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CAMPBELL BIOLOGY IN FOCUS © 2014 Pearson Education, Inc. Urry Cain Wasserman Minorsky Jackson Reece Lecture Presentations by Kathleen Fitzpatrick and Nicole Tunbridge 39 Motor Mechanisms and Behavior
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Page 1: Biology in Focus - Chapter 39

CAMPBELL BIOLOGY IN FOCUS

© 2014 Pearson Education, Inc.

Urry • Cain • Wasserman • Minorsky • Jackson • Reece

Lecture Presentations by Kathleen Fitzpatrick and Nicole Tunbridge

39Motor Mechanisms and Behavior

Page 2: Biology in Focus - Chapter 39

© 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

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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

Imprinting

Learningand

problemsolving

Cognition

Social learning

Spatial learning

Associative learning


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