LECTURE PRESENTATIONS For CAMPBELL BIOLOGY, NINTH EDITION Jane

LECTURE PRESENTATIONS For CAMPBELL BIOLOGY, NINTH EDITION Jane

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. Most common APB Topics Go o Kn d to ow !

Sign stimulus and fixed action pattern (FAPs) and the associated releasers (cues) Stickleback fish in Figure 51.2 Imprinting (Figure 51.7) Big Ideas 1.B.1: Organisms share many conserved core processes and features that evolved and are widely distributed among organisms today.

2011 Pearson Education, Inc. Big Ideas 2.A.1: All living systems require constant input of free energy. Strategies for regulating body temperature (e.g. basking behavior of ectotherms) See Fig. 40.10, 40.11 2011 Pearson Education, Inc.

Big Ideas 2.E.3: Timing and coordination of behavior are regulated by various mechanisms and are important in natural selection. Communication among, between organisms Innate behaviors that are inherited Learning that occurs through interaction with others and environment

2011 Pearson Education, Inc. Big Ideas 3.B: Expression of genetic information involves cellular and molecular mechanisms. Signal transduction and gene transcription, translation pheromones leading to reproduction or as alarm system (see Figure 51.6) 2011 Pearson Education, Inc.

Big Ideas 3.D: Cells communicate by generating, transmitting, and receiving signals. Function of NTs in acquiring, processing, and acting on environmental information (S.A.M.E) Signal transduction 2011 Pearson Education, Inc.

Big Ideas 3.E.1: Individuals act on information and communicate it to others.

Predator warnings (Fixed Action Pattern; see Fig. 51.2) Coloration (Fixed Action Pattern; see Fig. 51.2) Territorial marking in animals Bee dances (Fig. 51.5) Fight or flight (FOF) response Bird songs Herd, flock, and schooling behavior in animals Colony and swarming behavior in insects

2011 Pearson Education, Inc. Big Ideas 4.A: Interactions within biological systems lead to complex properties. Competition, parasitism, predation, mutualism, commensalism 2011 Pearson Education, Inc.

Big Ideas 4.C.3: The level of variation in a population affects population dynamics. Not all animals in a population stampede 2011 Pearson Education, Inc. TYUs Level 1: 1

Level 2: 4, 6 Level 3: 10 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 2011 Pearson Education, Inc. Figure 51.1 A behavior is the nervous systems response to a stimulus and is carried out by the muscular or the hormonal system Behavior is subject to natural selection

2011 Pearson Education, Inc. Concept 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 animals experience during growth and development influence the response? 2011 Pearson Education, Inc. 3. How does the behavior aid survival and reproduction? 4. What is the behaviors evolutionary history? 2011 Pearson Education, Inc.

Behavioral ecology integrates proximate and ultimate explanations for animal behavior Proximate causation addresses how a behavior occurs or is modified, including Tinbergens questions 1 and 2 Ultimate causation addresses why a behavior occurs in the context of natural selection, including Tinbergens questions 3 and 4

2011 Pearson Education, Inc. 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.

Figure 51.2 (a) (b) 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 Earths magnetic field 2011 Pearson Education, Inc.

Figure 51.3 Behavioral Rhythms Some animal behavior is affected by the animals 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 animals behavior Communication is the transmission and reception of signals

2011 Pearson Education, Inc. Forms of Animal Communication Animals communicate using visual, chemical, tactile, and auditory signals Fruit fly courtship follows a three step stimulusresponse chain 2011 Pearson Education, Inc.

1. A male identifies a female of the same species and orients toward her Chemical communication: he smells a females chemicals in the air Visual communication: he sees the female and orients his body toward hers 2011 Pearson Education, Inc. 2. The male alerts the female to his presence

Tactile communication: he taps the female with a foreleg Chemical communication: he chemically confirms the females 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. Figure 51.4 (a) Orienting (b) Tapping

(c) Singing 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 Figure 51.5a (a) Worker bees Figure 51.5b

(b) Round dance (food near) Figure 51.5c (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 2011 Pearson Education, Inc.

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 fishs 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 #2: Learning establishes specific links between experience and behavior Innate behavior is developmentally fixed and does not vary among individuals 2011 Pearson Education, Inc. Experience and Behavior Cross-fostering studies help behavioral ecologists to identify the contribution of environment to an

animals behavior A cross-fostering study places the young from one species in the care of adults from another species 2011 Pearson Education, Inc. 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 2011 Pearson Education, Inc. Table 51.1 In humans, twin studies allow researchers to compare the relative influences of genetics and

environment on behavior 2011 Pearson Education, Inc. Learning A Few Categories of Learned Behavior Learning is the modification of behavior based on

specific experiences Imprinting Spatial (Latent) Learning Associative Learning Cognition & Problem Solving Development of Learned Behaviors

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 geese is a nearby object that is moving away from the young geese 2011 Pearson Education, Inc. Figure 51.7 (a) Konrad Lorenz and geese (b) Pilot and cranes

Figure 51.7a (a) Konrad Lorenz and geese 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. Figure 51.7b (b) Pilot and cranes Learning A Few Categories of Learned Behavior

Learning is the modification of behavior based on specific experiences Imprinting Spatial (Latent) Learning

Associative Learning Cognition & Problem Solving Development of Learned Behaviors 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 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 animals surroundings For example, Clarks nutcrackers can find food hidden in caches located halfway between particular landmarks 2011 Pearson Education, Inc. Learning

A Few Categories of Learned Behavior Learning is the modification of behavior based on specific experiences Imprinting Spatial (Latent) Learning

Associative Learning Cognition & Problem Solving Development of Learned Behaviors 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 bells 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 Figure 51.9a Figure 51.9b Figure 51.9c Learning A Few Categories of Learned Behavior

Learning is the modification of behavior based on specific experiences Imprinting Spatial (Latent) Learning Associative Learning

Cognition & Problem Solving Development of Learned Behaviors 2011 Pearson Education, Inc. 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 Food

Lid Decision chamber Stimulus Entrance (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 2011 Pearson Education, Inc.

Learning A Few Categories of Learned Behavior Learning is the modification of behavior based on specific experiences

Imprinting Spatial (Latent) Learning Associative Learning Cognition & Problem Solving Development of Learned Behaviors 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 Figure 51.12a Figure 51.12b

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. Application: Case Study #1

2011 Pearson Education, Inc. Animal Behavior: Learned Behavior The Rat (Neotoma magister) Rats, like many other animals, have an amazing capacity to learn and therefore adapt to their surroundings.

Bart Weetjens has worked with, and trained, rats for many years. Please explain how Mr. Weetjens rats are exceptional. Moreover, explain how humans and these pests can live in harmony. Application: Case Study #2 2011 Pearson Education, Inc.

The Crow (Corvus brachyrhynchos) For many, crows evoke a creepy foreboding feeling. For others, like Joshua Klein, they are exceptional examples of animal intelligence. Mr. Klein has researched and worked with crows for twenty years. In this time frame, he has unearthed great problem solving skills. Please

summarize his research (methods and conclusions). Lastly, explain how humans and these pests can live in harmony. Concept 51.3: Selection for individual survival and reproductive success can explain most behaviors Behavior enhances survival and reproductive success in a population

2011 Pearson Education, Inc. 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 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 lowdensity 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.

Figure 51.13 Mean path length (cm) 7 Low population density 6

High population density 5 4 3 2 1 0 R1

R2 R3 K1 D. melanogaster lineages K2

K3 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. 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. 125 60 Average number of drops 50

100 40 Average number of drops 30 75 Total flight height 20

Drop height preferred by crows 5.23 m 10 0 50

25 2 3 5 7 Drop height (m) 15

Total flight height (number of drops drop height in m) Figure 51.14 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 (a) Monogamous species (b) Polygynous species

(c) Polyandrous species Figure 51.15a (a) Monogamous 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 or polyandrous 2011 Pearson Education, Inc. In polygyny, one male mates with many females The males are usually more showy and larger than the females

2011 Pearson Education, Inc. Figure 51.15b (b) Polygynous species In polyandry, one female mates with many males The females are often more showy than the males

2011 Pearson Education, Inc. Figure 51.15c (c) Polyandrous species 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 checks (monogamy) Consider bird species where checks 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. Figure 51.16 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 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 Figure 51.19 Experimental Groups of Parental Pairs Both parents ornamented

Males ornamented Control Group Females ornamented Parents not

ornamented Offspring Offspring Mate preference of female offspring: ornamented male Mate preference of female offspring:

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. Figure 51.20a Control Sample Male guppies with varying degrees of coloration

Female guppies prefer males with more orange coloration. Figure 51.20b 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, an often ritualized contest that determines which competitor gains access to a resource 2011 Pearson Education, Inc. Figure 51.21 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 individuals strategy and the strategy of other individuals 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 nonterritorial, mimic females, and use sneaky strategies to mate

2011 Pearson Education, Inc. Figure 51.22 Like rock-paper-scissors, each strategy will outcompete one strategy, but be outcompeted by the other strategy The success of each strategy depends on the frequency of all of the strategies; this drives

frequency-dependent selection 2011 Pearson Education, Inc. 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 SOUND RECORDINGS Chrysoperla plorabunda parent: Volley period Standard repeating unit

Vibration volleys crossed with Chrysoperla johnsoni parent: Volley period Standard repeating unit

RESULTS F1 hybrids, typical phenotype: Volley period Standard repeating unit Figure 51.23a

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.

Figure 51.24 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. Case Study: Variation in Prey Selection

The natural diet of western garter snakes varies by population Coastal populations feed mostly on banana slugs, while inland populations rarely eat banana slugs Studies have shown that the differences in diet are genetic The two populations differ in their ability to detect and respond to specific odor molecules produced by the banana slugs

2011 Pearson Education, Inc. Figure 51.25 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 N BRITAIN W GE RM S A

N NY Young W from SW Germany E S Adults from

E Britain and offspring of British adults Figure 51.26a EXPERIMENT Scratch

marks Figure 51.26b RESULTS N BRITAIN W

E GE RM S AN N Y Young from SW Germany

W E S Adults from Britain and offspring of British

adults Altruism Natural selection favors behavior that maximizes an individuals 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 Beldings 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 2011 Pearson Education, Inc. Figure 51.27 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. Hamiltons 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) 2011 Pearson Education, Inc. Figure 51.28

Parent B Parent A OR /2 (0.5) probability

/2 (0.5) probability 1 1 Sibling 1

Sibling 2 Natural selection favors altruism when rB > C This inequality is called Hamiltons rule Hamiltons 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 sisters action The brother can now father two children, so B2 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 Beldings 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. Mean distance (m) moved from birthplace

Figure 51.29 300 Male 200 100 Female 0

1 2 3 4 12 13 14 15 25 26 Age (months)

Figure 51.29a Naked mole rats living within a colony are closely related Nonreproductive 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 2011 Pearson Education, Inc. Reciprocal altruism is limited to species with

stable social groups where individuals meet repeatedly, and cheaters (who dont 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

occurs 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 2011 Pearson Education, Inc. 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

Cognition Associative learning Spatial learning Social learning Figure 51.UN02

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