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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 (immediate interaction with
the environment) or ultimate causes
(evolutionary differences)
• 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
• 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
• In the experiment, Tinbergen placed a circle of
pinecones around a nest opening
Nest
1
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
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
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
• Behavioral ecologists are especially
interested in the ultimate causes of
behavior, which are evolutionary 
Natural selection preserves behaviors that
enhance fitness
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)
both genes and the environment influence the
development of behavioral phenotypes- just
like any other traits
• 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
37.3 Innate behavior often appears as fixed action
patterns
• 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
• 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
• 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
– 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
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
Table 37.4
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
• Not all examples of imprinting involve parentoffspring 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
• Imprinting plays an important role in song
development for many kinds of birds
Figure 37.5B
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
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
• 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
37.7 Animal cognition includes problem-solving
behavior
• Some animals exhibit problemsolving behavior
– Examples: chimpanzees and
ravens
Figure 37.7A, B
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
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
• In the absence of environmental cues, these
rhythms continue
– But they become out of phase with the
environment
12:12 (natural)
Constant
darkness
Figure 37.9A
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
TYPES of ANIMAL MOVEMENT
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
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 (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
• 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
• The indigo
bunting learns
a star map and
navigates by
fixing on the
North Star
Paper
Ink pad
Funnelshaped
cage
Figure 37.11B
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
• 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
• 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
• 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
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
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
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
• Chickens establish a “peck order”
• Resources are often partitioned based upon
the dominance hierarchy
Figure 37.15
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
• Many animals exhibit territorial behavior
– It is a form of social behavior that partitions
resources
• 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
• 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
• Territoriality can enhance fitness if the benefits
of possessing a territory outweigh the energy
costs of defending one
• 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