BIO 201 CLEVELAND STATE UNIVERSITY

CHAPTER 22-26 OBJECTIVES

All notes, writings, and content contained herein are derived or copied from Biology, 8 Edition by Campbell & Reece, 2008 Published by Pearson Education, Inc. This guide is for private use only, and is not to be sold or otherwise distributed for profit. Inquiries, questions or comments about these notes should be directed to Thomas Trotter at thomastrotter22@hotmail.com

th

BIO 201 OBJECTIVES

|2

CHAPTER 22
DESCENT WITH MODIFICATION

THE HISTORICAL CONTEXT FOR EVOLUTIONARY THEORY

1. Explain the mechanism for evolutionary change proposed by Charles Darwin in On the Origin of Species.
During Darwins voyage he studied many adaptations, or characteristics of organisms that enhance their survival and reproduction in specific environments. He proposed that these adaptations arise from a mechanism called natural selection, or a process in which individuals with certain inherited traits leave more offspring than individuals with other traits.

2. Define Evolution and Adaptation. a. EVOLUTION: a change in the genetic composition of a population from generation to generation.
Or, simply, descent with modification. b. ADAPTATION: characteristics of organisms that enhance their survival and reproduction in specific environments.

3. Describe the theories of catastrophism, gradualism and uniformitarianism. a. CATASTROPHISM: states that events in the past occurred suddenly and were caused by
mechanisms different from those operating in the present. A meteor strike is a catastrophic mechanism, such as the one that killed most dinosaurs. b. GRADUALISM: states that profound changes take place through cumulative effect of slow but continuous processes. Earths geologic features are explained via gradualism, as they are still occurring today. c. UNIFORMITARIANISM: states that the mechanisms of change are constant over time. Natural selection is one such mechanism, though is associated with evolution.

4. Explain the mechanism for evolutionary change proposed by Jean Baptiste Lamarck. Explain why modern biology has rejected Lamarcks theories.
Lamarck proposed that parts of the body that are used the most get bigger over time, and those parts that are not used deteriorate. Another principle, that of inheritance of acquired characteristics stated that an organism could pass these modifications to its offspring. Modern genetics refutes this mechanism as there is no way to pass on acquired characteristics, or even develop them through use and disuse.

THE DARWINIAN REVOLUTION

1. Describe how Darwins observations on the voyage of the HMS Beagle led him to formulate and support his theory of evolution.
Darwin observed many species on his trip to the Galapagos Islands, but it was the birds he collected that led him to natural selection. He noted many adaptations in these birdsadaptations that were not found anywhere else in the worldand began to perceive that new species could arise from an ancestral form by the gradual accumulation of adaptations to different environments. In particular, it was the finches and their varying beaks that led him to this discovery.

BIO 201 OBJECTIVES

|3

2. Explain what Darwin meant by descent with modification.

Darwin thought that there was unity in lifethat all organisms descended from an ancestor that lived in the distant past. These descendants lived in various habitats over millions of years, and had accumulated diverse modifications, or adaptations, that fit them to specific ways of life. The mechanism by which this works is natural selection. 3. Explain what evidence convinced Darwin that species change over time. I believe that the species he discovered on the Galapagos Islands were what led him to this conviction. Particularly the birdswhich he hypothesized had been colonized by organisms that had strayed from South America and then diversified, giving rise to new species on various islands.

4. Describe observations and two inferences that led Darwin to propose natural selection as a mechanism for evolutionary change.
Darwin discussed selective breeding of domesticated plants and animals. Humans have modified other species (think dogs) over many generations by selective breedinga process called artificial selection. There are a total of four observations and two inferences made by Darwin, as listed below: a. Observation 1: Members of a population often vary greatly in their traits. b. Observation 2: Traits are inherited from parents to offspring. c. Observation 3: All species are capable of producing more offspring than their environment can support. d. Observation 4: Owing to lack of food or other resources, many of these offspring do not survive. e. Inference 1: Individuals whose inherited traits give them a higher probability of surviving and reproducing in a given environment tend to leave more offspring than other individuals. f. Inference 2: This unequal ability of individuals to survive and reproduce will lead to the accumulation of favorable traits in the population over generations.

5. Explain how an essay by the Rev. Thomas Malthus influenced Charles Darwin.
Darwin realized that the capacity to over reproduce was characteristic of all species after reading this essay. The essay was about human sufferingdisease, famine and warand how it was an inescapable consequence of the human populations potential to increase faster than food supplies and other resources.

6. Explain why an individual organism cannot evolve.

Natural selection occurs through interactions between individual organisms and their environment. However, individuals do not evolveit is the population that evolves over time. This has to do with heritable traits being passed on to offspring, as discussed below.

7. Explain why natural selection can act only on heritable traits.

There is little evidence that acquired traitsor those traits that an individual acquires over their lifetimecan be inherited by their offspring. Rather, it is the heritable traits that are either amplified or diminished via natural selectionor those traits that can be passed on to offspring. This has everything to do with the mountains of evidence that support natural selection and evolution.

THE EVIDENCE OF EVOLUTION

1. Describe the experiments that supported Endlers hypothesis that differences in color patterns in male guppies are due to selective pressure based on predation. So, these guppies use exotic color patterns to attract females for matting purposes. However, their bright colors also attract predators. So, Endler went out and studied these fish. He found that in ponds where predation was high, guppies with drab colors were more frequent. He found that in

BIO 201 OBJECTIVES

|4

ponds where predation was low, bright colors were more frequent. Therefore, he hypothesized that intense predation caused natural selection in male guppies, favoring either drab or bright colors depending on the intensity of predation. 2. Describe how natural selection favors the evolution of drug-resistant pathogens. So, when we take drugs to kill a virus, there may be a few drug-resistant viruses floating around. What happens is that these drug-resistant one are able to reproducein fact, they are the only ones able to reproducethus lowering the effect (or completely negating the effect) of the drug in question. HIV is a good example of this, as the viruses that survive the early doses reproduce, passing on the alleles that enable them to resist the drug. In this way, the frequency of resistant viruses increases rapidly in the population. 3. Explain how the fossil record may be used to test our current understanding of evolutionary patterns. So, the fossil record is a great tool into understanding evolutionary changes that have occurred over time in various groups of organisms. This record can be used to test hypotheses arising from other kinds of evidence (such as the hypothesis that early land vertebrates evolved from a group of fishes and that early amphibians evolved from descendants of early land vertebrates), by using radioactive dating techniques to determine the age of the fossils. Based on these dates, we can tell which species came first. 4. Explain how the existence of homologous and vestigial structures can be explained by Darwins theory of natural selection. First of all, homology is similarity in characteristics among related species resulting from common ancestry. Variations on a structural themesuch as arms, forelegs, flippers and wings of different animalsthat were present in a common ancestor are homologous structures. Furthermore, vestigial structures are remnants of features that served important functions in the organisms ancestors. When we compare the stages of development in different animal species we can see even more anatomical homologies, which even further support the idea that these differences arose via natural selection. Then we come to convergent evolution, which is the independent evolution of similar features in different lineages. 5. Explain how evidence from biogeography supports the theory of evolution by natural selection. Islands generally have species of plants and animals that are endemic, which means they are found nowhere else in the world. However, most of the island species are closely related to mainland species, suggesting that islands are colonized by species from the nearest mainland. 6. Explain the problem with the statement that Darwinism is just a theory. Distinguish between the scientific and colloquial use of the word theory. The colloquial use of the word theory is more closely associated with a hypothesis. A scientific theory is more comprehensive, and accounts for many observations and explains and integrates a wide variety of phenomenon.

BIO 201 OBJECTIVES

|5

CHAPTER 23
THE EVOLUTION OF POPULATIONS

GENETIC VARIATION, THE SUBSTRATE FOR NATURAL SELECTION

1. Explain the statement It is the population, not the individual that evolves. This can be seen in the finches that Darwin studied. A finchs beak does not change over the course of its lifethat is, it does not get bigger or smaller depending on its environmentrather, it is the population over time that changes. 2. Explain how Mendels particulate hypothesis of inheritance provided necessary support for Darwins theory of evolution by natural selection. Mendel proposed a particulate model of inheritance, which stated that organisms transmit discrete heritable unites (now called genes) to their offspring. Darwin did not know about genes, but it was this paper that set the stage for understanding genetic differences on which evolution is based. 3. Explain how quantitative and discrete characters contribute to variation within a population. First of all, discrete characters are those that are mainly determined by a single gene locus wither different alleles that produced distinct phenotypes (such as the color of a flower). Quantitative characters usually result from the influence of two or more genes on a single phenotypic character. These characters can be measured at both the whole-gene level and molecular level. 4. Distinguish between average heterozygosity and nucleotide variability. Explain why average heterozygosity tends to be greater than nucleotide variability. Well, average heterozygosity is the average percent of loci that are heterozygous, and is often estimated by surveying the protein products of genes using gel electrophoresis. Nucleotide variability is measured by comparing the DNA sequence of two individuals in a population and then averaging the data from many such comparisons. 5. Define a cline. A cline is a type of geographic variationwhich accounts for the differences in genetic composition of separate populations. A cline is a graded change in character along a geographic axisthat is, one population of fish up north (in cold water) will be different than a population of the same fish down south (in warm water).

MUTATION AND SEXUAL RECOMBINATION

1. Explain why the majority of point mutations (in total DNA) are harmless. First of all, a point mutation is the change of as little as one base in a gene, and they are relatively harmless (most of the time). This is because most of the DNA in eukaryotic genomes does NOT code for protein products. Also, there is redundancy in the genetic code, and even a point mutation in a gene that encodes a protein will have no effect on the proteins function if the amino acid composition is not changed. Moreover, even if there is a change in an amino acid, this may not affect the proteins shape and function.

BIO 201 OBJECTIVES

|6

2. Explain why mutation has little quantitative effect on allele frequencies in a large population. This is because it is sexual reproduction that produces most of the genetic variation in a population from the unique combination of alleles that each individual receives. There are three mechanisms for this, crossing over, independent assortment of chromosomes, and fertilization. Even though an individuals genome is the accumulation of past mutations, it is these three mechanisms that affect allele frequency in a large population. Also, mutation rates tend to be low in plants and animals, averaging about one mutation in every 100,000 genes per generation, and they are often even lower in prokaryotes. 3. Describe the significance of transposons in the generation of genetic variability. Transposable elements are an important source of variation. Transposons are mobile segments of DNA usually associated with viruses. It is thought that viruses evolved from transposons. The activity of these transposons results in genetic variability (theres not much in the chapter on this, so do you best!). Note: horizontal gene transfer is a process that works via transposons. This would inevitably mix genetic codes, producing greater variability. 4. Explain how sexual recombination generates genetic variability. As mentioned above, it is crossing over, independent assortment of chromosomes, and fertilization that generates genetic variability in respects to sexual recombination. During meiosis, homologous chromosomes, one inherited from each parent, trade some of their alleles by crossing over. These homologous chromosomes and the alleles they carry are then distributed at random into gametes. Then, because myriad possible mating combinations exist in a population, fertilization brings together gametes of individuals that are likely to have different genetic backgrounds.

THE HARDY-WEINBERG PRINCIPLE

1. Define the terms population, species, and gene pool. a. POPULATION: a group of individuals of the same species that live in the same area and interbreed, producing fertile offspring. b. SPECIES: a population or group of populations whose members have the potential to interbreed in nature and produce viable, fertile offspring, but do not produce viable, fertile offspring with members of other such groups. c. GENE POOL: all of the alleles for all the loci in a all individuals of a population. If only one allele exists for a particular locus in a population, that allele is said to be fixed and all individuals are homozygous. If there are two or more, then individuals can either be heterozygous or homozygous. 2. Explain why meiosis and random fertilization alone will not alter the frequency of alleles or genotypes in a population. Well, you also need natural selection and evolution for the frequency to change. This is known as the Hardy-Weinberg principle, as is described below. 3. List the five conditions that must be met for a population to remain in Hardy-Weinberg equilibrium. First of all, the Hardy-Weinberg principle states that frequencies of alleles and genotypes in a population will remain constant from generation to generation, provided that only Mendelian segregation and recombination of alleles are at work. Such a gene pool is said to be in HardyWeinberg equilibrium. There are five conditions that must be met for this to occur, as listed below:

BIO 201 OBJECTIVES

|7

a. No mutations. As weve already discussed, mutations can modify the gene pool by altering,
deleting, or duplicating entire genes.

b. Random mating. If everyone mated with their family, random mixing of gametes would not
occur.

c. No natural selection. d. Extremely large population size. e. No gene flow.

4. Write the Hardy-Weinberg equation. Use the equation to calculate allele frequencies when the frequency of homozygous recessive individuals in a population is 25%. Below is the equation to use when calculating allele frequencies (the Hardy-Weinberg equation): + 2 + =1

Where p2 is the expected frequency of homozygous dominant, 2pq is the expected frequency of heterozygous, and q2 is the expected frequency of homozygous recessive. In order to calculate when homozygous recessive is at 25%, we need to do a bit of math! Since we know what q2 is, we now need to know what p is. = 1 Now that we have an equation for p we can plug in the numbers and get the answer. First we calculate what q is from q2. = 0.25 = 0.25 = 0.5 = 1 0.5 = 0.5 = 2 0.5 0.5 = 0.5

NATURAL SELECTION, GENETIC DRIFT AND GENE FLOW

1. Explain the following statement: Only natural selection leads to the adaptation of organisms to their environment. This is because selection results in alleles being passed to the next generation in proportions different from their proportions in the present generation. Also, those with traits that are better suited to their environment tend to produce more offspring that those with traits that are less well suited. 2. Explain the role of population size in genetic drift. The smaller the size the more likely it is that chance alone will cause a deviation from the predicted results. Genetic drift is the process by which allele frequencies fluctuate unpredictably from one generation to the next, and when you have a small population the effects are amplified. 3. Distinguish between the bottleneck effect and the founder effect. Well, the founder effect is when few individuals become isolated from a larger population, then establish a new population whose gene pool differs from the source population. The bottleneck effect occurs when there is a sudden change in the environment, such as a fire or flood that drastically reduces the size of a population. This causes certain alleles to either be overrepresented or under-represented, and can have substantial effects on the gene pool until the population becomes large enough that chance events have less effect. 4. Describe how gene flow can act to reduce genetic differences between adjacent populations. Well, gene flow is the transfer of alleles into or out of a population due to the movement of fertile individuals or their gametes. An example would be insects carrying pollen from one population of

BIO 201 OBJECTIVES

|8

plants to another population. Because alleles are exchanged among populations, gene flow tends to reduce the genetic differences between populations. If it is extensive enough, gene flow can result in neighboring populations combining into a single population. 5. Define relative fitness. This is the contribution an individual makes to the gene pool of the next generations, relative to the contributions of other individuals. 6. Distinguish among directional, disruptive, and stabilizing selection. Give an example of each mode of selection. a. DIRECTIONAL SELECTION: this occurs when conditions favor individuals exhibiting one extreme of a phenotypic range, thereby shifting the frequency curve for the phenotypic character in one direction or another. Example: black bears in Europe became bigger during each frigid glacial period, and then shrunk during the warmer inter-glacial periods. b. DISRUPTIVE SELECTION: this occurs when conditions favor individuals at both extremes of a phenotypic range over individuals with intermediate phenotypes. Example: the black-bellied seedcracker finches in Cameroon. They either have very large beaks (for hard seeds) or very small seeds (for soft seeds), and not many in-between. c. STABALIZING SELECTION: occurs when conditions favor the intermediate phenotype and acts against either extreme. Example: human birth weight. Babies that are either too small or too large suffer higher rates of mortality. 7. Explain how diploidy can protect a rare recessive allele from elimination by natural selection. A considerable amount of genetic variations is hidden from selection in the form of recessive alleles. These alleles can persist by propagation in heterozygous individuals, and is exposed to natural selection only when both parents carry the same recessive allele and two copies end up in the same zygote (this happens rarely). 8. Describe how heterozygote advantage and frequency dependent selection promote balanced polymorphism. Balanced polymorphism or balanced selection occurs when natural selection maintains two or more forms in a population. Heterozygote advantage occurs if individuals who are heterozygous at a particular locus have greater fitness than do both kinds of homozygotes. In frequency-dependent selection, the fitness of a phenotype declines if it becomes too common in the population. 9. Define neutral variations. Explain why natural selection does not act on these alleles. In humans, many of the nucleotide differences in noncoding sequences appear to confer no selective advantage or disadvantage and therefore are considered neutral variation. Natural selection does not act on these alleles because they have little to no effect on reproductive success. 10. List and explain at least four reasons why natural selection cannot produce perfect organisms. 1. Selection can act only on existing variations. Natural selection favors only the fittest phenotypes among those currently in the population, which may not be the ideal traits. 2. Evolution is limited by historical constraints. Each species has a legacy of descent with modification from ancestral forms. It does not scrap the ancestral anatomy and build from scratch. 3. Adaptations are often compromises. Since organisms do many things, evolution must make compromises. For instance, a seal spends a lot of time on land. It could move around better to escape predators if it had legs, but then it wouldnt be able to swim as well. 4. Chance, natural selection, and the environment interact. You get the point.

BIO 201 OBJECTIVES

|9

CHAPTER 24
THE ORIGIN OF SPECIES

WHAT IS A SPECIES?
1. Define Ernst Mayrs biological species concept.
This is the primary definition of species used in the textbook, and states: a species is a group of populations whose members have the potential to interbreed in nature and produce viable, fertile offspringbut do not produce viable, fertile offspring with members of other such groups. Reproductive compatibility unites members of a biological species.

2. Distinguish between prezygotic and postzygotic reproductive barriers.

To begin, both are forms of reproductive isolationthe existence of biological factors that impede members of two species from producing viable, fertile offspring. Prezygotic barriers block fertilization from occurring. This is accomplished in one of three ways: by impeding members of different species from attempting to mate, by preventing an attempted mating from being completed successfully, or by hindering fertilization if mating is completed successfully. Postzygoic barriers contribute to reproductive isolation after fertilization has occurred. Developmental errors may reduce survival among hybrid embryos, or problems after birth may cause hybrids to be infertile or may decrease their chance of surviving long enough to reproduce.

3. Describe several prezygotic reproductive barriers and give an example of each.

There are five such examples listed in the book, so here you go: 1. HABITAT ISOLATION: two species occupy different habitats within the same area may not encounter each other. Example: a mountain may be in the way. 2. TEMPORAL ISOLATION: species that breed during the different times of the day, different seasons, or different years cannot mix their gametes. 3. BEHAVIORAL ISOLATION: courtship rituals that attract mates and other behaviors unique to a species are effect reproductive barriers, even between closely related species. 4. MECHANICAL ISOLATION: mating is attempted, but morphological differences prevent its successful completion. 5. GAMETIC ISOLATION: sperm of one species may not be able to fertilize the eggs of another species.

4. Explain a possible cause for hybrid breakdown.

First of all, hybrid breakdown is a postzygotic reproductive barrier in which some first generation hybrids are viable and fertile, but when they mate with one another or with either parent species, offspring of the next generation are feeble or sterile. One possible cause could be the amount of recessive alleles present in the next generation.

5. Explain how hybrid breakdown maintains separate species even if fertilization occurs.
Youre on your own for this one. It may have something to do with the next generation of offspring being feeble or sterile, but not a lot is said in the book about this.

6. Describe some limitations of the biological species concept.

(1)There is no way to evaluate the reproductive isolation of fossils. (2)The concept does NOT apply to organisms that reproduce asexually. (3)Gene flow occurs between many pairs of species that are morphologically and ecologically distinct, despite the definition claiming that a species is designated by the absence of gene flow.

BIO 201 OBJECTIVES

| 10

7. Define and distinguish among the following: ecological species concept, phylogenetic species concept, and morphological species concept.
These are alternate definitions for what a species is (and, of course, there are problems with all of them). Okay, lets get to it! a. MORPHOLOGICAL SPECIES CONCEPT: this characterizes a species by body shape and other structural features. b. ECOLOGICAL SPECIES CONCEPT: this views a species in terms of its ecological niches, the sum of how members of the species interact with the nonliving parts of the environment. For example, two species of frogs differ in the foods they eat or the climate they can tolerate. c. PHYLOGENETIC SPECIES CONCEPT: this defines a species as the smallest group of individuals that share a common ancestor, forming one branch on the tree of life. The phylogenetic history of a species is traced by comparing its characteristics, such as morphology and molecular sequences, with those of other organisms.

MODES OF SPECIATION
1. Distinguish between allopatric and sympatric speciation.
In allopatric speciation, gene flow is interrupted when a population is divided into geographically isolated subpopulations. For example, a mountain range or lake divides a population. In contrast, sympatric speciation occurs in populations that live in the same geographic area, and gene flow is reduced by such factors as polyploidy, habitat differentiation, and sexual selection. Sympatric speciation is much less common than allopatric speciation.

2. Define allopatric speciation. Describe the mechanisms that may lead to genetic divergence of isolated gene pools.
As mentioned above, geographic isolation causes an interruption in gene flow. Once the population has become isolated, mutations, natural selection and genetic drift alter allele frequency. There are numerous examples in the book of this happening, including the monkey flower Mimulus guttatus, which evolved to be more copper tolerant in populations near copper mines.

3. Explain how reproductive barriers evolve. Describe an example of the evolution of a prezygotic barrier and the evolution of a postzygotic barrier.
Reproductive barriers evolve in many diverse ways. The bottleneck effect, the founder effect, polyploidy (defined below) and allopatric/sympatric speciation all contribute to the evolution of reproductive barriers. A prezygotic barrier may evolve from any of the isolating factors described above (habitat, temporal, behavior, etc). A post zygotic barrier may evolve from reduced hybrid viability, reduced hybrid fertility or hybrid breakdown.

4. Define sympatric speciation and explain how polyploidy can cause reproductive isolation.
First of all, polyploidy is a condition in which a species may originate from an accident during cell division, resulting in extra sets of chromosomes. There are two types of polyploidy (discussed below), and both are more common in plants.

5. Distinguish between autopolyploid and an allopolyploid speciation and describe examples of each. a. AUTOPOLYPLOID: is an individual that has more than two chromosome sets that are derived from
a single species. This results from a failure in cell division (going from diploid to tetraploid). This causes the tetraploid to be reproductively isolated from the diploid (they can only produce viable, fertile offspring with other tetraploidsor triploids, though with reduced fertility).

BIO 201 OBJECTIVES

| 11

b. ALLOPOLYPLOID: is when two different species interbreed and produce a hybrid offspring. Most
hybrid species are sterile because the set of chromosomes from one species cannot pair during meiosis with the set of chromosomes from the other species. However, in subsequent generations, various mechanisms can change a sterile hybrid into a fertile polyploidy, called an allopolyploid. Allopolyploids are fertile only when mating with each otherthey cannot breed with either parent species.

6. Explain how habitat differentiation has led to sympatric speciation in North American maggot flies.
This type of sympatric speciation occurs when genetic factors enable a subpopulation to exploit a habitat or resource not used by the parent population. The N.A. maggot flys original habitat was the native hawthorn tree, but about 200 years ago, some populations colonized apple trees that had been introduced by European settlers. As apples mature more quickly than hawthorn fruit, natural selection has favored apple-feeding flies with rapid development. These apple-feeding populations now show temporal isolation from the hawthorn-feeding, providing a prezygotic restriction to gene flow between the two populations.

HYBRID ZONES
1. Define the term hybrid zone. Describe the three outcomes for hybrid zones over time.
A hybrid zone is a region in which members of different species meet and mate, producing at least some offspring of mixed ancestry. This can occur when allopatric populations come back into contact with one another. There are three outcomes for hybrid zones over time, listed below: a. REINFORCEMENT: or the strengthening of reproductive barriers. This is due to the hybrid species being less fit and therefore natural selection strengthens prezygotic barriers to reproduction. b. FUSION: or the weakening of reproductive barriers. This occurs when two species contact one another in a hybrid zone, but the barriers to reproduction are not strong. This allows so much gene flow to occur that reproductive barriers weaken further and the gene pools of the two species become increasingly alike. This can cause two hybridizing species to fuse into a single species. c. STABILITY: or the continued formation of hybrid individuals. Nothing dramatic happens, and the world is at peace.

2. Describe examples that illustrate rapid and gradual speciation events.

In a punctuated, or rapid, pattern, new species change most as they branch from a parent species and then change little for the rest of their existence. The wild sunflower is an example of this. In gradual speciation, other species diverge from one another much more gradually over time. You are an example of this.

3. Explain how a small number of genetic changes may lead to speciation in plants and animals.
The example in the book goes back to the monkey flower. There are two closely related species of this flower and the reproductive barrierparticularly the barrier of pollinator choicebetween them is influenced by a relatively small number of genes. This barrier is affected by at least two loci in the flower which influences flower color, thus attracting different pollinators.

BIO 201 OBJECTIVES

| 12

CHAPTER 25
THE HISTORY OF LIFE ON EARTH

THE ORIGIN OF LIFE

1. Describe evidence that suggests that RNA was the first genetic material. Explain the significance of the discovery of ribozymes.
RNA is thought to be the first genetic material due to its ability to carry out enzyme-like catalytic functions, called ribozymes. These ribozymes are capable of self-replication (thanks to natural selection) due to the fact that single-stranded RNA molecules are more stable and replicate faster with fewer errors than other sequences. This means that an RNA molecule will leave more descendant molecules, provided it exists in the right environment.

2. Describe how natural selection may have worked in an early RNA world.
See the above question. As an elaboration, in certain environments, RNA molecules will assume a variety of specific three-dimensional shapes. Now, due to replication errors, the descendants of an RNA molecule are not always exact. These errors can sometimes cause the descendant RNA molecule to assume a different, more stable shape, making that molecule more adept at self-replication than the ancestral sequence.

3. Describe how natural selection may have favored the proliferation of stable protobionts with self-replicating, catalytic RNA.
These protobionts with catalytic RNA in them would be different from those that carried RNA without such capabilitiesmainly that they could exploit their environment and resources better. Now, if the protobiont could grow, split, and pass its catalytic RNA molecules to its daughters, those daughters could also exploit their resources better. This would cause the population of protobionts with catalytic RNA to increase.

MAJOR EVENTS IN THE HISTORY OF LIFE ON EARTH

1. Explain why the fossil record provides an incomplete chronicle of evolutionary change.
Many of Earths organisms probably did not die in the right place at the right time to be preserved as fossils. Of those fossils formed, many were destroyed by later geologic processes, and only a fraction of the others have been discovered. As a result, the known fossil record is biased in favor of species that existed for a long time, were abundant and widespread in certain kinds of environments, and had hard shells, skeletons, or other parts that facilitated their fossilization.

2. Explain how radiometric dating can be used to determine the absolute age of rock strata. Explain how magnetism can be used to date rock strata.
Radiometric dating is based on the decay of radioactive isotopes. Radioactive isotopes decay to daughter isotopes at a constant rate, expressed by the half-life, or the time required for 50% of the parent isotope to decay. Each radioactive isotope has a characteristic half-lifeone that is NOT affected by temperature, pressure, or other such environmental variables. Carbon-14/Carbon-12 ratio is used to determine the age of fossils up to about 75,000 years old. Dating the rocks around the fossils works for those that are older than 75,000 years old (most organisms do not use radioisotopes that have long half-lives, such as uranium-238, to build their bones or shells). Now, the magnetism of rocks can also provide dating

BIO 201 OBJECTIVES

| 13

information. During the formation of rocks, ion particles align themselves with Earths magnetic field. When the rock hardens, the ions are frozen in time. We know that Earths magnetic field has reversed repeatedly in the past, so to the reversals in one location can be matched with corresponding patters elsewhere.

3. Describe the major events in Earth's history from its origin until 2 billion years ago. In particular, note when Earth first formed, when life first evolved, and what forms of life existed in each eon. Archaean Eon
4.6 billion years ago: Earth forms. 3.5 billion years ago: oldest fossils of prokaryotic cells appear.

Proterozoic Eon
2.1 billion years ago: oldest fossils of eukaryotic cells appear. 635 million years ago: diverse algae and soft-bodied invertebrate animals appear.

Phanerozoic Eon (we live in this eon)

Consists of the Paleozoic, Mesozoic and Cenozoic eras. Started 542 million years ago. Encompasses multicellular eukaryotic life. 251 million years ago: dinosaurs evolve. 65.5 million years ago: major radiation of mammals, birds and pollinating insects. 1.8 million years ago: humans appear. Study the images from the textbook and notes.

4. Name the reactions that have produced O2 on Earth. Describe the accumulation of atmospheric O2 over time and the effects of this accumulation on living things.
Most atmospheric oxygen is of biological origin, produced during the water-splitting step of photosynthesis. Oxygen probably dissolved in the surrounding water when it was first produced. When it reached a high enough level it began to react with dissolved iron, precipitating iron oxide, which accumulated as sediment. Once all the dissolved iron precipitated, additional oxygen dissolved in the water until the seas and lakes became saturated. After this, oxygen began to gas out of the water and enter the atmosphere. O2 increased gradually from about 2.7 to 2.2 billion years ago, but then shot up relatively rapidly to more than 10% of its present level. This is when most prokaryotic group probably died, and when cellular respiration evolved.

5. Explain the endosymbiotic theory for the evolution of the eukaryotic cell. Describe the evidence that supports this theory.
Endosymbiosis is a model which posits that mitochondria and plastids (chloroplasts and related organelles) were formerly small prokaryotes that began living within larger cells. The term endosymbiont refers to a cell that lives within another cell, call the host cell. Evidence: the inner membranes of both organelles (mitochondria and plastids) have enzymes and transport systems that are homologous to those found in the plasma membrane of living prokaryotes. They replicate by a splitting process that is similar to that of certain prokaryotes. They have a single, circular DNA molecule that, like the chromosomes of bacteria, is not associated with histones or large amounts of other proteins. They have ribosomes (which are more similar to prokaryotic ribosomesin terms of size, nucleotide sequence, and sensitivity to certain antibiotics) and other cellular machinery needed to transcribe and translate their DNA into proteins.

BIO 201 OBJECTIVES

| 14

6. State the evidence that suggests that the common ancestor of multicellular eukaryotes lived 1.5 billion years ago.
It is a rough agreement with the fossil record. The oldest known fossils of multicellular eukaryotes are of relatively small algae that lived about 1.2 billion years ago. Larger and more diverse multicellular eukaryotes do not appear in the fossil record until about 565 million years ago.

7. Explain the possible significance of Snowball Earth in the history of life on Earth.
The Snowball Earth hypothesis suggests that, during severe ice ages, most life would have been confined to areas near deep-sea vents and hot springs or to equatorial regions of the ocean that lacked ice cover. The fossil record of the first major diversification of multicellular eukaryotes corresponds roughly to the time when snowball Earth thawed. Life could spread and diversify with all that ice gone.

8. Briefly describe the Cambrian explosion

This is the sudden appearance of many phyla of living animals. It happened during the Cambrian period (535-525 million years ago). Prior to the Cambrian explosion, all large animals were soft-bodied. Afterwards, animals developed new defensive adaptations such as heavy body armor and sharp spines.

9. Describe the key evolutionary adaptations that arose as life colonized land.
The gradual evolutionary venture out of aquatic environments was associated with adaptations that made it possible to reproduce on land and that helped prevent dehydration. Plants developed roots and leaves that aid in the absorption of minerals from the soil.

CONTINENTAL DRIFT, MASS EXTINCTION, AND ADAPTIVE RADIATION

1. Describe the mass extinctions that ended the Permian and Cretaceous periods.
a. PERMIAN PERIOD: this extinction (251 million years ago) claimed about 96% of marine animal species and drastically altered life in the ocean. Terrestrial life was also affected. It occurred at the same time of enormous volcanic eruptions in what is now Siberia. These eruptions spewed enough lava, ash and CO2 to warm the global climate by an estimated 6C. The temperature differences between the equator and the poles would have slowed the mixing of ocean water, which in turn would have reduced the amount of oxygen available to marine organisms. b. CRETACEOUS PERIOD: this extinction (65 million years ago) claimed about 50% of all marine species and eliminated many families of terrestrial plans and animals, including most of the dinosaurs. It is thought that a giant asteroid or comet collided with Earth, blotting out the sunlight with its huge cloud of debris. This would have severely disturbed the global climate. Evidence for this comes from a thin layer of clay enriched in iridium (a element that is rare on Earth) that separates sediments from the Mesozoic and Cenozoic eras, and from a giant crater off the Yucatan cost of Mexico.

2. A sixth mass extinction may be currently underway. Explain this statement.

Not much to explain. People smarter than you or I think we are causing a mass extinction via habitat destruction and modification to the global environment.

3. Define adaptive radiation. Describe, with suitable examples, three circumstances under which adaptive radiation may occur.
Adaptive radiation is a period of evolutionary change in which groups of organisms form many new species whose adaptations allow them to fill different ecological roles, or niches, in their communities. Adaptive radiation can be a direct consequence of colonization of an isolated region, evolutionary innovation, vacant ecological niches, and an adaptive radiation in a group of organisms (such as plants) that another group uses as food. Large scale adaptive radiation occurred after each of the big five mass extinction events. After dinosaurs disappeared, mammals expanded greatly in both diversity and size,

BIO 201 OBJECTIVES

| 15

filling the ecological roles once occupied by terrestrial dinosaurs. Regional adaptive radiations occur over a more limited geographic area. Such radiations can be initiated when a few organisms make their way to a new, often distant location in which they face relatively little competition from other organisms. The Hawaiian archipelago is a great showcase for thisas are the finches from the Galapagos Islands.

THE EVOLUTION OF COMPLEX STRUCTURES

1. Explain why evolutionary change is not goal-directed.
An evolutionary trend does not imply that there is some intrinsic drive toward a particular phenotype. Evolution is the result of the interactions between organisms and their current environments; if environmental conditions change, an evolutionary trend may cease or even reverse itself. The cumulative effect of these ongoing interactions between organisms and their environments is enormous: it is through them that the staggering diversity of life has arisen. The classic example of this is the modern horse. Its evolutionary history is filled with evolutionary dead ends.

BIO 201 OBJECTIVES

| 16

CHAPTER 26
PHYLOGENY AND THE TREE OF LIFE

PHYLOGENIES SHOW EVOLUTIONARY RELATIONSHIPS

1. Distinguish between phylogeny and systematics.
Systematics is the reconstruction and study of phylogenies. A phylogeny is the evolutionary history of a species or group of species. In contrast, systematics is a discipline focused on classifying organisms and determining their evolutionary relationships. Data like fossils and molecules and genes are used to infer evolutionary relationships. Taxonomy is the process by which organisms are named and classified.

2. Explain the following characteristics of the Linnaean system of classification: a. Binomial nomenclature: The first part of the binomial is the name of the genus to which the species
belongs. In our case it is Homo. The second part, called the specific epithet, is unique for each species within a genus. In our case it is sapiens. b. Hierarchical classification: species are grouped into a hierarchy of increasingly inclusive categories, as seen below:

Domain Kingdom Phylum Class Order Family Genus Species

3. Explain the justification for the proposal to replace Linnaean classification with phylocode designations for monophyletic taxa.
The categories of the Linnaean classification system provide little information about phylogenythat is, the evolutionary relationships between similar species. Such difficulties have led some sytematists to propose that classification be based entirely on evolutionary relationships. PhyloCode is an approach that only names groups that include a common ancestor and all of its descendants.

4. Explain the statement: A phylogenetic tree represents a hypothesis about evolutionary relationships.
A phylogenetic tree depicts the evolutionary relationships among species as a series of dichotomies, or two way branch points. Each branch point represents the divergence of two evolutionary lineages from a common ancestor.

5. Explain why it is crucial to distinguish between homology and analogy before selecting characters to use in the reconstruction of phylogeny. Describe how homology and analogy can be distinguished from each other.
Homologies are similarities (can be structural, such as arms, legs, wings etc) in species due to a shared ancestry. Analogies (or homoplasies) are similarities due to convergent evolution rather than shared

BIO 201 OBJECTIVES

| 17

ancestry. In order to classify a species correctly, one must first distinguish between the two, or the phylogenetic tree will be incorrect. As an example: a bat has wingsa structural trait it shares with birds due to convergent evolutionand is therefore not grouped with birds. However, bird wings and bat wings are homologous in respect to vertebrate forelimbs. This is because birds and bats have a common ancestor that had forelimbs.

THE CONSTRUCTION OF PHYLOGENETIC TREES

1. Define a clade. Distinguish between a monophyletic clade and paraphyletic and polyphyletic groupings of species. a. CLADE: In the approach to systematics called cladistics, common ancestry is the primary criterion
used to classify organisms. Using this methodology, biologists place species into groups called clades, each of which includes an ancestral species and all of its descendants. If these conditions are met, you have a monophyletic clade. b. PARAPHYLETIC GROUPING: This group consists of an ancestral species and some, but not all, of its descendants. c. POLYPHYLETIC GROUPING: This group includes taxa with different ancestorsor a set of species that does not include the most recent common ancestor of the set.

2. Distinguish between shared ancestral characters and shared derived characters. Synapomorphies and symplesiomorphies. a. A shared ancestral character is a character that originated in an ancestor of a taxon. In contrast, a
shared derived character is a character shared by all species of a taxon but not found in an ancestor. Our backbone is an ancestral character (all mammals have a backbone, as did our ancestor), and our hair is a derived character (all mammals have hair, but our shared ancestor did not). b. Synapomorphies are shared, derived character states. Symplesiomorphies are share, ancestral character states.

3. Explain why shared derived characters are useful in establishing a phylogeny, while shared ancestral characters are not.
Shared derived characters are unique to particular clades. Because all features of organisms arose at some point in history of life, it should be possible to determine the clade in which each shared derived character first appeared and use that information to infer evolutionary relationships.

4. Describe the evidence that suggests there is a universal tree of life.

The reconstruction of the tree of life is based largely on sequence comparisons of rRNA genes, which code for the RNA parts of ribosomes. Because ribosomes are fundamental to the workings of the cell, rRNA genes have evolved so slowly that homologies between distantly related organisms can still be detected making these genes very useful for determining evolutionary relationships among deep branches in the history of life. Also, horizontal gene transfer, as seen below:

5. Explain why the occurrence of horizontal gene transfer has led to the suggestion that the history of life should be represented by a ring rather than a tree.
Some biologists hypothesize that the first eukaryote may have arisen through a fusion between an ancestral bacterium and an ancestral Archaean. Because phylogenetic trees are based on the assumption that genes are passed vertically from one generation to the next, the occurrence of such horizontal transfer events helps to explain why universal trees built using different genes can give inconsistent results (and also explains why the tree is ring-like).