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Gene’s Eye View of Evolution (Richard Dawkins, Oxford University): The gene as the fundamental unit of selection: • • • • The evolutionary importance of the individual has been highly overrated Individuals are ephemeral, mere blips on the evolutionary landscape; DNA is potentially immortal We are survival machines - robot vehicles blindly programmed to preserve the selfish molecules known as our genes. Organisms are simply VEHICLES for the transmission of REPLICATORS. A Replicator has the property of being able to make copies of itself. Types of Replicators • Passive Replicator - a replicator that has no influence over its probability of being copied • Active Replicator - a replicator whose nature has some influence over its probability of being copied MAKES POSSIBLE ORGANIC EVOLUTION 1 Properties of Successful Replicators • Stability: the universe is populated by stable entities (= Longevity) • Speed of Replication: (= Fecundity) • Accuracy of Replication (= Fidelity) Kinds of Replicators • Dead-end Replicators: not transmitted to the next generation, e.g., somatic cells in metazoans • Germ-line Replicators: theoretically immortal; alleles • ON EARTH, THE REPLICATOR IS DNA 2 Mutation: Mistakes in Replication • No replicator is capable of making perfect copies of itself • Mistakes in the replication process allow for evolution in the face of environmental change Replicators: Airy-fairy or Fact? HIV: An Active Replicator 3 “Life Cycle” of HIV Replicator/Vehicle Relations Benign or malevolent? • Importance of transmission mode —Vertical vs. horizontal 4 Science & Religion Defined • Science: systematized knowledge derived from observation, study, and experimentation carried out in order to determine the nature or principles of what is being studied. • Religion: the belief in a divine or superhuman power or powers to be obeyed and worshipped as the creator(s) and ruler(s) of the universe. What Distinguishes Science from Religion? • Scientific theories are falsifiable (capable of being able to be shown untrue or misleading). To be considered scientific, a theory, in principle, must be constructed in such a way that it is capable of being disproved. • Religion is based on faith. 5 Scientific Method • • • • Observation Question Hypothesis ---> Predictions Test of Predictions —Strong inference: randomized, manipulative experiment —Weak inference: correlational analysis gives a result consistent with the hypothesis • If predictions independently validated --> Theory What is Theory? • Theory: a scheme or system of ideas and statements held as an explanation or account of a group of facts or phenomena; a hypothesis that has been confirmed or established by observation or experiment, and is propounded or accepted as accounting for the known facts. • Because theory is a set of statements, it usually does not stand or fall on the basis of a single critical test, as simple hypotheses do. Rather, theories evolve as they are confronted with new phenomena or observation; parts of the theory are discarded, modified or added. 6 Historical Overview of Evolutionary Thought • Ancient Greeks (Plato & Aristotle): Typological View of Nature - individual variation as the imperfect manifestation of ethos. Ideas are an eternal, unchanging essence (Ethos are eternal ideas that exist in the mind of God) • Christian Theology: Scala Naturae or the Great Chain of Being. The ladder of life consists of gradation from inanimate material through plants, through lower animals and humans to angels and other spiritual beings. • Natural Theology: adaptations of organisms as evidence for the Creator’s benevolence; e.g., John Ray’s (1701) “The Wisdom of God Manifested in the Works of the Creation” First articulated theory of evolution Jean-Baptiste Lamarck, 1809 Philosophie Zoologique • Organisms continually arise by spontaneous generation • Organisms develop adaptations to a changing environment through the use and disuse of organs • Acquired characteristics are inherited 7 Lamarck vs. Darwin 8 What good, Lamarck? • If nothing else, demonstrates that theory in evolutionary biology can be falsified-Lamarck’s ideas were resoundingly rejected! —The notion of continual spontaneous generation was challenged by geologists and the principle of uniformitarionism, i.e., the same processes are responsible for shaping past and current events —Many empirical studies have demonstrated, that, in effect, you cannot inherit a good sun tan ON THE ORIGIN OF SPECIES BY NATURAL SELECTION 1859 • Competition • Individual Variation • Heritability • Blending Inheritance (hereditary factors mix in the blood) 9 Particulate Theory of Inheritance Evolution: Post-Darwin • Darwin’s Theory of Evolution by Natural Selection suffered from a lack of understanding of inheritance and the basis of genetic variation; Darwin believed in blending inheritance. • NeoDarwinism-The Modern Synthetic Theory of Evolution-The Modern Synthesis • Marriage of Darwin’s principle of natural selection with Mendel’s discovery of particulate inheritance • Hereditary information is encoded in genetic material called genes 10 Blending versus Particulate Inheritance: Fate of a Favorable Mutation Founders of the Modern Synthesis Ronald A. Fisher: theoretical population geneticist; founder of modern statistics & experimental design J.B.S. Haldane: theoretical population geneticist; Haldane's rule of speciation Sewall Wright: theoretical population geneticist; believed that genetic drift was an major force in evolution Theodosius Dobzhansky: Drosophila geneticist who integrated population genetic theory with empirical data on genetic variation Julian Huxley: Ornithologist who used genetic principles to explain macroevolutionary patterns Ernst Mayr: Systematist; perhaps best known for the "biological species concept" George G. Simpson: Paleontologist who used genetic principles to explain macroevolutionary patterns 11 The Modern Synthesis • Genetic variation arises by chance due to imperfections in DNA replication (mutation) and recombination (crossing over of homologous chromosomes during meiosis) • Populations evolve by changes in gene frequency brought about by random genetic drift, gene flow and especially natural selection • Most beneficial mutations cause slight changes in the phenotype and therefore evolutionary change tends to be slow and gradual • Speciation (new species formation) occurs gradually when populations are reproductively isolated by geographic barriers The Molecular Genetic Revolution • The Double Helix (Watson & Crick,1953) • The Central Dogma (DNA-->RNA-->Protein) • Genetic engineering • DNA fingerprinting • Genome sequencing Selfish genetic elements: a pervasive feature of organismal genomes 12