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Chapter 51 Animal Behavior

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Whale Bubble Net BBC

Bubble net Animation

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Behavior is what an animal does and how it does it.

Ethology- is the study of how animals behave in their natural habitat.

What is Behavior?

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Mechanistic basis for behavior:

Identify the stimuli that triggers the behavior or
Study the psychological, neuronal, and hormonal changes that underlie the behavior

E.g., MIGRATION

Hawaii

Distribution & Migration Map of Pacific Golden Plover

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Proximate Cause

External stimuli- changes in day length and temp
Internal stimuli - hormone levels

Winter plumage

Breeding plumage

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Ultimate Causation - addresses the evolutionary significance for a behavior and why natural selection favors this behavior.

Why did a behavior evolve?
Is it adaptive?
Does it contribute to reproductive success?

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Example: birds that migrate have a selective advantage over birds that don't/didn't, selected for over time, could be due to long term climate changes, glaciation, disease, taking advantage of food sources, etc.

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Play as a behavior has no apparent external goal, but may facilitate social development or practice of certain behaviors and provide exercise.

Practice and exercise may explain the ultimate bases of play

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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.

ULTIMATE 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.

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Nature versus Nurture:�

Nature

Behaviors are:

Innate
Hard-wired
Instinctual
Genetically determined

Nurture

Behaviors are:

Learned
Flexible
Not genetically determined

Behavior results from both genes and environmental factors

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To some extent ALL behavior has a Genetic Basis�

some is totally genetic - which implies heritable
some is learned but relies ENTIRELY on genetically based mechanisms
Genes code for RNA and proteins. They do not directly code for a behavior.

In general, behavior is a response to some environmental stimulus

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Mixed-Up Brothers of Bogata

Switched at birth! Two sets of identical twins were raised as two sets of Fraternal Twins.

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Innate Behaviors/ Instinct– inherited, highly stereotypes (same each time for many individuals)

The behavior is performed correctly the 1^st time without prior experience (not learned)
Triggered by some external Sign Stimulus
Once started, run to completion with no further input (FAP)
Breeding crosses produce hybrid behaviors�

�

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Directed Movement- Kinesis and Taxis

Reflex: movement of a body part in response to stimulus

Fixed Action Pattern (FAP)

Categories of Innate behaviors

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Directed Movements

Kinesis: simple change in activity or turning rate in response to a stimulus

Taxis: automatic movement, oriented movement +/- from stimulus (eg. phototaxis, chemotaxis, geotaxis)

Kinesis increases the chance that a sow bug will encounter and stay

in a moist environment.

Positive rheotaxis keeps trout facing into the current, the direction

from which most food comes.

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FAPs

Stereotyped and often complex series of movements, response to a specific sign stimulus/ releaser, hardwired, however, not purely genetic as may improve with experience

examples:

- courtship behavior� - rhythms - daily (circadian); annual (circannual)

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Japanese Crane

Courtship rituals

Fruit Fly Courtship

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Nobel Prize in Physiology or Medicine 1973

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Herring gull experiment by Niko Tinbergen

Releaser Stimuli- stimuli that release FAP

E.g., Chick and red dot on gull parents beak triggers feeding response- parent regurgitates food

Laysan albatross feeding chick

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Tinbergen: Male three-spined stickleback shows aggression at models with red undersides

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Lorenz: Egg rolling behavior in geese is a Fixed Action Pattern

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Learning

Behavior is modified based on experience

Categories:

Habituation
Associative Learning
Classical Conditioning

- Operant Conditioning

- Imprinting

Latent Learning
Insight Learning

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Habituation �

Decline in response to a harmless, repeated stimulus�

Prevents animal from wasting energy/attention on irrelevant stimuli��

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Associative Learning- Classical Conditioning

Classical Conditioning - animal learns to perform an "old" response to a new stimulus

Pavlov's dog�- place dried meat powder in dog mouth - salivation�- associate with bell - salivation to bell
Stimulus first, behavior second �(but of course there is an expectation of reward)

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Associative learning: Operant Conditioning.

This is called trial-and-error learning - an animal learns to associate one of its own behaviors with a reward or a punishment.

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Operant Conditioning �a. Skinner Box - levers, reward, electric

shocks

b. Behavior first, reward second ��

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What type of learning?

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Trial & Error & Observational Learning

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Observational Learning

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Social learning: learning by observing others

Vervet monkeys learning correct use of alarm calls.

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Imprinting �

1. A strong association learned during a specific developmental period "Sensitive Period" or "Critical Period"�

2. learning a releaser for

an innate behaviors

California condors raised

in captivity are reared by

Condor puppets

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Goose imprinting by Conrad Lorenz

Geese imprint on the first moving (with sound) object that they see after birth

There is a selection of a specific period of time (critical period- within hours of hatching) for social attachment and mate recognition in geese (to ensure geese imprint on the same species)

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Latent Learning- learning is not evident until later

Mice Wander through a Maze unrewarded
Mice developed a cognitive map of the maze
Learning became evident when mice could run the maze quickly when motivated by food.

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Insight, reasoning

manipulating concepts in the mind to arrive at adaptive behavior
mental trial-and-error
internal memory stores are used as additional sensory/information source

All examples of tool-using:

chickadees/tits and opening milk bottles
Egyptian Vulture - uses rocks
Cocos Finch - uses splinters of wood
North American Gulls, Northwestern Crow - smash clams on sandy beaches

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Cognition is the ability of an animal’s nervous system to perceive, store, process, and�use information gathered�by sensory receptors.

http://www.wimp.com/incrediblecrows/

Insight Learning

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Spatial Learning

Cognitive Map: internal representation of spatial relationship among objects in an animal’s surroundings

Birds use spatial maps to relocate nut caches

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Niko Tinbergen

Hypothesis: digger wasps use visual Landmarks to keep track of her nests

Move pine cones

Visual cue is arrangement pattern rather than objects themselves

Cognitive Map

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Communication

Pheromones – chemicals emitted by members of one species that affect other members of the species (eg. Queen bee, fruit fly, fish, termites, trees, humans)
Visual signals – eg. Warning flash of white of a mockingbird's wing
Tactile (touch) – eg. Male fruit fly taps female fly
Auditory signals – screech of blue jay or song of warbler

Courtship behavior of fruit flies

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Honeybee dance language

Used to inform other bees about distance and direction of travel to food sources

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Eating and

Reproduction

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Foraging: food-obtaining behavior�

Recognize, search for, capturing, and consuming food

Optimal Foraging Behavior- Minimize costs / Maximize benefits

What is the Cost?

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Energy costs and benefits in foraging behavior

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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.

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Fig. 51.7

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Sexual selection

Sexual Dimorphism
Ornaments correlate in general with health and vitality

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Natural selection favors mating behavior that maximizes the quantity or quality of mating partners

Courtship behavior consists of patterns that lead to copulation and consists of a series of displays and movements by the male or female.

Vogelkop Bowerbird

The bower is a cone-shaped hutThe bower is a cone-shaped hut-like structure some 100 cm high and 160 cm in diameter, with an entrance usually propped up by two column-like sticks. A front "lawn" of some square metersThe bower is a cone-shaped hut-like structure some 100 cm high and 160 cm in diameter, with an entrance usually propped up by two column-like sticks. A front "lawn" of some square meters area is cleaned of debris and laid out with mossThe bower is a cone-shaped hut-like structure some 100 cm high and 160 cm in diameter, with an entrance usually propped up by two column-like sticks. A front "lawn" of some square meters area is cleaned of debris and laid out with moss. On this, and in the entrance of the bower, decorations such as colourful flowers or fruit, shining beetle elytra, dead leaves and other conspicuous objects are collected and artistically arranged.

Males go to great lengths to ensure that their displays are in prime condition, replacing old items as needed, as well as trying to outdo their neighbours by finding more spectacular decorations, and arranging them appropriately. As opposed to other species of bowerbirds, such as the Satin BowerbirdMales go to great lengths to ensure that their displays are in prime condition, replacing old items as needed, as well as trying to outdo their neighbours by finding more spectacular decorations, and arranging them appropriately. As opposed to other species of bowerbirds, such as the Satin Bowerbird, there is no fixed preference for items of a certain colour, more important being the "novelty value" of the items instead, which can lead to fashion-like trends if males find rare or unusual items; such rare finds are prime targets for theft by neighboring males. Females visit bowers and, depending on whether they like the "treasure trove" on display, will mate with the attendant males. The bower, indeed the male, play no part in nesting and raising the young.

The songs and mimicryThe songs and mimicry skill of this bird are well known among the indigenous peoplesThe songs and mimicry skill of this bird are well known among the indigenous peoples. In September 1872, Odoardo BeccariThe songs and mimicry skill of this bird are well known among the indigenous peoples. In September 1872, Odoardo Beccari became the first naturalist to see the home grounds of this bowerbird in the Arfak MountainsThe songs and mimicry skill of this bird are well known among the indigenous peoples. In September 1872, Odoardo Beccari became the first naturalist to see the home grounds of this bowerbird in the Arfak Mountains of Irian Jaya.

Because of its unadorned and plain plumage, this bowerbird is relatively safe from persecution. A common species within its limited habitat range, the Vogelkop Bowerbird is evaluated as Least Concern on the IUCN Red List of Threatened Species.

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Mating systems differ among species.

Promiscuous: no strong bond pairs between males and females.
Monogamous: one male mating with one female.
Polygamous: an individual of one sex mating with several of the other sex.

Polygyny where a single male mates with many females.
Polyandry one female mates with several males.

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In polygamous relationships, an individual of one sex mates with several individuals of the other sex.
Species with polygamous mating systems are usually sexually dimorphic: males and females have different external morphologies.
Polygamous relationships can be either polygynous or polyandrous. In polygyny - one male mates with many females. The males are usually more showy and larger than the females.

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Fig. 51-20b

Polygynous species – Male larger and more dominant

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In polyandry = one female mates with many males.
The females are often more showy than the males.
Polyandry is a rare mating system.

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Fig. 51-20c

Polyandrous species – female has multiple partners

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Parental investment refers to the time and resources expended for raising of offspring.

It is generally lower in males
Females usually invest more time into parenting (fecundity, egg size, care of offspring)
Females are usually more discriminating in terms of the males with whom they choose to mate.

Females look for more fit males (i.e., better genes), the ultimate cause of the choice.

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Needs of the young are an important factor constraining evolution of mating systems.
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).

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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).

Females can be certain that eggs laid or young born contain her genes; however, paternal certainty depends on mating behavior.
Certainty of paternity influences parental care and mating behavior.

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Paternal certainty is relatively low in species with internal fertilization because mating and birth are separated over time.
Certainty of paternity is much higher when egg laying and mating occur together, as in external fertilization.
In species with external fertilization, parental care is at least as likely to be by males as by females.

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Paternal care by a male jawfish

Eggs

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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.

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For example, each side-blotched lizard has a blue, orange, or yellow throat, and each color is associated with a specific strategy for obtaining mates. There is a genetic basis to throat color and mating strategy.
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.

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Male polymorphism in the side-blotched lizard (Uta stansburiana)

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Agonistic behavior�

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Agonistic behavior is a contest involving threats.

Submissive behavior.
Ritual: the use of symbolic activity.
Generally, no harm is done.

Fig. 51.19

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Inclusive fitness can account for the evolution of altruistic social behavior

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.

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Altruism

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.

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Inclusive Fitness

Altruism can be explained by inclusive fitness.
Inclusive fitness is the total effect an individual has on proliferating its genes by producing offspring and helping close relatives produce offspring.

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Hamilton’s Rule and Kin Selection.

William Hamilton proposed a quantitative measure for predicting when natural selection would favor altruistic acts.
Hamilton’s rule states that natural selection favors altruistic acts.

Worker bees

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Hamilton’s Rule and Kin Selection

Natural selection favors altruism when:

rB > C

Three key variables in an altruistic act:

Benefit to the recipient (B)
Cost to the altruist (C)
Coefficient of relatedness (the fraction of genes that, on average, are shared; r)

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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 (don’t reciprocate) are punished. Reciprocal altruism has been used to explain altruism between unrelated individuals in humans.

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Social Learning

Social learning is learning through the observation of others and forms the roots of culture.
Culture is a system of information transfer through observation or teaching that influences behavior of individuals in a population.
Culture can alter behavior and influence the fitness of individuals.

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Case Study: Mate-Choice Copying

In mate-choice copying, individuals in a population copy the mate choice of others.
This type of behavior has been extensively studied in the guppy Poecilia reticulata.
Females who mate with males that are attractive to other females are more likely to have sons that are attractive to other females.

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Case Study: Social Learning of Alarm Calls

Vervet monkeys produce distinct alarm calls for different predators.
Infant monkeys give undiscriminating calls but learn to fine-tune

them by the time

they are adults.

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Sociobiology places social behavior in an evolutionary context

Social behavior is any kind of interaction between two or more animals, usually of the same species.

Orcas chasing Dusky Dolphin

Orca and Weddell Seal

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Competitive social behaviors often represent contests for resources

Sometimes�cooperation occurs.

Fig. 51.18

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Reconciliation behavior often happens between conflicting individuals.

Fig. 51.20

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Dominance hierarchies involve a ranking of individuals in a group (a “pecking order”).

Alpha, beta rankings exist.

The alpha organisms control the behavior of others.

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Territoriality is behavior where an individual defends a particular area, called the territory.

Territories are typically used for feeding, mating, and rearing young and are fixed in location.

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Schooling

https://drive.google.com/a/piedmont.k12.ca.us/file/d/0BwLXmoaDjNd2Yll2dWktUUhScG0tRXB5UF9NMkhFZ1hpQl8w/view?usp=sharing

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Evolution and Human Culture

No other species comes close to matching the social learning and cultural transmission that occurs among humans.
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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Learning

Imprinting

Learning

and

problem solving

Cognition

Spatial learning

Social learning

Associative learning

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Questions

What type of learning is associated with a critical period?

What is an example of this type of learning?

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What type of learning? Why?

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Scarecrows are only effective for a certain period of time. Why?

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Proximate? Ultimate?

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WIS/ WIM

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Describe the Conditions that would favor:

Reciprocal Altruism

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Songbird repertoires�provide us with�examples.

Why has natural�selection favored�a multi-song�behavior?

Fig. 51.5

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It may be advantageous for males attracting females.

Fig. 51.6

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You should now be able to:

Distinguish between the following pairs of terms: kinesis and taxis, classical and operant conditioning.
Suggest a proximate and an ultimate cause for imprinting in newly hatched geese or other behaviors.
Explain how associative learning may help a predator avoid toxic prey.

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Describe how cross-fostering experiments help identify the relative importance of environmental and genetic factors in determining specific behaviors.
Describe optimal foraging theory.
Define and distinguish among promiscuous, monogamous, and polygamous mating systems.

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Define altruistic behavior.
Distinguish between kin selection and reciprocal altruism.
Define social learning and culture.

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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.

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Western garter snake from a coastal habitat eating a banana slug

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Certainty of paternity can influence mating systems and parental care.

If the male is�unsure if offspring�are his, parental�investment is�likely to be lower.
Exceptions do�exist.

Male sea spider cares for eggs