CAMPBELL BIOLOGY IN FOCUS
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Urry • Cain • Wasserman • Minorsky • Jackson • Reece
Lecture Presentations by Kathleen Fitzpatrick and Nicole Tunbridge
39Motor Mechanisms and Behavior
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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
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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 39.2Muscle
Bundle ofmuscle fibers
Nuclei
Z lines
Myofibril
Single muscle fiber(cell)
Plasmamembrane
Sarcomere
Thick filaments(myosin)
Z line Sarcomere
Thin filaments(actin) Z line
M line
TEM0.5 m
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Figure 39.2a
Muscle
Bundle ofmuscle fibers
Nuclei
Z lines
Myofibril
Single muscle fiber(cell)
Sarcomere
Plasmamembrane
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Figure 39.2b
Sarcomere
Thick filaments(myosin)
Z line Sarcomere
Thin filaments(actin)
Z line
M line
TEM0.5 m
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Figure 39.2c
Sarcomere
TEM0.5 m
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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
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Figure 39.3
SarcomereRelaxed muscle
Contracting muscle
Fully contracted muscle
Z ZZZ MM
Contractedsarcomere
0.5 m
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Figure 39.3a
SarcomereZZ M
0.5 m
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Figure 39.3b
SarcomereZZ M
0.5 m
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Figure 39.3c
Contractedsarcomere
0.5 m
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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
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Figure 39.4
Thinfilaments Thick filament
Thin filament movestoward center of sarcomere.
Cross-bridge
Myosin head (lowenergy configuration)
Myosin head (lowenergy configuration)
Thin filament
Thick filament
Myosin head (highenergy configuration)
ATP
Myosin-binding sites
ATP
Actin
ADP P iADP
P i
ADPP i
5
1
2
3
4
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Figure 39.4a
Myosin head (lowenergy configuration)
Thin filament
Thick filament
ATP
1
ATP
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Figure 39.4b
2
Myosin head (highenergy configuration)
Myosin-binding sitesActin
ADPP i
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Figure 39.4c
3
Cross-bridgeADP
P i
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Figure 39.4d
4Thin filament movestoward center of sarcomere.
Myosin head (lowenergy 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
Myosin-binding site
TropomyosinActin Troponin complex
(a) Myosin-binding sites blocked
(b) Myosin-binding sites exposed
Ca2-binding sites
Ca2
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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
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Figure 39.6 Synaptic terminal
Ca2 releasedfrom SR
Ca2
1
T tubuleSarcoplasmicreticulum (SR)
Plasma membraneof muscle fiber
Myofibril
Axon ofmotor neuron
Sarcomere
Mitochondrion
Synaptic terminal of motor neuronT tubule
Plasmamembrane
Sarcoplasmicreticulum (SR)
Synaptic cleft
ACh
Ca2 pump
Ca2
CYTOSOLATP
2
3
4
5
6
7
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Figure 39.6a
Synaptic terminal
Ca2 releasedfrom SR
T tubule
Sarcoplasmicreticulum (SR)
Plasma membraneof muscle fiber
Myofibril
Axon ofmotor neuron
Sarcomere
Mitochondrion
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Figure 39.6b
Ca2
1 Synaptic terminal of motor neuronT tubule
Plasmamembrane
Sarcoplasmicreticulum (SR)
Synaptic cleft
ACh
Ca2 pump
Ca2
CYTOSOL
ATP
2
3
4
5
6
7
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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
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Figure 39.7
Synaptic terminals
Spinal cord
Motor neuroncell body
Motor neuronaxon
Motorunit 1
Motorunit 2
Nerve
Muscle fibers
Muscle
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
TetanusTe
nsio
n Summation oftwo twitches
Singletwitch
TimeActionpotential Pair of
actionpotentials
Series of actionpotentials at
high frequency
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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 slow-twitch 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
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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
Biceps
Human forearm(internal skeleton)
Exte
nsio
n
Grasshopper tibia(external skeleton)
Biceps
Triceps
Triceps
Extensormuscle
Flexormuscle
Extensormuscle
Flexormuscle
Flex
ion
Contracting muscle Relaxing muscleKey
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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 39.11Longitudinalmuscle relaxed(extended)
Longitudinalmusclecontracted
Circular musclecontracted
Circular musclerelaxed
Bristles Head end
Head end
Head end
1
2
3
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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 ClavicleScapula
Sternum
CarpalsPelvic girdleUlnaRadiusVertebraHumerusRib
PhalangesMetatarsalsTarsalsFibulaTibiaPatellaFemur
MetacarpalsPhalanges
Typesof joints
Ball-and-socketjoint
Hinge jointPivot joint
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Figure 39.13
Ball-and-socket joint
Hinge joint
Pivot joint
Ulna
Radius
Humerus
Ulna
Head ofhumerusScapula
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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
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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 stimulus-response 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
(a) Worker bees (b) Round dance (food near)
(c) Waggle dance (food distant)
Location A Location B Location C
Beehive
A
BC
30
30
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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)
Beehive
A
BC
30
30
Location A Location B Location C
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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 (b) Pilot and cranes
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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 39.18Experiment
Pinecone
Results
Nest
NestNo nest
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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
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Figure 39.19b
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Figure 39.19c
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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
Low population densityHigh population density
D. melanogaster lineages
Mea
n pa
th le
ngth
(cm
)
R1 R2 R3 K1 K2 K3
7
6
5
4
3
2
1
0
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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 (c) Polyandrous species
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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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Needs of the young are an important factor constraining evolution of mating systems
Mating Systems and Parental Care
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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 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
Sexual Selection and Mate Choice
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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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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
Case Study: Variation in Prey Selection
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Figure 39.25
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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
Altruism
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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
Sibling 1 Sibling 2
½ (0.5)probability
OR
½ (0.5)probability
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Kin selection is the natural selection that favors this kind of altruistic behavior by enhancing reproductive success of relatives
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Figure 39.UN01
Flying Running
Swimming
Body mass (g) (log scale)10−3 103 1061
102
10−1
10
1
Ener
gy c
ost (
cal/k
gm
)(lo
g sc
ale)
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Figure 39.UN02
Sarcomere
Relaxedmuscle
Contractingmuscle
Fully contracted muscle
Contractedsarcomere
Thinfilament
Thickfilament
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Figure 39.UN03
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