Animal Behavior 480

• John Laundré

• Office is 303 Illick (office hours posted)

• Tel: 4938

• Email:

• TA’s

• Amanda Dillon

• Ayesha Prasad

• William Helenbrook

Logistics

• Lectures Mon, Wed and Fri and a Recitation once a week

• Grade based on 2 lecture exams (25% each), Attendance (10%), Recitation participation and attendance (25%) and a behavioral profile (15%).

• Lecture exams: multiple guess/short answer.

Logistics cont.

• Behavioral profile: Each student will select an animal species. They will gather information on aspects of this species’ behavior (basic behavior, sexual behavior, behavioral ecology, etc. You will then write a 5-10 (No more than 10!) summary of the important aspects of its behavior.

• Will give more details later on format

• Recitations: Review of classic and recent literature in animal behavior. Each week students will present and discuss assigned articles. Your grade will depend on your participation and the quality of your presentation.

Logistics cont.

Animal Behavior – what is it?

• Study of behavior: All aspects of behavior: instinctive, innate, learned, ecological behavior, etc.

• But what is Behavior????• Animal vs plant!! We move, they don’t?

• Why do animals “behave”?

What is animal behavior?

• Way animals interact with each other, the environment, etc.

• But why? Why do we react/interact?• Basic reason is survival: individual/species• What is basis to survival?• To better understand what is behavior and

why animals behave, need to go to basics:

• Energy!!

Why Energy?

• All systems, cells to ecosystems, function on energy flow.

• Energy machines, capturing, processing, passing energy through them.

• Basis of all biology is Energy• Examples

Energy

• Species: Energy capturing machines

• Death: loss of ability to capture energy

• Populations trap energy via reproduction

• Populations lose energy via mortality

• Communities energy flow through trophic levels

• Communities: Food webs/chains

Energy

• Ecosystems: energy transfer among systems: aquatic to terrestrial, etc.

• Take home message: Whole biological world on all levels functions on the basis of energy capture, processing, and transfer, the base being the

individual or species.

Energy Flow• Variety of factors that affect this energy flow.

One as you will see is behavior!• So we may talk about behavior, really are

talking about affecting the flow of energy through animals.

• Animal Behavior then are those actions that an animal performs to maintain energy flow! (either in itself or the population)

Energy flow

• The study of animal behavior is the analysis and cataloging of those actions (behaviors) AND trying to understand how they aid in the individual’s survival (energy flow).

• Much of what they do, depends on the energy available.

• To understand behavior, we need to understand energy flow!

Energy flow

• So before we get into aspects of behavior, need to review, from an energy standpoint,

Energy flow

• The first step is to look at just how much or different types of energy machines there are in the world.

• We then need to ask, why are there so many or so few?

• Or basically, what are the base factors that affect how much energy is available to a species

Energy and behaivor

• Once we know that, we can then ask what are the behaviors that influence the use of this energy flow.

• So, to start with, how much energy is flowing through species or… just how many species are there?

# species x 1000.

How much energy is flowing or many species are there?

9,150X 1000Speciesof “animals”All of them “behaving”!!

So 13,600,000 species plus or minus – 9-10 million animals !

Why so many?

Why doesn’t just one animal type work the best?Are they uniformly distributed?

If not, why?

And if not, how could this affect their behavior?

What controls energy flow?

• First step is to understand what affects energy flow.

• Abiotic

• Biotic factors

Abiotic Controls of energy flow

• Abiotic Disturbance – Fire, hurricanes, wind, earthquakes, floods,

volcanoes, freezing rain, lightning, tree fall gap etc.

• Climate • Soil

• Variation and distribution of microclimates

• Salinity, Turbidity, Tides (aquatic)

How does climate impact

Energy?

How come so many and not distributed uniformly?

- Energy machines.

- Energy comes from the sun.

- Represent different methods for trapping energy under different conditions.

- So… the conditions to trap energy are different over the planet!

- This will affect a species anatomy, physiology AND behavior!

Why different conditions?? Earth is a ball!!

And is not directly facing the sun!

The amount of energy received changes with the seasons.

• More energy at the equator.

• Changes in other places as rotate around sun.

• What is the result of all this?

• Obvious, less energy in polar areas

- Seasons (periods of cold in north and south)

• So areas of high diversity would be in the tropics, more energy!!

• But it is not all! This ball at an angle to the sun causes profound interactions.

here

Climatic

What is result?

• Bands of different precipitation and temperature.

• Bands of different groups of energy machines, Ecosystems.

Few examples:Humid hot tropical forest

Hot

Wet

Tropical Dry forests.

Tropical grasslands

Deserts

Temperate grasslands

Temperate shurblands

Temperate forests

Boreal Forests

Tundra

Nice general pattern but not uniform. Why?

Bodies of water.

                                     

  Lake effect snow produced as cold winds blow clouds over warm waters

Impact of bodies of water on temperature and precipitation

Ocean currents.

Mountains

Islands of cool and moist conditions.

Impact of slope

and aspect

Principal air currents on planet

Impact of mountains on regional climate

Soils.

Each area has a basic biodiversity based on all these conditions.

• Where it is in the planet.

• Local conditions

• History!???

Disturbances.

Result

Disturbance

• any relatively discrete event in time that disrupts ecosystem, community or population structure and changes resources, substrate availability or the physical environment

Disturbances

Glaciers .

Fires.

Disturbances.

Disturbances.

Floods.

Huricanes.

Disturbances.

What is the result?

• 14 biomes in the world.

• 867 distinct ecoregions.

What does all this have to do with behavior??

• In all of these we have animal energy machines making a living capturing energy under the unique conditions that exist.

• Not surprising there is a grand diversity of animals!

• How do they do it? How do they capture energy?

• Job of biology is to understand how these particular machines work and why!

Morphology/Anatomy/physiology

• Deals with basic animal structure and function.

• What are characteristics of a “animal”?

• We learn how this basic energy machine has been modified.

• Fixed in short term, can change on evolutionary time scale.

Animal Ecology

• Interactions of ecosystem: energy flow from plants to animals, one animal to another. Outcomes of interactions: competition, predation, etc.

• Most mass action, treat organisms like balls.

Animal Behavior

• That final level: It is the INTERFACE of the individual machine with its environment.

• Not just inanimate balls! Most of the outcomes of ecological processes DEPEND on the behavior of the individual.

• Ecological processes are the integration of all individual actions.

Animal Behavior

• Examples: Competition: outcome depends on behavioral interactions of competitors.

• Predator-prey relationships: all depends on predator-prey BEHAVIOR.

• Etc. Etc.

• Behavior is the key.

•

So lets begin

• Although talk about individual behaviors, remember the basic driving force behind them is the capture of energy under the unique conditions an INDIVIDUAL faces.

What do we look at first?

• Lets start with Chapter 1!!

• This will help us try and define how we go about studying behavior.

Three foundations

• What are the bases of animal behavior, where do they come from??

1)Natural selection, obviously

2)Individual learning

3)Cultural transmission

here

Natural Selection

• Initially skeptical about behavior being under natural selection forces

• Requires a genetic basis!

• Reluctant to say behavior is inherited!

• Ramifications!!

• Now clear that some behaviors genetically controlled, will look at more later

Natural Selection

• What this means is that if a behavior enhances survival, can be selected for

• Can be passed on from one generation to another.

Natural selection examples

• Main link between genes and behavior is many behaviors depend on physical characteristics.

• Book example: Cricket on Hawaii that evolved soundless wings under parasite pressure.

• Common mole rats and aggression

• Male bird colors and courting behaviors

Cricket mating behavior

Feather color and behavior

Individual Learning

• Animals not plants! Most have sensory organs and brains to take in information and process it.

• This enables a certain amount of learning (changing behavior based on experience).

• Helpful to individual but not directly transfers new behavior to offspring

• Indications that the ability to learn has a genetic basis.

• Chap 4

Cultural transmission

• Learned behaviors good/or bad for you but how benefit species?

• Evidence that such behaviors can be transmitted to others (humans obvious)

• This is kind of an “evolution” without direct gene role!

• Can be rapid and even further modified.

• Chap 5

How do we study behavior?

• Three approaches:

• 1) Conceptual

• 2) Theoretical

• 3) Empirical

• Often a combination of the three

Conceptual approaches

• Put two and two together

• Rethink old ideas

• The old light bulb over the head

• Dreams and drugs

• Idea is that one way or other, we come up with concept or mental model of how we think it is… and then test it!

Theoretical approaches• Theoryland of models: x + y3 = behavior

• Try to reduce natural phenomena to math model

• Advantage: objective (sort of), gives testable predictions.

• Disadvantage: often simplistic (to some this is an advantage!)

• Only as good as inputs, garbage in garbage out.

Theoretical approaches

• Overall good because stimulates thought but must not let models take on a life of their own!

• Have to be based in reality!

• If it doesn’t make biological sense, it likely is not a good model!

Empirical approach

• Based on initial observations:

• Behavior more than others are based on observations!

• We can see them!!

• How many of you have seen an atom??

Empirical approach

• Basically initial observations help develop ideas/predictions, which we then test

• Test either with additional observations

• Test with experimental designs

• Two major divisions between early European (observations) and American (manipulations) ethologists

• Now excepted both have a role

Difference between observations and experiments

• Observations use existing conditions. At most, “manipulate” by making observations under different conditions.

• Experiments set up controls and “manipulations” where purposely change things to test specific predictions

• Some overlap of the two

General areas of behavior

• “pure” behavior: usually presented in the context of physiology or immediate stimuli

• Often not in any environmental context.

• Examples: Early ethological studies of Lorenz and others/behavioral physiologists in the US: lab rat maze experiments

• Good at investigating “roots” of behavior.

General areas of behavior

• Behavioral Ecology:

• Looks at behaviors in their evolutionary/ecological context.

• How do behaviors help animal interface with environment/others

• Social behaviors big

• Reproductive behaviors big

• Predator-prey behaviors also

Four basic questions

Tinbergen (1963)

• Immediate stimuli “what causes it”

• Development “How does it develop”

• Survival function “What good is it”

• Phylogeny “How does it change”

• Additional: Ecological function “How does it affect others”

Combined

• Obviously connected:

• First look at some of these base behaviors

• Then look at more behavioral ecology

• The first to start with are:

• Ultimate and Proximate Factors (chap 2 & 3)

Causes/origins of behavior

• Depends on the eye of the beholder!

Ultimate and Proximate Factors??

• What is difference???

• Proximate: Immediate causation

• Ultimate: Forces that shape evolutionary development.

• Overall: Ultimate factors involve natural selection/evolution that give rise to a trait

Ultimate and Proximate factors

Ultimate factors

• Deals with the evolution of behavior

• Don’t want to spend much time on it

• Well recognized now that behavior has evolutionary background

• Like other traits

• See variations in behavior/some “better” than others.

Behavioral variations

• Variations are the substance of evolution

• Natural selection for traits

• Behavior leads to survival advantage and then passed on in standard way.

• Chap 2 gives various real-life and hypothetical examples

• Pigeons/group hunting/ravens, etc.

Pigeons

Group hunting

Ravens

Evolution of behaviors

• Pretty much accepted.

• Inherited same way other traits

• Of interest from evolutionary perspective

• Sociobiology/selfish genes

• Interesting area of study

• BUT…

Proximate factors

• We will concentrate on proximate factors

• How is it that…

• What is it that…

• Basically the “proximate” reasons for a behavior…that has evolved over time!

Proximate factors

• Immediate causation

• Examples: Hormones and behavior

• Neurobiological impacts

• Molecular genetics impacts

• Developmental stages

• These make up Chap 3 and will discuss

here

Proximate factors

Hormones and Proximate causation

• Hormones: chemical substances of endocrine system

• Review: come from adrenal gland, pituitary gland, thyroid gland, pancreas, gonads, hypothalamus.

• Review: circulate via blood stream

• Review: Long term communication vs nervous system

Hormones

• Review: have their impact by changing cell metabolism and DNA expression.

• Large number of them

• We know they have impact on physical features: growth, stress, sexual maturity, menstrual cycle, etc.

• Often cell specific: only certain cells of body respond: lock and key system

Hormones

• First question, before how they affect behavior is: how are they released?

• Environmental cues: day length is a biggie

Temperature/rainfall, variety of things.

Overall view• Three interactive systems: input, processing,

output• Hormones changeprobabilityof specificoutputs• Change behaviors

Hormones and behavior

• Can change modify a behavior

• Can trigger a behavior

• Can prime animals for specific behavior

• Affect organization of behavioral systems

Changing behavior

• Testosterone and aggression: High levels of testosterone related to breeding season produce higher levels of aggression in males

• Can be a positive

feedback…

Triggering/priming behavior

• Fight or flight response

• Complex sequence of hormonal changes

• 1) Starts with individual senses a “stressor”

• 2) Hypothalamus initiates 2 responses first in Adrenal gland

• 3) Adrenal gland secretes epinephrine and norepinephrine.

Fight or flight

• 4) Causes increase in blood sugar and delivery to vital organs (Brain, skeletal muscles, heart

• 5) shut down of blood to non-essential organs.

• Allows quick behavioral response to stressor.

Fight or flight

• The second is what supplies the sugar• 1) Hypothalamus secrets several hormones e.g.

Corticotropin-releasing hormone (CRH) and others with long names!!

• 2) CRH stimulates anterior pituitary gland to increase ACTH (Adrenocorticotropic hormone)

• ACTH stimulates Adrenal gland to secrete cortisol , which coverts noncarbohydrates to sugars!

Organization of behavioral systems

• Basically: the basis of your behavioral repertoire

• In mice, who is your womb neighbor makes a difference!

• Male with two female

neighbors will be exposed

to less testosterone!

Behavioral organization

• These males will be less aggressive, less sexually active than other males.

• Exhibited more parental care behavior

Neurobiological basis of behavior

• Ok, brief overview of role of hormones

• What about other communication system?

• Nervous system: rapid electrical impulses.

• What roles does it play?

Nervous system• Obviously in the sensory part: via sight,

sound, touch, etc.

• But what more?

Nervous system

• Assume all have some knowledge of anatomy of nervous system.

• To understand the role of the nervous system need to start at the stimulus being detected and trace it to the behavioral reaction.

• Book uses touch as the sensory input device but works for all

Reception of stimulus

• Stimulus must produce response, initiate wave of electrical activity in Dendrites which then moves down axon

• To produce response stimulus must exceed nerve cell’s threshold

• Below which, no reaction, above which will get firing of nerve cell.

Passage of stimulus

• Once stimulated, nerve impulse travels along nerve pathway.

• Rapidly reach effector organ, which then produces appropriate response.

• What is the appropriate effector organ and what is the appropriate response?

• Good questions! Don’t want to bite when you should fly or visa versa!!

Central control

• Obviously some stimuli will require immediate reaction (behavior). Reflex behaviors, bypass brain.

• Others some integration is needed, decision making, here brain plays role in producing the appropriate behavior

Nervous and Endocrine systems

• Not surprisingly, nervous impulses stimulate endocrine glands to secrete hormones!

• Allows for external input, e.g. sunlight, to be transmitted to appropriate glands.

Again, How is it all done??

• Can say brain controls flow of information, etc. but what exactly are the pathways?

• Obviously too numerous, and unknown, to discuss in detail.

• Object of the physiological behavior research first started in U.S.

• Give a couple of examples to show types of research possible.

Neurobiology and learning in voles

• Book example P. 93

• Observation of behavior: Male meadow voles polygamous and range across large home range.

• Hypothesis: Males should have better navigation skills.

• Test: in mazes, male meadow voles seem to do better than females.

Learning in voles

• Hypothesis: Monogamous species with similar home range size, should be no difference!

• Tested Prairie voles: Indeed no difference!

• Question: what is neurological basis of this difference?

Neurology of voles

• How do we test for this?• Known that hippocampus in cerebral cortex

important in spatial navigation: Proximate cause.• Hypothesis: hippocampus should be bigger in

male meadow voles vs females but should be no difference in prairie voles.

• Results: yes in meadow voles, no in prairie voles!

• Evidence that there is neurological basis of navigation behavior.

Other sites in brain

• Others have shown that Parietal and prefrontal cortex areas have roles in spatial learning.

• Seems to be related to number of dendritic spines, more = better.

• May be genetic (male vs female)• May be experience!! Found animals

AFTER learning trials had more spines than animals that did not participate!!

Next molecular genetics and behavior!

• Lets go deeper, lower than cells to molecular level.

• Different from ultimate-evolutionary cause

• Here looking at gene that currently exists and how it might trigger a behavior.

• Use existing variation to help test: Look to see if difference in behavior between genotypes!

An example

• ultraviolet vision in birds P.100• Behavioral importance: In many birds,

courtship involves bright, often UV colors in males.

• Females use UV vision ability to discern and select mates.

• In zebra finch, UV ability is controlled by a single amino acid!, change it and they loose UV ability!

Will the reverse work?

• Showed that species without UV ability, can change a single amino acid and give it to them!!

• Further work has suggested that other groups (flies) similar,

• Some species of birds loose and gain ability as gene is gained and lost.

What does this mean?

• In reality makes sense.

• We know that what we are is an expression of our genes.

• So any physical feature that is important in a behavior, is ultimately the result of gene expression.

• Change gene, change expression, change behavior!!

• Molecular-cellular-organ individual behavior

• So what is the real “Proximate cause”??

• Depends on level your interested, they all are!

Change topic

• Enough on Proximate causes, interesting and a lot of work.

• What we will look at now is the behaviors themselves! Will drift back to proximate causes but main focus will be behaviors

• Best to start with the behavior that enables many others: Learning behavior

Learning behavior (Chap 4)

• Identified earlier as a source of gaining behaviors

• Is a behavior in itself!