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Evolution and the Diversity of Life What is Evolution? In the simplest biological terms evolution is defined as change over time. However, it is much more than that. Who was Charles Darwin? • Charles Darwin first published an explanation of how species changed over time, or evolved. • Darwin argued that contemporary species arose from ancestors •Through a process of “descent with modification,” with natural selection as the mechanism. How did he explain evolution? • Natural Selection: Occurs in natural populations when organisms with certain traits pass those traits on to the next generation. • The following are required for natural selection to take place:` 1. Organisms can change over generations. 2. Individuals with certain heritable traits produce more surviving offspring than others. 3. The result of natural selection is evolutionary adaptation. Natural Selection • Natural selection is the mechanism for evolution. • It only acts on populations. • It changes the frequency of genes in a population – not in an individual. What traits are usually selected? • Traits that allow the organism to survive and reproduce. http://evolution.berkeley.edu/evolibrary/article//bergstrom_02 Peppered Moth Example • Originally White • Now more black than white • Causes – Industrial Pollution • Soot covered trees How does natural selection happen? Evolution only occurs when there is a change in gene frequency within a population over time. Change Over Time Evidence for evolution Fossils evidence of an organism that lived some time in the past. The Fossil record The ordered sequence of fossils as they appear in rock layers which reveals the appearance of organisms in a historical sequence. Fossils Fossils show a succession from simple forms early in the fossil record to complex forms that appear much later. Fossils and Geological Time • Scientists will date the rocks in an attempt to determine the age of the fossil. • Two ways to do this: – Relative dating: When you compare the depth of the layer of rock the fossil was formed in relative to other layers of rock. – Radiometric dating: Uses isotopes. Relative Dating • Relative dating uses layers to determine age by order of appearance. • Does not give you a specific age in years, just relative to the layers above and below the fossil. • Compare rock layers to layers of clothing in your laundry hamper…Oldest on the bottom (unless disturbed). Radiometric Dating • Technique involves using radioactive isotopes, which are atoms with unstable nuclei that break down (decay) over time, giving off radiation. Biogeography Is the study of the geographic distribution of species. Convergent and Divergent Evolution • Divergent – Organisms with the same ancestral genetic heritage migrate to different habitats and evolve into species with different external forms and structures, but continue to use the same type of habitats • Ex) Ostrich • Convergent – The process by which species evolve in different places of times and, although they have different genetic heritages, develop similar external forms and structures as a result of adaptation to similar environments • Ex) shapes of sharks Divergent Evolution Convergent Evolution • occurs when similar environmental pressures and natural selection – Produce similar (analogous) adaptations in organisms from different evolutionary Figure 25.5 lineages Homologous Structures – structural features with a common evolutionary origin – shared by related species. Although used for such different functions as throwing, swimming, and flying, the same basic structural plan is evident in them all. In each case, the bone shown in color is the radius. Analogous StructuresStructural features which serve the same function in different species, but they evolved independently. Ex: Butterfly wings, Bat wings Bird wings • FIGURE 16.2 The Bones Are Homologous, the Wings Are Not The supporting bone structures of both bat wings and bird wings are derived from a common tetrapod (four-limbed) ancestor and are thus homologous. However, the wings themselves—an adaptation for flight— evolved independently in the two groups. Vestigial Structurescharacteristics of organisms that have lost all or most of their original functions. Humans have a vestigial tailbone. Vestigial toes in the horse. Vestigial limbs in whales and snakes. Structural Adaptations A. Mimicry: A structural adaptation that protects an organism by copying the appearance of another species. – Example: Gopher snake B. Camouflage – A structural adaptation that allows an organism to blend in with its environment. – Example: Snowshoe hare Gopher Snakes? The one on the left is a rattlesnake, the one on the right is a gopher snake. Snowshoe Hare Summer Winter Embryology – Early embryos of very different organisms closely resemble each other. Morphology: the study of animal form. How animal parts function together to make them run, fly, swim, eat, survive. Functional morphology: how a structure works. Ecological morphology: how a structure is used in nature. Molecular Data: Causes of genetic variation in populations at the molecular level (DNA, RNA and proteins). • Can be used for comparison with all organisms. • Provides tons of information, including tracking mutations. Molecular clocks help track evolutionary time Molecular Clocks • The molecular clock – Is a yardstick for measuring the absolute time of evolutionary change based on the observation that some genes and other regions of genomes appear to evolve at constant rates Neutral Theory • Neutral theory states that – Much evolutionary change in genes and proteins has no effect on fitness and therefore is not influenced by Darwinian selection – And that the rate of molecular change in these genes and proteins should be regular like a clock Difficulties with Molecular Clocks • The molecular clock – Does not run as smoothly as neutral theory predicts Applying a Molecular Clock: The Origin of HIV • Phylogenetic analysis shows that HIV – Is descended from viruses that infect chimpanzees and other primates • A comparison of HIV samples from throughout the epidemic – Has shown that the virus has evolved in a remarkably clocklike fashion Molecular Clock of Disease • Research a zoonotic disease – Determine when, where, and how a disease first entered a human population. – Phylogenetic Analysis – Present and Global diversity – How would the evolution of your disease help epidemiologist to identify how disease is spread and how to prevent future outbreaks? Evolutionary Tree/ Cladogram: Adaptations and Natural Selection Vocabulary Gene Pool: The total number of genes present in a population. Allelic Frequency: the frequency of an allele for a specific trait. Genetic equilibrium: when the allelic frequency of alleles remains the same over time. Vocabulary Darwinian Fitness: the contribution an individual makes to the gene pool of the next generation relative to the contributions of other individuals. Changes in Populations Gene variations: Mutations-changes in DNA Gene flow-movement of genes from one population to another Reproduction-introduces new gene combinations. • Genetic variations in populations – Contribute to evolution Inherited Variation • Individuals possessing alleles suited for their environment will reproduce and population will continue because of their success. • Population genetics provides a foundation for studying evolution • Microevolution – Is change in the genetic makeup of a population from generation to generation The Modern Synthesis • 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 individuals that are capable of interbreeding and producing fertile offspring MAP AREA • Fairbanks Fortymile herd range • Whitehorse • The gene pool – Is the total aggregate of genes in a population at any one time – Consists of all gene loci in all individuals of the population 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 A finite supply of environmental resources • As population increases, each individual has a harder time getting resources needed to survive. • Birth rate decreases, mortality increases, overall growth rate slows. • Mutation and sexual recombination produce the variation that makes evolution possible • Two processes, mutation and sexual recombination – Produce the variation in gene pools that contributes to differences among individuals Mutation – Are changes in the nucleotide sequence of DNA – Cause new genes and alleles to arise Point Mutations • A point mutation – Is a change in one base in a gene – Can have a significant impact on phenotype – Is usually harmless, but may have an adaptive impact Mutations That Alter Gene Number or Sequence • Chromosomal mutations that affect many loci – Are almost certain to be harmful – May be neutral and even beneficial • Gene duplication – Duplicates chromosome segments 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 Natural Selection • Differential success in reproduction – Results in certain alleles being passed to the next generation in greater proportions 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 CWCW CRCR CRCR Only 5 of 10 plants leave offspring CRCW CWCW CRCR CRCR CRCW CWCW CRCR CRCW CRCW CRCR CWCW CRCW CRCR CRCR CRCW Generation 1 p (frequency of CR) = 0.7 q (frequency of CW) = 0.3 Only 2 of 10 plants leave offspring CRCR CRCR CRCR CRCR CRCR CRCR CRCR CRCR CRCW CRCW Generation 2 p = 0.5 q = 0.5 Figure 23.7 CRCR CRCR Generation 3 p = 1.0 q = 0.0 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 • Understanding the bottleneck effect – Can increase understanding of how human activity affects other species (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 • Natural selection – primary mechanism of adaptive evolution • Natural selection – Accumulates and maintains favorable genotypes in a population • Polymorphism • Phenotypic polymorphism – Describes a population in which two or more distinct morphs for a character are each represented in high enough frequencies to be readily noticeable • Genetic polymorphisms – Are the heritable components of characters that occur along a continuum in a population • Genetic variation – Occurs in individuals in populations of all species – Is not always heritable (a) Map butterflies that emerge in spring: orange and brown (b) Map butterflies that emerge in late summer: black and white • Measuring Genetic Variation • Population geneticists – Measure the number of polymorphisms in a population by determining the amount of heterozygosity at the gene level and the molecular level • Average heterozygosity – Measures the average percent of loci that are heterozygous in 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 8.11 9.12 10.16 13.17 1 2.19 3.8 4.16 9.10 11.12 13.17 15.18 6 7.15 19 XX 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 EXPERIMENT Researchers observed that the average size Mean height (cm) 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. Atitude (m) 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. 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. Sierra Nevada Range Great Basin Plateau 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 Changes in Populations Reproductive success/fitness: Individuals best suited for their environment leave more offspring, changing the genetic makeup of a population. 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 What Is a Species? – A population or group of populations whose members have the potential to interbreed and produce fertile offspring. How does speciation occur? • Speciation occurs only when there are reproductive barriers between the isolated population and its parent population. Geographic Isolation • A population becomes divided into smaller populations due to unfavorable habitats which keeps them from mating. • Also called Allopatric Speciation Behavioral Isolation • Two populations acquire different mating rituals and behaviors. • Also called Sympatric Speciation Punctuated Equilibrium Species most often diverge in spurts of relatively sudden change. Changes in Populations Selection advantage: The environment determines which genetic variations are favorable, those are passed on to the population as a whole. Directional, Disruptive, and Stabilizing Selection • Selection – Favors certain genotypes by acting on the phenotypes of certain organisms • Three modes of selection are – Directional – Disruptive – Stabilizing • The three modes of selection Original population Original population Evolved population (a) Directional selection shifts the overall makeup of the population by favoring variants at one extreme of the distribution. In this case, darker mice are favored because they live among dark rocks and a darker fur color conceals them from predators. Phenotypes (fur color) (b) Disruptive selection favors variants at both ends of the distribution. These mice have colonized a patchy habitat made up of light and dark rocks, with the result that mice of an intermediate color are at a disadvantage. (c) Stabilizing selection removes extreme variants from the population and preserves intermediate types. If the environment consists of rocks of an intermediate color, both light and dark mice will be selected against. The Preservation of Genetic Variation • Various mechanisms help to preserve genetic variation in a population Diploidy • Diploidy – Maintains genetic variation in the form of hidden recessive alleles Balancing Selection • Balancing selection – Occurs when natural selection maintains stable frequencies of two or more phenotypic forms in a population – Leads to a state called balanced polymorphism Heterozygote Advantage • Some individuals who are heterozygous at a particular locus – Have greater fitness than homozygotes • Natural selection – Will tend to maintain two or more alleles at that locus • The sickle-cell allele – Causes mutations in hemoglobin but also confers malaria resistance – Exemplifies the heterozygote advantage Frequencies of the sickle-cell allele 0–2.5% 2.5–5.0% Distribution of malaria caused by Plasmodium falciparum (a protozoan) 5.0–7.5% 7.5–10.0% 10.0–12.5% >12.5% • Frequency-Dependent Selection • In frequency-dependent selection – The fitness of any morph declines if it becomes too common in the population Neutral Variation • Neutral variation – Is genetic variation that appears to confer no selective advantage Sexual Selection • Sexual selection – Is natural selection for mating success • Intrasexual selection – Is a direct competition among individuals of one sex for mates of the opposite sex • Intersexual selection – Occurs when individuals of one sex (usually females) are choosy in selecting their mates from individuals of the other sex – May depend on the showiness of the male’s appearance The Evolutionary Enigma of Sexual Reproduction – Produces fewer reproductive offspring than asexual reproduction, a so-called reproductive handicap Asexual reproduction Female Generation 1 Sexual reproduction Female Generation 2 Male Generation 3 Generation 4 • If sexual reproduction is a handicap, why has it persisted? – It produces genetic variation that may aid in disease resistance Why Natural Selection Cannot Fashion Perfect Organisms • Evolution is limited by historical constraints • Adaptations are often compromises • Mendelian inheritance – Preserves genetic variation in a population Generation 1 CW CW genotype CRCR genotype Plants mate Generation 2 All CRCW (all pink flowers) 50% CR gametes 50% CW gametes Come together at random Generation 3 25% CRCR 50% CRCW 50% CR gametes 25% CWCW 50% CW gametes Come together at random Generation 4 25% CRCR Figure 23.4 50% CRCW 25% CWCW Alleles segregate, and subsequent generations also have three types of flowers in the same proportions Preservation of Allele Frequencies • In a given population where gametes contribute to the next generation randomly, allele frequencies will not change Hardy-Weinberg Equilibrium • Hardy-Weinberg equilibrium – Describes a population in which random mating occurs – Describes a population where allele frequencies do not change • A population in Hardy-Weinberg equilibrium Gametes for each generation are drawn at random from the gene pool of the previous generation: 80% CR (p = 0.8) 20% CW (q = 0.2) Sperm CR (80%) CW (20%) CR (80%) pq p2 64% CRCR CW (20%) Eggs p2 16% CRCW 16% CRCW qp 4% CW CW 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: 16% CR from 64% CR from + CRCW homozygotes CRCR homozygotes 4% CW from CWCW 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: Figure 23.5 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 – And p2 and q2 represent the frequencies of the homozygous genotypes and 2pq represents the frequency of the heterozygous genotype Conditions for HardyWeinberg Equilibrium • The Hardy-Weinberg theorem – Describes a hypothetical population • In real populations – Allele and genotype frequencies do change over time Organizing Life’s Diversity Taxonomy- the identification, naming, and classification of species. Classification – grouping of organisms based on similarities. Aristotle:developed first biological classification system • Divided all organisms into two groups: plants or animals • Each animal according to where it lived – Land – Water – Air • Each plant according to – Size – Structure Carolus von Linneaus mid-1700’s • Determined levels of classification – He used similarities in structure to determine relationships among organisms. • Binomial Nomenclature: Two-word naming system (LATIN) Taxonomic Hierarchy – From largest to smallest: • Kingdom-broad • Phylum • Class • Order • Family • Genus • Species King Phillip Cut Open Five Green Snakes Naming organisms First word: genus, Second word: species Ex: scientific name for leopard is Panthera pardus Dichotomous keys • Dichotomous key: a tool that helps find the identity of organisms/items. – Ex: trees, flowers, mammals, fish and rocks. • Keys consist of a series of choices that lead the user to the correct name of a given item. • Dichotomous means "divided into two parts". Dichotomous keys always give two choices in each step. Examples: Where do living things come from? • One theory held in the past is referred to as Spontaneous generation. This theory held that nonliving material can produce life. • Fancesco Redi disproved this theory with an experiment now known as the Redi Experiment. This experiment helped to disprove the spontaneous generation of large organisms, but not microorganisms. Redi’s (1626-1697) Experiments Evidence against spontaneous generation: 1. Unsealed – maggots on meat 2. Sealed – no maggots on meat 3. Gauze – few maggots on gauze, none on meat The Origin of Life • Likewise, Louis Pasteur designed an experiment to disprove the spontaneous generation of microorganisms. Louis Pasteur (1822-1895) Pasteur's Problem • Hypothesis: Microbes come from cells of organisms on dust particles in the air; not the air itself. • Pasteur put broth into several special S-shaped flasks • Each flask was boiled and placed at various locations Pasteur's Experiment Step 1 • S-shaped Flask • Filled with broth • The special shaped was intended to trap any dust particles containing bacteria Pasteur's Experiment Step 2 • Flasks boiled • Microbes Killed Pasteur's Experiment Step 3 • Flask left at various locations • Did not turn cloudy • Microbes not found • Notice the dust that collected in the neck of the flask Pasteur's Experimental Results Question: So what do Pasteur's experimental results mean? So what now? The Theory of Biogenesis • Pasteur’s S-shaped flask kept microbes out but let air in. • Proved microbes only come from other microbes (life from life) - biogenesis Figure 1.3 The Origin of Life • Biogenesis – All living organisms must come from other living organisms. • Simple Prokaryotic cells were probably the first cells and over time the would have evolved into Eukaryotic cells. Evidence for this is seen in mitochondrial DNA. Table 14.1 Mass Extinctions and Explosive Diversifications of Life – The fossil record reveals an episodic history, • With long, relatively stable periods punctuated by briefer intervals when the turnover in species composition was much more extensive. Copyright © 2007 Pearson Education Inc., publishing as Pearson Benjamin Cummings – Extinction is inevitable in a changing world and occurs all the time. • However, extinction rates have not been steady. – Extinctions typically eliminate various species of organisms • And are followed by explosive diversifications of organisms. How many extinctions have happened? Are we in an extinction now? Classification and Phylogeny – The goal of classification is to reflect phylogeny, the evolutionary history of a species. Sorting Homology from Analogy – Homologous structures • What are they? • Are one of the best sources of information about phylogenetic relationships. – Convergent evolution • Involves superficially similar structures in unrelated organisms based on natural selection. – Analogy • Is similarity due to convergence. Molecular Biology as a Tool in Systematics – Molecular systematics • Compares DNA and amino acid sequences between organisms. • Can reveal evolutionary relationships. The Cladistic Revolution – Cladistics • Is the scientific search for clades, • Clades are distinctive branches in the history of life. – Cladistics • Has ______________ traditional classification of some organisms. Arranging Life into Kingdoms: A Work in Progress – Linnaeus designed a two-kingdom system of classification, • Which was replaced by a five-kingdom system in the mid-20th century. • Now we use a six-kingdom system, which may change again! Copyright © 2007 Pearson Education Inc., publishing as Pearson Benjamin Cummings – In the late 20th century, • Molecular studies and cladistics led to the development of a three-domain system. Classification Schemes Tools! • Dichotomous Key: Allows people to identify organisms in the real world based on a series of two choices at each step. Identify This Tree http://www.uwsp.edu/cnr/leaf/Treekey/tkframe.htm Dichotomous Key Activity! • Are you ready to try this in more detail?