LECTURE PRESENTATIONS

For CAMPBELL BIOLOGY, NINTH EDITION

Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson

Chapter 51

Animal Behavior

Lectures by
Erin Barley
Kathleen Fitzpatrick

© 2011 Pearson Education, Inc.

Learning Targets - See Unit 8
ENE-3.D.1 Organisms respond to changes in their environment through
behavioral and physiological mechanisms.

ENE-3.D.2 Organisms exchange information with one another in response to

internal changes and external cues, which can change behavior.

IST-5.A.1 Individuals can act on information and communicate it to others.

IST-5.A.2 Communication occurs through various mechanisms.

a. Organisms have a variety of signaling behaviors that produce
changes in the behavior of other organisms and can result in differential
reproductive success.
b. Animals use visual, audible, tactile, electrical, and chemical signals
to indicate dominance, find food, establish territory, and ensure
reproductive success.

IST-5.A.3 Responses to information and communication of information are vital

to natural selection and evolution.
a. Natural selection favors innate and learned behaviors that
increase survival and reproductive fitness.
b. Cooperative behavior tends to increase the fitness of the individual
and the survival of the population.

EVO-1.O.1 An adaptation is a genetic variation that is favoured by selection

and is manifested as a trait that provides an advantage to an organism in a
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?

© 2011 Pearson Education, Inc.

 In scientific terms, what is a behavior?

• A behavior is the nervous system’s response to

a stimulus and is carried out by the muscular or
the hormonal system
• Behavior is subject to natural selection

Video: Albatross Courtship Ritual

Video: Blue-footed Boobies Courtship Ritual

Video: Giraffe Courtship Ritual

© 2011 Pearson Education, Inc.
Concept 51.1: 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?
3. How does the behavior aid survival and
reproduction?
4. What is the behavior’s evolutionary history?
© 2011 Pearson Education, Inc.
 • Behavioral ecology is the study of the ecological
and evolutionary basis for animal behavior
• 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

© 2011 Pearson Education, Inc.

Animal Communication
Split into FIVE groups
• You will be assigned a behavior or a learning that establishes
behavior.
• You will then use an example to act out or model and explain a
scenario representative of your assignment.
• Consider and discuss: How might the behavior or mode of
communication increase or decrease evolutionary fitness?

1. Fixed action pattern and Migration (1138-1139)

2. Animal signals and communication (1139-1141)
3. Experience & Behavior: cross-fostering, learning, Imprinting (1141-
1142)
4. Experience & Behavior: Spatial and cognitive maps, associative
(1142-1144)
5. Cognition and problem solving, social learning (1144-1146)
6. Behavior and evolution, Genetic basis of behavior (Ms. Noll)
 Figure 51.2

The male stickleback

fish attacks other male
sticklebacks that invade
its nesting territory.

(a)
The realistic model
produced no
aggressive behavior.
The red underside
models did produce
aggressive behavior.

Explain the behavior

using Tinbergen’s
questions.
(b)
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

© 2011 Pearson Education, Inc.

 • 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

© 2011 Pearson Education, Inc.

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 clock that is an integral part of
their nervous system
– The position of the North Star
– The Earth’s magnetic field

© 2011 Pearson Education, Inc.

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

© 2011 Pearson Education, Inc.

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

Forms of Animal Communication

• Animals communicate using visual, chemical,
tactile, and auditory signals
• Fruit fly courtship follows a three step stimulus-
response chain
© 2011 Pearson Education, Inc.
 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

2. The male alerts the female to his presence

• Tactile communication: he taps the female with a
foreleg
• Chemical communication: he chemically confirms
the female’s identity
© 2011 Pearson Education, Inc.
 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

© 2011 Pearson Education, Inc.

 • 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

© 2011 Pearson Education, Inc.

Figure 51.5

(a) Worker bees (b) Round dance

(food near)
(c) Waggle dance
(food distant)

A
30°
C
B Beehive

30°

Location A Location B Location C

Pheromones
• Many animals that communicate through odors
emit chemical substances called pheromones
• 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, inducing a fright response among fish in
the area
© 2011 Pearson Education, Inc.
Figure 51.6
(a) Minnows
before alarm

(b) Minnows
after alarm
 • Pheromones can be effective at very low
concentrations
• 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

© 2011 Pearson Education, Inc.

Concept 51.2: Learning establishes specific
links between experience and behavior
• Innate behavior is developmentally fixed and
does not vary among individuals

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
© 2011 Pearson Education, Inc.
Explain the influence of environment on the California and white-footed
mice. What does this suggest about nature vs. nurture?
 A cross-fostering study with rats

High-interaction mother Low-interaction mother

Pups become
relaxed adults
Pups become
fearful adults

Female pups become Female pups become

high-interaction mothers low-interaction mothers

Cross-fostering
experiment

Pups become relaxed adults Pups become fearful adults

 • In humans, twin studies allow researchers to
compare the relative influences of genetics and
environment on behavior

Learning
• Learning is the modification of behavior based
on specific experiences

© 2011 Pearson Education, Inc.

Imprinting
• Imprinting is a behavior that includes learning
and innate components and is generally
irreversible
• It is distinguished from other learning by a
sensitive period
• A sensitive period is a limited developmental
phase that is the only time when certain
behaviors can be learned

© 2011 Pearson Education, Inc.

• 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 (a) Konrad Lorenz and geese

geese is a nearby object that is

moving away from the young
geese
Video: Ducklings
(b) Pilot and cranes
© 2011 Pearson Education, Inc.
 • 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

© 2011 Pearson Education, Inc.

Spatial Learning and Cognitive Maps
• Spatial learning is a more complex modification
of behavior based on experience with the spatial
structure of the environment
• Niko Tinbergen showed how digger wasps use
landmarks to find nest entrances

Video: Bee Pollinating

© 2011 Pearson Education, Inc.
Figure 51.8
EXPERIMENT

Nest
Pinecone

RESULTS

Nest No nest
 • A cognitive map is an internal representation of
spatial relationships between objects in an
animal’s surroundings
– For example, Clark’s nutcrackers can find food
hidden in caches located halfway between
particular landmarks

© 2011 Pearson Education, Inc.

Associative Learning
• In associative learning, animals associate one
feature of their environment with another
– For example, a white-footed mouse will avoid
eating caterpillars with specific colors after a bad
experience with a distasteful monarch butterfly
caterpillar

© 2011 Pearson Education, Inc.

 • Classical conditioning is a type of associative
learning in which an arbitrary stimulus is
associated with a reward or punishment
– For example, a dog that repeatedly hears a bell
before being fed will salivate in anticipation at the
bell’s sound

© 2011 Pearson Education, Inc.

 • Operant conditioning is a type of associative
learning in which an animal learns to associate
one of its behaviors with a reward or punishment
• It is also called trial-and-error learning
– For example, a rat that is fed after pushing a
lever will learn to push the lever in order to
receive food
– For example, a predator may learn to avoid a
specific type of prey associated with a painful
experience

© 2011 Pearson Education, Inc.

Figure 51.9
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”

© 2011 Pearson Education, Inc.

Figure 51.10

Decision
chamber
Food

Lid Stimulus

Entranc
e

(a) Color maze (b) Pattern maze

 • 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

Video: Chimp Cracking Nut

© 2011 Pearson Education, Inc.
Development of Learned Behaviors
• Development of some behaviors occurs in distinct
stages
– For example a white-crowned sparrow
memorizes the song of its species during an
early sensitive period
– The bird then learns to sing the song during a
second learning phase

© 2011 Pearson Education, Inc.

Social Learning
• 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
– For example, vervet monkeys give and respond
to distinct alarm calls for different predators

© 2011 Pearson Education, Inc.

Figure 51.11
Figure 51.12 Vervet monkeys learning correct use of alarm calls.
Why do whales sing? How does this
benefit whales?
 • 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

© 2011 Pearson Education, Inc.

Associated Learning Targets
IST-5.A.2 Communication occurs through various
mechanisms.

a. Organisms have a variety of signaling behaviors

that produce changes in the behavior of other organisms
and can result in differential reproductive success.
b. Animals use visual, audible, tactile, electrical, and chemical
signals to indicate dominance, find food, establish territory,
and ensure reproductive success.

IST-5.A.3 Responses to information and communication of

information are vital to natural selection and evolution.

a. Natural selection favors innate and learned

behaviors that increase survival and reproductive fitness.
b. Cooperative behavior tends to increase the fitness of the
individual and the survival of the population.
Concept 51.3: Selection for individual
survival and reproductive success can
explain most behaviors
• Behavior enhances survival and reproductive
success in a population

Foraging Behavior
• Natural selection refines behaviors that enhance
the efficiency of feeding
• Foraging, or food-obtaining behavior, includes
recognizing, searching for, capturing, and eating
food items
© 2011 Pearson Education, Inc.
Evolution of Foraging Behavior – what can explain the data
below? Genetics to selection?

7
Low population density
Mean path length (cm)

6 High population density

5
4
3
2
1
0
R R R K K K
1 2 D.3 melanogaster lineages
1 2 3

74 Generations of Drosophila melanogaster larvae

Evolution of Foraging Behavior
• 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

© 2011 Pearson Education, Inc.

 • 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

© 2011 Pearson Education, Inc.

Optimal Foraging Model
• Optimal foraging model views foraging behavior
as a compromise between benefits of nutrition
and costs of obtaining food
• The costs of obtaining food include energy
expenditure and the risk of being eaten while
foraging
• Natural selection should favor foraging behavior
that minimizes the costs and maximizes the
benefits

© 2011 Pearson Education, Inc.

 • Optimal foraging behavior is demonstrated by the
Northwestern crow
• A crow will drop a whelk (a mollusc) from a height
to break its shell and feed on the soft parts
• The crow faces a trade-off between the height
from which it drops the whelk and the number of
times it must drop the whelk

© 2011 Pearson Education, Inc.

Explain how this data shows minimizing of cost and maximizing benefit.

60 125

Total flight height (number of drops × drop height in m)

50
100

40
Average number of

30 Average number of drops 75

Total flight height
drops

20
Drop height
preferred 50

10 by crows = 5.23 m

0 25
2 3 5 7 15
Drop height (m)
 • Researchers determined experimentally that
the total flight height (which reflects total energy
expenditure) was minimized at a drop height of
5m
• The average flight height for crows is 5.23 m

© 2011 Pearson Education, Inc.

 How about
Cicadas…….
Animal Behavior - Foraging & Surviving

What are some

shorebird
strategies?

Go to this link
Balancing Risk and Reward
• Risk of predation affects foraging behavior
– For example, mule deer are more likely to feed in
open forested areas where they are less likely to
be killed by mountain lions

© 2011 Pearson Education, Inc.

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

© 2011 Pearson Education, Inc.

Mating Systems and Sexual Dimorphism
• The mating relationship between males and
females varies greatly from species to species

• In many species, mating is promiscuous, with no

strong pair-bonds or lasting relationships

• In monogamous relationships, one male mates

with one female
• Males and females with monogamous mating
systems have similar external morphologies

© 2011 Pearson Education, Inc.

Figure 51.15

(c) Polyandrous species

(a) Monogamous species

(b) Polygynous species

 • 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 (male has multiple female mates) or
polyandrous (female has multiple male mates)

© 2011 Pearson Education, Inc.

 • In polygyny, one male mates with many females
• The males are usually more showy and larger
than the females

• In polyandry, one female mates with many males

• The females are often more showy than the
males

Quick Google Search - Look up 2 examples of each!

Mating Systems and Parental Care

• Needs of the young are an important factor
constraining evolution of mating systems
© 2011 Pearson Education, Inc.
 • 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)

© 2011 Pearson Education, Inc.

 • 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

© 2011 Pearson Education, Inc.

 • 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

© 2011 Pearson Education, Inc.

Sexual Selection and Mate Choice
• Sexual dimorphism results from sexual selection,
a form of natural selection
• In intersexual selection, members of one sex
choose mates on the basis of certain traits
• Intrasexual selection involves competition
between members of the same sex for mates

© 2011 Pearson Education, Inc.

 Mate Choice by Females
• 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

© 2011 Pearson Education, Inc.

Figure 51.17

Male stalk-eyed flies - competing for female attention.

 • Another example of mate choice by females
occurs in zebra finches

• Female chicks who imprint on ornamented

fathers are more likely to select ornamented
mates
• Experiments suggest that mate choice by female
zebra finches has played a key role in the
evolution of ornamentation in male zebra finches

© 2011 Pearson Education, Inc.

Figure 51.18
 Is sexual selection influenced by imprinting? Justify.
Experimental Groups of Parental Pairs Control Group

Both parents Males Females Parents not

ornamented ornamented ornamented ornamented

Offspring Offspring

Mate preference of female offspring: Mate preference of female offspring:

ornamented male none
 • Mate-choice copying is a behavior in which
individuals copy the mate choice of others
– For example, in an experiment with guppies, the
choice of female models influenced the choice of
other females

© 2011 Pearson Education, Inc.

Figure 51.20
Control Sample

Male guppies
with varying
degrees of
coloration

Female guppies prefer

males with more orange
coloration.

Experimental Sample

Female model
in mock
courtship with
less orange
male
Female guppies prefer males that
are associated with another female.
 Male Competition for Mates
• Male competition for mates is a source of
intrasexual selection that can reduce variation
among males

• Such competition may involve agonistic behavior

(cause physical damage), an often ritualized contest
that determines which competitor gains access
to a resource

© 2011 Pearson Education, Inc.

Figure 51.21
 Video: Chimp Agonistic Behavior

Video: Snake Ritual Wrestling

Video: Wolves Agonistic Behavior

© 2011 Pearson Education, Inc.
Applying Game Theory
• In some species, sexual selection has driven the
evolution of alternative mating behavior and
morphology in males
• The fitness of a particular phenotype (behavior or
morphology) depends on the phenotypes of
other individuals in the population
• Game theory evaluates alternative strategies
where the outcome depends on each individual’s
strategy and the strategy of other individuals
(Similar to frequency-dependent selection)

© 2011 Pearson Education, Inc.

 • For example, each side-blotched lizard has a
blue, orange, or yellow throat
• Each color is associated with a specific strategy
for obtaining mates

- Orange-throat males are

the most aggressive and
defend large territories
- Blue-throats defend small
territories
- Yellow-throats are non
territorial, mimic females,
The success of each strategy
and use “sneaky”
depends on the frequency of all
strategies to mate
of the strategies; this drives
frequency-dependent selection
Concept 51.4: Inclusive fitness can account
for the evolution of behavior, including
altruism
• Animal behavior is governed by complex
interactions between genetic and environmental
factors

• Selfless behavior can be explained by inclusive

fitness

© 2011 Pearson Education, Inc.

Genetic Basis of Behavior
• A master regulatory gene can control many
behaviors
– For example, a single gene controls many
behaviors of the male fruit fly courtship ritual
• Multiple independent genes can contribute to a
single behavior
– For example, in green lacewings, the courtship
song is unique to each species; multiple
independent genes govern different components
of the courtship song

© 2011 Pearson Education, Inc.

Figure 51.23

EXPERIMENT RESULTS
SOUND RECORDINGS F1 hybrids, typical phenotype:
Chrysoperla plorabunda parent: Volley
Volley period period

Standard Vibration Standard

repeating unit volleys repeating unit
crossed
with

Chrysoperla johnsoni parent:

Volley period

Standard repeating unit

• 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

© 2011 Pearson Education, Inc.

Genetic Variation and the Evolution of
Behavior
• When behavioral variation within a species
corresponds to environmental variation, it may be
evidence of past evolution

© 2011 Pearson Education, Inc.

• The natural diet of western
Genetic
garter snakes varies by
population
Variation &
the Evolution
• Coastal populations feed mostly
of Behavior -
on banana slugs, while inland
populations rarely eat banana Case Study:
slugs Variation in
• Studies have shown that the Prey Selection
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
© 2011 Pearson Education, Inc.
Case Study: Variation in Migratory Patterns
• Most blackcaps (birds) that breed in Germany
winter in Africa, but some winter in Britain
• Under laboratory conditions, each migratory
population exhibits different migratory behaviors

• The migratory behaviors are regulated by

genetics

© 2011 Pearson Education, Inc.

Figure 51.26
EXPERIMENT

Scratch
marks

RESULTS

BRITAIN Adults from

W E Britain and
GE offspring
RM S of British
A adults
N NY

Young W E
from SW
Germany
S
Altruism
• Natural selection favors behavior that maximizes
an individual’s survival and reproduction

• These behaviors are often selfish

• On occasion, some animals behave in ways that

reduce their individual fitness but increase the
fitness of others

• This kind of behavior is called altruism, or

selflessness
© 2011 Pearson Education, Inc.
 – 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, non-
reproductive individuals may sacrifice their lives
protecting their reproductive queen and kings from
predators

© 2011 Pearson Education, Inc.

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

© 2011 Pearson Education, Inc.

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 (avg. # extra offspring) (B)
– Cost to the altruistic (# fewer offspring) (C)
– Coefficient of relatedness (the fraction of
genes that, on average, are shared; r)

© 2011 Pearson Education, Inc.

Figure 51.28
Parent A Parent B

Siblings have an r value of

0.5. This weakens with
OR hereditary distance.

/2 (0.5)
1 1
/2 (0.5)
probability probability

Sibling 1 Sibling 2
 • 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

© 2011 Pearson Education, Inc.

 • 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)

© 2011 Pearson Education, Inc.

 • Kin selection is the natural selection that favors
this kind of altruistic behavior by enhancing
reproductive success of relatives

• An example of kin selection and altruism is the

warning behavior in Belding’s ground squirrels
• In a group, most of the females are closely
related to each other
• Most alarm calls are given by females who are
likely aiding close relatives

© 2011 Pearson Education, Inc.

Figure 51.29

300
Mean distance (m)

Male
moved from
birthplace

200

100
Female
0
1 2 3 4 12 13 14 15 25 26

Age (months)
 • Naked mole rats living within a colony are closely
related

• Non reproductive individuals increase their

inclusive fitness by helping the reproductive
queen and kings (their close relatives) to pass
their genes to the next generation

© 2011 Pearson Education, Inc.

Reciprocal Altruism
• Altruistic behavior toward unrelated individuals
can be adaptive if the aided individual returns the
favor in the future
• This type of altruism is called reciprocal altruism

• Reciprocal altruism is limited to species with

stable social groups where individuals meet
repeatedly, and cheaters (who don’t reciprocate)
are punished
• Reciprocal altruism has been used to explain
altruism between unrelated individuals in
humans
© 2011 Pearson Education, Inc.
 • In game theory, a tit-for-tat strategy has the
following rules:
– Individuals always cooperate on first encounter
– An individual treats another the same way it was
treated the last time they met
• That is, individuals will always cooperate, unless
their opponent cheated them the last time they met

© 2011 Pearson Education, Inc.

 • Tit-for-tat strategy explains how reciprocal
altruism could have evolved
• Individuals who engage in a tit-for-tat strategy
have a higher fitness than individuals who are
always selfish

© 2011 Pearson Education, Inc.

Evolution and Human Culture
• No other species comes close to matching the
social learning and cultural transmission that
occur among humans
• Human culture is related to evolutionary theory in
the distinct discipline of sociobiology
• Human behavior, like that of other species, results
from interaction between genes and environment

• However, our social and cultural institutions may

provide the only feature in which there is no
continuum between humans and other animals
© 2011 Pearson Education, Inc.
Figure 51.UN01

Imprinting

Learning and
problem solving

Spatial learning
Cognition

Associative learning Social learning

 Killer Whales- How do
Cooperative they cooperate and
how does it benefit
Behavior their species?
Swarming
Manakin Bird

● What are some individual behaviors?

● What are some group behaviors?
● How do they contribute to survival?

26:50 - 31:08
Migration
Group feeding
Schools of Fish