Download Ch39Motor Mechanisms and Behavior

CAMPBELL BIOLOGY IN FOCUS
Urry • Cain • Wasserman • Minorsky • Jackson • Reece
39
Motor Mechanisms
and Behavior
Lecture Presentations by
Kathleen Fitzpatrick and Nicole Tunbridge
© 2014 Pearson Education, Inc.
Overview: The How and Why of Animal Activity
 Fiddler crabs feed with their small claw and wave
their large claw
 Why do male fiddler crabs engage in claw-waving
behavior?
 Claw waving is used to repel other males and to
attract females
Video: Albatross Courtship
Video: Boobies Courtship
Video: Giraffe Courtship
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Figure 39.1
© 2014 Pearson Education, Inc.
 A behavior is an action carried out by muscles
under control of the nervous system
 Behavior is subject to natural selection
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Concept 39.1: The physical interaction of protein
filaments is required for muscle function
 Muscle activity is a response to input from the
nervous system
 Muscle contraction is an active process; muscle
relaxation is passive
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 Muscle cell contraction relies on the interaction
between protein structures
 Thin filaments consist of two strands of actin coiled
around one another
 Thick filaments are staggered arrays of myosin
molecules
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Vertebrate Skeletal Muscle
 Vertebrate skeletal muscle moves bones and the
body and is characterized by a hierarchy of smaller
and smaller units
 A skeletal muscle consists of a bundle of long fibers,
each a single cell, running parallel to the length of
the muscle
 Each muscle fiber is itself a bundle of smaller
myofibrils, which contain thick and thin filaments
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 Skeletal muscle is also called striated muscle
because the regular arrangement of myofibrils
creates a pattern of light and dark bands
 The functional unit of a muscle is called a sarcomere
and is bordered by Z lines
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Figure 39.2
Muscle
Bundle of
muscle fibers
Nuclei
Single muscle fiber
(cell)
Plasma
membrane
Z lines
Myofibril
Sarcomere
TEM
Thick filaments
(myosin)
Thin filaments
(actin)
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Z line
M line
Sarcomere
0.5 m
Z line
Figure 39.2a
Muscle
Bundle of
muscle fibers
Nuclei
Single muscle fiber
(cell)
Plasma
membrane
Z lines
Myofibril
Sarcomere
© 2014 Pearson Education, Inc.
Figure 39.2b
Sarcomere
TEM
Thick filaments
(myosin)
Thin filaments
(actin)
© 2014 Pearson Education, Inc.
Z line
M line
Sarcomere
0.5 m
Z line
Figure 39.2c
Sarcomere
TEM
© 2014 Pearson Education, Inc.
0.5 m
The Sliding-Filament Mechanism of Muscle
Contraction
 According to the sliding-filament model, filaments
slide past each other longitudinally, causing an
overlap between thin and thick filaments
Video: Cardiac Muscle
© 2014 Pearson Education, Inc.
Figure 39.3
Relaxed muscle
Z
Sarcomere
M
Z
Z
M
Z
Contracting muscle
0.5 m
Fully contracted muscle
Contracted
sarcomere
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Figure 39.3a
Z
Sarcomere
M
Z
0.5 m
© 2014 Pearson Education, Inc.
Figure 39.3b
Z
Sarcomere
M
Z
0.5 m
© 2014 Pearson Education, Inc.
Figure 39.3c
0.5 m
Contracted
sarcomere
© 2014 Pearson Education, Inc.
 The sliding of filaments relies on interaction between
actin and myosin
 The “head” of a myosin molecule binds to an actin
filament, forming a cross-bridge and pulling the thin
filament toward the center of the sarcomere
 Muscle contraction requires repeated cycles of
binding and release
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 Glycolysis and aerobic respiration generate the
ATP needed to sustain muscle contraction
Video: Myosin and Actin
© 2014 Pearson Education, Inc.
Figure 39.4
1
Thin
filaments
Thick filament
Thin filament
Myosin head (low
energy configuration)
ATP
5
ATP
2
Thick
filament
Thin filament moves
toward center of sarcomere.
Actin
ADP
Myosin head (low
energy configuration)
ADP
Pi
ADP
P
i
4
© 2014 Pearson Education, Inc.
Myosinbinding sites
Pi
Cross-bridge
Myosin head (high
energy configuration)
3
Figure 39.4a
Thin
filament
1
ATP
ATP
Myosin head (low
energy configuration)
Thick
filament
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Figure 39.4b
2
Myosinbinding sites
Actin
ADP
Pi
© 2014 Pearson Education, Inc.
Myosin head (high
energy configuration)
Figure 39.4c
3
ADP
Pi
© 2014 Pearson Education, Inc.
Cross-bridge
Figure 39.4d
4
Thin filament moves
toward center of sarcomere.
Myosin head (low
energy configuration)
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The Role of Calcium and Regulatory Proteins
 The regulatory protein tropomyosin and the
troponin complex, a set of additional proteins, bind
to actin strands on thin filaments when a muscle
fiber is at rest
 This prevents actin and myosin from interacting
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Figure 39.5
Ca2-binding sites
Tropomyosin
Actin
Troponin complex
(a) Myosin-binding sites blocked
Ca2
Myosinbinding site
(b) Myosin-binding sites exposed
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 For a muscle fiber to contract, myosin-binding sites
must be uncovered
 This occurs when calcium ions (Ca2) bind to the
troponin complex and expose the myosin-binding
sites
 Contraction occurs when the concentration of Ca2
is high; muscle fiber contraction stops when the
concentration of Ca2 is low
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 The stimulus leading to contraction of a muscle
fiber is an action potential in a motor neuron that
makes a synapse with the muscle fiber
 The synaptic terminal of the motor neuron releases
the neurotransmitter acetylcholine
 Acetylcholine depolarizes the muscle, causing it to
produce an action potential
Animation: Muscle Contraction
© 2014 Pearson Education, Inc.
Figure 39.6
Axon of
motor neuron
Synaptic terminal
T tubule
Sarcoplasmic
reticulum (SR)
Mitochondrion
Myofibril
Plasma membrane
of muscle fiber
Ca2 released
from SR
Sarcomere
1 Synaptic terminal of motor neuron Plasma
Synaptic cleft T tubule
membrane
Sarcoplasmic
reticulum (SR)
2
ACh
Ca2 pump
3
Ca2
ATP
4
CYTOSOL
7
6
Ca2
5
© 2014 Pearson Education, Inc.
Figure 39.6a
Synaptic terminal
T tubule
Sarcoplasmic
reticulum (SR)
Axon of
motor neuron
Mitochondrion
Myofibril
Plasma membrane
of muscle fiber
Sarcomere
© 2014 Pearson Education, Inc.
Ca2 released
from SR
Figure 39.6b
1 Synaptic terminal of motor neuron
Synaptic cleft
T tubule
Sarcoplasmic
reticulum (SR)
2
ACh
Ca2 pump
3
Ca2
ATP
4
CYTOSOL
7
6
Ca2
5
© 2014 Pearson Education, Inc.
Plasma
membrane
 Action potentials travel to the interior of the muscle
fiber along transverse (T) tubules, infoldings of the
plasma membrane
 The action potential along T tubules causes the
sarcoplasmic reticulum (SR), a specialized
endoplasmic reticulum, to release Ca2
 The Ca2 binds to the troponin complex on the thin
filaments, initiating muscle fiber contraction
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 When motor neuron input stops, the muscle cell
relaxes
 Transport proteins in the SR pump Ca2 out of the
cytosol
 Regulatory proteins bound to thin filaments shift
back to the myosin-binding sites
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 Amyotrophic lateral sclerosis (ALS; also known as
Lou Gehrig’s disease) interferes with the excitation
of skeletal muscle fibers; this disease is usually
fatal
 Myasthenia gravis is an autoimmune disease that
attacks acetylcholine receptors on muscle fibers;
treatments exist for this disease
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Nervous Control of Muscle Tension
 Contraction of a whole muscle is graded, which
means that the extent and strength of its contraction
can be voluntarily altered
 There are two basic mechanisms by which the
nervous system produces graded contractions
 Varying the number of fibers that contract
 Varying the rate at which fibers are stimulated
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 In vertebrates, each motor neuron may synapse with
multiple muscle fibers, although each fiber is
controlled by only one motor neuron
 A motor unit consists of a single motor neuron and
all the muscle fibers it controls
© 2014 Pearson Education, Inc.
Figure 39.7
Spinal cord
Motor
unit 1
Motor
unit 2
Synaptic terminals
Nerve
Motor neuron
cell body
Motor neuron
axon
Muscle
Muscle fibers
Tendon
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 Recruitment of multiple motor neurons results in
stronger contractions
 A twitch results from a single action potential in a
motor neuron
 More rapidly delivered action potentials produce a
graded contraction by summation
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Figure 39.8
Tension
Tetanus
Summation of
two twitches
Single
twitch
Action
potential
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Time
Pair of
action
potentials
Series of action
potentials at
high frequency
 Tetanus is a state of smooth and sustained
contraction produced when motor neurons deliver
a volley of action potentials
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Types of Skeletal Muscle Fibers
 There are several distinct types of skeletal muscles,
each of which is adapted to a particular set of
functions
 They are classified by the source of ATP powering
the muscle activity and the speed of muscle
contraction
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Table 39.1
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 Oxidative and glycolytic fibers are differentiated
by their energy source
 Oxidative fibers rely mostly on aerobic respiration
to generate ATP
 These fibers have many mitochondria, a rich blood
supply, and a large amount of myoglobin
 Myoglobin is a protein that binds oxygen more
tightly than hemoglobin does
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 Glycolytic fibers use glycolysis as their primary
source of ATP
 Glycolytic fibers have less myoglobin than oxidative
fibers and tire more easily
 In poultry and fish, light meat is composed of
glycolytic fibers, while dark meat is composed of
oxidative fibers
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 Fast-twitch and slow-twitch fibers are differentiated
by their speed of contraction
 Slow-twitch fibers contract more slowly but sustain
longer contractions
 All slow-twitch fibers are oxidative
 Fast-twitch fibers contract more rapidly but sustain
shorter contractions
 Fast-twitch fibers can be either glycolytic or oxidative
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 Most human skeletal muscles contain both slowtwitch and fast-twitch muscles in varying ratios
 Some vertebrates have muscles that twitch at rates
much faster than human muscles
 In producing its characteristic mating call, the male
toadfish can contract and relax certain muscles
more than 200 times per second
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Figure 39.9
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Other Types of Muscle
 In addition to skeletal muscle, vertebrates have
cardiac muscle and smooth muscle
 Cardiac muscle, found only in the heart, consists of
striated cells electrically connected by intercalated
disks
 Cardiac muscle can generate action potentials
without neural input
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 In smooth muscle, found mainly in walls of hollow
organs such as those of the digestive tract,
contractions are relatively slow and may be initiated
by the muscles themselves
 Contractions may also be caused by stimulation
from neurons in the autonomic nervous system
© 2014 Pearson Education, Inc.
Concept 39.2: Skeletal systems transform
muscle contraction into locomotion
 Skeletal muscles are attached in antagonistic pairs,
the actions of which are coordinated by the nervous
system
 The skeleton provides a rigid structure to which
muscles attach
 Skeletons function in support, protection, and
movement
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Figure 39.10
Human forearm
(internal skeleton)
Extensor
muscle
Flexion
Biceps
Grasshopper tibia
(external skeleton)
Flexor
muscle
Triceps
Extension
Biceps
Extensor
muscle
Flexor
muscle
Triceps
Key
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Contracting muscle
Relaxing muscle
Types of Skeletal Systems
 The three main types of skeletons are
 Hydrostatic skeletons (lack hard parts)
 Exoskeletons (external hard parts)
 Endoskeletons (internal hard parts)
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Hydrostatic Skeletons
 A hydrostatic skeleton consists of fluid held under
pressure in a closed body compartment
 This is the main type of skeleton in most cnidarians,
flatworms, nematodes, and annelids
 Annelids use their hydrostatic skeleton for
peristalsis, a type of movement on land produced
by rhythmic waves of muscle contractions
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Figure 39.11
Longitudinal
muscle relaxed
(extended)
Circular muscle Circular muscle Longitudinal
contracted
muscle
relaxed
contracted
Bristles
Head end
1
Head end
2
3
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Head end
Exoskeletons
 An exoskeleton is a hard encasement deposited on
the surface of an animal
 Exoskeletons are found in most molluscs and
arthropods
 Arthropods have a jointed exoskeleton called a
cuticle, which can be both strong and flexible
 About 30–50% of the arthropod cuticle consists of a
polysaccharide called chitin
 An arthropod must shed and regrow its exoskeleton
when it grows
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Endoskeletons
 An endoskeleton consists of a hard internal
skeleton, buried in soft tissue
 Endoskeletons are found in organisms ranging from
sponges to mammals
 A mammalian skeleton has more than 200 bones
 Some bones are fused; others are connected at
joints by ligaments that allow freedom of movement
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Figure 39.12
Skull
Shoulder girdle
Clavicle
Scapula
Sternum
Rib
Humerus
Vertebra
Radius
Ulna
Pelvic girdle
Carpals
Phalanges
Metacarpals
Femur
Patella
Tibia
Fibula
Tarsals
Metatarsals
Phalanges
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Types
of joints
Ball-and-socket
joint
Hinge joint
Pivot joint
Figure 39.13
Ball-and-socket joint
Head of
humerus
Scapula
Hinge joint
Humerus
Ulna
Pivot joint
Ulna
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Radius
Size and Scale of Skeletons
 An animal’s body structure must support its size
 The weight of a body increases with the cube of its
dimensions, while the strength of that body
increases with the square of its dimensions
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 The skeletons of small and large animals have
different proportions
 In mammals and birds, the position of legs relative
to the body is very important in determining how
much weight the legs can bear
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Types of Locomotion
 Most animals are capable of locomotion, or active
travel from place to place
 In locomotion, energy is expended to overcome
friction and gravity
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Flying
 Active flight requires that wings develop enough lift
to overcome the downward force of gravity
 Many flying animals have adaptations that reduce
body mass
 For example, birds have no teeth or urinary bladder,
and their relatively large bones have air-filled regions
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Locomotion on Land
 Walking, running, or hopping on land requires an
animal to support itself and move against gravity
 Diverse adaptations for locomotion on land have
evolved in vertebrates
 For example, kangaroos have large, powerful muscles
in their hind legs, suitable for hopping
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Figure 39.14
© 2014 Pearson Education, Inc.
Swimming
 In water, friction is a bigger problem than gravity
 Fast swimmers usually have a sleek, torpedo-like
shape to minimize friction
 Animals swim in diverse ways
 Paddling with their legs as oars
 Jet propulsion
 Undulating their body and tail from side to side or
up and down
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Concept 39.3: Discrete sensory inputs can
stimulate both simple and complex behaviors
 Niko Tinbergen identified four questions that should
be asked about animal behavior
1. What stimulus elicits the behavior, and what
physiological mechanisms mediate the response?
2. How does the animal’s experience during growth
and development influence the response?
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3. How does the behavior aid survival and
reproduction?
4. What is the behavior’s evolutionary history?
 Behavioral ecology is the study of the ecological
and evolutionary basis for animal behavior
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 Behavioral ecology integrates proximate and ultimate
explanations for animal behavior
 Proximate causation addresses “how” a behavior
occurs or is modified, including Tinbergen’s
questions 1 and 2
 Ultimate causation addresses “why” a behavior
occurs in the context of natural selection, including
Tinbergen’s questions 3 and 4
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Fixed Action Patterns
 A fixed action pattern is a sequence of unlearned,
innate behaviors that is unchangeable
 Once initiated, it is usually carried to completion
 A fixed action pattern is triggered by an external
cue known as a sign stimulus
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 Tinbergen observed male stickleback fish
responding to a passing red truck
 In male stickleback fish, the stimulus for attack
behavior is the red underside of an intruder
 When presented with unrealistic models, the attack
behavior occurs as long as some red is present
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Figure 39.15
(a)
(b)
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Migration
 Environmental cues can trigger movement in a
particular direction
 Migration is a regular, long-distance change in
location
 Animals can orient themselves using
 The position of the sun and their circadian clock, an
internal 24-hour activity rhythm or cycle
 The position of the sun or stars
 Earth’s magnetic field
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Behavioral Rhythms
 Some animal behavior is affected by the animal’s
circadian rhythm, a daily cycle of rest and activity
 Behaviors such as migration and reproduction are
linked to changing seasons, or a circannual rhythm
 Daylight and darkness are common seasonal cues
 Some behaviors are linked to lunar cycles, which
affect tidal movements
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Animal Signals and Communication
 In behavioral ecology, a signal is a behavior that
causes a change in another animal’s behavior
 Communication is the transmission and
reception of signals
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Forms of Animal Communication
 Animals communicate using visual, chemical,
tactile, and auditory signals
 Fruit fly courtship follows a three-step stimulusresponse chain
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1. A male identifies a female of the same species and
orients toward her
 Chemical communication: he smells a female’s
chemicals in the air
 Visual communication: he sees the female and
orients his body toward hers
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2. The male alerts the female to his presence
 Tactile communication: he taps the female with a
foreleg
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3. The male produces a courtship song to inform the
female of his species
 Auditory communication: he extends and vibrates
his wing
 If all three steps are successful, the female will
allow the male to copulate
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 Honeybees show complex communication with
symbolic language
 A bee returning from the field performs a dance to
communicate information about the distance and
direction of a food source
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Figure 39.16
(b) Round dance (food near)
(a) Worker bees
(c) Waggle dance (food distant)
A
30
C
B
Beehive
Location A
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Location B
Location C
Figure 39.16a
(a) Worker bees
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Figure 39.16b
(b) Round dance
(food near)
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Figure 39.16c
(c) Waggle dance
(food distant)
A
30
C
B
Beehive
Location A
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Location B
Location C
Pheromones
 Many animals that communicate through odors
emit chemical substances called pheromones
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 For example,
 A female moth can attract a male moth several
kilometers distant
 A honeybee queen produces a pheromone that
affects the development and behavior of female
workers and male drones
 When a minnow or catfish is injured, an alarm
substance in the fish’s skin disperses in the water,
causing nearby fish to seek safety
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 Animals use diverse forms of communication
 Nocturnal animals, such as most terrestrial
mammals, depend on olfactory and auditory
communication
 Diurnal animals, such as humans and most birds,
use visual and auditory communication
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Concept 39.4: Learning establishes specific links
between experience and behavior
 Innate behavior is developmentally fixed and does
not vary among individuals
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Experience and Behavior
 Cross-fostering studies help behavioral ecologists to
identify the contribution of environment to an
animal’s behavior
 A cross-fostering study places the young from one
species in the care of adults from another species
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 Studies of California mice and white-footed mice
have uncovered an influence of social environment
on aggressive and parental behaviors
 Cross-fostered mice developed some behaviors
that were consistent with their foster parents
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Table 39.2
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 In humans, twin studies allow researchers to
compare the relative influences of genetics and
environment on behavior
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Learning
 Learning is the modification of behavior based
on specific experiences
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Imprinting
 Imprinting is the establishment of a long-lasting
behavioral response to a particular individual
 Imprinting can only take place during a specific time
in development, called the sensitive period
 A sensitive period is a limited developmental phase
that is the only time when certain behaviors can be
learned
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 An example of imprinting is young geese following
their mother
 Konrad Lorenz showed that when baby geese
spent the first few hours of their life with him, they
imprinted on him as their parent
 The imprint stimulus in greylag geese is a nearby
object that is moving away from the young geese
Video: Ducklings
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Figure 39.17
(a) Konrad Lorenz and geese
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(b) Pilot and cranes
Figure 39.17a
(a) Konrad Lorenz and geese
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Figure 39.17b
(b) Pilot and cranes
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 Conservation biologists have taken advantage of
imprinting in programs to save the whooping crane
from extinction
 Young whooping cranes can imprint on humans in
“crane suits” who then lead crane migrations using
ultralight aircraft
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Spatial Learning and Cognitive Maps
 Spatial learning is the establishment of a memory
that reflects the spatial structure of the environment
 Niko Tinbergen showed how digger wasps use
landmarks to find nest entrances
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Figure 39.18
Experiment
Nest
Pinecone
Results
Nest
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No nest
 A cognitive map is an internal representation of
spatial relationships between objects in an animal’s
surroundings
 For example, Clark’s nutcrackers can find food
hidden in caches located halfway between particular
landmarks
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Associative Learning
 In associative learning, animals associate one
feature of their environment with another
 For example, a blue jay will avoid eating butterflies
with specific colors after a bad experience with a
distasteful monarch butterfly
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Figure 39.19
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Figure 39.19a
© 2014 Pearson Education, Inc.
Figure 39.19b
© 2014 Pearson Education, Inc.
Figure 39.19c
© 2014 Pearson Education, Inc.
 Animals can learn to link many pairs of features of
their environment, but not all
 For example, pigeons can learn to associate danger
with a sound but not with a color
 For example, rats can learn to avoid illness-inducing
foods on the basis of smells, but not on the basis of
sights or sounds
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Cognition and Problem Solving
 Cognition is a process of knowing that may
include awareness, reasoning, recollection, and
judgment
 For example, honeybees can distinguish “same”
from “different”
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 Problem solving is the process of devising a
strategy to overcome an obstacle
 For example, chimpanzees can stack boxes in order
to reach suspended food
 For example, ravens obtained food suspended from
a branch by a string by pulling up the string
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 Social learning is learning through the observation
of others and forms the roots of culture
 For example, young chimpanzees learn to crack palm
nuts with stones by copying older chimpanzees
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 Culture is a system of information transfer through
observation or teaching that influences behavior of
individuals in a population
 Culture can alter behavior and influence the fitness
of individuals
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Concept 39.5: Selection for individual survival and
reproductive success can explain most behaviors
 Behavior enhances survival and reproductive
success in a population
 Foraging is a behavior essential for survival and
reproduction that includes recognizing, capturing,
and eating food items
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Evolution of Foraging Behavior
 Natural selection refines behaviors that enhance
the efficiency of feeding
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 In Drosophila melanogaster, variation in a gene
dictates foraging behavior in the larvae
 Larvae with one allele travel farther while foraging
than larvae with the other allele
 Larvae in high-density populations benefit from
foraging farther for food, while larvae in low-density
populations benefit from short-distance foraging
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 Natural selection favors different alleles depending
on the density of the population
 Under laboratory conditions, evolutionary changes in
the frequency of these two alleles were observed
over several generations
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Figure 39.20
Mean path length (cm)
7
Low population density
6
High population density
5
4
3
2
1
0
R1
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R2
R3
K1
D. melanogaster lineages
K2
K3
Mating Behavior and Mate Choice
 Mating behavior includes seeking or attracting
mates, choosing among potential mates, competing
for mates, and caring for offspring
 Mating relationships define a number of distinct
mating systems
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Mating Systems and Sexual Dimorphism
 The mating relationship between males and
females varies greatly from species to species
 In some species there are no strong pair-bonds
 In monogamous relationships, one male mates
with one female
 Males and females with monogamous mating
systems have similar external morphologies
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Figure 39.21
(a) Monogamous species
(b) Polygynous species
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(c) Polyandrous species
Figure 39.21a
(a) Monogamous species
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Figure 39.21b
(b) Polygynous species
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Figure 39.21c
(c) Polyandrous species
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 In polygamous relationships, an individual of one
sex mates with several individuals of the other sex
 Species with polygamous mating systems are
usually sexually dimorphic: males and females
have different external morphologies
 Polygamous relationships can be either polygynous
or polyandrous
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 In polygyny, one male mates with many females
 The males are usually more showy and larger than
the females
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 In polyandry, one female mates with many males
 The females are often more showy than the males
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Mating Systems and Parental Care
 Needs of the young are an important factor
constraining evolution of mating systems
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 Consider bird species where chicks need a
continuous supply of food
 A male maximizes his reproductive success by staying
with his mate and caring for his chicks (monogamy)
 Consider bird species where chicks are soon able to
feed and care for themselves
 A male maximizes his reproductive success by seeking
additional mates (polygyny)
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 Certainty of paternity influences parental care and
mating behavior
 Females can be certain that eggs laid or young born
contain her genes; however, paternal certainty
depends on mating behavior
 Paternal certainty is relatively low in species with
internal fertilization because mating and birth are
separated over time
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 Certainty of paternity is much higher when egg laying
and mating occur together, as in external fertilization
 In species with external fertilization, parental care is
at least as likely to be by males as by females
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Figure 39.22
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Sexual Selection and Mate Choice
 Sexual dimorphism results from sexual selection, a
form of natural selection
 In intersexual selection, members of one sex choose
mates on the basis of certain traits
 Intrasexual selection involves competition between
members of the same sex for mates
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 Female choice is a type of intersexual competition
 Females can drive sexual selection by choosing
males with specific behaviors or features of anatomy
 For example, female stalk-eyed flies choose males
with relatively long eyestalks
 Ornaments, such as long eyestalks, often correlate
with health and vitality
Video: Chimp Agonistic
Video: Snakes Wrestling
Video: Wolves Agonistic
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Figure 39.23
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 Male competition for mates is a source of
intrasexual selection that can reduce variation
among males
 Such competition may involve agonistic behavior, an
often ritualized contest that determines which
competitor gains access to a resource
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Concept 39.6: Inclusive fitness can account for
the evolution of behavior, including altruism
 The definition of fitness can be expanded beyond
individual survival to help explain “selfless”
behavior
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Genetic Basis of Behavior
 Differences at a single locus can sometimes have a
large effect on behavior
 For example, male prairie voles pair-bond with their
mates, while male meadow voles do not
 The level of a specific receptor for a neurotransmitter
determines which behavioral pattern develops
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Figure 39.24
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Genetic Variation and the Evolution of Behavior
 When behavioral variation within a species
corresponds to environmental variation, it may
be evidence of past evolution
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Case Study: Variation in Prey Selection
 The natural diet of western garter snakes varies by
population
 Coastal populations feed mostly on banana slugs,
while inland populations rarely eat banana slugs
 Studies have shown that the differences in diet are
genetic
 The two populations differ in their ability to detect
and respond to specific odor molecules produced
by the banana slugs
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Figure 39.25
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Altruism
 Natural selection favors behavior that maximizes an
individual’s survival and reproduction
 These behaviors are often selfish
 On occasion, some animals behave in ways that
reduce their individual fitness but increase the fitness
of others
 This kind of behavior is called altruism, or
selflessness
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 For example, under threat from a predator, an
individual Belding’s ground squirrel will make an
alarm call to warn others, even though calling
increases the chances that the caller is killed
 For example, in naked mole rat populations,
nonreproductive individuals may sacrifice their lives
protecting their reproductive queen and kings from
predators
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Figure 39.26
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Inclusive Fitness
 Altruism can be explained by inclusive fitness
 Inclusive fitness is the total effect an individual has
on proliferating its genes by producing offspring and
helping close relatives produce offspring
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Hamilton’s Rule and Kin Selection
 William Hamilton proposed a quantitative measure
for predicting when natural selection would favor
altruistic acts among related individuals
 Three key variables in an altruistic act
 Benefit to the recipient (B)
 Cost to the altruistic (C)
 Coefficient of relatedness (the fraction of genes
that, on average, are shared; r)
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 Natural selection favors altruism when
rB  C
 This inequality is called Hamilton’s rule
 Hamilton’s rule is illustrated with the following
example of a girl who risks her life to save her
brother
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 Assume the average individual has two children. As
a result of the sister’s action
 The brother can now father two children, so B  2
 The sister has a 25% chance of dying and not being
able to have two children, so C  0.25  2  0.5
 The brother and sister share half their genes on
average, so r  0.5
 If the sister saves her brother, rB ( 1)  C ( 0.5)
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Figure 39.27
Parent A
Parent B
OR
½ (0.5)
probability
½ (0.5)
probability
Sibling 1
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Sibling 2
 Kin selection is the natural selection that favors this
kind of altruistic behavior by enhancing reproductive
success of relatives
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Energy cost (cal/kgm)
(log scale)
Figure 39.UN01
Flying
Running
102
10
1
Swimming
10−1
10−3
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1
103
Body mass (g) (log scale)
106
Figure 39.UN02
Sarcomere
Relaxed
muscle
Contracting
muscle
Thin
filament
Fully contracted
muscle
Contracted
sarcomere
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Thick
filament
Figure 39.UN03
Imprinting
Learning
and
problem
solving
Cognition
Associative learning
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Spatial learning
Social learning