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Darwinian Evolution How do organisms respond to a changing environment? Darwinian Revolution November 24, 1859 Charles Darwin (1809-1882) publishes “On the Origin of Species by Means of Natural Selection” Darwin’s publication focused biologists’ attention on the great diversity of organisms. He presented evidence that species today were the descendants of ancestral species He proposed a mechanism for the evolutionary process; called natural selection Darwinian Revolution The period into which Darwin was born was one of great progress in all areas of science. Natural history was no exception and for some time exploration of the fossil record had already suggested that the Earth was far more ancient than Christian texts claimed. Fossils suggested exotic forms of life never seen nor described in biblical texts. This generated enormous debate among the aristocratic classes. Not only had the cultural histories of Western civilization always held species to be constant and unchanging but this view was common to both religion (Old Testament) and philosophy (Aristotle). Darwinian theory challenged these traditional views in a profound manner- it questioned not only creation (and with it the very role of God) but even provided a rational scientific explanation for how creationism should be replaced. Linnaeus (classification) Hutton (gradual geologic change) Lamarck (species can change) Malthus (population limits) Cuvier (fossils, extinction) Lyell (modern geology) Darwin (evolution, nutural selection) Mendel (inheritance) American Revolution 1750 Wallace (evolution, natural selection) French Revolution U.S. Civil War 1800 1850 1900 1795 Hutton proposes his theory of gradualism. 1798 Malthus publishes “Essay on the Principle of Population.” Before Origin of Species Carolus Linnaeus (1707-1778) Interpreted adaptations as evidence that the Creator had designed each species for a specific purpose Was a founder of taxonomy, classifying life’s diversity “for the greater glory of God” Paleontology, the study of fossils, was pioneered by French scientist Georges Cuvier (1769-1832). Cuvier opposed gradualism. Instead he advocated catastrophism and speculated that each boundary between geological layers represented a catastrophe This attention to taxonomic diversity and study of fossils helped to lay the groundwork for Darwin’s ideas Before Origin of Species Gradualism is the idea that profound change can take place through the cumulative effect of slow but continuous processes Geologists James Hutton (1726-1797) and Charles Lyell (1797-1875) perceived that changes in Earth’s surface can result from slow continuous actions still operating today. These ideas exerted a strong influence on Darwin’s thinking Before Origin of Species Jean-Baptiste Lamarck (1744-1829) hypothesized that an organism could pass on characteristics it acquired during its lifetime to its offspring. He even suggested that the environment gives rise to these changes in animal development. The mechanisms he proposed, however, were unsupported by evidence Today this is called Lamarckian Evolution. Voyage of the Beagle Even as a boy Charles Darwin had a consuming interest in nature Soon after receiving his B.A. degree Darwin was accepted on board the HMS Beagle, which was about to embark on a voyage around the world Darwin’s interest in the geographic distribution of species was kindled by the Beagle’s stop at the Galápagos Islands near the equator west of South America. ure 22.5 England NORTH AMERICA EUROPE ATLANTIC OCEAN PACIFIC OCEAN Galápagos Islands HMS Beagle in port SOUTH AMERICA AUSTRALIA Andes Darwin in 1840, after his return AFRICA Cape of Good Hope Cape Horn Tierra del Fuego Tasmania New Zealand Voyage of the Beagle Darwin began to perceive adaptation to the environment and the origin of new species as related processes (a) Cactus eater. The long, sharp beak of the cactus ground finch (Geospiza scandens) helps it tear and eat cactus flowers and pulp. Figure 22.6a–c (c) Seed eater. The large ground finch (Geospiza magnirostris) has a large beak adapted for cracking seeds that fall from plants to the ground. (b) Insect eater. The green warbler finch (Certhidea olivacea) uses its narrow, pointed beak to grasp insects. On The Origin of Species In 1844, Darwin wrote a long essay on the origin of species and natural selection but, anticipating the uproar it would cause, delayed publication. In June 1858 Darwin received a manuscript from Alfred Russell Wallace who had developed a theory of natural selection similar to his own. Darwin quickly finished The Origin of Species and published it the next year. Sirenia Hyracoidea (Manatees (Hyraxes) and relatives) Barytherium Mammuthus Stegodon Platybelodon Mammut Deinotherium Elephas Loxodonta Loxodonta maximus africana cyclotis (Africa) (Asia) (Africa) Moeritherium Millions of years ago Years ago Darwin developed two main ideas * Evolution explains life’s unity and diversity * Natural selection is a cause of adaptive evolution In the Darwinian view, the history of life is like a tree with multiple branchings from a common trunk to the tips of the youngest twigs that represent the diversity of living organisms Natural Selection From Wikipedia..... Natural selection is a natural law by which genetically heritable traits become more or less common in a population over successive generations. Natural genetic variation within a population of organisms means that some individuals will survive and reproduce more successfully than others in their current environment Natural selection acts on the phenotype but the inheritable, genetic basis of any phenotype which gives a reproductive advantage will become more common in a population (see allele frequency)* * Allele frequency to follow later. Don’t forget to google “Darwin Awards” Personal Favourite Natural Selection (a) A flower mantid Natural selection can produce an increase over time Malaysia in the adaptation of organismsin to their environment (b) A stick mantid in Africa Top: Flower Mantid (Malaysia) Bottom: Stick Mantid (Africa) Natural Selection If an environment changes over time natural selection may result in adaptation to these new conditions Reznick and Endler transplanted guppies from pike-cichlid pools to killifish pools EXPERIMENT and measured the average age and size of guppies at maturity over an 11-year period (30 to 60 generations). Pools with killifish, but not guppies prior to transplant Predator: Killifish; preys mainly on small guppies Experimental transplant of guppies Guppies: Larger at sexual maturity than those in “pike-cichlid pools” Predator: Pike-cichlid; preys mainly on large guppies Guppies: Smaller at sexual maturity than those in “killifish pools” Figure 22.12 Natural Selection RESULTS 185.6 161.5 67.5 76.1 Males Females Age of guppies at maturity (days) Weight of guppies at maturity (mg) After 11 years, the average size and age at maturity of guppies in the transplanted populations increased compared to those of guppies in control populations. 85.7 92.3 48.5 58.2 Males Females Control Population: Guppies from pools with pike-cichlids as predators Experimental Population: Guppies transplanted to pools with killifish as predators CONCLUSION Reznick and Endler concluded that the change in predator resulted in different variations in the population (larger size and faster maturation) being favored. Over a relatively short time, this altered selection pressure resulted in an observable evolutionary change in the experimental population. Natural Selection In humans, the use of drugs artificially selects for pathogens that through chance mutations are resistant to the drugs’ effects. Natural selection is a cause of adaptive evolution Percent of HIV resistant to 3TC Researchers have developed numerous drugs to combat HIV- but using these medications selects for viruses resistant to the drugs Patient No. 1 Patient No. 2 Patient No. 3 Weeks Natural Selection One common misconception about evolution is that individual organisms evolve, in the Darwinian sense, during their lifetimes Natural selection acts on individuals, but populations evolve. It is variation that allows populations to evolve. Members of a population may show no variation for a particular trait. Members of a population may show two or more variations for a particular trait. Galahs are monomorphic in respect to plumage. Lupins are polymorphic for colour. Population Genetics How do genes affect groups? Populations & Gene Pools MAP AREA CANADA ALASKA Population genetics is the study of how populations change genetically over time. It integrates Mendelian genetics with the Darwinian theory of evolution by natural selection and focuses on populations as units of evolution Beaufort Sea Porcupine herd range N TE OR RR TH IT WE O S RI T ES Fortymile herd range ALASKA YUKON • Fairbanks • Whitehorse A population is a localized group of individuals that are capable of interbreeding and producing fertile offspring The gene pool is the total aggregate of genes in a population at any one time- this includes all the alleles present in the population Allele Frequencies Allele frequency is a measure of the relative frequency of an allele in a population. Most genes show only one allele in most of the population; this is called the wild-type allele. For variation to occur, however, more than one allele of a gene must exist; this is called polymorphism. Non-evolving populations Allele frequencies stay constant if... * There is no random mating * Matings are fertile * There are no mutations * There is no migration * The population is large Evolving Populations Allele frequencies are subject to various “change agents”.You need to know the types of change agents and the consequences for the population. Evolving Populations: Mutations Mutations are the origin of genetic variation. While often harmful to the individual they can also be neutral or beneficial. Point Mutations Block Mutations Substitution Insertion Deletion Insertion Deletion Duplication Inversion Translocation Aneuploidy Evolving Populations: Gene Flow Gene flow involves the migration of individuals between populations. This may result in very rapid changes in allele frequencies. Immigration may increase the variety of alleles present in the population Emigration may decrease the variety of alleles present in the population Evolving Populations: Genetic Drift Genetic Drift describes how allele frequencies can fluctuate unpredictably from one generation to the next. This process ends to reduce genetic variation and has greater effects in small populations. CW CW CR C R CR C R Only 5 of 10 plants leave offspring CR C W CW CW CR C R CR C R CR C W CW CW CR C R CR C W CR C W CR C R CW CW CR C W CR C R CR C R CR C W Generation 1 p (frequency of CR) = 0.7 q (frequency of CW) = 0.3 Only 2 of 10 plants leave offspring CR C R CR C R CR C R CR C R CR C R CR C R CR C R CR C R CR C W CR C W Generation 2 p = 0.5 q = 0.5 CR C R CR C R Generation 3 p = 1.0 q = 0.0 Evolving Populations: Bottleneck Effect Blam! In the bottleneck effect a sudden change in the environment may drastically reduce the size of a population The surviving 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. Figure 23.8 A Original population Bottlenecking event Surviving population Evolving Populations: Founder Effect The founder effect occurs when a small, unrepresentative sample becomes isolated from a larger population. Captive breeding programs in zoos represent a founder effect. Care must be taken to avoid inbreeding and maximise genetic diversity. Speciation How do new species arise? The Biological Concept of Species Organisms belong to the same species if they can interbreed and produce viable, fertile offspring. * All human races belong to the same species as we can reproduce. * Domestic dogs also belong to the same species as they can interbreed eg. Spoodles... * Horses and donkeys, however, are separate species; even though they can interbreed the offspring are infertile. * Obviously a whale and a gold fish are different species. We won’t have much success trying to breed a gold whale by normal standards. * Also, the biological species concept cannot be applied to asexual organisms. The Biological Concept of Species (a) Similarity between different species. The eastern meadowlark (Sturnella magna, left) and the western meadowlark (Sturnella neglecta, right) have similar body shapes and colorations. Nevertheless, they are distinct biological species because their songs and other behaviors are different enough to prevent interbreeding should they meet in the wild. (b) Diversity within a species. As diverse as we may be in appearance, all humans belong to a single biological species (Homo sapiens), defined by our capacity to interbreed. Speciation Speciation is a process whereby a parent species splits into daughter species. For this to happen the gene pool of the original species must divided into separate gene pools and remain reproductively isolated for long enough that the individuals are eventually unable to breed between the populations. These isolating mechanisms can be classified in two ways. Pre-reproductive Isolation Impede mating between species or hinder fertilization Geographic: Temporal: Behavioural: Morphological: individuals may be separated by seas, mountains, distances or habitat individual may breed during different times of year or day individual may have different courtship patterns individuals may have different reproductive structures so that mating is physically impossible. Post-reproductive Isolation These do not prevent mating but do prevent fertile young from being produced. Gamete mortality: gametes do not survive Zygote mortality: zygote forms but does not survive Hybrid sterility: offspring are born infertile Pre-Reproductive Isolation Prezygotic barriers impede mating or hinder fertilization if mating does occur Habitat isolation Behavioral isolation Temporal isolation Individuals of different species Mechanical isolation Mating attempt HABITAT ISOLATION TEMPORAL ISOLATION BEHAVIORAL ISOLATION (b) MECHANICAL ISOLATION (g) (d) (e) (a) (f) (c) Post-Reproductive Isolation Gametic isolation Reduce hybrid fertility Reduce hybrid viability Hybrid breakdown Viable fertile offspring Fertilization REDUCED HYBRID VIABILITY GAMETIC ISOLATION REDUCED HYBRID FERTILITY HYBRID BREAKDOWN (k) (j) (m) (l) (h) (i) Allopatric Speciation Allopatric speciation occurs where gene flow is interrupted or reduced because a population is divided into two or more geographically isolated subpopulations Once geographic separation has occurred one or both populations may undergo evolutionary change during the period of separation A. harrisi A. leucurus Allopatric Speciation EXPERIMENT Diane Dodd, of Yale University, divided a fruit-fly population, raising some populations on a starch medium and others on a maltose medium. After many generations, natural selection resulted in divergent evolution: Populations raised on starch digested starch more efficiently, while those raised on maltose digested maltose more efficiently. Dodd then put flies from the same or different populations in mating cages and measured mating frequencies. Initial population of fruit flies (Drosphila Pseudoobscura) Some flies raised on starch medium Mating experiments after several generations Some flies raised on maltose medium Allopatric Speciation RESULTS 22 9 8 20 Male Male Maltose Starch Female Starch Maltose Mating frequencies in experimental group CONCLUSION Different Same populations population When flies from “starch populations” were mixed with flies from “maltose populations,” the flies tended to mate with like partners. In the control group, flies taken from different populations that were adapted to the same medium were about as likely to mate with each other as with flies from their own populations. Female Different Same population populations 18 15 12 15 Mating frequencies in control group The strong preference of “starch flies” and “maltose flies” to mate with like-adapted flies, even if they were from different populations, indicates that a reproductive barrier is forming between the divergent populations of flies. The barrier is not absolute (some mating between starch flies and maltose flies did occur) but appears to be under way after several generations of divergence resulting from the separation of these allopatric populations into different environments. Extinction A variety of causes can lead to extinction. * Loss of habitat * Competition for resources with a better adapted organism * Catastrophic events can produce a mass extinction of species. For example: oMeteorite strike 65 mya (Cretaceous period). This explosion caused major fires, tidal waves and threw up enough dust to block enough sunlight to cool the planet. o Quaternary Period– Earth’s orbit changed raising temp in Antarctic Ocean * Human activity increases extinction, habitat destruction/salination, introduction of foreign plants and animals, competing with or attacking native flora and fauna, chemical use