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BIOLOGY CONCEPTS & CONNECTIONS Fourth Edition Neil A. Campbell • Jane B. Reece • Lawrence G. Mitchell • Martha R. Taylor CHAPTER 37 Behavioral Adaptations to the Environment From PowerPoint® Lectures for Biology: Concepts & Connections Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings Tracking Jaguars • The study of an animal's behavior is key to understanding its evolution and ecological roles • In the 1980s, zoologist Alan Rabinowitz studied jaguar behavior and ecology in Belize Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Rabinowitz monitored the activities of six adult jaguars over extended periods – He fitted the animals with radio transmitter collars to trace their movements in the dense jungle Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Rabinowitz made many important discoveries regarding jaguar breeding, hunting, and signaling – Jaguars are solitary hunters – They shun contact with other jaguars except during the breeding season – Male jaguars help care for their young • Many of these behaviors provide clues about jaguar evolution Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings CLASSIC CONCEPTS IN BEHAVIOR 37.1 Behavioral biologists study the actions of animals in their natural environments • Behavioral biology is the study of what animals do when interacting with their environment • Behavior can be interpreted in terms of proximate causes, or immediate interaction with the environment Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Nobel laureates Karl von Frisch, Konrad Lorenz, and Niko Tinbergen were among the first experimentalists in behavioral biology • Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • A classic Tinbergen experiment deals with the nesting behavior of the digger wasp – The female wasp often excavates and cares for four or five separate nests • Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • In the experiment, Tinbergen placed a circle of pinecones around a nest opening Nest 1 Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings Figure 37.1, Part 1 • After the female flew away, Tinbergen moved the pinecones a few feet to one side of the nest opening – When the female wasp returned, she flew to the middle of the circle of pinecones rather than to the actual nest opening Nest 2 Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings No Nest Figure 37.1, Part 2 • Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening – This time the wasp flew to the stones Nest 3 Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings No Nest Figure 37.1, Part 3 • The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves • This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Behavioral ecologists are especially interested in the ultimate causes of behavior, which are evolutionary • Natural selection preserves behaviors that enhance fitness Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings 37.2 Behavior results from both genes and environmental factors • Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning) • In biology, the nature-versus-nurture debate is not about either/or – It is about how both the genes and the environment influence the development of phenotypic traits Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • The gathering of nest materials by lovebirds has genetic and environmental components Single long strip carried in beak (Fischer’s lovebird) Several short strips tucked under feathers (peach-faced lovebird) Tucking failure Strip in beak Hybrid behavior Figure 37.2 Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings 37.3 Innate behavior often appears as fixed action patterns • Sign stimuli trigger innate, essentially unchangeable fixed action patterns (FAPs) – A sign stimulus is often a simple clue in an animal’s environment • The genetic programming underlying FAPs ensures that such activities are performed correctly without practice – Such as many parent-offspring interactions Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner – This is a fixed action pattern – She carries this sequence to completion, even if the egg slips away during the process Figure 37.3A Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Several key events in the life cycle of the European cuckoo are determined by fixed action patterns – Egg-laying behavior 1 2 3 Figure 37.3B Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings – The behavior of the cuckoo hatchling ejecting the host eggs from the nest – The feeding behavior of a foster mother to the cuckoo chick Figure 37.3B Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings 37.4 Learning ranges from simple behavioral changes to complex problem solving • Learning is a change in behavior resulting from experience • Habituation is one of the simplest forms of learning – An animal learns not to respond to a repeated stimulus that conveys little or no information – For example, birds eventually become habituated to scarecrows and no longer avoid nearby fruit trees Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings Table 37.4 Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings 37.5 Imprinting is learning that involves both innate behavior and experience • Imprinting is irreversible learning limited to a sensitive period in an animal's life • Imprinting enhances fitness by enabling rapid learning Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • In Konrad Lorenz's most famous study, he used the graylag goose to demonstrate imprinting – Lorenz took over the maternal role for a group of goslings Figure 37.5A Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Not all examples of imprinting involve parentoffspring bonding – Although newly hatched salmon do not receive any parental care, they seem to imprint on the complex mixture of odors unique to the freshwater stream where they hatch – This allows the salmon to find their way back to the stream to spawn after spending a year or more at sea Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Imprinting plays an important role in song development for many kinds of birds Figure 37.5B Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings 37.6 Many animals learn by association and imitation • Associative learning is learning that a particular stimulus or response is linked to a reward or punishment – These ducks have learned to associate humans with food handouts – They congregate rapidly whenever a person approaches the shoreline Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings Figure 37.6A • Trial-and-error learning is a common form of associative learning – An animal learns to associate one of its own behavioral acts with a positive or negative effect Figure 37.6B Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Imitation is learning by observing and mimicking the behavior of others – This form of learning is not limited to a sensitive period – Many predators, including cats and coyotes, seem to learn some of their basic hunting tactics by observing and imitating their mother Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings 37.7 Animal cognition includes problem-solving behavior • Some animals exhibit problemsolving behavior – Examples: chimpanzees and ravens Figure 37.7A, B Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Cognition is the ability of an animal's nervous system to perceive, store, process, and use information • The study of animal cognition is called cognitive ethology – Consciousness, or awareness, is one area of study in cognitive ethology Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Answers to questions about animal thinking may profoundly affect how we interact with other animals and how we view ourselves • Scientists face unique challenges in determining the differences between human consciousness and that of other animals • Are humans unique, or are we simply at one end of a consciousness continuum? Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings ECOLOGICAL ROLES OF BEHAVIOR 37.8 An animal's behavior reflects its evolution • Behavior is an evolutionary adaptation that enhances survival and reproductive success • Behavior evolves as natural selection finetunes an animal to its environment – The hunting and reproduction behaviors of jaguars – Nest location by digger wasps – Imprinting of goslings Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings 37.9 Biological rhythms synchronize behavior with the environment • Animals exhibit a great variety of rhythmic behavior patterns • Circadian rhythms are patterns that are repeated daily – Sleep/wake cycles in animals and plants • Circadian rhythms appear to be timed by an internal biological clock Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • In the absence of environmental cues, these rhythms continue – But they become out of phase with the environment Figure 37.9A Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Long-term isolation has been used to study human circadian rhythms • Body rhythms affect our general well-being, work efficiency, and decision-making ability Figure 37.9B Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings 37.10 Animal movement may be oriented to stimuli or landmarks • Movement in a directed way enables animals to – avoid predators – migrate to a more favorable environment – obtain food – find mates and nest sites Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • A simple type of animal movement is kinesis – Random movement in response to a stimulus • Another simple type of animal movement is taxis – A more or less automatic movement directed toward or away from some stimulus – Examples include rheotaxis, chemotaxis, and phototaxis • Some animals use landmarks to find their way within an area Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings 37.11 Movement from place to place often depends on internal maps • Many animals formulate cognitive maps – Internal representations of spatial relationships among objects in their surroundings • Some animals undertake long-range migrations – Examples: whales, birds, monarch butterflies • Animals navigate using the sun, stars, landmarks, or Earth's magnetism Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Migrating gray whales use coastal landmarks to stay on course Arctic Ocean FEEDING GROUNDS Siberia Alaska NORTH AMERICA Pacific Ocean Atlantic Ocean Baja California BREEDING GROUNDS Figure 37.11A Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • The indigo bunting learns a star map and navigates by fixing on the North Star Paper Ink pad Funnelshaped cage Figure 37.11B Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings 37.12 Behavioral ecologists use cost/benefit analysis in studying feeding behavior • Animals are generally selective and efficient in their food choices – Some animals, such as gulls, are feeding “generalists” – Other animals, such as koalas, are feeding “specialists” Figure 37.12A, B Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • The mechanism that enables an animal to find particular foods efficiently is called a search image • Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energy – This is the theory of optimal foraging Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Whenever an animal has food choices, there are a number of tradeoffs – A bass can get more usable energy from minnows, but crayfish are easier to catch – However, it may take more time to eat a crayfish because of its tough exoskeleton Figure 37.12C Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Jaguars in Belize have many prey options – Armadillos, only about 5kg of food for the jaguar, are abundant and easy to catch – 200 kg tapirs are less abundant and run quickly – In optimal foraging, the armadillo is the preferred prey Figure 37.12D Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground, where it is exposed to predators Figure 37.12E Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings SOCIAL BEHAVIOR AND SOCIOBIOLOGY 37.13 Sociobiology places social behavior in an evolutionary context • Social behavior is defined as the interaction among members of a population • The discipline of sociobiology studies social behavior in the context of evolution Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings 37.14 Rituals involving agonistic behavior often resolve confrontations between competitors • Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population • Agonistic behavior can directly affect an individual's evolutionary fitness – The victor often gains first or exclusive access to mates Figure 37.14 Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings 37.15 Dominance hierarchies are maintained by agonistic behavior • Many animals live in social groups maintained by agonistic behaviors • Dominance hierarchy is the ranking of individuals based on social interactions Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Chickens establish a “peck order” • Resources are often partitioned based upon the dominance hierarchy Figure 37.15 Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings 37.16 Talking About Science: Behavioral biologist Jane Goodall discusses dominance hierarchies and reconciliation behavior in chimpanzees • Dr. Jane Goodall is one of the world's best-known biologists • She has studied the behavior of chimpanzees in their natural habitat, in East Africa, since the early 1960s Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings Figure 37.16A • Dr. Goodall's research indicates that dominance hierarchies and reconciliation behaviors are integral parts of the lives of many primates – For example, a chimpanzee that has threatened another member of its group may use a hand gesture to invite reconciliation Figure 37.16B Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings 37.17 Territorial behavior parcels space and resources • Humans tend to space themselves out when they are close to others – They establish what we might call personal territories Figure 37.17A Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Many animals exhibit territorial behavior – It is a form of social behavior that partitions resources Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • A territory is an area that individuals defend and from which other members of the same species are usually excluded – The size of the territory varies with species, the function, and the available resources – Territories are typically used for feeding, mating, and/or rearing young Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Territoriality is often maintained by agonistic behavior – These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other Figure 37.17B Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Territorial rights are proclaimed continually in a variety of ways – Bird songs – Noises, such as the bellowing of sea lions and the chattering of squirrels – Defecation in open areas – Scent markers, such as urine Figure 37.17C Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings 37.18 Mating behavior often involves elaborate courtship rituals • Courtship behaviors advertise the species, sex, and physical condition of potential mates • Many species court in pairs, isolated from the rest of the population – Example: the common loon Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • The courtship and mating of the common loon involves the male and female loons swimming side by side while performing a series of displays – The courting birds frequently turn their heads away from each other – The birds then dip their beaks in the water 1 2 Figure 37.18A, Part 1 & 2 Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings – The loons then submerge their heads and necks – Prior to copulation, the male invites the female onto land by turning his head backward with his beak held downward 3 4 Figure 37.18A, Part 3 & 4 Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • The loons then copulate 5 Figure 37.18A, Part 5 Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Some species perform courtship rituals in groups – Mates are chosen from a group of candidates • The male sage grouse performs by strutting about and erecting his tail feathers in a bright, fanlike display – Researchers hypothesize that there is a connection between a male’s display and the quality of his genes Figure 37.18B Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings 37.19 Complex social organization hinges on complex signaling • In behavioral ecology, a signal is a form of communication that causes a change in behavior in another animal • Social behavior depends on signaling – Sounds – Scents – Displays – Touches Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • This ring-tailed lemur is communicating aggression with its prominent tail – Prior to this display, it smeared its tail with odorous secretions from glands in its forelegs – By waving its scented tail over its head, the lemur transmits both visual and chemical signals Figure 37.19A Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Honeybees perform dances that seem to communicate the direction and distance of nectar to other members of the colony Figure 37.19B Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings – The round dance indicates that food is nearby, in an unspecified direction Figure 37.19C Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings – The waggle dance probably indicates both distance and direction of food (1) 30º (3) (2) (1) Beehive (2) Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings (3) Figure 37.19D 37.20 Altruistic acts can be explained by evolution • Some animals exhibit altruism – Behavior that reduces an individual's fitness while increasing the fitness of another individual • Examples of altruistic behavior can be found in honeybee social systems – Worker bees spend their lives laboring on behalf of the fertile queen bee Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Naked mole rats also exhibit altruism – The reproducing female (queen) mates with one to three males (kings) – The rest of the colony consists of nonreproductive females and males, who care for and protect the queen, kings, and new offspring Figure 37.20A, B Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Some altruistic acts can be explained in terms of kin selection – An animal can increase the survival of genes like its own by helping relatives • In reciprocal altruism, a favor may be repaid later by the beneficiary or another member of the social system – This explains altruistic acts by nonrelatives Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings 37.21 Talking About Science: Edward O. Wilson promoted the field of sociobiology and is a leading conservation activist • Evolutionary biologist Edward O. Wilson promoted a relatively new area of research called sociobiology Figure 37.21 Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Sociobiology is based on the concept that social behavior evolves, like anatomical traits, as an expression of genes • Sociobiologists thus believe that natural selection underlies many human behaviors Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings • Wilson believes that there is a biological basis for some of the human behaviors that have created our current biodiversity crisis – Caring only about a small piece of land, a limited band of kinfolk, and a time span of a few generations Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings 37.22 Connection: Both genes and culture contribute to human social behavior • Human behavior has a genetic basis but is quite variable and influenced by learning and culture • One of our most important cultural attributes is education – Education fosters behavior that ultimately evolved because it has adaptive value in the human species Figure 37.22 Copyright © 2003 Pearson Education, Inc. publishing as Benjamin Cummings