Behavioral biology studies animal behavior and its causes. Niko Tinbergen performed experiments on digger wasps to demonstrate that they use landmarks to locate their nests. He moved pinecones around nests and the wasps flew to the arrangement of landmarks, showing they can learn and use new landmarks. Behavior results from both innate responses and learning from the environment.

1. • Behavioral biology is the study of what
animals do when interacting with their
environment
• Behavior can be interpreted in terms of
proximate causes (immediate interaction with
the environment) or ultimate causes
(evolutionary differences)
37.1 Behavioral biologists study the actions of
animals in their natural environments
CLASSIC CONCEPTS IN BEHAVIOR

2. • Early insights into the nature of behavior came
from studies by Nobel laureates Karl von
Frisch, Konrad Lorenz, and Niko Tinbergen
• They were among the first experimentalists in
behavioral biology
• Tinbergen and Lorenz performed
experimental studies of innate behavior and
simple forms of learning

3. • 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

4. • In the experiment, Tinbergen placed a circle of
pinecones around a nest opening
Figure 37.1, Part 1
1
Nest

5. Nest
• 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
Figure 37.1, Part 2
2
No Nest

6. • 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
Figure 37.1, Part 3
3
No Nest

7. • 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

8. • Behavioral ecologists are especially
interested in the ultimate causes of
behavior, which are evolutionary 
Natural selection preserves behaviors that
enhance fitness

9. • Animal behavior often involves a combination
of genetic programming (innate behavior) and
environmental experiences (learning)
both genes and the environment influence the
development of behavioral phenotypes- just
like any other traits
37.2 Behavior results from both genes and
environmental factors

10. • The gathering
of nest
materials by
lovebirds has
genetic and
environmental
components
Figure 37.2
Single long strip carried in beak
(Fischer’s lovebird)
Several short strips tucked under feathers
(peach-faced lovebird)
Hybrid behavior
Tucking
failure
Strip in
beak

11. • Sign stimuli (often a simple cue in an animal’s
environment) trigger innate, essentially
unchangeable fixed action patterns (FAPs)
• The genetic programming underlying FAPs
ensures that such activities are performed
correctly without practice
37.3 Innate behavior often appears as fixed action
patterns

12. • 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

13. • Several key events in the life cycle of the
European cuckoo are determined by fixed
action patterns
– Egg-laying behavior
Figure 37.3B
1 2 3

14. – 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

15. • 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
37.4 Learning ranges from simple behavioral
changes to complex problem solving

16. Table 37.4

17. 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; it enhances
fitness by enabling rapid
learning
• Example: Lorenz used the
graylag goose to demonstrate
imprinting. He took over the
maternal role for a group of
goslings

18. • Not all examples of imprinting involve parent-
offspring bonding
– Although newly hatched salmon do not receive
any parental care, they imprint on the complex
mixture of odors unique to the freshwater
stream where they hatch
– This allows salmon to find their way back to
the stream to spawn after spending a year or
more at sea

19. • Imprinting plays an important role in song
development for many kinds of birds
Figure 37.5B

20. • 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
37.6 Many animals learn by association and
imitation
Figure 37.6A

21. • 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

22. • 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

23. • Some animals exhibit problem-
solving behavior
– Examples: chimpanzees and
ravens
37.7 Animal cognition includes problem-solving
behavior
Figure 37.7A, B

24. • Behavior is an evolutionary adaptation that
enhances survival and reproductive success
• Behavior evolves as natural selection fine-
tunes an animal to its environment
– The hunting and reproduction behaviors of
jaguars
– Nest location by digger wasps
– Imprinting of goslings
37.8 An animal's behavior reflects its evolution
ECOLOGICAL ROLES OF BEHAVIOR

25. • 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
37.9 Biological rhythms synchronize behavior with
the environment

26. • In the absence of environmental cues, these
rhythms continue
– But they become out of phase with the
environment
Figure 37.9A
12:12 (natural)
Constant
darkness

27. • Movement in a directed way enables animals
to
– avoid predators
– migrate to a more favorable environment
– obtain food
– find mates and nest sites
37.10 Animal movement may be oriented to stimuli
or landmarks

28. 1. kinesis- simplest type of animal movement -
random movement in response to a stimulus
2. taxis- another simple type
– A more or less automatic movement directed
toward or away from some stimulus
– Examples include rheotaxis (current) chemotaxis,
and phototaxis
3. Some animals use landmarks to find their way
within an area
TYPES of ANIMAL MOVEMENT

29. • Many animals formulate cognitive maps
– Internal representations of spatial relationships among
objects in their surroundings (wasp example)
• Some animals undertake long-range migrations
– Examples: whales, sea turtles, birds, monarch
butterflies
• Animals navigate using the sun, stars, temperature
gradients, landmarks, or Earth's magnetism
37.11 Movement from place to place often depends
on internal maps

30. • Migrating gray whales use coastal landmarks to
stay on course
Figure 37.11A
FEEDING
GROUNDS
Siberia
Arctic Ocean
Alaska
NORTH
AMERICA
Pacific
Ocean
Baja California
BREEDING GROUNDS
Atlantic
Ocean

31. • The indigo
bunting learns
a star map and
navigates by
fixing on the
North Star
Figure 37.11B
Paper
Ink pad
Funnel-
shaped
cage

32. • 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”
37.12 Behavioral ecologists use cost/benefit
analysis in studying feeding behavior
Figure 37.12A, B

33. • 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

34. • 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

35. • 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

36. • Social behavior is defined as the interaction
among members of a population
• The discipline of sociobiology studies social
behavior in the context of evolution
37.13 Sociobiology places social behavior in an
evolutionary context
SOCIAL BEHAVIOR AND SOCIOBIOLOGY

37. • 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
37.14 Rituals involving agonistic behavior often
resolve confrontations between competitors
Figure 37.14

38. • Many animals live in social groups maintained
by agonistic behaviors
• Dominance hierarchy is the ranking of
individuals based on social interactions
37.15 Dominance hierarchies are maintained by
agonistic behavior

39. • Chickens establish a “peck order”
• Resources are often partitioned based upon
the dominance hierarchy
Figure 37.15

40. • Humans tend to space themselves out when
they are close to others
– They establish
what we might
call personal
territories
37.17 Territorial behavior parcels space and
resources
Figure 37.17A

41. • Many animals exhibit territorial behavior
– It is a form of social behavior that partitions
resources

42. • 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

43. • 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

44. • Territoriality can enhance fitness if the benefits
of possessing a territory outweigh the energy
costs of defending one

45. • 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