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Behavior Development b. Learning. Learning is a change in an animal’s behavior linked to a particular experience it has Brain properties change by gene and environmental interactions . Learning. Forms of learning Imprinting Specialized learning Variation in learning behavior

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  • Learning is a change in an animal’s behavior linked to a particular experience it has
  • Brain properties change by gene and environmental interactions
  • Forms of learning
    • Imprinting
    • Specialized learning
  • Variation in learning behavior
    • Environmental influences
  • Why would this behavior be adaptive?
  • Occurs when a young animal’s early social interactions lead to learning
  • Functions
    • Recognition of parents in animals with preccocial young (ex geese)
    • Recognition of an appropriate sexual partner
  • Dependence of recognition on recognition
  • Young animal must see model to recognize it
  • Learning is flexible- model does not need to look like parent or even same species
konrad lorenz
Konrad Lorenz
  • Imprinting in Greylag Geese
  • Imprint on humans
  • Later preferred Humans as mates
great tits blue tits
Great Tits & Blue Tits

Imprinting has different effects among species

imprinting has different effects among species
Imprinting has Different Effects Among Species
  • When cross imprinted
  • Some individuals became imprinted on opposite species
  • None of the Great Tits mated with a member of it’s own species
    • Formed a mate preference based on imprinted foster parent
  • Most of Blue Tits mated within it’s own species
    • Exhibited a different developmental interaction
specialized learning
Specialized Learning
  • Memory formation to aid in finding food
clark s nutcracker1
Clark’s Nutcracker
  • Whack cones of Whitebark pine
    • Also eats limber pine seeds
  • Seed pouch under tongue
  • Distribute seeds in Caches
    • Cache more than they will retrieve
      • Hides up to 38,000 of seeds per season
      • Up to 5,000 separate caches
      • Up to 20 miles away
    • Buries seeds in fall downslope
    • In winter will retrieve
clark s nutcracker3
Clark’s Nutcracker
  • Nutcrackers able to relocate caches with up to 80% accuracy
  • Recall memory for months
  • Use landmarks to relocate caches
seeds germinate in clusters
Seeds germinate in clusters

Un-retrieved caches create new stands

learning varies among members of the same species
Learning varies among members of the same species
  • Chickadees in Alaska require fewer inspections to locate food stores that the same species in Colorado
  • What causes variation among individuals?
environmental differences
Environmental Differences
  • Individuals can learn based on their interactions as young
  • Interactions with siblings can shape behavior
  • Kin recognition
    • Used to identify closely related individuals from potential rivals
  • Cues such as olfaction & sight can be used as recognition cues
polistes paper wasps
Polistes Paper Wasps
  • Paper wasps use both olfactory cues to recognize individuals from the same nest
  • Females are also able to recognize facial markings
    • Individuals with altered face markings were attacked more frequently
belding s ground squirrels2
Belding’s Ground Squirrels
  • Lives in subalpine and alpine communities
    • Meadows
  • Social ground squirrel
    • Females remain, males disperse
    • Closely related females help raise and protect each others offspring
  • Prey species lifestyle
    • Aerial predators such as hawks
    • Ground predators such as weasels
    • Colonial living aids in protection
kin recognition
Kin Recognition
  • Kin recognition critical
    • Helps identify closely related for assistance
    • Ability to recognize intruders
    • Prevents inbreeding
  • Strong selection pressure favoring genes that code for recognition behaviors
kin recognition1
Kin Recognition
  • Signals used to identify kin include
  • Scent
  • Appearance
ground squirrel musical chairs
Ground Squirrel Musical Chairs
  • Newborn Ground squirrels were moved from their nests into 4 groups
  • Siblings reared apart
  • Siblings reared together
  • Non-siblings raised apart
  • Non-siblings raised together
belding s ground squirrel torture
Belding’s Ground Squirrel Torture!
  • After raised in their respective groups ground squirrels were placed in an arena to test their ability to recognize each other
  • Recognition was measure using levels of aggression
  • Aggression indicates less kin recognition
what would you predict
What would you predict?
  • Siblings reared apart
  • Siblings reared together
  • Non-siblings raised apart
  • Non-siblings raised together
individuals learn based on their olfactory interactions as young
Individuals learn based on their olfactory interactions as young
  • Ground squirrels raised together learned each others smell and were less aggressive towards each other
    • Independent of whether they were siblings
  • Ground squirrels raised separately tended to be more aggressive toward each other
have we met
Have we met?
  • Biological sisters raised apart had fewer aggressive interactions than nonsiblings raised apart
  • Indicates siblings have a secondary learned behavior for recognizing kin
armpit effect
Armpit Effect
  • Animals have ability to recognize relatives they have never met before
  • Individuals can learn their own olfactory profile
  • Self recognition provides a reference to compare other individual’s smells to
    • Individuals who smell similar are more closely related, whereas individuals that smell less similar are less closely related
chemical communication
Chemical Communication
  • Oral and dorsal glands
  • Nasal investigation
  • Scent mark behavior
  • 5 odors that are individually distinct
scent discrimination
Scent Discrimination
  • Individuals learn to recognize their own scent
  • Spend more time sniffing less closely related relatives
ultimate causation
Ultimate Causation
  • Kin Recognition by using scent discrimination allows Belding’s ground squirrels in order to
    • Helps identify closely related for assistance
    • Ability to recognize intruders
    • Prevents inbreeding
genetic differences
Genetic Differences
  • Some behavioral phenotypes can be determined by a genetic component
  • Alleles that code for behavioral differences can be selected for or against by natural selection to maximize fitness
western terrestrial garter snake thamnophis elegans1
Western Terrestrial Garter SnakeThamnophis elegans
  • Highly variable in habitat
    • Can be found near water or away from water
  • Feeds on a wide array of food sources
    • Slugs, worms, leeches, tadpoles, frogs, fish, insects, lizards, small birds
western terrestrial garter snake populations
Western Terrestrial Garter Snake Populations
  • Coastal and inland snake populations exhibit variation in their diet preference
  • Inland population
    • Lives in arid habitats near lakes and streams
    • Feed primarily on fish and frogs
  • Coastal populations
    • Lives in moist coast ranges
    • Feed primarily on banana slugs

Is variation in prey preference genetic?

garter snake prey preference experiment
Garter Snake Prey Preference Experiment
  • Pregnant snakes from both coastal and inland populations were brought into lab
  • Immediately after birth offspring were isolated
    • Controlling for environmental learning influences
  • After several days each snake offspring was offered a segment of Banana slug
garter snake populations exhibit prey preference
Garter Snake populations exhibit Prey Preference
  • Feeding score indicates how many days a snake ate the offered slug
  • A score of 10 indicates slug was eaten every offering
  • Coastal slugs exhibited greater likelihood of eating slug than inland garter
the tadpole vs slug taste test
The Tadpole vs Slug Taste Test
  • Newborn garter snakes are offered cotton swabs with different prey “juice”
  • Snakes were offered swab for 1 minute
  • Preference was measured by number of tongue flicks


tongue flicking in snakes
Tongue Flicking in Snakes
  • Tongue flicking is a sensory-gathering behavior
    • Olfaction
  • Tongue flicks are used during prey trailing, foraging, mate searching
  • Chemical molecules gathered by the tongue are delivered to the Vomeronasal organ on roof of mouth
tongue flicking in snakes1
Tongue Flicking in Snakes
  • Ability to triangulate chemical cues in the environment is maximized by:
    • Waving tongue in the air to detect vertical gradients
    • Forked tongue increases sensitivity to chemical gradients in the environment
    • Tongue flicking rate

vomeronasal organ
Vomeronasal Organ
  • AKA Jacobsen’s organ

the tadpole vs slug taste test1
The Tadpole vs Slug Taste Test
  • There was no difference in preference to toad tadpole samples
    • i.e. Both populations had about the same number of tongue flicks to the tadpole covered cotton swab
  • Coastal garter snakes showed higher preference to slug samples
    • i.e. Coastal population had a significantly higher number of tongue flicks to the slug covered cotton swab
  • Genetic crosses of both populations indicated that this prey preference was being driven by a variation in alleles
how could prey preference variation occurred
How Could Prey Preference Variation Occurred?
  • Remember the bell shaped curve?
  • A rare slug feeding allele could have become more prevalent in the coastal population
  • Hypothesis- Garter snakes with rare slug eating allele were able to gain a fitness advantage because they could acquire more energy from eating a slug than a snake eating a tadpole

Selection Pressure

genetic differences1
Genetic Differences
  • Genetic differences in behavior are hard to understand because most phenotypes are polygenic
  • Polygenic Inheritance means multiple genes effect 1 characteristic
    • Ex. Skin pigmentation
  • But sometimes a single gene can effect a behavior
single gene effect on behavior development
Single Gene Effect on Behavior Development
  • When a single gene effects a behavior scientists can manipulate the gene to test the effects of genes on behavior
    • Inactivate the gene
    • Knockout experiments
  • Aids in understanding the role of genes on the development
knockout experiments
Knockout Experiments
  • Laboratory mice
  • fosB genes
  • Codes for proteins involved in gene transcription
  • If fosB gene is inactivated transcription can not occur
  • Scientists artificially alter the code of the fosB gene
knockout experiments1
Knockout Experiments
  • fosB genes involved in influencing maternal behavior
  • Females with active fosB genes tends to offspring
  • Females with inactivated fosB genes does not tend offspring

Ultimate causation??

How would a natural mutation effect mice?

developmental homeostasis
Developmental Homeostasis
  • The capacity of developmental mechanisms within individuals to produce adaptive traits, despite potentially disruptive effects of mutant genes and suboptimal environmental conditions
  • Behavioral development is redundant
    • There is a “back-up” plan
  • Allows animals to adapt and develop normally when they face unusual environments
developmental homeostasis mediates suboptimal environments
Developmental Homeostasis Mediates Suboptimal Environments
  • Belding’s ground squirrels
  • Baby squirrels were raised in isolation
    • No environmental sensory input
  • When hear audio recording of alarm call react normally
    • Stop behavior and look around
  • Illustrates that there is redundancy in development of behavior
developmental homeostasis mediates effects of mutations
Developmental Homeostasis Mediates Effects of Mutations
  • Developmental homeostasis helps counteract deleterious mutations
    • Use of a “back-up” plan
  • Mutations thought to contribute to asymmetry in body development
  • Developmental homeostasis aids individuals in developing symmetrical bodies despite mutations
why is symmetry so important
Why is Symmetry so Important?
  • Symmetry is thought to be a signal of mate quality
  • In many species, symmetrical individuals are mated with more frequently than non-symmetrical males
  • Symmetry is adaptive!!
adaptive value of learning
Adaptive Value of Learning
  • Modification of behavior based on experience
  • Behavior modified to maximize fitness
  • Behavior modification and its mechanisms are costly
cost benefit analysis of behavior
Cost- Benefit Analysis of Behavior
  • Benefit of a behavior has to outweigh the cost
  • Discriminating behavior in mate selection behavior of Thinnine wasps
cost benefit analysis
Cost- Benefit Analysis
  • Male invests lots of energy in process of “mating” with orchid
  • Potentially loses mating opportunities
  • Males change behavior by remembering locations of orchids and avoiding scents from those locations
    • Cost of processing stimulus
    • Cost of storing memory (develop brain)
  • The benefit of learning behavior is that male increases his efficiency at finding females
    • Benefit in reproductive potential outweighs cost of learning behavior
sex differences in learning behavior
Sex Differences in Learning Behavior
  • Males and females may vary in their ecological pattern
  • Variations can lead to different cost-benefit analysis formulas between sexes
vole ecology
Vole Ecology
  • Polygynous Meadow vole
    • Males have multiple mates
    • Male territory has to encompass all female territories
    • Male territory 4X size that of female
    • Consequence- male has to be able to navigate greater area than female
  • Monogamous Prairie vole
    • Male and female live together
    • Male and female have same size territory
    • Consequence- males and females navigate same area
vole torture
Vole Torture
  • Meadow voles (polygynous)
    • Males made fewer errors than females
    • Suggests males learn more readily
  • Prairie voles (monogamous)
    • Males and females had no significant difference in error rates
    • Suggests males and females learn equally
cost benefit analysis1
Cost-Benefit Analysis
  • Recall learning behavior is costly
  • Behavior modification and its mechanisms are costly
    • Requires genes, gene expression, brain power
mammalian hippocampus
Mammalian Hippocampus
  • Long term memory
  • Spatial relationships like navigation
  • What would you expect when comparing hippocampus size in male and female meadow voles?
vole hippocampus
Vole Hippocampus
  • Male meadow voles have larger hippocampus than the female
  • Pine voles (monogamous)
    • Male and female similar sized hippocampus
not so fast
Not so fast!
  • Brown Headed Cowbird: Interspecific Brood Parasite
reproduction requires lots of energy reproductive effort
Reproduction Requires Lots of Energy (reproductive effort)
  • Save energy by eliminating parental care
  • Allows more energy for creating additional gametes for further reproductive events.
  • More reproductive events = higher fitness
  • Cowbirds produce 30-40 eggs per season
don t put all of your eggs in one basket
Don’t Put All of Your Eggs in One Basket!
  • Reduces risk of predation by having many nesting sites
  • Predators include mice, ground squirrels, weasel, badgers, deer, and hawks
  • Increase fitness by targeting multiple nests
    • Parasitizes smaller birds
female cowbird ecology
Female Cowbird Ecology
  • Female must find host nest
  • Female monitors nest to lay eggs when host is laying eggs
    • Must remember location to return
  • Female lays up 40 eggs- must be able to locate sufficient number of nests
  • Female needs high spatial processing abilities
male cowbird ecology
Male Cowbird Ecology
  • Males do not need high spatial processing abilities
    • Males do not need to find nests
    • Males spatial processing limited to finding mates and food
Sex Differences in Learning Behavior-Variation in male and female ecology result in different cost-benefit analysis formulas between sexes
  • Female Cowbirds exhibit larger hippocampus volume
operant conditioning
Operant Conditioning
  • A type of learning based on trial and error, in which an action, becomes more frequently performed if rewarded
  • Involves an operant (a voluntary action) and consequence that comes from that action
skinner box
Skinner Box
  • B.F.Skinner, psychologist
  • Lever is accidentally pressed as rat explores box
    • Food is dispensed
  • Positive reinforcement will increase frequency of behavior
negative reinforcement
Negative Reinforcement
  • Novel taste followed by nausea results in avoidance of food source
  • Adaptive behavior for avoiding toxic foods