Behavioral Ecology Chapter 55 Background and early studies What is behavior? • The way an animal responds to stimuli in its environment • Behavior is a pattern of responses to a stimulus How can we explain behavior? • How it works physiologically Proximate answer • The adaptive value of the behavior Ultimate answer • So, behavioral scientists study what behavior an organism does, how it does it and why it does it. Behavior • Behavior is influenced by both genetic and environmental factors (behavior is an expression of genetic and environmental cues) Nature versus nuture argument Gene by environment interaction is important Genes and behavior • Almost all behavior has a genetic basis • So, behavior is subject to evolution through natural selection! • Behavior is essential to understanding an animal’s evolution and ecological interactions History of behavioral ecology • Konrad Lorenz--imprinting and instinct • Karl von Frisch--honeybee dances & communication • Niko Tinbergen--gull behavior and learning+instinct • All three shared Nobel Prize in physiology/medicine in 1972 Konrad Lorenz • Imprinting – learning that is limited to a specific time period in an animal’s life and that is generally irreversible • Critical period (sensitive period)– a limited phase in an individual animal’s development when learning of particular behaviors can take place Filial imprinting Konrad Lorenz • Wrote book “On Aggression” where he asserts that human aggressive impulses are to a degree innate, and draws analogies between human and animal territorial behavior These assertions have engendered considerable controversy Karl von Frisch • Honeybees communicate: Direction to food source Distance to food source Quality of food source Two types of dances • Round dance • Waggle dance Distance to food source • The orientation of the straight run in the waggle dance conveys the direction of the food source, relative to the position of the sun Distance to food source • The duration or tempo of the straight runs conveys the distance between nest and target: Dance tempo slows down with increasing distance to the food source. The farther away the target, the longer the straight run part of the dance Quality of food source • The intensity of the dance conveys the quality of the food source Robot bees Tinbergen - gull feeding behavior Gull feeding behavior Learning • This related experiment with artificial beaks shows learning by gull chicks Tinbergen • Published famous paper on the 4 questions in behavioral biology Tinbergen’s four questions Tinbergen’s four questions Tinbergen’s four questions 1) Mechanisms Genetic Hormonal Physiological In grouse, males begin to exhibit mating behaviors when sex hormones rise Tinbergen’s four questions 2) Development Instincts Learning Instincts x learning In grouse, females seem to learn mate choice from their mothers Tinbergen’s four questions 3) Adaptiveness Survival Reproduction Male grouse that attract the most females have higher reproductive success. However, display is costly to survival Tinbergen’s four questions 4) Phylogenetic history Development of behavior through ancestry The grouse family generally shows this mate choice behavior, where males are elaborate Development of behavior Innate or instinctive behavior • Performed without having been learned • Usually triggered by simple sign stimuli • Response to the stimulus is a stereotyped motor program (hard-wired) Preset paths in nervous system Innate behaviors • Innate behaviors are developmentally fixed • Not completely genetic, as a physical environment is necessary for the gene to be expressed • The key to innate behaviors is that they are expressed under a wide variety of environmental conditions Instinctive behavior • Sign stimuli are often nonspecific • Innate releasing mechanism - a neural component of the organism that provides the motor program • Fixed action pattern – a sequence of behavioral acts that is essentially unchangeable and usually carried to completion once started Instinctive behavior • When a goose sees an egg outside the nest (sign stimulus), it begins a repeated movement of dragging the egg with its beak and neck Stickleback • Male stickleback fish will attack anything with a red underside Supernormal stimuli • Given a choice, animals respond to a larger stimuli than a smaller one Supernormal stimuli Behavior is genetic Genetic basis of behavior • In some cases, single genes control behavior Adaptive behavior • Behavior that promotes reproductive success • Frequency of individuals expressing this behavior will be maintained or increase in successive generations due to natural selection Learning influences behavior • Animals alter their behavior as a result of previous experiences, which is learning Feeding in gull hatchlings Types of learning • Nonassociative learning Does not require an animal to form an association between 2 stimuli or between a stimulus and response Habituation • A type of nonassociative learning • An individual learns NOT to respond to a stimulus that has neither good nor bad consequences • Pigeons in cities learn that people are no threat and do not flee from them • Deer become increasingly tame in parks • Ability to ignore signals is adaptive Types of learning • Associative learning Requires an association between 2 stimuli or between a stimulus and response Behavior is “conditioned” through the association Two major types: Classical conditioning Operant conditioning Differ in the way associations are established Classical conditioning • Pair presentation of two different stimuli causes association between the two stimuli • Also called Pavlovian conditioning Operant conditioning • A behavior becomes associated with its consequences • “Trial-and-error” learning What type of conditioning is this? What has the coyote learned? Instinct and learning • Innate predispositions toward forming certain associations Pigeons can learn to associate food with colors, but not with sound • Learning is possible only within the boundaries set by instinct • In nature, adaptation by learning is important to survival Spatial learning • Through experience with an environment, an organism creates a mental map • Birds remember the location of thousands of places where they have stored food Animal Cognition • Do animals other than humans possess cognition? Cognition - the ability of an animal’s nervous system to perceive, store, process, and use information gathered by sensory receptors In other words, do they process information in a way that suggests thinking? Insight Learning • An animal solves a problem without trial-and-error attempts at a solution • Captive chimpanzees show insight learning when they solve a novel problem, as when they stack boxes to reach food that is out of reach Instincts and learning • Instinct = behavior exhibited without feedback from environment • Learned behavior = behavior that can be altered in response to information acquired from environment • Instinct and learning are not exclusive - they can interact! Innate song Bird Song: Instinct + Learning • Bird comes hard-wired to listen for certain acoustical cues; instinctively pays attention to particular sounds • Genetic template guides birds to learn the appropriate song • Which dialect the bird sings depends on what song it hears • • Exposed to own species song during development Not exposed to song New information shows that social interaction may override the genetic template! Importance of social interactions