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Jan 7: History--Darwin Charles Darwin: the first ethologist Darwin’s theory of natural selection accounts for two major phenomena that puzzled 19th century biologists Continuity vs. diversity of form • Organisms are so varied and yet the variation is overlaid on a smaller number of basic themes • This results from nature of evolutionary process: descent with modification Complexity and adaptation • Organisms appear to be complex machines designed to solve particular problems • Natural selection was proposed as the agent of design Jan 7: History--Darwin’s insight Darwin’s key insight for ethology: behavioral traits are a part of an organisms evolved phenotype Phenotype = Morphology + Physiology + Behavior Jan 6: History--Difficulties in study of behavior Some major difficulties in the study of behavior (and how Darwin and others have dealt with them) • Anthropomorphism or overly complex hypotheses about other species’ behavior • Fluidity of behavior--how to partition the phenotype to identify discrete evolved traits • Plasticity of behavior under influence of environment and experience--again, what exactly is the trait? Jan 7: History -- Anthropomorphism Anthropomorphism: if animals act like us they must think like us What’s the problem? • Underlying mechanisms may be different--probably are simpler Principle of Parsimony Morgan’s Canon: "In no case may we interpret an action as the outcome of the exercise of a higher psychical faculty, if it can be interpreted as the outcome of one which stands lower in the psychological scale." ( Conwy Lloyd Morgan 1894) Morgan’s Canon makes the same point as “Occam’s Razor: "Entities are not to be multiplied without necessity" Jan 7: History -- Fluidity of behavior Fluidity of behavior: how to partition the phenotype A complex behavioral repertoire can often be dissected into components that: • Are consistent in form (stereotyped) • Are consistent among individuals of the same species (species-specific) • Develop without the need for specific experience (innate) Behavioral traits as evolved “organs” Jan 7: History -- Plasticity of behavior Plasticity: changes in behavior resulting from experience • Darwin: “habits” (learned) vs. “instincts” (unlearned) • Early psychologists: notion of “instinct” was rejected in favor of environmental or cultural influences on behavior • Early ethologists: learning occurs, but under tight constraints--animals have an evolved “instinct to learn” • The debate about “nature” vs. “nurture” remains one of the most controversial issues in the behavioral sciences Jan 7: History -- Quick history 1 Quick history of behavioral biology since 1900 North America: behaviorist psychology (Thorndike, Watson, Skinner) • Strong emphasis on environmental, or cultural explanations for behavioral traits) Eddie Bauer American Legend • Animals studied mainly as models of human behavior • In psychology nowadays, genetic influences on behavior are given more credit, but animals are still studied mainly as experimental models (e.g., in neuroscience) Jan 6: History -- quick history 2 Quick history of behavioral biology since 1900-cont’d Europe: ethology: more direct intellectual descent from Darwin • Comparative ethology by comparative anatomists (Heinroth, Huxley, Lorenz): use natural behavioral repertoires of animals to study phylogenetic relationships • This broadens Darwin's insight that behavioral traits are evolved, instinctive species-specific "organs" Example: courtship behavior in ducks Jan 6: History -- quick history 2 Quick history of behavioral biology since 1900-cont’d • Konrad Lorenz: continued this tradition, and also began to think about internal aspects of behavior • “Fixed action pattern” (FAP) as unit of behavioral phenotype • Lorenz also hypothesized about motivational triggers of FAP Jan 6: History -- Nobel Prize Quick history of behavioral biology since 1900-cont’d Karl von Frisch: Developed sophisticated experimental approach to study of sensory systems, orientation, and communication (honey bees and other species) Lorenz, Tinbergen, and von Frisch were awarded the 1973 Nobel Prize in Medicine or Physiology “... for their discoveries concerning organization and elicitation of individual and social behaviour patterns.” Jan 6: History -- quick history 3 Quick history of behavioral biology since 1900-cont’d The rise of experimental ethology: “physiology without breaking the skin” Niko Tinbergen: used rigorous experiments to test ideas about the features of environment that animals respond to • Earliest example: Egg-rolling by goose • Pioneered use of physical models to probe mechanisms of behavior Jan 6: History--4 questions Tinbergen’s four questions • How? physiological causation What are the sensory, neural, and motor events giving rise to the behavior? • How? ontogeny How does the trait develop over the course of the animal’s lifespan? Northern mockingbird • How? phylogeny From what ancestral traits did the behavior arise during the evolutionary history of the species? • Why? adaptive function What is the survival value of the trait, which favors its maintenance in the population by natural selection? Alcock calls these “proximate questions” Alcock calls these “ultimate questions” Jan 6: History -- quick history 3 Models used by Tinbergen to study how herring gull chick knows where to peck to get food Important concepts: sign stimulus and supernormal stimulus