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33 BEHAVIOR AND THE ENVIRONMENT EXTENDED LECTURE OUTLINE Some Behavior is Genetically Determined (p. 732) 33.1 Approaches to the Study of Behavior (p. 732; Fig. 33.1) A. Animals respond to their environment in many ways. B. Behavior encompasses the way an animal responds to stimuli in its environment. C. Animals with nervous systems have far more complex behaviors. 1. A variety of sense organs enable the perception of environmental stimuli. D. Explaining Behavior 1. Proximate causation is analyzed by measuring hormone levels or recording the impulse activity of certain neurons. 2. Ultimate (evolutionary) causation is used to determine how an animal’s behavior influences survival or reproductive success. E. A Controversial Field of Biology 1. Whether behavior is a result of nature [instinct] or nurture [learning] is controversial. 2. We now know that both instinct and learning play a part in behavior. 33.2 Instinctive Behavioral Patterns (p. 733; Fig. 33.2) A. Ethology is the study of animal behavior in natural conditions. 1. Karl von Frisch, Konrad Lorenz, and Niko Tinbergen won the Nobel prize in Physiology for their contribution to ethology. B. An Example of Innate Behavior 1. Konrad Lorenz studied egg retrieval in geese. a. According to his research, egg retrieval behavior is triggered by a sign stimulus which is the appearance of an egg outside of the nest. b. The innate releasing mechanism in the goose’s brain responds to the sign stimulus by providing a fixed action pattern. 2. Niko Tinbergen studied sign stimuli in the mating behavior of male stickleback fish. a. He was able to produce the aggressive display in males by challenging them with unfish-like models, so long as the sign-stimulus was present. 33.3 Gene Effects on Behavior (p. 734; Fig33.3) A. Behavioral genetics is the investigation of behavior by studying genetics. B. Studies of Genetic Hybrids 1. William Dilger found that a hybrid species of lovebird exhibited the behavior of carrying nest material that is intermediate between the parents. C. Studies of Twins 1. A study of 50 sets of human twins revealed many similarities in behavior even though the twins were raised in different environments. 2. Results show that genes play a key role in behavior. D. A Detailed Look at How One Gene Affects Behavior 1. In 1996 a new gene in mice was discovered that determines whether or not female mice nurture their young. Behavior Can Also Be Influenced By Learning (p. 734) 33.4 How Animals Learn (p. 734; Fig. 33.4) A. Animals can later alter their behavior as a result of learning. B. Nonassociative learning is the simplest. 1. One form is sensitization in which repeating a stimulus produces a greater response. 2. Another function is habituation in which a decrease in response to a repeated stimulus occurs. a. Being able to ignore unimportant stimuli is critical to confronting a barrage of stimuli. 197 33.5 33.6 C. Associative learning involves a change in behavior that involves an association between two stimuli. 1. The two major types are classical conditioning and operant conditioning. D. Classical Conditioning 1. The paired presentation of two stimuli causes an animal to form an association. 2. Pavlov caused a dog to salivate when hearing a bell by classical conditioning. E. Operant Conditioning 1. The animal learns to associate its behavior response with reward or punishment. 2. B.F. Skinner conditioned a rat to touch a lever for food reward. F. Imprinting 1. Imprinting forms preferences to other individuals that will influence behavior later in life. 2. In filial imprinting, social attachments form between parents and offspring. Instinct and Learning Interact to Determine Behavior (p. 736; Fig. 33.5) A. Learning preparedness demonstrates that learning is possible only within the boundaries set by instinct. 1. Pigeons can learn to associate food with colors but not with sounds. B. Innate programs [instincts] have evolved because each of them reinforces an adaptive response. 1. The seed a pigeon eats has a distinct color but makes no sound whereas the approach of a predator generates noise but involves no distinctive color. C. Behavior Often Reflects Ecological Factors 1. The genetic component of behaviors has evolved to match an animal to its habitat. D. The Interaction Between Instinct and Learning 1. Peter Marler’s study in song birds suggests that birds have a genetic template that guides them to learn the appropriate song. a. Song acquisition depends on learning, but only the song of the correct species can be learned. Animal Cognition (p. 737; Fig. 33) A. Evidence of Conscious Planning 1. Birds in urban areas remove foil caps from milk bottles to get at the cream. 2. Macaques learn to float grain on water to separate it from sand. 3. Chimpanzees strip a twig of its leaves to use as a tool. 4. Sea otters use rocks to break open clams. B. Problem Solving 1. A chimp figured out how to stack boxers to reach a banana during a 1920s experiment. 2. In Bernard Heinrich’s experiment with a raven, the raven had devised a solution when presented with a novel problem. Evolutionary Forces Shape Behavior (p. 738) 33.7 33.8 Behavioral Ecology (p. 738; Fig. 33.7) A. Investigation of animal behavior is divided into three sorts of questions: the study of its development, the study of its physiological basis, and study of its function. B. Behavioral ecology is the study of how natural selection shapes behavior. C. All genetic differences need not have survival value. D. Niko Tinberger observed that after gull nestlings hatched, the parents quickly removed the shells. 1. He concluded through experimentation that eggshell removal reduces predation of unhatched eggs and increases the survival of offspring. E. Other behaviors enhance energy intake, reduce exposure or increase resistance to disease, enhance the ability to acquire a mate or in some other way, increase an individual’s fitness. A Cost-Benefit Analysis of Behavior (p. 739; Fig. 33.8) A. Behavioral ecologists examine the evolutionary advantage of a behavior by asking if it provides an evolutionary benefit greater than its cost. B. Foraging Behavior 1. The choice of what food to select and how far to go to seek it is called foraging behavior. 198 It involves a trade off between a food’s energy content and the cost of obtaining it. The optimal foraging theory predicts that animals will select food items that maximize their net energy intake per unit of foraging time. 4. Many animals alter their foraging behavior when predators are present. C. Territorial Behavior 1. Animals often move over a home range during the day. 2. Territoriality is the behavior exhibited to defend an individual’s portion of the home range. 3. Territories are defended by displays that advertise occupation and by overt aggression. 4. An animal is territorial if there is an adaptive value to this behavior. Migratory Behavior (p. 740; Figs. 33.9) A. Long-range two-way annual movements are called migrations. 1. It is particularly common in birds. B. The Compass Sense 1. Migrating birds have the ability to detect the earth’s magnetic field and to orient themselves accordingly. 2. Experiments have indicated that inexperienced starlings migrate using compass sense. C. The Map Sense 1. Not much is known about map sense. 2. Young birds move with a flock of experienced older birds and learn to recognize cues. Reproductive Behaviors (p. 742; Figs. 33.10; Table 33.1) A. Reproductive success is influenced by a number of factors that in turn are directly affected by its behavior. B. Sexual Selection is the competition for mating opportunities. 1. Intrasexual selection leads to evolution of structures used in combat with other males. a. Combat for mates is a form of agonistic behavior. 2. Intersexual selection or mate choice leads to the evolution of complex courtship behaviors. C. The Benefits of Mate Choice 1. Females benefits by choosing the male that can best care for offspring. 2. Females choose males with the best territories to maximize reproductive success. 3. Females select vigorous males for their good genetic makeup. D. Mating Systems 1. The typical number of mates an animal has during its breeding season is called the mating system. 2. There are three principal mating systems: monogamy, polygyny, and polyandry. 3. Mating systems have evolved to maximize reproductive fitness. E. Reproductive Strategies 1. A set of behaviors that has evolved to maximize that species reproductive success. 2. 3. 33.9 33.10 Social Behavior (p. 744) 33.11 Communication Within Social Groups (p. 744; Fig. 33.11, 33.12, 33.13) A. Many animals live in social groups in which information is communicated between group members. 1. Mammalian “guards” give an alarm call when a predator appears. 2. Social insects secrete alarm pheromones that trigger attack behavior. 3. Ants deposit trail pheromones between the nest and a food source. 4. Honeybees have an extremely complex dance language which directs nest mates to nectar. B. The Dance Language of the Honeybee 1. A scout bee performs a behavior pattern [waggle dance] on a vertical comb. 2. The path during the dance resembles a figure eight. 199 3. 33.12 33.13 On the straight part of the path, the bee vibrates or waggles her abdomen. a. The scout bee indicates the direction of food by representing angles between the food, hive, and sun. b. The distance is indicated by the tempo of the dance. C. Primate Language 1. Vocalizations of African vervet monkeys distinguish eagles, leopards, and snakes. 2. Chimps and gorillas learn to recognize a large number of symbols and use them to communicate abstract concepts. 3. All human languages share basic structural similarities. Altruism and Group Living (p. 746; Fig. 33.14, 33.15) A. Altruism is the performance of an action that benefits another individual at a cost to the actor. B. One would expect alleles for altruism to be a disadvantage. 1. One explanation is the evolution of the trait for the good of the species. a. Natural selection operates on individuals; not on the species. 2. Group selection can occur but is rare. C. Reciprocity 1. Reciprocal altruism occurs when “partnerships” form in which mutual exchanges of altruistic acts occur because it benefits both. a. Cheaters (non-reciprocators) are discriminated against and are cut off receiving future aid. D. Kin Selection 1. Natural selection will favor any strategy that increases the net flow of an individual’s alleles to the next generation. 2. Selection that favors altruism directed toward relatives is called kin selection. F. Examples of Kin Selection 1. Ground squirrels give alarm calls when they spot a predator. a. Researchers found that females are more likely to give alarm cells than females with no kin nearby. 2. The white-fronted bee-eater lives in colonies of 100-200 birds. a. Kin selection is important in determining helping behavior. b. Helpers are usually males which are related to other birds in the colony. c. When birds have the choice of helping different parents, they usually choose the parents to which they are most closely related. Vertebrate Societies (p. 748; Fig. 33.16) A. Vertebrate social groups are usually less rigidly organized and cohesive compared to insect societies. B. They also display more conflict and aggression among group members than do insect societies. C. Vertebrate societies, like insect societies, have particular types of organization. D. African weaver birds exemplify the relationship between ecology and social organization. 1. One set of species lives in the forest and builds camouflaged, solitary nests. a. Males and females are monogamous. b. They forage for insects to feed their young. 2. Another group nests in colonies in trees on the savanna. a. They are polygynous and feed in flocks on seeds. 3. The feeding and nesting habits of these two sets of species are correlated with their mating systems. E. The naked mole rat is the exception to the general rule that vertebrate societies are not organized like insects. 1. They form large underground colonies with a far ranging system of tunnels and a central nesting area. 2. There is a division of labor in the colony as in insect societies. 3. There is a reproductive division of labor, similar to insects, where breeding is done by a “queen.” 200 33.14 Human Social Behavior (p. 749; Fig. 33.18) A. The most profound lesson of biology is that we humans are animals that are close relatives of the chimpanzee. B. Genes and Human Behavior 1. Ethology, the study of human behavior in natural conditions, has provided much evidence that behaviors evolve. 2. Sociobiology is the study of social behavior and was pioneered by E.O. Wilson of Harvard University. 3. Genes Impact Human Behavior a. Human facial expressions, similar the world over, have deep genetic basis as infants born blind still smile and frown. b. Learning has a huge impact on how humans behave. 4. Diversity is the Hallmark of Human Culture a. The variation in social behavior of other species is vanishingly small compared to the enormous diversity of human cultures. b. Both heredity and learning play key roles in determining how we behave. LEARNING OBJECTIVES Differentiate between the physiological mechanisms in animals that produce behaviors vs. the evolutionary forces responsible for animal behavior. Describe the ethological approach to studying animal behavior. Describe the studies that support the conclusion that genes play a key role in many animal behaviors. Describe habituation/sensitization with conditioning/imprinting. Describe the interaction between instinct and learning using Peter Marler’s experiment with whitecrowned sparrows. Describe cognitive ability in animals using the problem solving ability of the raven. Describe the contribution of Tinbergen to behavioral ecology. Describe the optimal foraging theory. Explain the concept of territoriality. Differentiate between compass sense and map sense. Explain how mate choice, breeding system, and parenting behaviors maximize reproductive success. Describe the waggle dance of the honeybee and the message it represents. Explain how kin selection can increase the frequency of altruism. Compare insect societies with vertebrate societies and describe how the naked mole rats are the exception. Describe how human behavior is similar to behavior in other animals. KEY TERMS proximate causation (p. 732) ultimate causation (p. 732) ethology (p. 733) sign stimulus (p. 733) innate releasing mechanism (p. 733) fixed action pattern (p. 733) behavioral genetics (p. 734) learning (p. 735) nonassociative learning (p. 735) associative learning (p. 735) classical conditioning (p. 735) operant conditioning (p. 735) imprinting (p. 735) 201 cognitive behavior (p. 737) behavioral ecology (p. 738) foraging behavior (p. 739) optimal foraging (p. 739) territoriality (p. 739) migrations (p. 740) sexual selections (p. 741) mate choice (p. 741) mating system (p. 742) alarm call (p. 744) alarm pheromone (p. 744) trail pheromones (p. 744) dance language (p. 744) altruism (p. 746) group selection (p. 746) reciprocal altruism (p. 746) kin selection (p. 747) sociobiology (p. 749) CHANGES TO THE NEW EDITION Refer to the Johnson instructor web site at http://www.mhhe.com/biosci/genbio/tlw4 for a complete list of changes to this edition. CRITICAL THINKING QUESTIONS 1. 2. 3. Describe two advantages of sound communication over visual communication. Discuss how studies of human twins can help determine the genetic components of human behavior. Determine whether fraternal or maternal twins separated at birth would provide the most insight into determining the genetic components of human behavior and explain. FILMS/MEDIA SUGGESTIONS (Telephone and fax numbers and/or web sites for the sources of the following materials are listed in the Appendix.) Biologists at Work. The wide variety of what biologists do for a living is the focus of this video. 1994. 42 minutes. Hawkhill Video, #117. Evolution. This two-part video teaches the basics of modern evolutionary theory. Part I explains how the theories of evolution were developed. Part II shows evolution by natural selection. 1994. 38 minutes. Hawkhill Video, #108. Heredity. This three-video set includes a wide variety of topics ranging from traits, genes, and alleles to multiple genes, chromosomes, and sex linkage. 36 minutes each. CLEARVUE/eav, WW5VH 1270 Evolution. This two-part video teaches the basics of modern evolutionary theory. Part I explains how the theories of evolution were developed. Part II shows evolution by natural selection. 1994. 21 minutes. Hawkhill Video, #108. 202 Tales from the Hive. This NOVA presentation portrays life in a bee colony. 2000. 60 minutes. Carolina Biological Supply, CE-49-2512 Primates. This Discovery Channel production shows the numerous types of primates living in exotic places. Family order, social life, and tool use are shown. 52 minutes. Films for the Humanities and Sciences, BLV11927 203