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    Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

    PowerPoint Lectures forBio logy, Seventh Edit ion

    Neil Campbell and Jane Reece

    Lectures by Chris Romero

    Chapter 51

    Behavioral Ecology

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    Overview: Studying Behavior

    Humans have probably studied animalbehavior

    For as long as we have lived on Earth

    As hunters

    Knowledge of animal behavior was essential to

    human survival

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    Cranes are birds that have captivated peoples

    interest Possibly because they are large and their

    behavior is easily observed

    Figure 51.1

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    The modern scientific discipline of behavioral

    ecology Extends observations of animal behavior by

    studying how such behavior is controlled and

    how it develops, evolves, and contributes to

    survival and reproductive success

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    Concept 51.1: Behavioral ecologists distinguish

    between proximate and ultimate causes ofbehavior

    The scientific questions that can be asked

    about behavior can be divided into two classes

    Those that focus on the immediate stimulus

    and mechanism for the behavior

    Those that explore how the behavior

    contributes to survival and reproduction

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    What Is Behavior?

    Behavior

    Is what an animal does and how it does it

    Includes muscular and nonmuscular activity

    Figure 51.2

    Dorsal fin

    Anal fin

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    Learning

    Is also considered a behavioral process

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    Proximate and Ultimate Questions

    Proximate, or how, questions about behavior

    Focus on the environmental stimuli that triggera behavior

    Focus on the genetic, physiological, and

    anatomical mechanisms underlying abehavioral act

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    Ultimate, or why, questions about behavior

    Address the evolutionary significance of abehavior

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    Ethology

    Ethology is the scientific study of animal

    behavior Particularly in natural environments

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    Mid 20th-century ethologists

    Developed a conceptual framework defined bya set of questions

    These questions

    Highlight the complementary nature of

    proximate and ultimate perspectives

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    F ixed Action Patterns

    A fixed action pattern (FAP)

    Is a sequence of unlearned, innate behaviorsthat is unchangeable

    Once initiated, is usually carried to completion

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    A FAP is triggered by an external sensory

    stimulus Known as a sign stimulus

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    In male stickleback fish, the stimulus for attack

    behavior Is the red underside of an intruder

    Figure 51.3a

    (a)A male three-spined stickleback fish shows its red underside.

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    When presented with unrealistic models

    As long as some red is present, the attackbehavior occurs

    Figure 51.3b

    (b)The realistic model at the top, without a red underside, produces no

    aggressive response in a male three-spined stickleback fish. The

    other models, with red undersides, produce strong responses.

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    Proximate and ultimate causes for the FAP

    attack behavior in male stickleback fish

    Figure 51.4ULTIMATE CAUSE: By chasing away other male sticklebacks, a male decreases

    the chance that eggs laid in his nesting territory will be fertilized by another male.

    BEHAVIOR: A male stickleback fish attacks other male sticklebacks that invade its nesting

    territory.

    PROXIMATE CAUSE: The red belly of the intruding male acts as a sign stimulus

    that releases aggression in a male stickleback.

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    Imprinting

    Imprinting is a type of behavior

    That includes both learning and innatecomponents and is generally irreversible

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    Imprinting is distinguished from other types of

    learning by a sensitive period A limited phase in an animals development

    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

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    Konrad Lorenz showed that

    When baby geese spent the first few hours oftheir life with him, they imprinted on him as

    their parent

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    There are proximate and ultimate causes for

    this type of behavior

    Figure 51.5

    BEHAVIOR: Young geese follow and imprint on their mother.

    PROXIMATE CAUSE:During an early, critical developmental stage, the young

    geese observe their mother moving away from them and calling.

    ULTIMATE CAUSE:On average, geese that follow and imprint on their mother

    receive more care and learn necessary skills, and thus have a greater chance of

    surviving than those that do not follow their mother.

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    Conservation biologists have taken advantage

    of imprinting

    In programs to save the whooping crane from

    extinction

    Figure 51.6

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    Concept 51.2: Many behaviors have a strong

    genetic component

    Biologists study the ways both genes and the

    environment

    Influence the development of behavioralphenotypes

    Behavior that is developmentally fixed

    Is called innate behavior and is under strong

    genetic influence

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    Directed Movements

    Many animal movements

    Are under substantial genetic influence

    These types of movements

    Are called directed movements

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    Kinesis

    A kinesis

    Is a simple change in activity or turning rate inresponse to a stimulus

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    Sow bugs

    Become more active in dry areas and lessactive in humid areas

    Figure 51.7a

    Dry open

    area

    Moist site

    under leaf

    (a) Kinesis increases the chance that a sow bug will encounter and

    stay in a moist environment.

    T i

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    Taxis

    A taxis

    Is a more or less automatic, orientedmovement toward or away from a stimulus

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    Many stream fish exhibit positive rheotaxis

    Where they automatically swim in an upstreamdirection

    Figure 51.7b

    Direction

    of river

    current

    (b) Positive rheotaxis keeps trout facing into the current, the direction

    from which most food comes.

    Mi ti

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    Migration

    Many features of migratory behavior in birds

    Have been found to be geneticallyprogrammed

    Figure 51.8

    A i l Si l d C i ti

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    Animal Signals and Communication

    In behavioral ecology

    A signal is a behavior that causes a change inanother animals behavior

    Communication

    Is the reception of and response to signals

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    Animals communicate using

    Visual, auditory, chemical, tactile, andelectrical signals

    The type of signal used to transmit information

    Is closely related to an animals lifestyle and

    environment

    Ch i l C i ti

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    Chemical Communication

    Many animals that communicate through odors

    Emit chemical substances called pheromones

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    When a minnow or catfish is injured

    An alarm substance in the fishs skin dispersesin the water, inducing a fright response among

    fish in the area

    Figure 51.9a, b

    (a) Minnows are widely dispersed in an aquarium

    before an alarm substance is introduced.(b) Within seconds of the alarm substance being

    introduced, minnows aggregate near the

    bottom of the aquarium and reduce their movement.

    Auditory Communication

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    Auditory Communication

    Experiments with various insects

    Have shown that courtship songs are undergenetic control

    Charles Henry, Luca Martnez, and ent Holsinger crossed males and females of Chrysoperla plorabunda and Chrysoperla johnsoni, two

    morphologically identical species of lacewings that sing different courtship songs.

    EXPERIMENT

    SONOGRAMS

    Chrysoperla plorabundaparent

    Vibration

    volleysStandard

    repeating

    unit

    Chrysoperla johnsoniparent

    Volley period

    crossed

    with

    Standard repeating unit

    The researchers recorded and compared the songs of the male and female parents with

    those of the hybrid offspring that had been raised in isolation from other lacewings.

    Volley

    period

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    The F1hybrid offspring sing a song in which the length of the standard repeating unit is similar to that sung by the Chrysoperla plorabunda parent, but the

    volley period, that is, the interval between vibration volleys, is more similar to that of the Chrysoperla johnsoniparent.

    RESULTS

    The results of this experiment indicate that the songs sung by Chrysoperla plorabunda and Chrysoperla

    johnsoni are under genetic control.CONCLUSION

    Standard repeating unit

    Volley

    period

    F1hybrids, typical phenotype

    Genetic Influences on Mating and Parental Behavior

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    Genetic Influences on Mating and Parental Behavior

    A variety of mammalian behaviors

    Are under relatively strong genetic control

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    Research has revealed the genetic and neural basis

    For the mating and parental behavior of male prairievoles

    Figure 51.11

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    Concept 51.3: Environment, interacting with an

    animals genetic makeup, influences the

    development of behaviors

    Research has revealed

    That environmental conditions modify many ofthe same behaviors

    Dietary Influence on Mate Choice Behavior

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    Dietary Influence on Mate Choice Behavior

    One example of environmental influence on

    behavior

    Is the role of diet in mate selection by

    Drosophila mojavensis

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    Laboratory experiments have demonstrated

    That the type of food eaten during larval developmentinfluences later mate choice in females

    Figure 51.12

    William Etges raised a D. mojavensispopulation from Baja California

    and a D. mojavensis population from Sonora on three different culture media: artificial

    medium, agria cactus (the Baja host plant), and organ pipe cactus (the Sonoran host

    plant). From each culture medium, Etges collected 15 male and female Baja D. mojavensis

    pairs and 15 Sonoran pairs and observed the numbers of matings between males and

    females from the two populations.

    EXPERIMENT

    When D. mojavensis had been raised on artificial medium, females from the

    Sonoran population showed a strong preference for Sonoran males (a). When D. mojavensis had been raised on cactus medium, the Sonoran females mated with Baja and Sonoran

    males in approximately equal frequency (b).

    RESULTS

    The difference in mate selection shown by females that developed on

    different diets indicates that mate choice by females of Sonoran populations of D. mojavensis

    is strongly influenced by the dietary environment in which larvae develop.

    CONCLUSION

    100

    75

    50

    25

    0

    Artificial Organ pipe cactus Agria cactus

    Culture medium

    With Baja males

    With Sonoran

    males

    (b)

    Proportionofmatings

    bySonoranfemales

    (a)

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    Therese Markow and Eric Toolson proposed

    That the physiological basis for the observedmate preferences was differences in

    hydrocarbons in the exoskeletons of the flies

    Figure 51.13

    Social Environment and Aggressive Behavior

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    Social Environment and Aggressive Behavior

    Cross-fostering studies in California mice and

    white-footed mice

    Have uncovered an influence of social

    environment on the aggressive and parental

    behaviors of these mice

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    Influence of cross-fostering on male mice

    Table 51.1

    Learning

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    Learning

    Learning is the modification of behavior

    Based on specific experiences

    Learned behaviors

    Range from very simple to very complex

    Habituation

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    Habituation

    Habituation

    Is a loss of responsiveness to stimuli thatconvey little or no information

    Spatial Learning

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    Spatial Learning

    Spatial learning is the modification of behavior

    Based on experience with the spatial structureof the environment

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    In a classic experiment, Niko Tinbergen

    Showed how diggerwasps use landmarks

    to find the entrances

    to their nests

    After the mother visited the nest and flew away, Tinbergen

    moved the pinecones a few feet to one side of the nest.

    Figure 51.14CONCLUSION

    A female digger wasp excavates and cares for four

    or five separate underground nests, flying to each nest daily with food

    for the single larva in the nest. To test his hypothesis that the wasp

    uses visual landmarks to locate the nests, Niko Tinbergen marked onenest with a ring of pinecones.

    EXPERIMENT

    Nest

    When the wasp returned, she flew to the center of

    the pinecone circle instead of to the nearby nest. Repeating the

    experiment with many wasps, Tinbergen obtained the same results.

    RESULTS

    The experiment supported the hypothesis

    that digger wasps use landmarks to keep track of their nests.

    NestNo Nest

    Cognitive Maps

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    Cognitive Maps

    A cognitive map

    Is an internal representation of the spatialrelationships between objects in an animals

    surroundings

    Associative Learning

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    Associative Learning

    In associative learning

    Animals associate one feature of theirenvironment with another

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    Classical conditioning is a type of associative

    learning

    In which an arbitrary stimulus is associated

    with a reward or punishment

    Figure 51.15

    Before stimulus

    Influx of water alone

    Influx of alarm substances

    Influx of pike odor

    Day 1 Day 3

    Control

    group

    Control

    group

    Experimental

    groupExperimental

    group

    Relativeactivitylevel

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    Operant conditioning is another type of

    associative learning

    In which an animal learns to associate one of

    its behaviors with a reward or punishment

    Figure 51.16

    Cognition and Problem Solving

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    Cognition and Problem Solving

    Cognition is the ability of an animals nervous

    system

    To perceive, store, process, and use

    information gathered by sensory receptors

    7

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    7

    Problem solving can be learned

    By observing the behavior of other animals

    Figure 51.17

    Genetic and Environmental I nteraction in Learning

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    Genetic and Environmental I nteraction in Learning

    Genetics and environment can interact

    To influence the learning process

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    Concept 51.4: Behavioral traits can evolve by

    natural selection

    Because of the influence of genes on behavior

    Natural selection can result in the evolution of

    behavioral traits in populations

    Behavioral Variation in Natural Populations

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    p

    When behavioral variation within a species

    Corresponds to variation in the environment, itmay be evidence of past evolution

    Variation in Prey Selection

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    y

    Differences in prey selection in populations of

    garter snakes

    Are due to prey availability and are evidence of

    behavioral evolution

    Figure 51.18a, b

    (a) A garter snake (Thamnophis

    elegans)

    (b) A banana slug (Ario l imus

    cal i forn icus); not to scale

    Variation in Aggressive Behavior

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    gg

    Funnel spiders living in different habitats

    Exhibit differing degrees of aggressiveness indefense and foraging behavior

    Figure 51.19

    50

    40

    30

    20

    10

    0

    Timetoattack(seconds)

    Field Lab-raised

    generation 1

    Lab-raised

    generation 2

    Desert

    grassland

    population

    Riparian

    population

    60

    Population

    Experimental Evidence for Behavioral Evolution

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    p

    Laboratory and field experiments

    Can demonstrate the evolution of behavior

    Laboratory Studies of DrosophilaForaging Behavior

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    p

    Studies of Drosophilapopulations raised in high-

    and low-density conditions

    Show a clear divergence in behavior linked to

    specific genes

    Figure 51.20

    14

    12

    10

    8

    6

    2

    0

    A

    veragepathlength(cm)

    4

    L1 L2 L3 H1 H2 H3 H4 H5

    D. Melanogasterlineages

    Low population

    density

    High population

    density

    Migratory Patterns in Blackcaps

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    g y p

    Field and laboratory studies of Blackcap birds

    Have documented a change in their migratorybehavior

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    Birds placed in funnel cages

    Left marks indicating the direction they weretrying to migrate

    Figure 51.21a

    (a) Blackcaps placed in a funnel cage left marks indicating the

    direction in which they were trying to migrate.

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    Migratory orientation of wintering adult birds

    captured in Britain

    Was very similar to that of laboratory-raised birds

    Figure 51.21b

    (b) Wintering blackcaps captured in Britain and their laboratory-raised

    offspring had a migratory orientation toward the west, while

    young birds from Germany were oriented toward the southwest.

    N

    E

    S

    W

    Adults from

    Britain and

    F1 offspring

    of British

    adults

    N

    E

    S

    WYoung

    from SW

    Germany

    Mediterranean

    Sea

    BRITAIN

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    Concept 51.5: Natural selection favors

    behaviors that increase survival and

    reproductive success

    The genetic components of behavior

    Evolve through natural selection

    Behavior can affect fitness

    Through its influence on foraging and matechoice

    Foraging Behavior

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    Optimal foraging theory

    Views foraging behavior as a compromisebetween the benefits of nutrition and the costs

    of obtaining food

    Energy Costs and Benefits

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    Reto Zach

    Conducted a cost-benefit analysis of feedingbehavior in crows

    The crows eat molluscs called whelks

    But must drop them from the air to crack the

    shells

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    Zach determined that the optimal flight height

    in foraging behavior

    Correlated with a fewer number of drops,

    indicating a trade-off between energy gained

    (food) and energy expended

    Figure 51.22

    60

    50

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    30

    20

    10

    0

    Averagenum

    berofdrops

    2 3 5 7 15

    Average number of drops

    Drop height

    preferred

    by crows

    125

    100

    25

    75

    50

    Total flight height

    Totalflightheight(numb

    erofdrops

    d

    ropheight)

    Height of drop (m)

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    In bluegill sunfish

    Prey selection behavior is related to prey density

    Figure 51.23

    Low prey density High prey density

    33%33%

    33%

    32.5%32.5%

    35%

    2%40%

    57%

    100%

    50%35%

    14%

    33%

    33%33%

    Small prey

    Medium prey

    Large prey

    Small prey

    Medium preyLarge prey

    Small prey

    Medium prey

    Large prey

    Percentage available

    Predicted percentage in diet

    Observed percentage in diet

    Large prey at

    far distance

    Small prey at

    middle distanceSmall prey at

    close distance

    Risk of Predation

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    Research on mule deer populations

    Has shown that predation risk affects wherethe deer choose to feed

    Figure 51.24

    70

    60

    40

    30

    20

    10

    0

    Pre

    dationoccurrence(%)

    50

    Relativedeeruse

    Relative deer use Predation

    risk

    Open Forest edge

    Habitat

    Forest interior

    0

    5

    10

    15

    20

    Mating Behavior and Mate Choice

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    Mating behavior

    Is the product of a form of natural selection callsexual selection

    Mating Systems and Mate Choice

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    The mating relationship between males and

    females

    Varies a great deal from species to species

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    In many species, mating is promiscuous

    With no strong pair-bonds or lastingrelationships

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    In monogamous relationships

    One male mates with one female

    Figure 51.25a

    (a) Since monogamous species, such as these trumpeter swans, are

    often monomorphic, males and females are difficult to distinguish

    using external characteristics only.

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    In a system called polygyny

    One male mates with many females

    The males are often more showy and larger

    than the females

    Figure 51.25bAmong polygynous species, such as elk, the male (left) is

    often highly ornamented.

    (b)

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    In polyandrous systems

    One female mates with many males

    The females are often more showy than the males

    Figure 51.25c

    (c) In polyandrous species, such as these Wilsons phalaropes, females

    (top) are generally more ornamented than males.

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    The needs of the young

    Are an important factor constraining theevolution of mating systems

    The certainty of paternity

    Influences parental care and mating behavior

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    In species that produce large numbers of

    offspring

    Parental care is at least as likely to be carried

    out by males as females

    Figure 51.26

    Eggs

    Sexual Selection and Mate Choice

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    In intersexual selection

    Members of one sex choose mates on thebasis of particular characteristics

    Intrasexual selection

    Involves competition among members of one

    sex for mates

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    Mate Choice by Females

    Male zebra finches

    Are more ornate than females, a trait that may

    affect mate choice by the females

    Figure 51.27

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    Imprinting of female chicks on males with more

    ornamentation

    Affects mate

    selection as

    adults

    Figure 51.28

    Experimental Groups Control Group

    Parents notornamented

    Both parentsornamented

    Malesornamented

    Femalesornamented

    Results

    Females reared byornamented parents

    or ornamented fathers

    preferred ornamented

    males as mates.

    Females reared by

    ornamented mothers or

    nonornamented parentsshowed no preference

    for either ornamented or

    nonornamented males.

    Males reared by all experimental groups showed no

    preference for either ornamented or nonornamented

    female mates.

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    The size of eyestalks in stalk-eyed flies

    Affects which males the females choose tomate with

    Figure 51.29

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    Male Competition for Mates

    Male competition for mates

    Is a source of intrasexual selection that can

    reduce variation among males

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    Such competition may involve agonistic

    behavior

    An often ritualized contest that determines

    which competitor gains access to a resource

    Figure 51.30

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    Morphology affects the mating behavior

    In isopods of the same species that aregenetically distinct

    Figure 51.31

    Large Paracerceis males

    defend harems of females

    within intertidal sponges.

    Tiny males are

    able to invade

    and live within

    large harems.

    males mimic female morphology and

    behavior and do not elicit a defensive

    reponse in males and so are able to

    gain access to guarded harems.

    Applying Game Theory

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    Game theory evaluates alternative behavioral

    strategies in situations

    Where the outcome depends on each

    individuals strategy and the strategy of other

    individuals

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    Mating success of male side-blotched lizards

    Was found to be influenced by malepolymorphism and the abundance of different

    males in a given area

    Figure 51.32

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    Concept 51.6: The concept of inclusive fitness

    can account for most altruistic social behavior

    Many social behaviors are selfish

    Natural selection favors behavior

    That maximizes an individuals survival and

    reproduction

    Altruism

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    On occasion, some animals

    Behave in ways that reduce their individualfitness but increase the fitness of others

    This kind of behavior

    Is called altruism, or selflessness

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    In naked mole rat populations

    Nonreproductive individuals may sacrifice theirlives protecting the reproductive individuals

    from predators

    Figure 51.33

    Inclusive Fitness

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    Altruistic behavior can be explained by

    inclusive fitness

    The total effect an individual has on

    proliferating its genes by producing its own

    offspring and by providing aid that enables

    close relatives to produce offspring

    Hamiltons Rule and Kin Selection

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    Hamilton proposed a quantitative measure

    For predicting when natural selection wouldfavor altruistic acts among related individuals

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    The three key variables in an altruistic act are

    The benefit to the recipient

    The cost to the altruist

    The coefficient of relatedness

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    The coefficient of relatedness

    Is the probabilitythat two relatives

    may share the

    same genes

    Figure 51.34

    Parent A Parent B

    OR

    Sibling 1 Sibling 2

    1/2(0.5)

    probability

    1/2(0.5)

    probability

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    Natural selection favors altruism when the

    benefit to the recipient

    Multiplied by the coefficient of relatedness

    exceeds the cost to the altruist

    This inequality

    Is called Hamiltons rule

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    Kin selection is the natural selection

    That favors this kind of altruistic behavior byenhancing reproductive success of relatives

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    An example of kin selection and altruism

    Is the warning behavior observed in Beldingsground squirrels

    Male

    Female

    Age (months)

    Meandistance

    m

    ovedfrom

    n

    atalburrow

    (m)

    300

    200

    100

    00 2 3 4 12 13 14 15 25 26

    Figure 51.35

    Reciprocal Altruism

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    Altruistic behavior toward unrelated individuals

    Can be adaptive if the aided individual returnsthe favor in the future

    This type of altruism

    Is called reciprocal altruism

    Social Learning

    S

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    Social learning

    Forms the roots of culture

    Culture can be defined as a system of

    information transfer through observation or

    teaching

    That influences the behavior of individuals in a

    population

    Mate Choice Copying

    M t h i i

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    Mate choice copying

    Is a behavior in which individuals in apopulation copy the mate choice of others

    Thi t f b h i

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    This type of behavior

    Has been extensively studied in the guppyPoecilia reticulata

    Figure 51.36

    Male guppies

    with varying

    degrees of

    coloration

    Control Sample

    Female guppies prefer

    males with more orange

    coloration.

    Experimental Sample

    Female model

    engaged in

    courtship with

    less orange

    male

    Female guppies prefer less

    orange males that are associated

    with another female.

    Social Learning of Alarm Calls

    V t k

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    Vervet monkeys

    Produce a complex set of alarm calls

    I f t k i di i i ti l

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    Infant monkeys give undiscriminating alarm

    calls at first

    But learn to fine-tune them by the time they are

    adults

    Figure 51.37

    N th i

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    No other species

    Comes close to matching the social learning andcultural transmission that occurs among humans

    Figure 51.38

    Evolution and Human Culture

    H lt

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    Human culture

    Is related to evolutionary theory in the distinctdiscipline of sociobiology

    H b h i lik th t f th i

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    Human behavior, like that of other species

    Is the result of interactions between genes andenvironment

    However, our social and cultural institutions

    May provide the only feature in which there is

    no continuum between humans and other

    animals


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