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Animal Behavior 480
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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
Logistics cont.
• 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.
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
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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?
How much energy is flowing or many species are there?
9,150
X 1000
Species
of “animals”
All of them
“behaving”!!
# species x 1000.
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
• Climate
• Soil
• Variation and distribution of microclimates
• Salinity, Turbidity, Tides (aquatic)
• Abiotic Disturbance
– Fire, hurricanes, wind, earthquakes, floods,
volcanoes, freezing rain, lightning, tree fall gap
etc.
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?
here
• 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.
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.
Impact of bodies of water on temperature and precipitation
Lake effect snow produced as cold winds blow clouds over warm waters
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.
Result
Each area has a basic biodiversity based
on all these conditions.
• Where it is in the planet.
• Local conditions
• History!???
Disturbances.
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 .
Disturbances.
Fires.
Disturbances.
Floods.
Disturbances.
Huricanes.
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.
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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.
here
Three foundations
• What are the bases of animal behavior,
where do they come from??
1) Natural selection, obviously
2) Individual learning
3) Cultural transmission
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?
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Three approaches:
1) Conceptual
2) Theoretical
3) Empirical
Often a combination of the three
Conceptual approaches
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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
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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
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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!
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Proximate factors
Immediate causation
Examples: Hormones and behavior
Neurobiological impacts
Molecular genetics impacts
Developmental stages
• These make up Chap 3 and will discuss
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
change
probability
of specific
outputs
• Change
behaviors
Hormones and behavior
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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
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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
behavior
individual
• 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!