Transcript Learned behaviors

In This Lesson:
Unit 2
Animal Behavior
(Lesson 1 of 3)
Today is Tuesday,
nd
September 22 , 2015
Pre-Class:
What are those ants up there doing?
I know it’s a little hard to see, but really, why would ants
pile onto each other like that…especially with nothing
supporting them?
Also, get a paper towel for you/your partner.
Today’s Agenda
• Biostatistics: Chi-Squared Analysis
• Behavioral manifestations of evolution.
• Where is this in my book?
– Chapter 51.
By the end of the lesson…
• You should be able to calculate chi-squared
values (and determine significance) for a set of
data.
• You should be able to distinguish between
proximate and ultimate perspectives when
analyzing behavior.
• You should be able to separate innate behaviors
from learned behaviors, and taxis from kinesis.
• You should be able to describe social behaviors,
including examples.
Quick Heads-Up
• Just a quick note to let you know that we’ll be
interrupting these notes to do our Animal
Behavior lab (Investigation 12) and to learn
chi-squared data analyses.
• Find these online for more info:
– Fact Sheet – Unit 2 – Chi-Squared Tutorial
– Fact Sheet – Unit 2 – Chi-Squared Table
Before we start…
• Challenge questions!
The first thing you need to know…
• Let’s be honest: You’ll never know exactly
what it’s like to be your dog, for example.
• So, when people talk for their dog, or even
just feel a need to ascribe human
emotions/thoughts to them, they’re
anthropomorphizing.
• Anthropomorphosis is the ascription of
human behavior to animals. It’s wrong. Don’t
do it.
Think this dolphin is smiling?
http://www.marineland.net/images/DolphinHeader.jpg
Think this gorilla is sad?
http://upload.wikimedia.org/wikipedia/commons/b/b1/Eyes_of_gorilla.jpg
Think this bluebird is angry?
http://www.songbirdgarden.com/store/ProdImages/EK6600-1.jpg
Then you’re anthropomorphizing!
Not Anthropomorphism
• How Can You Tell if Your Goat is Happy? article
The second thing you need to know…
• Just like Mendelian geneticists studies the
phenotypes that emerge from a certain
genotype, behaviorists study the behavioral
phenotypes that emerge from those same
genotypes.
– Just like other traits, behaviors that improve
successful reproduction chances will also be
selected for.
• That ended in a preposition and I have no idea how to
fix it.
The Basic Behavior
• Before we even define behavior, let’s take a quick
look at the most basic behavior out there:
movement.
• Movement is often a simple response to a stimulus.
We call it either taxis or kinesis.
– Taxis is a directional response, like how a trout will turn
itself so that it’s facing upstream to catch food as it drifts
downstream.
– Kinesis, on the other hand, is non-directional movement.
• It’s still in response to a stimulus, just not directed.
• Woodlice, for a common example, move less when humidity goes
up. Non-directional.
Taxis or Kinesis?
• A photosynthetic microorganism swims toward
the light side of its petri dish.
– Taxis (phototaxis)
• Ants exit their colony and wander in search of
food.
– Kinesis
• Once food is found, other ants follow the trail of
pheromones to the first ant.
– Taxis (chemotaxis)
• Pillbugs under a recently overturned log run for
darkness.
– Taxis (phototaxis)
With that in mind…
• Behavior is quite simply anything an animal does in
response to a stimulus from its environment.
– A stimulus is simply a “cue,” which could be internal or
external. (plural = stimuli)
• An internal stimulus might be hunger.
• An external stimulus might be a baseball thrown at your face.
• These behaviors can be broken down into two general
categories:
– Innate behaviors are those with which an animal is born.
They are automatic and do not depend on experience.
– Learned behaviors emerge during an animal’s lifetime. They
are variable (they change) and depend on the present
stimuli of the environment.
Innate vs. Learned
• Mother birds will feed,
almost uncontrollably, the
gaping beak of a nestling
bird.
– They just do it…
– …even if the image is
printed on a piece of paper.
• However, a learned
behavior is something that
can often be trained.
– Dog house-breaking, for
example.
– More on this later.
http://britishwildlifehelpline.com/Chicks.JPG
Nature vs. Nurture
• All of this hints at the age-old debate of nature vs.
nurture.
– Is it something genetic, or is it learned?
– Homosexuality is, unfortunately, often brought up in this
context.
• To that end:
– Nurture: Male migratory birds kept in an outdoor aviary in
view of their migrating (free) male counterparts will begin
to exhibit homosexual behaviors like allopreening
(basically cuddling and grooming).
– Nature: Removing a particular gene in male fruit flies
causes them to attempt to mate with other males.
You observe a singing songbird:
• What can you ask about the
behaviors observed? You might ask…
– How does the bird sing?
– What prompts the singing?
– Why does the bird sing?
• Ask Maya Angelou that one.
– Numbers 1-2 are proximate questions.
They concern the present or near past.
– Number 3 is an ultimate question. It
concerns the evolutionary history of the
bird.
Proximate vs. Ultimate
• Proximate or Ultimate?
– How does photoperiod (day
length) affect breeding in cranes?
• Proximate. It’s a “small picture” sort
of question.
– Why do cranes perform a courtship
dance?
• Ultimate. It’s asking about something
that must have taken generations to
evolve.
– Why do cranes breed in spring?
• Ultimate. It’s the “big picture”
evolutionary question.
Expanding the debate…
• Niko Tinbergen proposed the following “four
questions” of animal behavior.
– Let’s use singing songbirds as an example again:
Dynamic View
Ontogeny
How did the bird’s anatomy
Proximate View
change as it grew to allow
song?
Ultimate View
Phylogeny
How did the bird’s
evolutionary ancestors
sing?
Static View
Mechanism
How does a bird’s
anatomy allow it to
sing?
Adaptation
How does song allow a
bird to survive and
reproduce?
Four Questions of Animal Behavior
Case in Point
• BBC – Vogelkop Bowerbird
The Study of Behavior
• Let’s take a look at the history of ethology, which
is a fancy word for the study of behavior, in
biology.
– At some point we needed to get to the typical cast of
old white guys in biology.
• The Big Names:
–
–
–
–
B.F. Skinner
Ivan Pavlov
Karl von Frisch
Konrad Lorenz
B.F. Skinner: Operant Conditioning
• Creator of the Skinner box.
– Sounds violent.
• A mouse learns that when it
pushes a lever, it gets a reward –
positive reinforcement.
– Learned behavior, obviously.
• If the mouse doesn’t push the
lever, it may receive an electric
shock.
• So operant conditioning is training
to associate a voluntary behavior
with a certain response.
– Operant conditioning has a large
share of trial-and-error learning as
the mouse must learn about the
lever’s function by simply trying it.
Operant Conditioning Words
• Reinforcement: A reward.
– Positive Reinforcement: Giving you something good.
• Example: A cookie!
– Negative Reinforcement: Taking away something bad.
• Example: If I play a screeching loud sound until you do
something I want.
• Punishment: A change that decreases the likelihood
of a certain behavior continuing.
• Example: An electric shock.
Operant Conditioning Demo
• I needs me a volunteer.
• This volunteer will have to exit the room briefly.
• When s/he returns, s/he will have to guess the
desired behavior determined by the class by
doing different things in order to get rewards.
• This is often termed “clicker training” because
trainers will make a click sound when the desired
behavior is performed, thus edging into a
different kind of conditioning.
Ivan Pavlov: Classical Conditioning
• Give a dog some food and it’ll salivate
automatically.
– An innate, unconditioned behavior.
• Ring a bell near a dog and it probably
won’t care for more than a second or
two.
• Ring a bell and give a dog food
enough times, however, and the dog
begins to associate the bell with food.
• Suddenly, you can simply ring a bell
and get a dog to salivate.
– That’s a learned, conditioned response.
• So, classical conditioning is pairing an
innate behavior with a neutral
stimulus.
Operant vs. Classical Conditioning
• TED: Peggy Andover – The Difference Between
Operant and Classical Conditioning
Learning
• Another principle in learning is
habituation/sensitization.
• Habituation is when the
response to a stimulus
decreases with exposure to that
stimulus.
– “The Boy Who Cried Wolf”
phenomenon, minus the actual
wolf at the end.
http://www.kcet.org/news/rewire/assets_c/2013/10/owl-pigeon-10-25-13-thumb-600x600-62607.jpg
Learning
• Sensitization, on the other hand, is the opposite.
– Suppose every 30 seconds you don’t do a behavior I
want, I turn on a red light, then deliver a small electric
shock.
– Soon, the sight of a red light may be such a powerful
stimulus that it almost seems to cause pain anyway!
• Sensitization is an increased response to a
stimulus.
Learning
• One last example of learned behavior: tool
use.
• Using tools is NOT an innate behavior.
Individuals need to learn to use them by trialand-error or from other individuals.
• Tool use was originally considered a primateonly thing, until…
– New Caledonian Crow video!
Just kidding!
• Here’s the real last example:
– The Big Bang Theory
Conditioning
• What do the works of Pavlov and Skinner have in common?
– They both concern learned behaviors.
– Collectively, they form what’s known as associative learning – the
pairing of stimuli with actions.
• Lorenz and von Frisch spent their time more with innate
behaviors.
– Lorenz: Birds.
– Von Frisch: Bees.
• Before I tell you a little story about the birds and the bees, I
need to tell you about the fixed action pattern.
• And before that I need to tell you about the chi-squared test.
And other stuff.
Fixed Action Pattern
• The Fixed Action Pattern (FAP) is like the holy
grail of innate behaviors.
• FAPs are so deeply rooted that they are
automatic behaviors that must be completed
once started.
• Case in Point: Niko Tinbergen’s Sticklebacks.
Stickleback FAPs
• Male sticklebacks are highly
territorial.
• Introduce a sign stimulus into their
territory and they will react
predictably.
– A sign stimulus is a simplified version
of a more complicated stimulus.
• Putting a slender oval model, made
of wood, that has a red “half” is the
only stimulus needed to
communicate an intruding male to
the stickleback.
– The stickleback will attack the model.
Stickleback FAPs
• If, instead, that sign stimulus is
slightly…uh…distended, and has
a greenish belly, it presents to
the stickleback as a female.
– The stickleback will court the
model.
– A lot like what we mentioned
before – momma birds instinctively
feeding the shape of a gaped
nestling beak.
Other Fixed Action Patterns
• Egg Rolling
– Greylag Geese
– Move an egg out of Mother Goose’s nest and
she’ll instinctively roll it back.
– Remove the egg while she’s rolling it and she’ll
keep making the rolling motion, even without the
egg.
• After she completes the FAP, she’ll look for the egg
again, but she can’t stop the behavior once started.
– Video!
Other Fixed Action Patterns:
Humans
• Yawning:
– Once started, a yawn cannot be stopped. Further,
reading the word yawn, hearing a yawn, or seeing
someone yawn may induce a yawn FAP.
• Eyebrow Flashing:
– Upon seeing someone familiar, humans often
quickly raise and lower their eyebrows. It’s an
involuntary reaction to someone familiar.
For more on fixed action patterns,
let’s turn to…
• …South Park!
The Supernormal Stimulus
• A supernormal stimulus is one that is
exaggerated beyond natural occurrence.
• Examples:
– Mother birds that feed models with the reddest
and widest beaks.
– Lipstick in humans being even more attractive
than realistically red lips (?).
– Birds that prefer to incubate the largest and most
gray-speckled eggs (Tinbergen again).
http://obsidianwings.blogs.com/.a/6a00d834515c2369e2017eea7923fc970d-pi
Soler, M., Martinez, J. G., Soler, J. J., & Møller, A. P. (1995). Preferential allocation of food by magpies Pica pica to great spotted cuckoo Clamator
glandarius chicks. Behavioral Ecology and Sociobiology, 37(1), 7-13.
Konrad Lorenz
• Lorenz noticed that baby birds
frequently would imprint upon
whatever moving object they saw first,
animate or not.
• Imprinting is when juvenile organisms
come to see another…thing…as a
parent.
– In the wild, typically they first saw their
mother in the nest.
– Lorenz demonstrated that baby birds
might follow humans or even inanimate
objects if they saw them first instead.
– Imprinting is also irreversible.
Imprinting’s Value
• Today, biologists often use
imprinting as a means of
conservation.
• Cranes, for example (why
do they keep coming up?),
have been imprinted on
pilots wearing crane
costumes and taught
migratory routes that
would keep them out of
danger.
Imprinting in Film
• Even the 1996 movie Fly
Away Home, starring a very
young Anna Paquin, depicted
the same thing!
– Don’t ask if we can watch it.
http://www.teachwithmovies.org/guides/fly-away-home.html
Critical Period
• Imprinting is also a great example of a behavior that is
learned, yet also time sensitive.
– Imprinting only occurs in the first few moments of a nestling’s
(visual) life.
• Therefore, imprinting only occurs during a critical period,
before or after which no imprinting can happen.
– In humans? Language learning. In birds? Songs.
Critical Period: Case in Point
• The cuckoo is a brood parasite.
– Mama Cuckoo lays her eggs in other birds’ nests to let them take
care of the young.
– Because of the surrogate mother bird’s fixed action pattern, she
doesn’t know the egg isn’t hers. She’ll incubate it just the same.
• Cuckoos, once hatched, don’t really have a “song.” They learn
the song of their surrogate mothers!
– Weird Fact: Cuckoos also lay eggs that nearly precisely match the
surrogate’s eggs, and parasitize the surrogate breed’s nests only.
• There are, in a way, different “versions” of female cuckoos.
– Tying it all together: Because cuckoos are actually taking
advantage of another bird’s FAP, they are engaged in what’s
known as code-breaking.
Cuckoo Photos
http://upload.wikimedia.org/wikipedia/commons/5/5c/Reed_warbler_cuckoo.jpg
http://whyevolutionistrue.files.wordpress.com/2011/03/f2_hauber_egg.jpg
Cuckoo Photo
• All your nest are belong to us:
http://www.pbs.org/wgbh/nova/sciencenow/0407/images/03-moth-06.jpg
One last thing on cuckoos…
• Cuckoos also provide us an opportunity to discuss the
concept of cross-fostering.
– This is an elegant method of determining whether something
is genetically or behaviorally driven.
• Shortly after birth (or as eggs), switch the young
around and have them raised by different parents.
– Cuckoos automatically do this, remember?
• So a cuckoo’s appearance, which is consistent with all
cuckoos, is genetic (“nature”).
• But, a cuckoo’s song is learned (“nurture”) because it
varies based on its environment.
Classic Experiment: Cross Fostering
• Rat pups that are licked by their mothers grow up to
be calm adults.
• Rat pups that are not licked grow up to be anxious
adults.
– Is it that mother rats with poor social skills passed on their
poor social skills to their high-anxiety pups?
– Alternatively, is the licking behavior the sole determinant
of rat “well-adjustedness?”
• Cross-foster!
– Sure enough, if you take rats born to a mother that does
not lick her young and have them raised by a mother that
does, they grow up to be calm just the same.
• Rat lickin’ video!
Back to innate behaviors…
• Unlike birds, a lot of bee behaviors are innate.
• Insect song (like crickets) is often completely
fixed.
– You can cross-foster all you want. Genetics
determines their patterns.
• Additionally, many innate behaviors are also
social behaviors, and this is where Karl von
Frisch comes in.
– Social behaviors are those that are exhibited
between individuals.
Social Behavior Summary Slide
• Communication and Language
• Agonistic Behavior
• Dominance Hierarchies
– This is the one your teacher studied in college.
• Represent!
• Cooperation
• Altruism
Communication and Language
• Bees (as discovered by Von
Frisch) do a “waggle dance” to
communicate food locations.
– Waggle dance video!
• Whales communicate via lowfrequency sound across ocean
basins.
• Communications are sometimes
learned (humans, birds),
sometimes innate (bugs).
– Case in point: French crows and
American crows have different
alarm calls.
• FYI, that photo is a blackbird.
Communication and Language
• There are other forms of communication too,
besides the auditory.
– Visual (fireflies, for example)
– Chemical (pheromones – like hormones released
outside the body)
Pheromone Examples
• Female mosquitoes use CO2
concentrations to locate victims by
their exhalations.
• Spiders sometimes use moth sex
pheromones (allomones, since
they’re from a different species) to
lure prey.
• Big cat territory marking.
Agonistic Behavior
• Agonistic behaviors are any fighting-related behaviors,
whether it’s aggression or submission.
– Think males head-butting each other for a mate.
• That’s how I met my wife.
– Typically, agonistic behaviors don’t result in long-term harm
(but they can).
– Typically, agonistic behaviors occur among males (but they
can be among females too).
• Camry vs. Ram video!
Agonistic Behavior Example
• Here’s a gorilla doing the classic
chest-beating agonistic behavior.
• Many times, animals will simply
intimidate one another and not
actually fight.
• For another
example…TRANSFORMER OWL!
(Southern White-Faced Owl)
• For yet another example…BUCK
FIGHT!
http://hoothollow.com/Image%20Super%20Folder/Images%20%20Gorilla%20and%20Rwanda%20Superfolder/2010%20Joe/Mountain_Gorilla_5x3.jpg
Aside: Screwworms
• Here’s a howler monkey.
• Here’s a parasitic fly that comes
from a maggot called a screwworm.
• They live in some of the same areas
(South and Central America).
• Screwworms cause nasty
infestations (and maybe death).
• These infestations start with an
open cut (as small as a tick bite),
into which the adult fly lays its eggs.
• The growing larvae then consume
the flesh of the host animal.
http://www.fao.org/docrep/003/t0756e/T0756E113.jpg
http://entnemdept.ufl.edu/creatures/livestock/secondary_screwworm01.jpg
http://animalwonder.com/wp-content/uploads/2014/09/HowlerMonkeyCostaRica.jpeg
Aside: Screwworms
• Back to howler monkeys.
• Since open cuts are dangerous, there has long been a
selective pressure against aggression in monkey
populations.
– Because a physical conflict might result in a cut that could
cause an infestation…
• So, it turns out howler monkeys engage in a whole lot
of nonviolent agonistic behavior and settle differences
without beating the monkey poo out of each other.
• It also turns out that parasites drive a lot of evolution.
Dominance Hierarchies
• Dominance hierarchies are
social rankings.
– Also called a “pecking order” (for
quite literal reasons).
– Typically among males.
– Most dominant males often
have the best access to
food/mates.
• Why am I painting the bird’s beak
in that picture?
• We set out to determine whether
males respond to supernormal
stimuli, but instead found stable
dominance hierarchies.
• Zebra finch video!
Cooperation
• Like the word suggests, cooperation is working
together for a common goal.
• Importantly, all cooperating individuals will
share in the benefit, so each has a personal
interest as well.
http://www.factzoo.com/sites/all/img/mammals/canids/wild-dogs-hunting.jpg
Altruism
• Okay, this is a weird one. Let’s slow down a bit.
• Altruistic behaviors are those in which the
“performer” of the behavior incurs a negative
impact on fitness, while the “receiver” of the
behavior gains fitness.
– WTF?
• Altruism is at first quite a dilemma for
evolutionary biologists as there’s no reason for
this to exist.
– Why would anyone lower their own fitness?
Altruism
• Let’s add a couple more examples:
– Vampire bat allofeeding.
• Not all vampire bats successfully enjoy a blood meal each
night.
• Those that did feed will sometimes regurgitate some of
their meal and feed a neighbor in their group.
– Ground squirrel guard duty.
• Some individuals act as lookouts.
• Doing so puts them at additional risk, and making an alarm
call further calls predatorial attention to toward them.
http://images.nationalgeographic.com/wpf/media-live/photos/000/005/cache/common-vampire-bat_505_600x450.jpg
Roberts, G. (1998). Competitive altruism: from reciprocity to the handicap principle. Proceedings of the Royal Society of London. Series B:
Biological Sciences, 265(1394), 427-431.
Sherman, P. W. (1985). Alarm calls of Belding's ground squirrels to aerial predators: nepotism or self-preservation?. Behavioral Ecology and
Sociobiology, 17(4), 313-323.
So…why do they do it?
• It’s called kin selection.
• Although the altruism “giver” slightly lowers
his/her chances of survival, the animals that
behave altruistically usually live in groups of
closely-related individuals.
• In other words, they’re still helping to ensure
the survival and passing-along of their genes,
even if they’re slightly different.
Aside: Altruism and Morals?
• What if there’s something else at play here?
• Recent research is indicating there may be
something “higher” going on.
– Altruism – NOVA
Aside: Mirror Self-Recognition
• This really doesn’t have much to do with
what we’re discussing but it’s so interesting
I want to talk about it anyway.
• It turns out that the only animals
conclusively shown to recognize themselves
in a mirror are primates, elephants, and
dolphins.
– In the case of elephants, this was proven by
marking the elephant in a spot s/he could not
see him/herself, but could see in a mirror.
– Upon confrontation with a giant expensive
elephant-proof mirror, elephants touched the
mark with their tongue, something they
wouldn’t do if they thought their reflection
was someone else.
http://www.pnas.org/content/suppl/2001/05/02/101086398.DC1/0863Movie2still.gif
http://scienceblogs.com/mixingmemory/wp-content/blogs.dir/455/files/2012/04/i-6b245d33867c2aac87febde729b32c24-mirrorelephant.JPG
Last Thing
• This is really hard to place in sequence with the
other notes, but some behaviors serve to help the
animal maintain homeostasis.
• An example you should know:
– Endothermic animals (mammals, birds) can regulate
their own body temperature – hence 98.6 °F being the
average human body temperature, for example.
– Ectothermic animals (amphibians, reptiles, fish) cannot
regulate their own body temperature.
• In response, these animals must move to warm areas when
they’re cold or cold areas when they’re hot.
• Key: Ectotherms’ respiration rate slows with decreasing
temperatures.
Aside: Endothermic vs. Ecothermic vs…?
Endothermic
Homeothermic
(constant body
temperature)
Poikilothermic
(seasonal or
environmental
temperature
adjustment)
http://minerva.union.edu/linthicw/endo.htm
Ectothermic
(produce their own heat)
(heat gained through
environment)
Birds and mammals
that can regulate
their own
temperature.
Tropical reptiles,
deep ocean
creatures.
Some birds and
mammals, many
insects.
Most fish and
amphibians.
Aside: Endothermic vs. Ecothermic vs…?
http://minerva.union.edu/linthicw/endo.htm
Chi-Squared Analysis
• Now that you know all these cool behavior-related
things, you’ll also need to know whether you’re
actually seeing a change in behavior.
• This is when we will explore the idea of a statistical
test that will quantify whether a result is significant.
Remember that?
– This is not just an animal behavior thing.
– This test can be extended to many different studies to
determine whether you’re actually getting a real change or
if just chance is at work.
– Here’s where I switch PowerPoints to “Chi-Squared
Tutorial,” found in the “Fact Sheets” section of the website.
Closure
• If I train my dog by putting up an invisible fence (one
that delivers a small electric shock when the dog passes
a certain point), what kind of training am I using?
– Operant conditioning.
• If, instead, I train the dog by offering rewards
periodically whenever the dog remains within the yard,
what kind of training am I using?
– Still operant conditioning.
– Classical conditioning would have me pairing another
stimulus to a natural one, like if I made a click sound prior to
giving the food.