ANIMAL BEHAVIOR

Outline
• What is behavior?
• Why study behavior?
• How to study behavior scientifically
– Behavioral Observation
– Tinbergen’s 4 questions
• Empirical examples of each the 4 questions
– Hypotheses and Predictions
– Experimental Design
 What is behavior?
• What organisms “do”
– First line of interaction between animal and
environment
• Allows animals to actively respond and make
adjustments to changes in the environment (biotic and
abiotic).

http://www.youtube.com/watch?v=BGPG
knpq3e0
 What is behavior?
• Numerous
types/categories of
behavior
– Feeding/Foraging
– Anti-predator behavior
– Dispersal/Migration
– Habitat selection
– Communication
– Social/cooperative
behaviors
– Reproduction
– Territoriality/Aggression
– Self-maintenance
– Etc…
 Why study behavior?
“Pure science” reasons
• Studying behavior with the desire to understand
how and why animals do what they do
• Behavior is critical to animals’ survival and
reproductive success
– Internal homeostasis
• Temperature
• Water/salt balance
• Energy balance and nutrition
– Predator avoidance
– Reproduction
 Why study behavior?
“Applied” reasons
• Studying behavior in order
to:
– Understand human
behavior, health
– Control pests, diseases, etc
– Improve captive animal
welfare
– Make conservation and
management decisions for
wild animals
 Why study behavior?
“Pure Science” often leads
to “applied” benefits:
• Practical applications of
behavioral research often
come out of studying
behavior from an
originally “pure science”
reason
How to study behavior scientifically
1. Begin with an observation
2. Pose question
-Tinbergen’s 4 questions
3. Propose hypothesis (or
alternative hypotheses)
Niko Tinbergen
4. Generate predictions
5. Test the predictions
 Tinbergen’s “0th” question
• Observation: What is the animal doing?
– Spend a great deal of time watching the animal
– Characterize the behavior
– Methods of behavioral observation
• E.g. Ethograms (time budget of behaviors)
– Must observe behavior before attempting to
explain the behavior
 Some things to consider when
observing behavior
• Avoid anthropomorphism
– Can lead to misinterpretation
– Important to separate our emotional reactions to
the animal, from the animal’s actions
• Observations should use neutral wording
– Don’t attribute any motivation, intention, or
purpose; e.g. squirrel “buries” vs “hides” nuts in
the ground.
 Some things to consider when
observing behavior
• Umwelt
– The entire sensory environment of the organism,
including how they perceive it and how they
interact with it
– Species in same environment can have different
umwelts
Umwelt: Human sensory abilities pale in
comparison to other animals
How to study behavior scientifically
1. Begin with an observation
2. Pose question
-Tinbergen’s 4 questions
3. Propose hypothesis (or
alternative hypotheses)
Niko Tinbergen
4. Generate predictions
5. Test the predictions
 Posing Questions:
Levels of Analysis
Proximate causes Ultimate causes
• The mechanisms • Evolutionary causes of
underlying the behavior the behavior
• Processes within the • Processes beyond the
animal’s lifetime animal’s lifetime
• The “how” of behavior (previous generations)
• The “why” of behavior
 Levels of Organization
Ecosystems
Communities Implications for
Populations higher levels
Individuals
Organ systems
Mechanisms at
lower levels
Organs
Cells
Organelles
Molecules
 Tinbergen’s “4 questions” about
behavior
1. How does it develop?
Proximate
level 2. What mechanisms
activate it?

Niko Tinbergen

3. How does it contribute to fitness?

Ultimate
level 4. How did it evolve?
Tinbergen’s “4 questions” about behavior:
Proximate questions
1. How does it develop? (PD)
– Genetic-developmental mechanisms
• Effects of heredity on behavior
• Effects of early experience and/or learning on behavior
• Development of sensory-motor systems via gene-
environment interactions
– Involves processes throughout the lifetime of the
animal, from embryo to adult
Tinbergen’s “4 questions” about behavior:
Proximate questions
Environmental cues
2. What mechanisms activate it?
(PC)
– Sensory-motor mechanisms Sensory
(immediate causal mechanisms)
• Nervous systems for detecting
environmental cues/stimuli (sensory) Response
• Hormone systems for adjusting system
responsiveness to stimuli
• Skeletal-muscular systems for
carrying out responses (motor)
– Involves processes that are Motor
occurring right around the time
the behavior is being performed
Behavior
Tinbergen’s “4 questions” about behavior:
Ultimate questions
3. How does it contribute to fitness? (UF)
– Selective processes shaping the history of a behavioral
trait
• How the behavior improves reproductive success
and survival, both in evolutionary past and current
usefulness
– Note: effect of trait is on individual’s fitness, not a
benefit to the group. (see “The Problem with Group
Selection” in the book, pg 21)
 Evolution of Behavior
• Variation in behavior
• Inheritance of behavioral
variants
• Differential reproduction of
individuals showing different
behavioral variants Charles Darwin

• Evolution of different behavior patterns, and

of mechanisms underlying those behaviors
 Tinbergen’s “4 questions” about behavior:
Ultimate questions
4. How did it evolve? (UH)
– Historical pathways leading
to behavioral trait
• Evolutionary stages, from
origin of trait to the present
• Involves analysis of
phylogenetic distribution of
the trait, comparing closely
related species
See box 1.1 in book for a
refresher on phylogenies
How to study behavior scientifically
1. Begin with an observation
2. Pose a question
– 1 of Tinbergen’s 4 questions
3. Propose hypothesis (or alternative
hypotheses)
4. Generate predictions
5. Test the predictions
 Proposing Hypotheses
• All behaviors are influenced by all 4 of the
aspects: development, immediate causal,
fitness function, and evolutionary history
• Alternative hypotheses must be at the same
question level (i.e. PD vs PD, UF vs UF, etc)
– Otherwise they cannot be potentially mutually
exclusive
• Note: “develop” and “evolve” are completely
different processes, on different time scales.
Why do cats have an aversion to water?
1. Mother cats teach their kittens to avoid
water.
2. Aversion to water is a trait that evolved early
in the feline lineage.
3. When cats see water, neurons in the
amygdala are excited which leads the cat to
back away from the water source.
4. Cats are born with an aversion to water.
5. When water contacts the cat’s fur, a reflex
response occurs in which the cat draws back
from the water.
6. Avoiding water prevents cats from getting
their fur logged with water which can lead to
hypothermia.
 Practice:

• Potential alternative hypotheses?

• Can you think of an alternative hypotheses for

each?

• Can you think of a prediction for each

hypothesis?
 Proximate example:
Homing behavior by beewolf wasp

Beewolf wasps :
• Females dig burrows in which they place honey bees
that they have stung. Then they lay an egg on the
paralyzed bee.
• The larval wasp then uses the bee’s body as a food
source while developing.
 Tinbergen’s observations about
beewolf homing behavior:
• Beewolf wasps hide their nest entrance each
time they depart
• They fly around the nest area before leaving
• They go directly to the entrance when they
return
 PROXIMATE QUESTION
• What cues do beewolf wasps use to locate their
hidden nest entrance?

HYPOTHESIS
• They use visual landmarks for orientation

PREDICTIONS
• Removal of landmarks will prevent the wasps from
finding the nest entrance
• Moving the landmarks will cause the wasps to orient
to a different location
 Test of predictions
• While the wasps are away from burrow,
remove any conspicuous landmarks in the
vicinity and then observe the wasps’ behavior
when they returned.

Time to find nest

?

control landmarks
removed
 Test of predictions
• When visual cues were displaced, this misled
the wasps to land in the wrong spot
ALTERNATIVE HYPOTHESES?
 Ultimate example:
Egg shell removal by nesting birds

Observation: Many nesting birds will remove egg

shells from their nest after the chicks hatch
 ULTIMATE QUESTION
• Why do birds remove egg shells from their nests?
(fitness benefit?)

HYPOTHESIS
• They remove egg shells to reduce risk of predation
on the newly-hatched chicks.

PREDICTION
• Eggs near broken shells will be eaten more
frequently than eggs far from broken egg shells
Black-backed Gull at nest
Egg and chick predators:

Other gulls

Eggs
Crows
 Test of Prediction
Number of intact eggs 140
120
100
80
60 Eggs taken by
predator
40
20 Eggs not taken
by predator
0
15 cm 100 cm 200 cm
Distance from egg shell to intact egg

• The further away the egg shells are from the nest,
the fewer number of intact eggs taken by predators
ALTERNATIVE HYPOTHESES?
 Another example:
monogamy in prairie voles

Read in the book about both proximate and ultimate

explanations of their monogamous behavior.
http://www.youtube.com/watch?v=5ddlKQwDIe8
Connection between
evolutionary history
and mechanisms of
behavior
 Hypotheses vs Predictions
• Hypotheses
– Answer to the Tinbergen question posed
• Alternative hypotheses
– Must be answers to the same Tinbergen question
• Predictions:
– If the hypothesis is true, then you would expect….
– Predicted result from a study or experiment
 Evaluate this experimental design
Jane wants to test the following hypothesis:
• Decreasing temperatures in the fall cause
sparrows to migrate south.

Experiment:
• Capture a sparrow, put it in a cage and expose
it to cold temperatures in a controlled lab
setting. Observe whether it shows migratory
behavior or not.
 Practice:
• Can you think of potential problems with this
experimental design?
• How would you fix it to make it better?
 Uncertainty in science
• Experimental results support hypotheses, they
do not prove them.
– More support leads to more certainty
– Ruling out alternative explanations can also
increase certainty
• Scientific understanding changes over time, as
new evidence arises.