Behavior Development b

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Behavior Development b
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Learning

• Learning is a change in an animals behavior
  linked to a particular experience it has
• Brain properties change by gene and environmental
  interactions

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Learning

• Forms of learning
• Imprinting
• Specialized learning
• Variation in learning behavior
• Environmental influences

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Imprinting

• http//www.youtube.com/watch?v7OynlzqtxmY
• Why would this behavior be adaptive?

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Imprinting

• Occurs when a young animals 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

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Konrad Lorenz

• Imprinting in Greylag Geese
• Imprint on humans
• Later preferred Humans as mates
• http//www.youtube.com/watch?v2UIU9XH-mUI
• http//www.youtube.com/watch?vSK65euK1FGU

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Great Tits Blue Tits
Imprinting has different effects among species
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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
  its own species
• Formed a mate preference based on imprinted
  foster parent
• Most of Blue Tits mated within its own species
• Exhibited a different developmental interaction

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Specialized Learning

• Memory formation to aid in finding food

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Clarks Nutcracker
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Clarks 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

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Clarks Nutcracker
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Clarks Nutcracker

• Nutcrackers able to relocate caches with up to
  80 accuracy
• Recall memory for months
• Use landmarks to relocate caches

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Nutcracker torture!
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White Bark Pine
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Seeds germinate in clusters
Un-retrieved caches create new stands
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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?

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Factors that Influence the Development of Learned
Behavior

• Environmental
• Genetic

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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

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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

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Beldings Ground Squirrels
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Beldings Ground Squirrels
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Beldings 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

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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

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Kin Recognition

• Signals used to identify kin include
• Scent
• Appearance

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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

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Beldings 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

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What would you predict?

• Siblings reared apart
• Siblings reared together
• Non-siblings raised apart
• Non-siblings raised together

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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

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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

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Kin Discrimination
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Why are Females more Discriminating in their
Sense of Smell?
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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 individuals smells to
• Individuals who smell similar are more closely
  related, whereas individuals that smell less
  similar are less closely related

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Chemical Communication

• Oral and dorsal glands
• Nasal investigation
• Scent mark behavior
• 5 odors that are individually distinct

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Scent Discrimination

• Individuals learn to recognize their own scent
• Spend more time sniffing less closely related
  relatives

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Ultimate Causation

• Kin Recognition by using scent discrimination
  allows Beldings ground squirrels in order to
• Helps identify closely related for assistance
• Ability to recognize intruders
• Prevents inbreeding

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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

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Western Terrestrial Garter SnakeThamnophis
elegans
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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

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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?
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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
• http//www.youtube.com/watch?v92InYz1cU2o

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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

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Garter Snake populations exhibit Prey Preference
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• What happens if Garter snakes are offered
  multiple prey options?

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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

What?
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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

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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

http//vimeo.com/1206868
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Vomeronasal Organ

• AKA Jacobsens organ

http//www.youtube.com/watch?vyzC8DZK0aT4
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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

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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
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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

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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

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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

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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?
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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

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Developmental Homeostasis Mediates Suboptimal
Environments

• Beldings 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

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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

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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!!

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Adaptive Value of Learning

• Modification of behavior based on experience
• Behavior modified to maximize fitness
• Behavior modification and its mechanisms are
  costly

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Cost- Benefit Analysis of Behavior

• Benefit of a behavior has to outweigh the cost
• Discriminating behavior in mate selection
  behavior of Thinnine wasps
• http//www.youtube.com/watch?v0nfgibIYbg8

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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

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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

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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

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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

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Cost-Benefit Analysis

• Recall learning behavior is costly
• Behavior modification and its mechanisms are
  costly
• Requires genes, gene expression, brain power

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Mammalian Hippocampus

• Long term memory
• Spatial relationships like navigation
• What would you expect when comparing hippocampus
  size in male and female meadow voles?

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Vole Hippocampus

• Male meadow voles have larger hippocampus than
  the female
• Pine voles (monogamous)
• Male and female similar sized hippocampus

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Not so fast!

• Brown Headed Cowbird Interspecific Brood Parasite

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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
• http//www.youtube.com/watch?v9O7Ruzaxgvoplaynex
  t1listPL807E25E0A3A38593

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Dont 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
• http//www.youtube.com/watch?vprvz7a5IBaYfeature
  related

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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

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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

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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

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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

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Skinner Box

• B.F.Skinner, psychologist
• Lever is accidentally pressed as rat explores box
• Food is dispensed
• Positive reinforcement will increase frequency of
  behavior

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Negative Reinforcement

• Novel taste followed by nausea results in
  avoidance of food source
• Adaptive behavior for avoiding toxic foods

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(No Transcript)
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Sources

• http//www.youtube.com/watch?vJbKXNjbgvqc
• http//www.birdsamore.com/videos/snake-eatingslug.
  htm