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Genes vs. Environment? Cake analogy… recipe + ingredients = cake genes + environment = trait Is a cake more recipe or more ingredients? Is the difference between 2 cakes due to differences in the recipe or ingredients? Natural Experiment: Cross-fostering of galah chicks by pink cockatoos Which species will the galahs behave like? Are behaviors results of social environment or genetic input? Depends on the behavior… Begging calls galah Alarm calls galah Contact calls cockatoo Flight pattern cockatoo Food preference cockatoo galah pink cockatoo Categorization of Behaviors: Innate Learned Begging calls Contact calls Alarm calls Flight pattern Food preference Learned Behaviors Studied by behaviorists (Skinner) Modified by specific experiences Two main types of associative learning… Classical Conditioning Famous example of Pavlov’s dogs Conditioning of response (salivation) by association of the unconditioned stimulus (food) with a conditioned stimulus (bell) (Basis for successful advertising) Operant Conditioning Association of voluntary activity with the consequences that follow (Skinner box) Can have positive and negative reinforcement Basis of dog training! Operant Conditioning After a behavior is performed: Add a stimulus reinforcement + (food) punishment _ (shock) Remove a stimulus Effect _ Increases behavior (alarm clock) + (grounded) Decreases behavior Basics of operant condition (Dog Training 101) The reinforcement must follow the desired behavior within 1 sec (with one exception…). Negative reinforcement must be consistent to extinguish a behavior. Inconsistent positive reinforcement only increases the frequency of the behavior! Limitations/exceptions of operant conditioning Nausea as punishment is effective even hours after the behavior and administered only once. Ultimate explanation for this? Some combinations of behaviors and consequences can not be associated Innate/instinctive behaviors Studied by ethologists (Tinbergen, Lorentz) Appears fully formed the first time it is performed Usually triggered by a simple cue… Pecking behavior by gull chicks Fixed Action Pattern (FAP): Any innate behavior that gets carried to completion once triggered by a cue. Sign stimulus or releaser: the cue that initiates a FAP Yawning FAP and releaser Consistent in length and appearance Difficult to stop once started!! Susceptible to cheaters.. Rove beetle & ant Beetle larvae produce ant pheromone, releasing “take to brood chamber” behavior Also get fed by stimulating regurgitation behavior in workers, both as larvae and as adults Supernormal Stimulus More effective releaser of behavior than biologically correct stimulus Previous example of long stick with dot releasing begging in gull chicks Many examples in nature and is basis for one theory on male ornamentation. Genetics of Behavior How to measure the genetic component of behavior. 1) Control environmental variation 2) Similarities between relatives & non-relatives 3) Selection experiments 4) Modern techniques. Heritability (H2): the proportion of variance in a trait attributable to genetic variation. Phenotype, Genotype, Environment P = G + E + (G x E) H2 = VG / VP VG H2 = VG + VE + V (G x E) 1) Control Environmental Variation By raising genetically variable individuals in the same environment, we can estimate amount of phenotypic variation that is genetic (VG). Do the same in very different environments to estimate total phenotypic variance (VP). How do we measure the environmental component of a behavior (VE)? Raise individuals that are genetically similar in different environments. Norm of reaction: range of phenotypic variation for a single genotype resulting from environmental differences. Migration routes of blackcap warbler populations: Change in blackcap warbler migratory behavior in Britain in 1950’s prompted investigations into the genetic control of migration. Hypothesis 1: Birds over-wintering in Britain had lost ability to migrate. Test: See if birds caught in Britain in winter and their offspring raised in lab show same pre-migratory activity as German birds raised in lab. Result: They do show the same activity Conclusion: Hypothesis 1 not supported, birds have not lost their migratory activity. Hypothesis 2: Birds over-wintering in Britain have a genetic difference which causes them to migrate in a different direction Test: Check migratory direction of offspring of British birds and German birds reared in the same laboratory conditions. Results: Conclusions: Appears to be a genetic difference in migratory behavior. Why have western mutants not arisen in other populations? Another test of the genetic basis hypothesis: Cross birds from southwestern Germany (brown arrows) to birds from Austria (white arrows). Offspring (blue arrows) flew in intermediate direction! 2) Similarities between relatives and non-relatives: Parent-offspring comparisons Twins, full-sibs, and half-sib comparisons But don’t forget about influence of similar environment!! Identical twins reared apart – opposite of controlled environment studies Coefficient of Relatedness: the proportion of alleles, on average, that two individuals share. C.R. Identical (monozyotic) twins Parent-offspring Fraternal (dizygotic) twins Full siblings Half siblings Step siblings non-relatives 1 0.5 0.5 0.5 0.25 0 0 IQ studies in humans Hypothesis: Differences in IQ scores are caused by differences in genes ONLY Conclusions? 3) Selection Experiment In a behaviorally variable population, only allow certain individuals to reproduce. If the differential reproduction results in a change in the behaviors in the population, then there MUST be genetic variation associated with the behavioral variation! The speed of the change gives an indication of how much of the behavioral difference was due to genetic differences Nest building behavior in mice 1) Measure the amount of cotton used by mice to make nests. 2) Create a “high” line with mice that used lots of cotton, “low” line with mice that used little cotton, and “control” line of mice picked at random. 3) Continue selection (within each line) over several generations. Other selection experiments: flight speed in Drosophila calling behavior in crickets Are there genes for behaviors? “anxiety and aggression” gene? http://www.sciencedaily.com/releases/2003/01/030123072840.htm “alocholism” gene? Gene Linked To Alcoholism Alcoholism tends to run in families, suggesting that addiction, at least in part, has an underlying genetic cause. Now, researchers at the University of Illinois at Chicago have discovered a gene linked to alcohol dependency. Laboratory mice deficient in the gene were found to consume excessive amounts of alcohol, preferring ethanol to water and evincing highly anxious behavior in a maze test. Drunken Fruit Flies Reveal Molecular Pathway Regulating Sensitivity To Alcohol (June 17, 1998) -- Researchers at UC San Francisco have identified a molecular pathway in intoxicated fruit flies that is responsible for regulating the flies' meandering, wobbling responses to alcohol. http://www.sciencedaily.com/releases/2004/05/040526063645.htm CNN Pinker says it's nature, not nurture. Genes, instincts determine thoughts, feelings, behavior According to Steven Pinker, every human exclamation, every chuckle, every expression of love stems not from life experience, but from millions of years of human development. http://www.cnn.com/2004/WORLD/americas/04/16/pinker/index.html CNN Caution urged for brain research on violence Research into brain patterns linked to violent behavior is important, but should be approached with caution. "The research ... provides a window into the many factors that influence behavior. Early experiences are important but not necessarily fixed." http://archives.cnn.com/2000/HEALTH/07/28/violent.brains/index.html 'Mood Genes: Hunting for Origins of Mania and Depression' By Samuel H. Barondes “sitter” and “rover” Drosophila larvae True-breeding parents “sitter” and “rover” Drosophila larvae True-breeding parents F1 = all rovers F2 = 3:1 rovers:sitters “larval feeding behavior” gene: dg2 cGMP-dependent protein kinase Alleles = enzyme variants with different catabolic efficiencies Produced only in cells in olfactory system and brain Affects reception and assessment of environmental cues? Changes in nervous system producing locomotion 4) Modern methods for finding genes that influence behavior: Induced-mutation experiments: Mutate individuals, look for variation in behavior, then figure out which gene was mutated to cause change Knock-out experiments: Inactivate a specific gene, then look for any changes in behavior 4) Modern methods for finding genes that influence behavior: QTL mapping: Using many markers (quantitative trait loci) throughout genome to identify chromosome region(s) that correlate with behavior. Gene expression (microarrays): Quantify variation in gene expression among individuals performing different behaviors. Additivity or dominance: the effect of one allele on another allele at the same genetic locus. Epistasis: the effect of one genetic locus on another independent locus. Pleiotropy: the effect of a single gene or two or more phenotypic traits. Polygenic: the effect of two or more genes on a single phenotypic trait. Phenotypic plasticity: the interaction between genetic factors and the environment (e.g, norm of reaction). Genomic Imprinting Effects (gene products) of a gene differ depending on whether inherited from mother or father What fitness benefit is gained by changing gene function depending on the parent it’s inherited from? Hypothesis: Benefits stems from conflict between the parents over investment in the offspring. Mothers may reduce investment in current offspring to invest in future offspring, while father only cares about current offspring, because mother’s future offspring may not be his. Proximate studies: genetic variation for behavioral variation is maintained. Ultimate studies: why is genetic variation maintained if selection usually acts to decrease variation!?! One of the biggest unanswered questions - what maintains genetic variation in traits under strong selection. Proximate studies: genetic variation for behavioral variation is maintained. Ultimate studies: why is genetic variation maintained if selection usually acts to decrease variation!?! Fitness benefits of genetic variants are not constant, but depend on environment, and the environment is variable? Back to rovers and sitters – why are there two feeding strategies? Hypothesis: Sitting is best strategy at low densities, while roving is best in high densities. Test: Raise random sets of larvae in different environments and measure average movement after several generations.