the flippers of a seal must not only allow it to walk on land, but also swim efficiently, their design is a compromise between these environments.
  - Similarly, human limbs are flexible and allow versatile movements, but at the cost of injuries, such as sprains, torn ligaments, and dislocations.
    - Better structural reinforcement would compromise agility.
Not all evolution is adaptive.
- Chance affects the genetic structure of populations to a greater extent than was once believed.
  - For example, founders of new populations may not necessarily be the best individuals, but rather those individuals that are carried into the open habitat by chance.
Selection can only edit existing variations.
- Selection favors only the fittest variations from those phenotypes that are available.
- New alleles do not arise on demand.

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Dr. Graeme Lindbeck .