What is behavior? Behavior everything an animal does & how it does it response to stimuli in its environment innate inherited, “instinctive” automatic & consistent learned ability to learn is inherited, but the behavior develops during animal’s lifetime variable & flexible change with experience & environment

Why study behavior? Evolutionary perspective… part of phenotype acted upon by natural selection lead to greater fitness? lead to greater survival? lead to greater reproductive success?

What questions can we ask? Proximate causes immediate stimulus & mechanism (physiological) “how” & “what” questions Ultimate causes evolutionary significance how does behavior contribute to survival & reproduction adaptive value “why” questions Proximate cause questions Male songbirds sing during the breeding season as a response to a high level of testosterone which binds to hormone receptors in the brain & triggers the production of song. Ultimate cause questions The male sings to defend territory from other males & to attract a female with which to reproduce. This is the evolutionary explanation for the male’s vocalization. The red–crowned cranes, like many animals, breed in spring and early summer. A proximate question about the timing of breeding by this species might be, “How does day length influence breeding by red–crowned cranes”? A reasonable hypothesis for the proximate cause of this behavior is that breeding is triggered by the effect of increased day length on an animal’s production of and responses to particular hormones. Indeed, experiments with various animals demonstrate that lengthening daily exposure to light produces neural and hormonal changes that induce behavior associated with reproduction, such as singing and nest building in birds. In contrast to proximate questions, ultimate questions address the evolutionary significance of a behavior. Ultimate questions take such forms as, Why did natural selection favor this behavior and not a different one? Hypotheses addressing “why” questions propose that the behavior increases fitness in some particular way. A reasonable hypothesis for why the red–crowned crane reproduces in spring and early summer is that breeding is most productive at that time of year. For instance, at that time, parent birds can find ample food for rapidly growing offspring, providing an advantage in reproductive success compared to birds that breed in other seasons.

Male Songbird: Proximate & Ultimate causes/explanations what triggers singing? (P or U?)  how does he sing? (P or U?)  why does he sing? (P or U?) Proximate cause questions Male songbirds sing during the breeding season as a response to a high level of testosterone which binds to hormone receptors in the brain & triggers the production of song. Ultimate cause questions The male sings to defend territory from other males & to attract a female with which to reproduce. This is the evolutionary explanation for the male’s vocalization.  how does daylength influence breeding?  why do cranes breed in spring?

Evolutionary perspective Adaptive advantage? innate behaviors automatic, fixed, “built-in”, no “learning curve” despite different environments, all individuals exhibit the behavior ex. early survival, reproduction, kinesis, taxis learned behaviors modified by experience variable, changeable flexible with a complex & changing environment

male sticklebacks exhibit aggressive territoriality Innate behaviors Fixed action patterns (FAP) sequence of behaviors essentially unchangeable & usually conducted to completion once started sign stimulus the releaser that triggers a FAP attack on red belly stimulus court on swollen belly stimulus

Innate: Directed movements Taxis change in direction automatic movement toward (positive taxis) or away from (negative taxis) a stimulus phototaxis chemotaxis Kinesis change in rate of movement in response to a stimulus Rolie polies The sow bugs become more active in dry areas and less active in humid areas. Though sow bugs do not move toward or away from specific conditions, their increased movement under dry conditions increases the chance that they will leave a dry area and encounter a moist area. And since they slow down in a moist area, they tend to stay there once they encounter it. In contrast to a kinesis, a taxis is a more or less automatic, oriented movement toward (a positive taxis) or away from (a negative taxis) some stimulus. For example, many stream fish, such as trout, exhibit positive rheotaxis (from the Greek rheos, current); they automatically swim or orient themselves in an upstream direction (toward the current). This taxis keeps the fish from being swept away and keeps them facing the direction from which food will come.

Complex Innate behaviors Migration “migratory restlessness” seen in birds bred & raised in captivity navigate by sun, stars, Earth magnetic fields Bird migration, a behavior that is largely under genetic control. Each spring, migrating western sandpipers (Calidris mauri), such as those shown here, migrate from their wintering grounds, which may be as far south as Peru, to their breeding grounds in Alaska. In the autumn, they return to the wintering grounds.

Innate & Learning: Imprinting Learning to form social attachments at a specific critical period both learning & innate components But how do the young know on whom—or what—to imprint? How do young geese know that they should follow the mother goose? The tendency to respond is innate in the birds; the outside world provides the imprinting stimulus, something to which the response will be directed. Experiments with many species of waterfowl indicate that they have no innate recognition of “mother.” They respond to and identify with the first object they encounter that has certain key characteristics. In classic experiments done in the 1930s, Konrad Lorenz showed that the most important imprinting stimulus in graylag geese is movement of an object away from the young. When incubator–hatched goslings spent their first few hours with Lorenz rather than with a goose, they imprinted on him, and from then on, they steadfastly followed him and showed no recognition of their biological mother or other adults of their own species. Again, there are both proximate and ultimate explanations Konrad Lorenz

Conservation Conservation biologists have taken advantage of imprinting by young whooping cranes as a means to teach the birds a migration route. A pilot wearing a crane suit in an Ultralight plane acts as a surrogate parent. teaching cranes to migrate Cranes also imprint as hatchlings, creating both problems and opportunities in captive rearing programs designed to save endangered crane species. For instance, a group of 77 endangered whooping cranes hatched and raised by sandhill cranes imprinted on the sandhill foster parents; none of these whooping cranes ever formed a mating pair–bond with another whooping crane. As a consequence, captive breeding programs now isolate young cranes and expose them to the sights and sounds of members of their own species. But imprinting can also be used to aid crane conservation Young whooping cranes imprinted on humans in “crane suits” have been taught to follow these “parents” flying ultralight aircraft along new migration routes. And importantly, such cranes have formed mating pair–bonds with other whooping cranes.

Critical period Sensitive phase for optimal imprinting some behavior must be learned during a receptive time period

Learned behavior Associative learning learning to associate a stimulus with a consequence operant conditioning trial & error learning associate behavior with reward or punishment ex: learning what to eat classical conditioning Pavlovian conditioning associate a “neutral stimulus” with a “significant stimulus”

Operant conditioning Skinner box mouse learns to associate behavior (pressing lever) with reward (food pellet)

Classical conditioning Ivan Pavlov’s dogs connect reflex behavior (salivating at sight of food) to associated stimulus (ringing bell)

Learning: Habituation Loss of response to stimulus “cry-wolf” effect decrease in response to repeated occurrences of stimulus enables animals to disregard unimportant stimuli ex: falling leaves not triggering fear response in baby birds ex: loss of feeling of wearing clothes

Learning: Problem-solving Do other animals reason? crow chimpanzee problem-solving sea otter tool use

Social behaviors Interactions between individuals develop as evolutionary adaptations communication / language agonistic behaviors dominance hierarchy cooperation altruistic behavior

Language Honey bee communication Did you watch the video ? Language Honey bee communication dance to communicate location of food source waggle dance View Waggle Dance AVI file: waggledance180x135.avi

Communication by song Bird song Insect song species identification & mating ritual mixed learned & innate critical learning period Insect song mating ritual & song innate, genetically controlled Red-winged blackbird

Social behaviors Agonistic behaviors threatening & submissive rituals symbolic, usually no harm done ex: territoriality, competitor aggression View Lifewire territoriality video: “lizards cost of defending-lifewire.swf” Review setting up a behavior experiment:

Social behaviors Dominance hierarchy social ranking within a group pecking order

Social behaviors Cooperation working together in coordination Pack of African dogs hunting wildebeest cooperatively White pelicans “herding” school of fish

Social behaviors Altruistic behavior reduces individual fitness but increases fitness of recipient kin selection increasing survival of close relatives passes these genes on to the next generation How can this be of adaptive value? Belding ground squirrel

Social interaction requires communication Pheromones chemical signal that stimulates a response from other individuals alarm pheromones sex pheromones

REVIEW of Proximate & Ultimate Proximate: what are the immediate causes and mechanisms of the behavior? Ultimate: explanation at the evolutionary level. What function does/did this behavior have in promoting survival and fitness? Example: why do male songbirds sing? Proximate explanation: increased daylight in the spring leads to increased testosterone production, which acts to activate a brain center that controls singing. Additional factors: role of experience in song-learning. Ultimate explanation: singing served to attract females and defend territory from other males. Males were selected for singing over evolutionary time

Proximate & Ultimate Example: Crested Owl. Nearly all owls hunt at night and many can capture prey in total darkness. Proximate questions: How do they do it? What neural circuits do they have to permit such hunting? Is this skill learned from parents or does it develop by individual practice alone? Ultimate questions: Does nocturnal hunting provide more food than daytime hunting? Are the prey species different? Does the owl make efficient choices about which prey to attack?

Proximate & Ultimate Example: Golden Bamboo Lemur. It feeds primarily on bamboo shoots -- but these contain liberal amounts of cyanide!! Proximate questions: What cues does it use to decide which shoots to eat? How does it know when it has had enough of the potentially deadly shoots? Ultimate questions: How did it evolve the ability to eat these shoots? Why does it eat these rather than less poisonous species of bamboo shoots?