Animal Behavior Exam 2

Studies of Site specific expression of genes how is it done

use staining techniques to locate cells with a genes product or couple gene with another so that activation of the gene is revealed

Prairie vole example of site specific expression

vasopressin receptors on the ventral pallidum in monogamous prairie voles can be visualized and you see more vasopressin receptors in the lateral septum in polygynous montane voles

prairie voles and meadow voles

experiementally increasing the V1a receptors in meadow voles in the ventral pallidum leads to increased monogamous behavior (site specific expression)

ZENK gene in zebra finches

example of site specific expression, at 40 days they start learning their songs, in the forebrain, ZENK is expressed in birds who are exposed to conspecific song and not in those that arent (genes AND environment)

song learning regions in birds regions of the brain how did we discover and visualize

study of site specific expression, actually could see how the genes affect phenotype for each region

quantitative trait locus analysis (QTL)

• used to identify/locate possible genes involved in polygenic (quantitative) traits

• looking for an association of a “marker gene” on a particular chromosome with variation in a specific trait

• if such an association shows up, it suggests that a gene influencing that trait is closely linked with the marker gene

• you can use a variety of modern molecular techniques to try to identify the specific gene that is associating with the marker

QTL mapping example

QTL mapping for anxiety in mice

QTLs for anxiety behavior have been found on all nineteen autosomal chromosomes as well as X chromosomes in mice

genome wide association studies

used in large populations to identify multiple genetic variations (SNPs), sequence the whole genome, shows us genes that are correlated and which ones are involved in phenotype

GWAS Study of Dog Behaviors what did they study

looked at 268 whole genome sequences from 130 modern breeds and investigated dog herding, predation, temperament, and trainability

Genes associated with hearding GWAS dog study

found 44 sites on 5 chromosomes associated with herding behavior in dogs, allows us to see which genes are involved in polygenic traits

microarray analysis

allows us to analyze the activity of a large set of genes by detecting certain products (messenger RNAs) made when those genes have been “turned on”

developmental sequence of worker honeybee behavior

only live 4-5 weeks, foraging is most dangerous is this due to the activation of different genes?

honeybee microarray analysis results

gene activity (in worker brain) of forager gene correlates with behavioral activity

“DNA is both inherited and environmentally responsible”

even if you remove foragers or nurses bees will switch expression to fill that gap regardless of age

conclusions regarding behavioral genetics

• there is an abundance of evidence that there are genes for behavior (in the proper sense)

  • these maybe studied using a variety of methods, both classical and molecular (gene therapy, knockout, GWAS, and QTL)

• With molecular methods, we are beginning to understand not only how differences in alleles correlate with differences in traits but increasingly also how genes exert their developmental influences on behavior

behavioral genetics belongs to which of Tinbergens categories

development

historically, discussion of behavior development were usually about whether a behavior was…

instinctive or learned, these were considered by both comparative psychologists, and ethologists, to be two mutually exclusive types that all behaviors fell into

traditional dichotomies and what is the correct frame of mind

Instinctive vs. Learned, Genetic vs. Environmental, Nature vs. Nurture (from shakespeare “on whose nature nurture could never stick”)

modern consensus in instinct and learning

behaviors fall along a continuum from entirely learned (human languages) to entirely instinctive (cricket songs) with many in the middle (bird songs)

the first ethological textbook was

Tinbergens The Study of Instinct

Lorenz’s list (first 3) of the characteristics of instinctive behaviors (not learned and not modifiable)

part of early ethologists ideas about instinct

1. instinctive behaviors are genetically programmed

2. instinctive behaviors are not learned

3. instinctive behaviors are not modifiable as a result of learning or experience

Lorenz’s second 3 characteristics of instinctive behaviors

4. instinctive behaviors are fixed action patterns FAPs (highly stereotyped, dont vary between individuals or over time with and individual

5. Instinctive behaviors are released by simple, specific stimuli

6. animals have an inherent need to perform instinctive behaviors; motivation builds up over time since last performance (even vacuum behaviors may occur)

vacuum behavior and example

behaviors done without stimuli when built up and not done for awhile

beaver making dam from christmas presents an decorations

how to determine whether a behavior was instinctive or learned?

carry out a deprivation (Kaspar Hauser) experiment

1. rear animal without exposure to the relevant stimulus

2. if it shows normal behavior the behavior was instinctive, if it does not show normal behavior, the behavior was learned

flaw in deprivation experiement

if it doesn’t work, 1. have you identified the correct stimulus and 2. it could be due to trauma of the experience or something else rather than lack of learning you dont really know

Sonograms of eastern pheobe example

example of deprivation experiment, the sonograms of the controls and deafened birds were the same

Harry Harlows monkeys what did it tell us

was the animal so messed up from the manipulation that it couldnt show the correct behavior?

• didnt develop the ability to socialize with other monkeys

• the monkeys that grew up without mothers didnt mother their own children

• monkeys who even received 15 minutes of socialization a day were fine

Harry Harlows traumatized monkeys

• effects of deprivation, especially social deprivation can cause even instinctive behaviors to be abnormal

• so if deprivation results in normal behavior, we know the behavior was instictive, but if the deprivation results in abnormal behavior, we dont know whether that reflects a lack of learning or is the result of a general traumatic effect

Cross fostering experiments; and mice example

when you swap children of two similar species

swapped oldfield mice and deer mice babies, the cross fostered mice acted like their parents

how are time of burrowing and length of burrows related in oldfield and deer mice?

they are separate but linked traits with clear dominance and recessive aspects

An experiment that undermined right ideas about instinctive behaviors

Jack Hailmans study of pecking behavior by laughing gull chicks about how “instinct is learned”

Hailmans Study what did he do

paid close attention to laughing gull chicks behavior right after hatching (Tinbergen usually waited two days before observing behavior)

the instinctive behavior changed a lot over the first few days due to learning

Hailman Study results

through practice, chicks get better at hitting what they are aiming at

through trial and error learning, the chicks learn to restrict their pecking to the correct stimulus (maturation PLUS learning from practice)

used different versions of “gull” looking things and the less it looked like a gull on future days the less it pecked at it but day 1 it pecked pretty uniformly besides like the super deformed one

a modern view of instincts

instinctive behaviors are ones that appear in a functional form the first time they occur, without any required exposure beforehand to a specific environmental stimulus. No other assumptions or attributes necessary

mot mot study what is it an example of and what is it

example of instinctive behavior, yellow and red ringed things triggered young mot mots to not peck, but stripes in line with the log it pecked 2/3 amount of times and pecked the same when the pattern wasnt uniform across the whole stick (pecked less on yellow and red ring side than blank side)

Ultimate explanations for instinctive behaviors (why instinct over learning)

1. nervous system economy

2. short life spans

3. constant environments

4. fitness consequences of getting things right the first time

No Squirrel Can Fly

traditional beliefs about learning among comparative psychologists

1. learning is unconstrained

2. all learning is reversible

3. for learning to occur, the relevant stimuli must be very closely linked in time

4. learning usually requires repeated exposure

unicorns really can repeat

study that challenged beliefs about learning by comparative psychologists

John Garcias study of taste aversion learning in rats

John Garcias rat study results

• rats acquired an aversion even if they became nauseous an hour later

• Aversions were acquired with only one exposure

• aversions were not reversible

• rats could acquire aversions to tastes if associated with nausea but could not learn other associations

Garcias results regarding not being able to learn other associations rats

used a shuttle box for shock (pain)

taste + nausea and sound + shock but not the other combos

do all species learn aversions?

No! vampire bats dont

Vampire bat study

the vampire bat consumed the same amount of fluid regardless of if it was an immediate, delayed, or asychronous toxin, or saline. Insectivorous bats learned taste aversion to the immediate and delayed toxin.

Why? blood is blood, and some insects are poisonous

Is learning an adaptation? Who referred to this as “preparedness’? How does evolution play into this?

Rats could only learn natural connections and vampire bats dont learn aversions. Martin Seligman.

Evolution seems to have resulted in different species having different learning capabilities, related to their environmental challenges, rather than more or less of a “general ability.”

food caching in birds black capped chickadee and pinyon jay

example of learning as a adaptation, after storage spent a lot more time looking in hoard sites compared to before

pinyon jays differed by sex, females deteriorated but males dont get worse and males also payed attention to where their mate stored their seeds

sphexish behavior

animals can therefore look extremely intelligent in one context but act like stupid robots in another context names after digger wasp Sphex (when Fabre stole their bug that they placed in their nest and put it away in the same spot the digger wasp didnt know where to look no matter how many times it showed up in the same spot)

sphexish behavior beewolves example

tinbergen found out its physical landmarks that the beewolves use to identify their nests

modern definition of learning

durable modification of behavior in response to specific experience, no other assumptions or attributes are necessary

stephens learning model

when is learning worth it?

types of one stimulus learning

sensitization and habituation

early example of habitiuation

spider reacting less strongly to a tuning fork that touches her web after repeated exposure

Wolfgang Schleits Study of Classical Ethology

studied previously studied “instinctive fear response” to birds of prey stimuli in young birds (geese, ducks, turkeys) and thought it may be due to habituation not instict

he found rapid habituation occurs to all stimuli in the first couple of days and attributed tinbergens results to differential habituation of the chicks

Eric Kandel aplysia habituation

gill withdrawal reflex, studied at the cellular level, the more water jets that were shot at the aplysia the less it withdrew,

sensory neuron comes to release less NT at synapse with the motor neuron thus motor neuron fails to order gill withdrawal

sensitization in aplysia eric kandel

develops only later in life, interneurons are involved, when the head was poked that caused the interneuron to release more NT and when the siphon was poked the sensory neuron released more NT causing more vigorous gill withdrawal

habituation

waning of response with repeated presentation of a neutral stimulus

sensitization

accentuation of response due to prior presentation of a noxious stimulus (more likely to respond)

Association learning

the animal learns to respond appropriately when two stimuli are reliably associated. Often two major classes of association are recognized.

the simplest scheme of learning lists only two schemes

one stimulus learning and association learning

classical pavlovian conditioning

the animal learns tan association between two external stimuli

operant skinnerian conditioning

trial and error learning, instrumental learning, the animal learns an association between its actions and their consequences

Classical Conditioning Paradigm

1. Unconditioned stimulus (US) → Unconditioned respose (UR)

2. Conditioned Stimulus (CS) → Unconditioned Response (UR) MUST BE IN THAT ORDER

3. CS Alone → CR (CR need not be the same as the UR! Could be something else)

Betta siamese fighting fish

Brightly colored, present mirror think themselves are a rival and threaten it. Can condition them to respond to themselves in the mirror. Then shine a light and show the mirror. Then you just need the light and they will respond to assume a rival is coming. Some responded when light turned on and some would wait until the light went off, some went over to mirror and waited until after light and mirror shows up then responded.

CR just has to be

something new and relevant to the situation

Raushers study of pipevine swallowtail

• pipeline swallowtail caterpillars eat only pipline leaves and females only lay eggs on pipevine leaves

• Problem: different species of pipevine, with differently-shaped leaves, are out at different times of the year - how does female know what to look for?

• Solution: Land randomly on leaves use innate reaction to pipevine leaf chemicals to identify current pipevine leaf chemicals; learn its shape; carry out efficient visual search (learn tase shape association)

• Its inherently rewaring to get the buzz of chemicals from the correct leaf or is it learning the relationship between leaf and reward (classical conditioning)

Operant Conditioning associate with who

Skinner and Skinner box

Reinforcement (operant conditioning)

Increases performance of the behavior

• provide positive stimulus (positive reinforcement)

• remove aversive stimulus (negative reinforcement)

Punishment (operant conditioning)

decrease performance of a behavior

• administer aversive stimulus (positive punishment)

• remove positive stimulus (negative reinforcement)

Errors in Entomolgy textbook

1. confused classical and operant conditioning

2. Pavlov got nobel prize for physiology

3. he got he reinforcement terminology wrong

Lewis’s study of nectaring by cabbage whites (butterflies)

Lotus corniculatus and vicia cracca flowers (very different one yellow one purple) - operant conditioning

• have to learn how to work the flower if using different flowers

• First exposure to VC: took 12s to obtain nectar

• Second exposure to VC: 2s to obtain nectar

• First exposure to LC: took 12 s to obtain nectar

• Second exposure to LC: took only 2s to obtain nectar!

• Re-expose to VC: took 12s to obtain nectar

• reexpose to LC: took 12 s

Conclusions: cabbage whites quickly learn how to operate a flower but can only remember one flower at a time

Latent learning rats

rats which are hungry but not thirsty are given forced runs in a t maze leading to food in one arm and water in the other if subsequently tested when thirsty what do they do?

• according to skinner they wouldnt learn unless they got a reinforcement (so should be random)

• But ran to water → despite the lack of immediate reward, they did learn the location of the water in their first run thorugh the maze their leanring was only demostrated after a latency period thus latent learning

other perspectives on rat study besides latent learning rat study

• some consider this just operant conditioning with a delayed reward other say that acquiriing useful information about he environment can be rewarding in itself

• some list spatial learning separately

insight learning what is it and who

ability to use advanced cognitive processes to determine the correct solution withotu reandom trial and error (as in operant conditioning)

advocated by Wolfgang Kohler, founder of gestalt psychology who carried out famous studies in Tenerife with Sultan chimp

Wolfgang Kohler chimps

chimps could have flashes of insight

• release chimp into room and give them a problem and watch them try to solve the problem ex. suspend banana from ceiling, jump and try to reach banana then get really angry by situation but then suddenly start stacking boxes to get banana → insight

criticism of chimp study

• the chimps had already done problem solving tasks in that room so had developed other approaches for new problems (shortening time to solve problem)

• trial and error has to be physical (mental trial and error of how to get it) “learning set learning” or “learning how to learn”

• perhaps just operant conditioning with mental (virtual) trial and error

• very active area is “cognitive ethology” to try to get in the mind of animals

most behaviors involve…

both innate and learned aspects

new technology in birdsong research

portable reel to reel tape recorders (to “capture” sound), sound spectrographs (sonographs), (which reproduce sounds on paper as measurable sonograms, allowing quantitative study of animal vocalizations)

Peter Marler WM Thorpes student, became a leading figure in this area

Why did ethologists suspect that songs were learned from birds and not instinctive?

songs were complex and varied between individuals (unlike calls, which are simple, repetitive vocalizations)

Two male winter wrong sonograms, what must it involve?

have syllables in songs, very complex and very different while they have a lot of the same syllabes differ in how they put them together→ must involve learning

Marler and Peters study of Swamp and Song sparrow

Swamp sparrow is repetative buzz, song sparrow has a bit more complex

• did deprivation study to find innate song

  • repepetive buzz swamp and some whistles for song but still had skeletal elements of more complex full song

• If you played tape recording of opposite species song or of same species (heterospecific)

  • still sang innate song

• Live (heterospecific) bird they learn some of the other species song

• It is easier to learn conspecific song

Conclusion of white crowned sparrow and strawberry finch

they learned both strawberry finch and the white crowned sparrow when exposed to strawberry finch live tutor

sensitive period for song learning in birds long billed marsh wren graph

another innate aspect of song learning, 2-9 weeks for long billed wren, there are certain times where learning is more likely to take place than another times

why was the graph bimodal and not bell curve? the sample size effect

two more treatments marler and peters and what was each manipulation missing

• early deafening by cutting the auditory nerve before sensitive period; should sing innate song but didnt just squack, Missing:

  • Missing:cant hear themselves still (which would be enough to get innate song)

• late deafening by cutting auditory nerve after senseitive period but before bird starts singing. outcome: permanent subsong (babbling); normal birds trotted out a lot of syllables until final order and lower syllables “crystallized”

  • Missing: cant hear what theyre trying to do so cant get feedback from themselves

auditory template model

• period during when birds are susceptible to other birds songs “template stage” and specifically look for songs that match their crude template(days 10-50) called the memorization phase

• day 150 they start singing their song called “plastic song” which starts up do to testosterone and then full song around day 200 called the motor phase

exceptions to auditory template model

• some “open learners” can add new songs during a new sensitive period each year

• species wiht the largest repertoires add songs continuously; many are mimics such as mockingbirds, mynahs

• at the other extreme, some birds have totally instinctive songs

Brood parasites what do they do, and what speiceis

• they deposit their young in others nests, host can be intra- (same species) or inter (different) specific

• mostly birds and social insects

• Cuckoos (Eurasia), Whydahs or widowbirds, and cowbirds

European cuckoos what is it an example of and what do they do?

brood parasite; parasitize numerous species of hosts

• Female cuckoo will spy on nests and pick up host egg and replace it with her egg (females can only lay 1 egg a day)

• They make different eggs that look like the host eggs

• Chicks hatch and the cuckoo will kick out the competition

how do cuckoos develop song?

completely instinctive since they grow up in other birds nests and males go back to africa after mating

whydahs what are they and what do they do

brood parasites; usually parasitize close relatives

• use different host races (species?) because have to mate with another bird of same type

• have to match the host species in looks as well as the egg and their gate

• Have evolved different looks based on the host species that they infiltrate

how do whydahs differ from cuckoos

dont expel host chicks and have to compete and mimic host chicks gape, their songs are totally learned and not instictive

how do whydahs learn song?

learn the hosts song and are totally learned for the males to sing it and females to prefer it

Indigobird (whydah) study new song

when given new input of bengalee finch to see if they would still learn the song and they did

can all birds distiguish between their eggs and parasite eggs or just hosts? how were cuckoo host species determined?

a mismatched egg will get thrown out

species can distinguish between eggs whether or not their are parasitized or not (hosts and non hosts did it) → there are other species that used to be hosts that outcompeted the cuckoo

how do cuckoos mate and whats the pattern?

• davies and collegues origionally thought it was based on natal habitat but DNA fingerprinting showed that females mate with males randomly with respect to habitat or host

• solution is unusual ZZ/ZW sex determination system (females are hetergametic)

• egg trait genes are on the W chromosome - males dont have this and are therefore irrelevant

brown-headed cowbirds: name origin, what type of species

• name comes from habit fo following cows (bison) to catch flushed insects

• a classic edge species (thrive w people and are in two diff habitats)

• very generalist parasite lacking egg or chick mimicry

what does deprivation experiment of brown headed cowbirds show?

effects of male dominance: isolated birds sing “potent songs” , but socially reared males sing less potent songs because the older, more dominant males attack them and they learn not to sing potent songs until they are older and become dominant themselves

so birds reared alone sing potent songs

audience feedback in cowbirds (California and Indiana cowbirds)

california and indiana cowbird populations have different innate songs. Indana females are unresponsive to potent california male songs. But after several weeks of being housed in the same room, in visual and acoustic contact, Indianan femles show copulation sociciataion postures to california males. California males learned Indiana songs by operant conditioning by response to females.

cowbirds instictive or learned song?

Both!! Can change based on interaction with other males or with females.

cuckoo rubenstein example eggs

foreign eggs were more likely to be rejected the more they differed in color from the hosts eggs but as foreign eggs increased in number, host required greater difference to reject them

imprinting definition

early learning by an animal of its parents and species characteristics, associated with Konrad Lorenz

Konrad Lorenz why did he do imprinting, his students, and what are the 2 forms

as a means to an end to be able to study them closely (thought it was a permanent thing), his student studied it as a unique form of learning in its own right

• filial (learn and attach to parents) and sexual (learn characteristics of the opposite sex for later sexual behavior)

Characteristics of imprinting

• occurs early in life

• requires a somewhat specific range of stimuli to be effective

• occurs only during a critical or sensitive period

• is sometimes not reversible

• an active not passive process

Early Stimuli (specific stimuli) Create (critial period) Active (active process) Impressions (is sometimes reversible)