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The Living World Fifth Edition George B. Johnson Jonathan B. Losos Chapter 39 Behavior and the Environment Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. 39.1 Approaches to the Study of Behavior • Behavior can be defined as the way an animal responds to stimuli in its environment proximate causation deals with how the behavior works • the field of psychology often focuses on this ultimate causation deals with why the behavior evolved • the field of ethology (evolutionary behavior) typically focuses on this Figure 39.1 Two ways to look at a behavior 39.1 Approaches to the Study of Behavior • The study of behavior has a long history of controversy one issue has been the question of whether an animal’s behavior is determined more by an individual’s genes, or by its learning and experience in other words, is behavior the result of nature (instinct) or nurture (learning)? 39.2 Instinctive Behavioral Patterns • Ethology is the study of animal behavior in natural conditions ethologists study the instinctive nature of animal behavior behavior in animals is often stereotyped and based on preset paths in the nervous system • the trigger for the behavior is a sign stimulus • the animal responds to this stimulus via the innate releasing mechanisms that provides the neural instructions • the response is called the fixed action pattern Figure 39.2 Sign stimulus and fixed action pattern 39.3 Genetic Effects on Behavior • Many animal behaviors are strongly influenced by genes passed from parent to offspring behavioral genetics investigates the inheritance of genes connected to behavior • for example, identical twins reared in different environments show many similarities, indicating that genes play a key role in determining human behavior 39.3 Genetic Effects on Behavior • In mice, the fosB gene determines whether or not female mice will nurture young • When a female lacks the fosB allele, she will ignore her newborn babies Figure 39.3 A gene alters maternal care 39.4 How Animals Learn • In many cases, animals alter their behavior as a result of previous experiences this is called learning • nonassociative learning – this is the simplest type of learning – it does not require an animal to form associations between two stimuli or between a stimulus and a response » sensitization occurs when repeating a stimulus produces a greater response » habituation is a decreased response to a repeated stimulus 39.4 How Animals Learn • Associative learning is a form of learning in which behavior is modified, or conditioned, because of an association classical conditioning occurs when paired stimuli are presented, causing the animal to form an association between the stimuli • for example, Ivan Pavlov’s work with dogs – unconditioned stimulus is meat powder, which produces salivating – conditioned stimulus is the sound of a bell, which the dog associates with the meat powder 39.4 How Animals Learn • Operant conditioning is a form of learning in which an animal learns to associate its behavioral response with a reward or punishment B. F. Skinner’s worked with rats exposed to an experimental cage called a “Skinner box” • the rats learned through trial-and-error to depress a lever in order to be fed 39.4 How Animals Learn • As an animal matures, it may form preferences or social attachments to other individuals this process is called imprinting and is sometimes considered a type of learning • in filial imprinting, social attachments form between parents and offspring – for example, young birds of some species begin to follow their mother or the first object they see after hatching Figure 39.4 An unlikely parent 39.5 Instinct and Learning Interact to Determine Behavior • Some animals have innate predispositions toward forming certain associations learning preparedness means that learning is only possible within the boundaries set by instinct for example, song development in birds is based on a genetic template that guides learning the appropriate song • only the song of the correct species can be learned Figure 39.5 Song development in birds involves both instinct and learning 39.6 Animal Cognition • In recent years, serious attention has been given by researchers to the topic of animal awareness do animals other than humans show cognitive behavior—that is, do the process information and respond in a manner that suggesting thinking? • for example, problem solving Figure 39.6 Problem solving by a raven 39.7 Behavioral Ecology • The investigation of animal behavior can be conveniently divided into three areas the study of its development the study of its physiological basis the study of its function • Behavioral ecology is the study of how natural selection shapes behavior Figure 39.7 The adaptive value of eggshell removal 39.8 A Cost-Benefit Analysis of Behavior • Behavioral ecologists examine the evolutionary advantage of behavior by asking if it provides an evolutionary benefit greater than its cost • Optimal foraging theory predicts that animals will select food items that maximize their net energy intake per unit of foraging time 39.8 A Cost-Benefit Analysis of Behavior • Territoriality is a behavior in which an individual defends a portion of its home range and uses it exclusively territories are defended by displays that advertise that the territories are occupied and by overt aggression the adaptive value of territoriality depends on the trade-off between the benefits and the costs Figure 39.8 The benefit of territoriality 39.9 Migratory Behavior • Many animals breed in one part of the world and spend the rest of the year in another migrations are long-range two-way annual movements • compass sense is an innate ability to move in a particular direction or bearing • map sense is learned ability to adjust a bearing depending on the animal’s location Figure 39.9 Starlings learn how to navigate 39.10 Reproductive Behaviors • Sexual selection is competition for mating opportunities intrasexual selection • occurs between members of the same sex • leads to the evolution of structures used in combat, such as antlers intersexual selection • is also called mate choice and occurs between members of the opposite sex • leads to the evolution of complex courtship behaviors and of ornaments Figure 39.10 The male peacock’s feather is a product of sexual selection 39.10 Reproductive Behaviors • The mating system is the typical number of mates an animal has during its breeding season monogamy is when one male mates with one female polygyny is when one males mates with more than one female polyandry is when one female mates with more than one male 39.11 Communication Within Social Groups • In animals living in groups, information is communicated between group members alarm call is given by an animal acting as a “guard” and warns the group of a predator alarm pheromone is secreted by social insects and triggers attack behavior by the group trail pheromone is secreted by social insects to lead colony members to a food source Figure 39.11 The chemical control of fire ant foraging 39.11 Communication Within Social Groups • Honeybees have an extremely complex dance language that directs nestmates to rich nectar sources Karl von Frisch unraveled the details of the communication system • a waggle dance is performed by a scout upon its return to the hive – the dance traces figure-eight and conveys information about the direction and distance of the food source Figure 39.12 The waggle dance of honeybees 39.11 Communication Within Social Groups • Some primates have a “vocabulary” that allows individuals to communicate the identity of specific predators • Language is the primary channel of human communication but nonverbal signals may also convey information Figure 39.13 Primate semantics 39.12 Altruism and Group Living • Altruism is the performance of an action that benefits another individual at a cost to the actor for example, helpers at the nest assist parent birds in raising young at first glance, altruistic behavior appears to be counter-selective for an individual but the evolution of this behavior may be explained by • reciprocity – mutual exchanges of altruistic behaviors benefit both participants, provided there are no cheaters • kin selection – selection that favors altruism towards relatives Figure 39.14 An altruistic act—or is it? Figure 39.15 Kin selection is common among vertebrates 39.13 Vertebrate Societies • For the most part, vertebrate social groups are less rigidly organized and cohesive than insect societies the organization of the group is most influenced by ecological factors • Naked mole rats are an exception to the rule and have a highly organized social structure with a reproductive division of labor Vertebrate Societies Figure 39.16 Savanna-dwelling African weaver birds Figure 39.17 Naked mole rats live in structured colonies Figure 39.18 A chimpanzee family group 39.14 Human Social Behavior • Sociobiology is the idea that complex social behaviors of animals, including humans, should evolve human populations exhibit a wide variety of cultural differences with respect to behavior human cultural diversity is no doubt largely determined by learning and experience Figure 39.19 A New York City street scene Inquiry & Analysis • What is the most energetically optimal mussel size for the crabs to eat, in mm? • Do shore crabs tend to feed on those mussels that provide the most energy? Graph of Energy Budget of a Shorecrab Mussel Diet