EVOLUTION AND NATURAL SELECTION CHAPTER 14 Darwin’s Voyage on HMS Beagle • Charles Darwin (1809– 1882). • English naturalist who first suggested an explanation for why evolution occurred. • Published in 1859, On the Origin of Species by Means of Natural Selection. Darwin’s Voyage on HMS Beagle • Charles Darwin’s work on evolution challenged established worldviews. • Darwin proposed a mechanism for evolutionary change called natural selection. • Darwin’s hypothesis for evolutionary change, after much testing, eventually became accepted as theory. Darwin’s Voyage on HMS Beagle • Darwin voyaged from 1831–1836 on the HMS Beagle, a ship mapping the world’s coastlines. • Darwin observed firsthand different plants and animals in various locales. • These observations played an important role in the development of his thoughts about the nature of life on earth. THE FIVE-YEAR VOYAGE OF HMS BEAGLE North America Asia North Pacific Ocean North Atlantic Ocean Canary Islands Cape Verde Islands Marquesas Indian Ocean South America Galápagos Islands Society Islands Africa Philippine Islands Keeling Islands Equator Madagascar Bahia Ascension Valparaiso St. Helena Rio de Janeiro Montevideo Buenos Aires Straits of Magellan Cape Horn Mauritius Bourbon Island Port Desire Falkland IIslands Tierra del Fuego Australia Friendly Islands Sydney Cape of Good Hope King George’s Sound Hobart South Atlantic Ocean New Zealand Darwin’s Evidence • Darwin made several observations that helped lead him to believe that species evolve rather than remain fixed. 1) Fossils of extinct organisms resembled those of living organisms. 2) Geographical patterns suggested that organismal lineages change gradually as individuals move into new habitats. 3) Islands have diverse animals and plants that are related to, yet different from, their mainland sources. FOSSIL EVIDENCE OF EVOLUTION Armadillo Glyptodont Four Galápagos finches and what they eat Large ground finch (seeds) Vegetarian finch (buds) Cactus finch (cactus fruits and flowers) Woodpecker finch (insects) • Darwin observed that, although all the finches shared a common ancestor, their beak sizes had evolved to suit their food. Darwin termed this “descent with modification.” THE THEORY OF NATURAL SELECTION • Thomas Malthus’ Essay on the Principle of Population (1798) provided Darwin with a key insight. • While human populations tend to increase geometrically, the capacity for humans to feed this population only grows arithmetically. GEOMETRIC AND ARITHMETIC PROGRESSIONS 54 Geometric progression 18 Arithmetic progression 6 2 4 6 8 • What happens to a population when this gap between population density and resource availability gets wider? EXPANDING ON MALTHUS • Darwin expanded Malthus’ view to include every organism. • All organisms have the capacity to overreproduce. • Only a limited number of these offspring survive and produce the next generation. THE THEORY OF NATURAL SELECTION • Darwin associated survivors with having certain physical, behavioral, or other attributes that help them to live in their environment. • By surviving, they can pass their favorable characteristics on to their own offspring. THE THEORY OF NATURAL SELECTION • Darwin envisioned the frequency of favorable characteristics increasing in a population through a process called natural selection. • Favorable characteristics are specific to an environment; they may be favored in one but not in another. • Organisms whose characteristics are best suited to their particular environment survive more often and leave more offspring. THE THEORY OF NATURAL SELECTION • Darwin’s selection concept is often referred to as the “survival of the fittest.” • Darwinian fitness does not always refer to the biggest or the strongest. • Fitness, in evolutionary theory, refers to organisms who, due to their characteristics, survive more often and leave more offspring. THE THEORY OF NATURAL SELECTION • Domesticated animals evolved through selective breeding for certain traits that breeders preferred. • The resulting differences between breeds of domesticated species are more extreme than what exists in nature. • Darwin termed this form of selection artificial selection, because breeders determined which traits were successful, rather than nature. THE THEORY OF NATURAL SELECTION • Darwin drafted his ideas in 1842 but hesitated to publish them for 16 years. • Another researcher, Alfred Russel Wallace, sent an essay to Darwin outlining a theory of evolution by natural selection. • Wallace had independently arrived at the same mechanism for evolution that Darwin had. • Lyell and Hooker arranged for a joint presentation of their ideas in London. • Darwin finally published On the Origin of Species in 1859. THE THEORY OF NATURAL SELECTION • Darwin’s 1859 publication ignited controversy. • But the scientific community soon accepted Darwin’s arguments. The Beaks of Darwin’s Finches • Darwin’s finches are a closely related group of distinct species. • All the birds are similar to each other except for the shape of their beaks. • Genetic differences account for the physical differences in the beaks. • Birds with larger beaks make more of a protein called BMP4. A DIVERSITY OF FINCHES ON A SINGLE ISLAND Cactus finch (Geospiza scandens) Warbler finch (Certhidea olivacea) Sharp-beaked finch (G. difficilis) Woodpecker finch (Cactospiza pallida) Small insectivorous tree finch (Camarhynchus parvulus) Small ground finch (G. fuliginosa) Cactus eater Large insectivorous tree finch (Camarhynchus psittacula) Medium ground finch (G. fortis) Insect eaters Seed eaters Vegetarian tree finch (Platyspiza crassirostris) Bud eater Large ground finch (G. magnirostris) The Beaks of Darwin’s Finches • Darwin supposed that the birds evolved from a single ancestor to become individual species who specialized in particular foods. The Beaks of Darwin’s Finches • Peter and Rosemary Grant studied the medium ground finch on the island of Daphne Major in the Galápagos. • They measured beak shape over many years and recorded feeding preferences. The Beaks of Darwin’s Finches • The finches preferred to feed on small, tender seeds. • The finches switched to larger, harder-tocrack seeds when the small seeds became hard to find. • Beak depth increased when only large, tough seeds were available. Beak depth EVIDENCE THAT NATURAL SELECTION ALTERS BEAK SIZE IN GEOSPIZA FORTIS Wet year Dry year Dry year Dry year 1977 1980 1982 1984 23 The Beaks of Darwin’s Finches • The Grants’ work with the medium ground finch is an example of evolution in action. • Average beak depth increased after a drought. • Only large-beaked birds were able to crush the bigger seeds and survive to make the next generation. • When wet periods returned, smaller beaks prevailed at handling the then more plentiful small seeds. HOW NATURAL SELECTION PRODUCES DIVERSITY Geospiza fuliginosa • Darwin’s finches on the Galápagos are an example of adaptive radiation. • In adaptive radiation, a cluster of species changes to occupy a series of different habitats within a region. • Each habitat offers different niches to occupy. • A niche represents how a species interacts both biologically and physically with its environment in order to survive. • Each species evolves to become adapted to that niche. Geospiza fortis Geospiza magnirostris Geospiza scandens Ground and cactus finches Geospiza conirostris Geospiza difficilis Camarhynchus parvulus Camarhynchus psittacula Camarhynchus Tree pauper finches Cactospiza heliobates Cactospiza pallida (woodpecker finch) Platyspiza Vegetarian crassirostris tree finch Certhidea fusca Certhidea olivacea Warbler finches THE EVIDENCE FOR EVOLUTION • There are many lines of evidence supporting Darwin’s theory of evolution. • The fossil record comprises the most direct evidence of macroevolution. • Fossils are the preserved remains, tracks, or traces of once-living organisms. • They are created when organisms or their traces become buried in sediment. • By dating the rocks in which the fossils occur, one can get an accurate idea of how old the fossils are. THE EVIDENCE FOR EVOLUTION • Fossils in rock represent a history of evolutionary change. • Fossils are treated as samples of data and are dated independently of what the samples are like. • Successive changes through time are a data statement. • Thus, the statement that macroevolution has occurred is a factual observation. TESTING THE THEORY OF EVOLUTION WITH FOSSIL TITANOTHERES Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. 50 45 40 Millions of years ago 35 THE EVIDENCE FOR EVOLUTION • The anatomical record also reflects evolutionary history. • Example: all vertebrate embryos share a similar set of developmental instructions and features. Pharyngeal pouches Pharyngeal pouches Tail Tail Reptile Bird Courtesy of Michael Richardson and Ronan O’Rahilly Human THE EVIDENCE FOR EVOLUTION • Homologous structures are derived from the same body part present in an ancestor. • For example, the same bones might be put to different uses in related species. Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Bat Human Horse Porpoise THE EVIDENCE FOR EVOLUTION • Analogous structures are similar-looking structures in unrelated lineages. Taking Flight To take to the air, three very different vertebrates lightened bones and transformed hands into wings. • These are the result of parallel evolutionary adaptations to similar environments. • This form of evolutionary change is referred to as convergent evolution. Pterosaur (extinct) Eastern bluebird Samoan flying fox (fruitbat) Wolf Mouse Marsupial mouse Two Worlds Marsupials evolved the same sort of adaptations in isolation in Australia that placental mammals did elsewhere. Tasmanian wolf Flying phalanger Flying squirrel THE EVIDENCE FOR EVOLUTION • Traces of our evolutionary past are also evident at the molecular level. • Organisms that are more distantly related should have accumulated a greater number of evolutionary differences than two species that are more closely related. • The same pattern of divergence can be seen at the protein level. MOLECULES REFLECT EVOLUTIONARY DIVERGENCE Human Macaque Dog 8 0 32 Bird 45 Frog 67 Lamprey 125 10 20 30 40 50 60 70 80 90 100 110 120 Number of amino acid differences between hemoglobin of vertebrate species and that of humans THE EVIDENCE FOR EVOLUTION • Evolutionary changes appear to accumulate at a constant rate. • This permits changes in an individual gene, compared over a broad array of organisms, to be dated from the time of divergence. • This dating is referred to as a molecular clock. • For example, changes have accumulated in the cytochrome c gene at a constant rate. Nucleotide substitutions THE MOLECULAR CLOCK OF CYTOCHROME C Human/kangaro 100 Human/ Dog/ cow 75 cow Rabbit/ rodent Human/rodent 50 Horse/ Llama/ donkey cow Horse/cow Sheep/goat Pig/ 25 cow Goat/cow 0 0 25 50 75 100 125 Millions of years ago GENETIC CHANGE IN POPULATIONS: THE HARDY-WEINBERG RULE • Population genetics is the study of the properties of genes in populations. • Gene pool is the sum of all of the genes in a population, including all alleles in all individuals. GENETIC CHANGE IN POPULATIONS: THE HARDY-WEINBERG RULE • Variation in populations puzzled many scientists. • Why don’t dominant alleles drive recessive alleles out of populations? GENETIC CHANGE IN POPULATIONS: THE HARDY-WEINBERG RULE • G.H. Hardy and W. Weinberg, in 1908, studied allele frequencies in a gene pool. • In a large population in which there is random mating, and in the absence of forces that change allele frequencies, the original genotype proportions remain constant from generation to generation. • Because the proportions do not change, the genotypes are said to be in Hardy-Weinberg equilibrium. • If the allele frequencies are not changing, the population is not evolving. GENETIC CHANGE IN POPULATIONS: THE HARDY-WEINBERG RULE • Hardy and Weinberg arrived at their conclusion by analyzing the frequencies of alleles in successive generations. • Frequency is the proportion of something compared to the total. • Knowing the frequency of the phenotype, one can calculate the frequency of the genotypes and alleles in the population. • Population of 1000 cats • 840 black 160 white • 840/1000 = 0.84 or 84% Proportion of black cats GENETIC CHANGE IN POPULATIONS: THE HARDY-WEINBERG RULE • By convention, the frequency of the more common of two alleles is designated by the letter p and that of the less common allele by the letter q. • Because there are only two alleles, the sum of p and q must always equal 1. • p+q=1 GENETIC CHANGE IN POPULATIONS: THE HARDY-WEINBERG RULE • In algebraic terms, the Hardy-Weinberg equilibrium is written as an equation. (p + q) x (p + q) = p2 + 2pq + q2 = 1 Individuals Individuals Individuals homozygous heterozygous homozygous for allele B for alleles B for allele b &b GENETIC CHANGE IN POPULATIONS: THE HARDY-WEINBERG RULE • Population of 1000 cats • 840 black, 160 white • But how many BB vs Bb? • Frequency of p & q Sperm p = 0.6 Phenotypes q = 0.4 B B p = 0.6 BB Genotypes Frequency of genotype in the population (number in a population of 1,000 cats) BB 360 cats 360/1,000 = 0.36 Bb bb 480 cats 480/1,000 = 0.48 160 cats 160/1,000 = 0.16 p2 = 0.36 b Frequency of alleles in the population (total of 2,000) 720 B 480 B + 480 b 720 B + 480 B = 1,200 B 1,200/2,000 = 0.6 B 320 b 480 b + 320 b = 800 b 800/2,000 = 0.4 b b Bb Bb pq = 0.24 pq = 0.24 bb Number of alleles in the population (2 per cat) q2 Eggs = 0.16 q = 0.4 GENETIC CHANGE IN POPULATIONS: THE HARDY-WEINBERG RULE • Used to calculate allele frequencies (p & q) in a simple way: • 1000 cats, 160 white (bb) so q2 = 0.16 • q = square root of 0.16 = 0.40. • Since p + q = 1; p = 1 – q = 0.60. • p2 = 0.36 x 1000; so 360 homozygous dominant (BB) • 2pq = 0.48 x 1000; 480 heterozygous (Bb) GENETIC CHANGE IN POPULATIONS: THE HARDY-WEINBERG RULE • The Hardy-Weinberg equilibrium only works if the following five assumptions are met: – 1. The size of the population is very large or effectively infinite. – 2. Individuals can mate with one another at random. – 3. There is no mutation. – 4. There is no immigration or emigration. – 5. All alleles are replaced equally from generation to generation (natural selection is not occurring). AGENTS OF EVOLUTION • Five factors can alter the proportions of homozygotes and heterozygotes enough to produce significant deviations from HardyWeinberg predictions. 1. Mutation 2. Nonrandom mating 3. Genetic drift 4. Migration 5. Selection AGENTS OF EVOLUTION • Mutation is a change in a nucleotide sequence in DNA. • Mutation rates are generally too low to significantly alter Hardy-Weinberg proportions. • Mutations must affect the DNA of the germ cells or the mutation will not be passed on to offspring. • However, no matter how rare, mutation is the ultimate source of variation in a population. AGENTS OF EVOLUTION • Nonrandom mating occurs when individuals with certain genotypes mate with one another either more or less commonly than would be expected by chance. • Sexual selection is choosing a mate often based on physical characteristics. AGENTS OF EVOLUTION • Migration is the movement of individuals between populations. • The movement of individuals can be a powerful force upsetting the genetic stability of natural populations. • The magnitude of the effects of migration is based on two factors: • The proportion of migrants in the population. • The difference in allele frequencies between the migrants and the original population. AGENTS OF EVOLUTION • Genetic drift describes random changes in allele frequencies. • In small populations, the frequencies of particular alleles may be changed drastically by chance alone. • In extreme cases, individual alleles of a given gene may be: • All represented in few individuals. • Accidentally lost if individuals fail to reproduce or die. AGENTS OF EVOLUTION • A series of small populations that are isolated from one another may come to differ strongly as the result of genetic drift. AGENTS OF EVOLUTION • Founder effect occurs when one of a few individuals migrates and becomes the founder of a new, isolated population at some distance from their place of origin. • The alleles that they carry will become a significant fraction of the new population’s genetic endowment. AGENTS OF EVOLUTION • Bottleneck effect occurs when a population is drastically reduced in size. • the surviving individuals constitute a random genetic sample of the original population. AGENTS OF EVOLUTION • Selection occurs if some individuals leave behind more progeny than others, and the likelihood that they will do so is affected by their individual characteristics. • In artificial selection, a breeder selects for the desired characteristics. • In natural selection, conditions in nature determine which kinds of individuals in a population are the most fit. SELECTION FOR COAT COLOR IN MICE Light coat color pocket mouse is vulnerable on lava rock Light coat color favored by natural selection because it matches sand color Dark coat color favored by natural selection because it matches black lava rock 54 AGENTS OF EVOLUTION • There are three types of natural selection: (b) Disruptive selection (c) Directional selection Number of individuals (a) Stabilizing selection 0 25 50 75 100 Body size (g) 125 Number of individuals Selection for midsized individuals 0 25 50 75 100 Body size (g) 50 75 100 Body size (g) 125 0 Selection for small and large individuals Peak gets narrower 0 25 0 25 50 75 100 Body size (g) 50 75 100 Body size (g) 125 Selection for larger individuals Peak shifts Two peaks form 125 25 125 0 25 50 75 100 Body size (g) 125 • The result is an increase in the frequency of the already common intermediate phenotype. • For example, human birthweight is under stabilizing selection. 20 100 70 50 30 20 10 15 10 7 5 3 2 5 2 3 4 5 6 7 8 9 10 Birth weight in pounds Percent infant mortality • Stabilizing selection is a form of selection in which both extremes are eliminated. Percent of births in population AGENTS OF EVOLUTION AGENTS OF EVOLUTION • Disruptive selection is a form of selection in which the two extremes in an array of phenotypes become more common in the population. • Selection acts to eliminate the intermediate phenotypes. • For example, beak size in African blackbellied seedcracker finches is under disruptive selection because the available seeds are only large or small. • Directional selection is a form of selection that occurs when selection acts to eliminate one extreme from an array of phenotypes. Average tendency to fly toward light AGENTS OF EVOLUTION 11 10 9 8 7 6 5 4 3 2 1 0 2 4 6 8 10 12 14 16 18 20 Number of generations • For example, in Drosophila, flies that fly toward light can be selected against and those that avoid light selectively bred, producing a population of flies with a greater tendency to avoid light. SICKLE-CELL ANEMIA • Sickle-cell anemia is a hereditary disease affecting hemoglobin molecules in the blood. Val 6 (a) • The disorder results from a single nucleotide change in the gene for b-hemoglobin. (b) Sickled red blood cells (c) Normal red blood cells • This causes the sixth amino acid in the chain to change from glutamic acid (very polar) to valine (nonpolar). • The hemoglobin molecules clump together and deform the red blood cell into a “sickle-shape”. SICKLE-CELL ANEMIA • People homozygous for the sickle-cell genetic mutation frequently have a reduced lifespan. • The sickled form of hemoglobin does not carry oxygen atoms well. • The red blood cells that are sickled do not flow smoothly through capillaries. • Heterozygous individuals make enough functional hemoglobin to keep their red blood cells healthy. SICKLE-CELL ANEMIA • The frequency of the sickle-cell allele is about 0.12 in Central Africa. • One in 100 people is homozygous for the defective allele and develops the fatal disorder. • In contrast, sickle-cell anemia strikes only roughly two African Americans out of every thousand. • If natural selection drives evolution, why has natural selection not acted against the defective allele in Africa and eliminated it from the population? SICKLE-CELL ANEMIA • The defective allele has not been eliminated from Central Africa because people who are heterozygous are much less susceptible to malaria. Sickle-cell allele in Africa 1–5% 5–10% 10–20% Falciparum malaria in Africa Malaria • The payoff in survival of heterozygotes makes up for the price in death of homozygotes. • This is called heterozygote advantage. • Stabilizing selection occurs because malarial resistance counterbalances lethal anemia. PEPPERED MOTHS AND INDUSTRIAL MELANISM • The peppered moth, Biston betularia, is a European moth that rests on tree trunks during the day. • Until the mid-19th century, almost every captured individual had light-colored wings. • Since then, individuals with dark-colored wings increased in frequency in industrialized areas. • Can Darwin’s theory explain this? • Yes it can! PEPPERED MOTHS AND INDUSTRIAL MELANISM • Industrial melanism describes the evolutionary process in which darker individuals come to predominate over lighter individuals since the industrial revolution as a result of natural selection. • Dark organisms are better concealed from their predators in habitats that have been darkened by soot and other industrial pollution. PEPPERED MOTHS AND INDUSTRIAL MELANISM Percentage of melanic moths • In England, the air pollution promoting industrial melanism began to reverse following enactment of the Clean Air legislation in 1956. – As a result, the frequency of the melanic (dark) form of Biston appears to decrease as 100 well. 90 80 70 60 50 40 30 20 10 0 59 63 67 71 75 79 Year 83 87 91 95 SELECTION ON COLOR IN GUPPIES • On the island of Trinidad, guppies are found in two very different stream environments. • In pools above waterfalls, the guppies are found along with the killifish, a seldom predator of guppies. • In pools below waterfalls, the guppies are found in pools along with the pike cichlid, a voracious predator of guppies. • Guppies can move between pools by swimming upstream during floods SELECTION ON COLOR IN GUPPIES • Guppy populations above and below waterfalls exhibit differences. Guppy (Poecilia reticulata) • Guppies in high-predation pools are not as colorful as guppies in low-predation pools. • Guppies in high-predation pools tend to reproduce at an earlier age and attain relatively smaller adult body sizes. • These differences suggest the action of natural selection. Pikecichlid (Crenicichla alta) Guppy (Poecilia reticulata) Killifish (Rivulus hartii) SELECTION ON COLOR IN GUPPIES • John Endler conducted experiments on guppies to determine whether predation risk was really the driving selective force in this system. • In a controlled laboratory setting, he created artificial pool environments in which he placed guppies in one of three conditions: • with no predator present • with killifish present (low predation risk) • with cichlid present (high predation risk) FIGURE 14.31 EVOLUTIONARY CHANGE IN SPOT NUMBER • After 10 guppy generations, he found that the guppies from no or low predation risk pools were both larger and more colorful than the guppies from the high predation risk pool. • He later found the same results in field experiments . No predation 14 Spots per fish 13 Low predation 12 11 10 High predation 9 8 0 4 8 Months 12 THE BIOLOGICAL SPECIES CONCEPT • Speciation is the macroevolutionary process of forming new species from pre-existing species. • It involves successive change. • First, local populations become increasingly specialized. • Then, if they become different enough, natural selection may act to keep them that way. THE BIOLOGICAL SPECIES CONCEPT • Ernst Mayr coined the biological species concept, which defines species as “groups of actually or potentially interbreeding natural populations which are reproductively isolated from other such groups”. • Populations whose members do not mate with each other and cannot produce fertile offspring are said to be reproductively isolated and, thus, members of different species. THE BIOLOGICAL SPECIES CONCEPT • Barriers called reproductive isolating mechanisms cause reproductive isolation by preventing genetic exchange between species. • Prezygotic isolating mechanisms prevent the formation of zygotes. • Postzygotic isolating mechanisms prevent the proper functioning of zygotes once they have formed. ISOLATING MECHANISMS • There are six different prezygotic reproductive isolating mechanisms: • • • • • • geographical isolation ecological isolation temporal isolation behavioral isolation mechanical isolation prevention of gamete fusion ISOLATING MECHANISMS • Geographical isolation occurs in cases when species exist in different areas and are not able to interbreed. ISOLATING MECHANISMS • Ecological isolation results from two species that occur in the same area but utilize different portions of the environment and are unlikely to hybridize. ISOLATING MECHANISMS • Temporal isolation results from two species having different reproductive periods, or breeding seasons, that preclude hybridization. • Behavioral isolation refers to the often elaborate courtship and mating rituals of some groups of animals, which tend to keep these species distinct in nature even if they inhabit the same places. ISOLATING MECHANISMS • Mechanical isolation results from structural differences that prevent mating between related species of animals and plants. • Prevention of gamete fusion blocks the union of gametes even following successful mating. ISOLATING MECHANISMS • If hybrid matings do occur, and zygotes are produced, postzygotic factors may prevent those zygotes from developing into normal individuals. • In hybrids, the genetic complements of two species may be so different that they cannot function together normally in embryonic development. • Even if hybrids survive the embryo stage, they may not develop normally. • Finally, many hybrids are sterile.