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Evolutionary Theory • Who are the two founders of evolutionary theory? • Explain the logic of natural selection. • What is the difference between genotype and phenotype? • What are the three main categories of adaptive problems? • What is the difference between proximate and ultimate explanations? Are they alternatives or complementary? • Give examples of Tinbergen’s four whys. • What are the three types of natural selection? • Explain the difference between adaptive behaviour and behavioural adaptation. • Why is the differential reproduction of organisms more important than their differential survival? • Compare sexual selection and natural selection. The 3 Premises of Natural Selection 1.The Principle of Variation Individuals in a population have variation in their physical and behavioural traits (i.e., phenotype) 2.The Principle of Inheritance Some of this variation is heritable (i.e., due to genotype) 3.The Principle of Adaptation Variation can help individuals to compete Nature and Nurture Nobody really believes that all of human behaviour is innate (i.e., can be explained by genes with no input from the environment). Similarly, nobody believes that the environment determines all behaviour without constraints of biology. Adaptive Problems in Human Evolution • Survival • Finding, choosing, processing appropriate food • Avoiding predators • Combating pathogens • Keeping warm • Reproduction • Choosing & wooing appropriate mates • Competing with same-sex rivals • Scheduling births • Caring for children • Sociality • Helping relatives & allies, cooperating & co-ordinating • Avoiding exploitation (selective allocation of help) • Learning & teaching: sharing information & skills • Political alliance & between-group conflicts Charles Darwin Alfred Russel Wallace Levels of Explanation • Proximate - How? • How do the environment and the individual’s genetic make-up interact to produce this behaviour? • What body systems (nervous, hormonal, etc.) are involved in this behaviour? • Ultimate - Why? • What is the function? • How does/did this trait help individuals increase their genetic representation in future generations? • What environmental pressures selected for this trait? • How has this behaviour changed over evolutionary history? Tinbergen’s Four Whys 1. Proximate or mechanistic cause The baby was crying Breasts were full of milk 2. Ontogenetic or developmental or cause Mother learned to nurse by observation Behaviours may be prepared by hormones 3. Phylogenetic or historical cause All mammals produce milk to feed offspring Explanations for how mammals evolved 4. Ultimate or functional cause Nursing promotes the survival of offspring Types of Natural Selection Evolutionary adaptation is a special and onerous concept that should not be used unnecessarily, and an effect should not be called a function unless it is clearly produced by design and not by chance. When recognized, adaptation should be attributed to no higher a level of organization than is demanded by the evidence. George Williams Evidence of Design Evidence of Design Gecko walking upside down on glass Lamellae on gecko toes Magnified hairs Gecko foot Autumn et al. (2002) Proc Nat Acad Sci Inferring Function from Structure Functions of Pregnancy Sickness miscarriages fetal deaths 20% 15% 10% 5% 0% NVP absent Nausea Only Vomiting (Flaxman & Sherman 2000) Does Adaptation = Adaptive? NO. Change in the environment is a major reason why an adaptation may not be currently adaptive. Behaviour resulting in an adaptive outcome is not necessarily the result of a psychological adaptation. This is why we need the criterion of complex design to identify adaptations. Basis of Sexual Selection Survival of the Fittest? Natural selection favours traits that help organisms to increase their genetic representation in future generations. Traits increasing reproduction will be favoured over traits increasing survival if the two are in conflict. Survival is only useful to an organism inasmuch as it is correlated with increased reproduction. Sexual selection is often seen as separate from natural selection because it commonly results in adaptations that reduce survival. Types of Sexual Selection Darwin noted that aggressive competition for mates is mainly a male affair, whereas mate choice is exercised mainly by females. But he had no explanation. Intersexual Selection Bateman (1948) was the first to recognize that males can often gain fitness more or less linearly by mating with more females, whereas females seldom profit similarly from extra mates. Members of one sex choose members of the other sex Williams (1966) and Trivers (1972) expanded on Bateman’s argument, noting that the sex difference in parental investment is not solely a matter of anisogamy. Anisogamy: an ovum is much larger than a sperm. Reproductive Variance in Fruit Flies Intrasexual Selection Members of one sex compete for access to members of the other sex Reproductive Variance in Humans Lifetime reproduction for birth cohorts of 100 ! Kung San of each sex, living as traditional hunter-gatherers in the Kalahari desert (Howell 1979) Males have more offspring per additional mate, more mates, and more variance in number of mates (Bateman 1948) Number of living children as a function of number of wives / co-wives for the Temne of Sierra Leone (Dorjahn 1958) Human Mating Systems Polygyny Marriage system: whether a culture allows men and/or women to have more than one official mate (708) Effective mating system: (4) Polyandry male fitness variance female fitness variance (137) Monogamy (Murdock 1967) Vocabulary • Adaptation • Adaptive • Alfred Russel Wallace • Anisogamy • Charles Darwin • Directional Selection • Disrupting Selection • Effective mating system • Fitness • Genotype • Heritable • Innate • Monogamy • Natural Selection • Naturalistic Fallacy • Ontogenetic • Operational Sex Ratio • Phenotype • Phylogenetic • Polyandry • Polygyny • Proximate • Reproductive Variance • Sexual Dimorphism • Sexual Selection • SSSM • Stabilising Selection • Ultimate Sexual Dimorphism