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Natural selection and the modern synthesis Evolution Biol 4974/5974 DF Tomback Biol. 4974/5974 Lecture #2 M. W. Strickberger (2000) Evolution, Jones and Bartlett, D. J. Futuyma (1998) Evolutionary Biology, Sinauer Associates Evolution and Natural Selection, and the Development of the Modern Synthesis “How extremely stupid not to have thought of that” ---Thomas Huxley Both science and religion have resulted from humans trying to figure out, “What’s it all about?” Learning goals Understand and be able to explain: • How 18th and 19th century scientific discoveries and developments paved the way for public acceptance of evolution. • The evolutionary mechanisms proposed by Lamarck, why they were a step forward in scientific thinking, and how they were discredited. • The mechanism of natural selection proposed by Darwin and Wallace, and why it is a “statistical phenomenon.” • Why natural selection was not immediately accepted when On the Origin of Species was published, and even after Mendel’s Laws were rediscovered. • The origins of the modern evolutionary synthesis and its “official beginning.” • The impact of the evolutionary synthesis on the natural sciences and related fields. 1 Natural selection and the modern synthesis Evolution Biol 4974/5974 DF Tomback Historical setting: timing for a theory of evolution Darwin’s timing in history was excellent for acceptance of the idea of evolution and a “material” mechanism. During the 18th and 19th centuries, various physical and natural sciences were flourishing. Linnaeus began the system of binomial classification (genus and species). Comparative anatomists such as Cuvier were describing the similarities and differences among species. The law of gravity was developed by Newton, who had also discovered calculus. Van Leeuwenhoek discovered the world of microorganisms after he built the first microscope. Fossils of extinct animals placed in historical sequence by geological strata (the first dinosaur bone was found in 1677 but identified as reptilian in 1824 by William Buckland at Oxford). Principles of geological change over time by the natural forces of slow erosion and deposition (uniformitarianism) developed by Charles Lyell. Led to age of the earth reevaluated by different methods, all resulting in ages greater than biblical 5,000 years. 2 Natural selection and the modern synthesis Evolution Biol 4974/5974 DF Tomback Dinosaurs even became part of “popular” culture! Dinner in a dinosaur (Iguanodon), Crystal Palace (1852-1854) P.W. Price (1996) Biological Evolution, Saunders College Publishing These scientific developments challenged previously held views that the world was unchanging By the early 19th century, the great age of earth, extinctions of species, and evolutionary change were generally accepted During Darwin’s life, more discoveries were made, which further supported his ideas about evolution M.W. Strickberger (2000) Mechanism of evolution By the early 19th century, evolution was gaining acceptance, but how evolution worked was unknown. Jean-Baptiste de Lamarck (17441829) proposed the first plausible evolutionary mechanism. He suggested that species were gradually modified over time, offering a connection from strange, extinct forms to contemporary forms. Lamarck 3 Natural selection and the modern synthesis Evolution Biol 4974/5974 DF Tomback D.J. Futuyma (1998) Lamarckism Change in organisms resulted from some internal force driving the organism towards “perfection” or progress. This was expressed as two “Laws”: The Principle of Use and Disuse The Inheritance of Acquired Characters National Geographic (1987) Field Guide to the Birds of North America Lamarckian evolution was formalized as orthogenesis”: It ascribed to evolution the drive towards progress---ever greater perfection. Cuvier (1769-1832) discredited Lamarckian evolution with logical criticism: No intermediate forms Hybrids are always sterile Plants and lower animals have no consciousness—how can they strive for perfection? At least Lamarck’s mechanism relied on natural processes that accounted for gradual change over time—a step forward in thinking. The recent discovery of stress-induced mutagenesis in bacteria (“directed evolution”) is not unlike Lamarckism. M.W. Strickberger (2000) Natural selection D.J. Futuyma (1998) In 1858, Charles Robert Darwin and Alfred Russel Wallace published papers introducing natural selection together in the Journal of the Linnaean Society. In 1859, Darwin published his detailed treatise on the subject, On the Origin of Species. 4 Natural selection and the modern synthesis Evolution Biol 4974/5974 DF Tomback Summary of natural selection High reproductive rates in most organisms. Finite food and space limits numbers. Also, predation, parasitism, disease, competition, climate. Variation among individuals. Certain variation results in higher survival and/or better reproduction. Thus, relatively greater contribution of offspring from some individuals. Their offspring may in turn have similar traits. Thus, certain “favored” variation spreads through the species over time. Leads to “descent with modification” or evolution over time. Figure 11. Schematic of the main conceptual arguments for evolution by natural selection given by Charles Darwin and Alfred Russel Wallace. Evolutionary mechanism? The public was receptive to Darwin’s ideas. Evolution not doubted by most. Natural selection was difficult for some to accept: Mechanism of inheritance not known. (Mendel’s work was published in 1866 but not widely read.) Between 1860 and the 1940’s scientists agreed on evolution, but thought the mechanism was some combination of natural selection and Lamarckism. Here’s how Darwin’s ideas prevailed…. 5 Natural selection and the modern synthesis Evolution Biol 4974/5974 DF Tomback The biological revolution In the 19th century, biology was developing rapidly, specializing, splitting into new fields, e.g., embryology, cytology, physiology. There were the experimental or lab-based biologists, who worked on “systems.” There were the “whole organism” biologists, who collected, described, observed, and compared--e.g., paleontologists, botanists, zoologists, systematists, working in field, lab, and museum. Each group asked different questions; had different perspectives, particularly regarding evolutionary processes. Rediscovery of Mendel’s Laws In 1900, Mendel’s laws were rediscovered, and genetics became a field of science. Ironically, genetics caused a further rift: The naturalists were evolutionists and studied natural populations. The new geneticists considered mutation as the major mechanism of evolution; they did not see how natural selection could change gene frequencies. D.J. Futuyma (1998) They didn’t read each others’ work or cross-communicate; multiple theories of evolutionary mechanisms flourished Scientific isolation Meanwhile, new genetic advances supported gradual evolutionary change, and mutationism fell out of favor, but these ideas were not communicated beyond genetics. Some naturalists and paleontologists still subscribed to orthogenesis. “At the end of the 1920s, the situation in biology did not seem at all promising for a meeting of the minds in the near future.” –Mayr and Provine 6 Natural selection and the modern synthesis Evolution Biol 4974/5974 DF Tomback The “quiet synthesis” Despite hostility among disciplines during the first 30 years of the 20th century, the foundation for a modern evolutionary synthesis was laid: abundant evidence in support of natural selection. A few pioneers developed the ideas that later were incorporated: Huxley, Ford, Haldane, Grinnell, and Baur. The “bridge builders” were biologists willing to read outside of their field. The synthesis occurred quietly, with enlightened “new biologists” arriving at similar conclusions about the mechanisms of evolution. The birth of the Modern Evolutionary Synthesis The official beginning: All participants agreed about the mechanisms of evolution. There were few arguments and most over small problems. No major revision has occurred since. Some of the participants: D.J. Futuyma (1998) 7 Natural selection and the modern synthesis Evolution Biol 4974/5974 DF Tomback D.J. Futuyma (1998) There has been some “fine-tuning” of the modern synthesis since the Princeton Conference. Ernst Mayr • Born July 5, 1904 • Died February 3, 2005 An astute naturalist and arguably the most important contributor to the modern evolutionary synthesis. Stephen Jay Gould 1941-2002 •Brought the science of evolution to the general reader through his brilliant essays. •Made contributions to our understanding of the processes of speciation and the relationship between ontogeny and phylogeny. •Introduced many terms to the field, including “punctuated equilibrium.” 8 Natural selection and the modern synthesis Evolution Biol 4974/5974 DF Tomback The impact of the modern synthesis on science Was the modern synthesis a “scientific revolution”? Yes and No. It was not drastically new—the seeds were sown in 1859. But the impact on modern science has been tremendous. The impact of the modern evolutionary synthesis All branches of the natural sciences have been influenced and organized by evolutionary principles—known as the “first principles.” Also, the influence extends to geology, paleontology, and anthropology, and now to sociology, psychology, economics, and business! Researchers are looking at evolution and human behavior. The impact of the modern evolutionary synthesis on various disciplines D.J. Futuyma (1998) 9 Natural selection and the modern synthesis Evolution Biol 4974/5974 DF Tomback Yet, public acceptance of human evolution in the U.S. ranks near bottom of 34 countries surveyed. Why? Miller et al. 2005. Public acceptance of evolution. Science 313: 765-766. Question asked: “Human beings, as we know them, developed from earlier species of animals.” Choices: definitely true, probably true, probably false, definitely false, did not know or uncertain. Only 14% of Americans chose “definitely true”. “Probably true” and “Not certain” accounted for 55%. Study questions 1. 2. 3. 4. Why was Darwin’s timing very good for acceptance of the idea that evolution had changed species? What other discoveries had set the stage for acceptance? Explain the evolutionary mechanism known as Lamarckism. What were the two fundamental principles involved? What were criticisms of Lamarckism? What was the contribution of Lamarck? Explain the basic process of natural selection as presented by Darwin and Wallace? Study questions, continued 5. 6. 7. 8. 9. Was natural selection immediately accepted by scientists as the primary evolutionary mechanism? If not, what did people believe? Did the rediscovery of Mendel’s laws unite biologists? Why or why not? How did some scientists become the “bridge builders”? How did this set the stage for the modern synthesis? When was the official beginning of the modern evolutionary synthesis? What is the scientific impact of the modern evolutionary synthesis? 10