1. 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

2. 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?

3. 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.

4. 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?

5. 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

6. 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

7. 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.

8. 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.

9. 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

10. 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.

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

12. 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”

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

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

15. 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

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

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

18. 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

19. 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

20. 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:

21. Social behaviors Dominance hierarchy social ranking within a group
pecking order

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

23. 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

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

25. 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

26. 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?

27. 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?