Reinforcement%20(something%20

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• Reinforcement (something good) Anything that
  INCREASES the probability of a behaviour
• Positive reinforcement something good is
  added (e.g., food)
• Negative reinforcement something bad is taken
  away (e.g., parent stops nagging when child
  cleans room)
• Punishment (something bad) Anything that
  DECREASES the probability of a behaviour
• Positive something bad is added (e.g., shock)
• Negative something good is taken away (e.g.,
  no TV)

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Other Aspects (Types) of Learning

• Learning set (problem-solving strategy)
• Improve across similar tasks, generalize
• E.g., Win-Stay, Lose-Shift strategy

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Other Aspects (Types) of Learning

• Latent learning (learn associations without
  reinforcement)
• e.g., through exploratory behavior

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Other Aspects (Types) of Learning

• Observational learning
• E.g., food-washing, separating grains from sand
  in Japanese macaques
• Controversy true imitation, or (unseen) trial
  and error?

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Other Aspects of Learning

• Imprinting

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Invertebrate Learning

• Many phyla show basic learning abilities (i.e.,
  habituation and classical conditioning able to
  learn reflexive associations)
• Some arthropods show operant conditioning
• E.g., Bees can solve discrimination (choice)
  tasks using visual and/or olfactory (odour) cues

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Vertebrate Learning

• All types of association learning (i.e.,
  habituation, classical, operant conditioning)
  demonstrated in almost all phyla
• In addition to traditional learning abilities,
  we will also consider higher mental functions
  (re intelligence, cognition) to come

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Nature vs. NurtureWhere Does Learning Fit In?

• Recall definition of learning relatively
  permanent change in behaviour (or potential for
  behaviour) that results from experience
• If learning results from experience (i.e.,
  nurture, environment, proximate factors), what
  role can evolution play?
• Ultimate factors (i.e., nature, genetics,
  selection)?

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Effects of Natureon Learning Ultimate Factors

• Biological constraints
• Levels of preparedness (to learn a behaviour)
• Sensory Umwelt, behavioural repertoire, etc.
• Learning as adaptive behaviour
• Selection pressure
• Changes to structure and/or function through
  selection (e.g., hippocampal size)

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Preparedness and Constraints

• Preparedness (evolutionarily sensible)
• e.g., learn to associate food with illness
• Unprepared (arbitrary, no conflict w/
  species-typical behaviour, requires training)
• e.g., learn to bar press for food
• Contra-prepared (conflict w/ species-typical
  behaviour)
• e.g., bar press to avoid shock, rather than run
  away

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Raccoon trained to deposit coins in piggy bank,
but contra-prepared to learn this
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Another Example Garcias Bright Noisy Water
(BNW) Expt.

• Grp 1 BNW shocked
• Grp 2 BNW nauseated (irradiation poisoning)
• Grp 3 Sucrose water shocked
• Grp 4 Sucrose water nauseated
• Then, all given choice of sweet water or BNW

H2O
Click!
H2O
(bright/noisy)
(sweet)
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Preparedness Constraints
Bright Noisy Water Sweet Water
Shocked Avoided BNW, drank sweet water No preference
Nauseated No preference Avoided sweet water, drank BNW

• Prepared to learn association between BNW
  shock, and between sweet water and nausea
  unprepared to learn other associations

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Human Examples?

• Easy to learn fear of insects, snakes, spiders,
  (but not chickens or sheep or flowers)
• Not born with fear born with preparedness to
  fear
• E.g., preparedness, sound and emotion

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Methods Constraints on Learning

• Stimuli used for testing must fit into sensory
  Umwelt of species being tested
• Must consider the testing apparatus, animals
  prior experience (e.g., wild vs. captive), age,
  gender, perceptual capabilities, etc.

I have not got my specs with me
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Learning As Adaptive Behaviour

• E.g., Spatial Memory
• Formation of cognitive maps of environment
• learning b/c requires gaining info thru
  experience (e.g., latent learning, exploratory
  behaviour)
• cognitive b/c involves a mental manipulation
  of that info, ability to act flexibly, e.g.,
  detours)

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Spatial Learning Functional Significance

• Animals must learn the layout of their
  environment to solve various problems, e.g.
• efficient foraging hummingbirds remember and do
  not return to flowers they have already drained
  of nectar
• food caching nutcrackers store and retrieve
  10,000s of seeds in 1000s of locations

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Spatial Learning Functional Significance (contd)

• Animals must learn the layout of their
  environment to solve various problems, e.g.
  (contd)
• Complex navigation birds migrate 1000s km and
  return to same nesting spot each year
• Predator avoidance often-predated species
  (e.g., rodents) know escape routes and hiding
  places

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Food Caching in Corvids
Dashed lines indicate relative capacity for food
transportation (size of gullet) morphological
adaptation (physical change resulting from
different selection pressures on each species)
Corvids e.g., crows, jays, nutcrackers
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Food Caching in Corvids

• Morphological adaptations (size of gullet) relate
  to ecological niche/importance of stored seed to
  diet
• Species that rely on stored seed have not only
  adapted physically, but have also specialized in
  learning and memory for food caches
• Thus, learning and spatial memory abilities may
  be considered an adaptation as well

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Food Caching Clarks Nutcrackers

• 1 NC will cache up to 33,000 seeds in 1 yr, in
  6000-8000 caches (locations)
• Find 90 of cached seeds, despite change in
  landscape caused by snow cover
• 80-100 of winter diet

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• Continue cache-recovery into next summer, 9-11
  months after hiding it
• Demonstrates long-term spatial memory, cognitive
  map of cache locations
• Young may learn to cache from parents

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Complex Navigation Leachs Storm-Petrels

• Great Island, Witless Bay colony (250,000 pairs)
• Open meadows vs. wooded areas w/ small, closely
  arranged burrows
• Nocturnal (forage at night)
• High spatial demand

Abbott, Walsh, Storey, Stenhouse Harley (1999)
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Abbott, Walsh, Storey, Stenhouse Harley (1999)

• Hippocampus brain region responsible for
  spatial memory
• Compared subpopulations of same species (LSP)
• Live in woods, close burrows (requires complex
  navigation) larger relative hippocampal volume
• Live in open meadow, sparse burrows smaller
  relative hippocampal volume

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Complex Navigation Leachs Storm-Petrels (LSP)

• Abbott et al. (1999) also compared relative
  hippocampal volume across species LSP compared
  to
• Northern Fulmar Diurnal, nests at open sites,
  lower densities
• Storing (e.g., nutcracker)
• Non-storing bird species

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Abbott et al. (1999) Results

• Increased spatial demands larger relative
  hippocampal volume

Relative Hippocampal Volume
Low
High
Non-storers
Northern Fulmar
Leachs Storm-Petrel
Storers (e.g., NC)
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Spatial Learning is Adaptive

• Ability to learn allowed these species (i.e., NC,
  LSP) to adapt to specific environment challenges
• Need to find food in winter/Need for home/shelter
• Large hippocampus correlated with increased
  spatial ability
• Hard to determine cause and effect relationship
• Large HPC allowed LSP to enter woods, or
• LSP entered woods, large HPC selected for

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Learning is Adaptive Example 2 Brood Parasitism

• Parasitic species (e.g., cowbirds)
• Female lays eggs in nest of host species (no care
  from true parent)
• Parasitic chicks hatch earlier, grow faster
• Preferentially fed, push others out of nest

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This adult Common Yellow-throat is feeding a
cowbird fledgling that's more than twice its
size. Photo by John Gavin, Source.
birds.cornell.edu/.../ tanager/parasitism.html
Thanks, Mommy!
Learning

• Parasitic chicks learn to recognize
  characteristics of host species
• They survived species is good host
• Will lay eggs in nests of same species
• Defense Hosts learn to recognize own eggs,
  reject others