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(CLASS 6 (detailed))
(Study guide ("stuff that may be on the test"))
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**effects of brain damage
**effects of brain damage
*adaptive control of thought theory
*adaptive control of thought theory
===CLASS 10===
canceled due "swinesque" flu symptoms
===CLASS 11===
*two neural “learning” systems display different developmental time-courses.
**developmental dissociation of habituation from sensitization
*development of sensory systems and simple associative learning mediating by those systems
*developmental dissociation may result from a caudal-to-rostral sequence of maturation
*the ability to form configural / relational associations emerges later
**evidence from studies of younger versus older rats
**evidence from studies of younger versus older human children
***facilitation of configural performance in younger children
*development of the hippocampus in rats and humans
**correlation with ability to form behavioral configural associations
***infantile amnesia
**effects of neonatal brain damage in general
***effects of neonatal hippocampal damage
==Disability accommodation==
==Disability accommodation==

Revision as of 13:53, 23 November 2009

Hartman lab banner rat.jpg

LLU PSYC 544: Foundations of Learning and Behavior


Autumn 2009 - 4 credits

  • Mondays and Wednesdays from 1:30-3:20 in CDR 201 (syllabus)


Course description

This class will introduce the major theories, methods, and applications in the science of learning, including simple stimulus learning, complex associative learning, and memory storage/retrieval, with a special emphasis on the biological basis of learning and memory.


  1. ISBN:019514175X [Eichenbaum]


Recommended readings

Be prepared for graduate school.


There will be 3 exams (worth 100 points each) with questions drawn from both the reading assignments and classroom discussions. For each test, the class average will be determined and adjusted to at least 70% (C). For example, if the class average is 65%, 5% will be added to all grades to bring the average up to 70%. Note that the average will never be adjusted down. Also, note that it is possible to end up with a score of higher than 100%. There will be no make-ups after test day. If a test is missed, you will receive a score of 0 for that test. However, there will be an optional comprehensive final exam. Your 3 highest test scores will be used to calculate your grade. Therefore, taking the final exam can only help your grade. Additionally, each student will write a paper on a topic of interest and present a talk on that topic (50 points paper / 5 points talk - details to be presented in class). Finally, 20 points will be assigned for classroom participation (attendance, discourse, not sleeping, etc – especially during the student presentations).

Grading Scale

  • 378-420 90% A
  • 336-377 80% B
  • 294-335 70% C
  • 252-293 60% D
  • 0-251 ouch F

Study guide ("stuff that may be on the test")



  • definitions of "learning" and "memory”
  • relationship between learning and memory
  • biological basis
  • evidence of learning
  • behaviors that may appear to be "learning"
    • Tolman and Honzick (1930) latent learning
  • methods of studying learning / memory
    • behavioral
    • cognitive
    • neuropsychological
  • orienting response
  • habituation / sensitization
    • parametric features of habituation
  • classical (Pavlovian) conditioning
    • associative learning
    • predicting / measuring the relationship between stimuli
      • Rescorla-Wagner model
    • critical features of a classical conditioning experiment
      • CS
      • US
    • extinction
    • spontaneous recovery
    • generalization
    • discrimination
    • the role of contiguity
    • phobias
  • operant / intrumental conditioning
    • associative nature
    • Thorndike and "instrumental" learning
      • law of effect
    • stimulus-response learning
    • Skinner and "operant" learning
    • reinforcers
      • theories of reinforcement
      • brain centers important for "reward"
      • Premack principle
      • biofeedback
      • positive vs. negative reinforcers
      • primary vs. secondary reinforcers
    • punishers
      • positive vs. negative punishment
    • reinforcement schedules
      • continuous / partial
      • interval / ratio
      • interval / fixed
    • shaping
      • chaining
  • similarities and differences between classical and operant conditioning


  • crash course in neurophysiology and neuroplasticity
    • neurotransmitters
    • synaptic relationships between neurons
    • polarized nature of cell membrane
    • post-synaptic potentials
      • excitatory
      • inhibitory
      • characteristics of the post-synaptic potential
      • axonal / dendritic relationships
      • role of the axon hillock
        • ion channels at the hillock
      • action potential characteristics
        • result of an action potential
        • refractory period
  • biological models of learning / memory in the Aplysia snail
    • neuronal circuit for the gill withdrawal reflex
    • habituation
      • behavioral habituation (and dis-habituation) of the reflex
      • presynaptic mechanism
      • postsynaptic mechanism
      • basis of short-term memory (function)
      • basis of long-term memory (structure)
      • basis of "forgetting"
    • sensitization
      • behavioral sensitization of the reflex
      • addition of facilitating interneurons to the circuit
      • presynaptic mechanism
      • postsynaptic mechanism
      • basis of short-term memory (function)
      • basis of long-term memory (structure)
        • prevention of long-term memory formation
    • classical conditioning
      • same circuit as sensitization
        • associative learning due to temporal relationship of the stimuli (“activity dependent”)
        • amplication of presynaptic facilitation
      • “reflexive” nature of classical conditioning
  • overview of cellular processes that alter neuronal function / structure (neuroplasticity)
    • Calcium
    • cAMP
    • CREBs
    • RNA transcription > production of proteins
  • The Hippocampus, NMDA Receptors, and Learning
    • hippocampal formation and association cortex
      • main areas
      • output pathways
      • role of hippocampal circuitry
      • tri-synaptic pathway
      • long-term potentiation of the tri-synaptic pathway
        • role of glutamate and its receptors
          • unique role of NMDA receptors
      • long-term depression


  • “Hebbian” rule
    • synaptic correlation > LTP
    • lack of synaptic correlation > LTD
  • structural changes in dendrites
    • role of dendritic spines
  • upregulation of receptors on dendrite
  • retrograde messengers
  • LTP and LTM
    • hippocampus
    • 1 trial learning
    • long-lasting
    • specific
    • associative
    • LTP and “real learning” use similar biochemical mechanisms
  • NMDA receptors, neuroplasticity, and susceptibility to excitotoxic damage
  • Effects of hippocampal damage in humans
    • patient H.M.
      • brain surgery
      • anterograde amnesia
      • retrograde amnesia
        • lateralization of hippocampal functions
        • damage limited to CA1
        • deficits in specific types of learning / memory
          • types of learning / memory that are spared after bilateral hippocampal damage

CLASS 5 (detailed)

  • memory for context and relations between stimuli - mediated in part by the hippocampal formation
  • memory for simple stimulus-stimulus associations (e.g., simple object discrimination) - mediated by other (more “primitive”?) neural systems
  • Complex vs Simple (explicit vs. implicit, declarative vs. procedural, recognition vs. habit, configural vs. simple)
  • Simple: procedural learning / skills, classical / operant conditioning, non-associative, priming (single presentation of a stimulus facilitates its recall)
  • “Priming” involves activating the necessary perceptual and identification processes involved in information processing before they are actually needed
  • diseases associated w/ procedural learning: Parkinson’s, cerebellar damage
  • Learning how – process by which knowledge is acquired
    • largely independent of conscious awareness
    • Cannot be verbally explained
    • Alterations in the connections between and within related brain structures are likely behind procedural skills
    • Performance of skills is task-specific
    • Improvement in performance or skills is only method of conveyance
    • H.M.’s / others implicit learning / memory is as good as “normal”
      • had no memory of ever performing the task before and required instructions each time he performed the task
  • “Declarative” ~ learning that – largely dependent on conscious awareness
    • (by definition) verbally explained
    • episodic
    • semantic
    • patients w/ damage to the medial temporal lobes have deficiencies with this type of learning
    • Contextual - involves “tying together” relations between many stimuli in the environment
      • Reaux Sham Beaux
  • Adaptive control of thought theory
    • Declarative stage:
      • Learning factual knowledge of the nature of task
    • Transition (Knowledge compilation stage):
      • after additional practice, performance speed increases
      • less conscious “guidance” is required
      • groups of rules / operations are chunked / compiled & stored in LTM
      • performance starts to depend more on LTM than working
      • working memory is now “freed up”
      • increases efficiency of task
    • Procedural stage: With more practice, task is performed almost automatically
      • Major variable – practice (expertise)
      • Optimal learning conditions: Spacing / Generation effects / Practice / Develop strategies
  • studying spatial working memory in the radial arm maze
    • Win-shift paradigm - each identical arm was baited at its end
    • most efficient strategy is to visit each arm only once per trial
    • during each trial, the rat had to remember which arms it had visited based on their relation with extra-maze cues
    • normal rats can learn and retain this strategy almost perfectly
    • bilateral disruption of the main hippocampal input (entorhinal cortex) and output (fimbria-fornix) pathways OR intrinsic hippocampal circuitry - performed at chance levels
    • lesions in other brain areas (cortex, caudate nucleus, amygdala) - no retention deficits
  • studying spatial memory in the Morris water maze
    • hippocampal-damaged rats were profoundly impaired on the relational, spatial-cue-based navigation, but normal during acquisition of the cued condition
  • studying spatial memory with the spatial alternation task
    • rats were trained to find water at the end of one arm of a T-maze
      • forced run - one arm contained water and the other arm was blocked
      • choice run - the opposite arm contained water and either arm could be chosen
        • correct response - choose the arm blocked on the previous forced run
          • only in the context of the previous (forced run) trial could the correct choice be made.
            • damage to bilateral hippocampal OR amygdaloid lesions
              • hippocampus - performed at chance levels
              • amygdala - displayed no impairment (performed at control levels)
  • removal of the hippocampus soon after birth in rats results in a loss of spatial abilities through adulthood
    • this suggests that there is no backup (redundant) system for this type of information processing
  • spatial learning requires only a small portion of functional tissue in the dorsal hippocampus

CLASS 6 (detailed)

  • we can learn more about a stimulus when it is easily perceived
    • increased experience with stimuli allows such enhanced perceptions to occur
    • “the more you know, the more you can learn”
    • perceptual fluency – the ability to quickly distinguish one stimuli from another
  • presentation of varying degrees of contrasting stimulus categories leads to effective understanding of items
    • once easy discrimination can be made, more subtle ones are easier
    • learning to discriminate between stimuli generally requires some type of “attention”
  • nervous system is a big mass of biological tissue composed of interconnected neurons
    • parallel and serial networks within massively parallel networks
    • hierarchy: Spinal cord / Hindbrain / Midbrain / Diencephalon / Basal ganglia / Neocortex
      • primary motor / sensory
      • secondary motor / sensory
      • association
  • individual cortical neurons arranged into layers and columns
  • 6 layers:
    • layer 4 is mostly sensory inputs (lots of dendrites)
    • layer 5/6 are mostly outputs (lots of long axons out)
    • columns are arranged “topographically” into “maps” of various parts of our body
    • each mapped area processes certain aspects of incoming data before passing it to other areas
  • sensations cause changes in spatiotemporal patterns of the cortical activity
    • we experience this as our “perception” of reality
  • receptors are specialized neurons for transducing information from the physical environment into chemical and electrical signals (sensations)
  • Receptive sheet
    • receptors innervate very specific regions
    • area which a sensory system innervates = receptive sheet
    • somatosensory receptive sheet is total body surface
    • visual receptive sheet is the retina
  • Receptive field - that portion of the environment to which a receptor (or cortical neuron) will respond
    • each rod or cone has a precise receptive field
    • each subsequent neuron also has a receptive field
    • convergence / divergence at each subsequent synapse creates larger / more complex receptive fields
    • increased coding and abstraction of “information
  • receptors send info about the external world through the thalamus (sensory relay station) to several areas of primary sensory cortex
  • What is the nature of “information” sent to the cortex?
    • Qualitative - “what is it?”
      • labeled line - chain of neurons that results in a sensation
    • Quantitative - “how much?”
      • frequency coding - rate of APs
      • population coding - # of receptors firing
    • Adaptation - response of a neuron to constant stimulation
      • phasic - quickly adapting
      • tonic - slowly adapting
  • By the time info gets to cortex, some elaborate processing is already taking place
    • distinct regions of cortex respond (increase firing rate) to small, specific region of the sensory field
    • cortical neurons have overlapping receptive fields
    • adjacent neurons represent adjacent parts of the field
    • receptive fields are organized in a “topographic map” of the sensory field
  • in addition to overlapping topographic map of the receptive sheet, certain neurons also respond preferentially to specific trigger features
  • individual columns arranged into microcircuits of ~1 mm sq hypercolumns
    • ocular dominance columns
    • orientation columns
    • “color” blob
      • responses of individual neurons are probabilistic (and relatively unimportant)
  • within each sensory area, there exists a hierarchy of larger circuits representing increasingly more complex processing of incoming sensory information
    • visual cortex encompasses almost all of the occipital lobe and parts of the parietal & temporal
    • serial and parallel connections
    • there is as much backward flow as forward flow
    • early, lower, “upstream” areas
      • identifying basic properties of stimuli
        • orientation
        • spatial frequency
        • speed
        • color
        • location
    • later, higher, “down-stream” levels of processing
      • “maps” get fuzzier higher in the cortical hierarchy
      • response properties of these cells more complex
      • arise from combinations of inputs from lower levels
        • what (ventral) streams - “categorization” (faces, hands, etc.)
        • where (dorsal / parietal) - localizing in 3D space
        • higher levels of processing networks connect with each other in association cortex (multimodal, association cortex)
        • these multimodal areas send outputs to areas in the frontal, temporal and parietal cortices AND the hippocampus
  • cells change firing patterns with experience
    • less activity for repeated stimuli
    • processing is “easier” / less effort
    • not only “cellular” mechanisms, but “network”-level mechanisms
  • Learning / Memory is completely intertwined with sensory / perceptual processing
    • early experience (both external and internal) guides the development of the general organization of cortical circuitry
    • experience driven “tuning” / modification
    • response properties are plastic
    • like muscles, using circuits changes their operation
    • experience alters the structure and function of the circuits
  • constantly changing dynamic chemical bath
    • this ball of cells detects stuff in the environment that causes transient changes in the pulsation frequencies
    • produces constantly changing dynamic electrical patterns
    • sensations cause changes in spatiotemporal patterns of the cortex – we experience this as our “perception” of reality
    • different networks naturally tend to “pulse” at different frequencies
    • areas that are active at the same time tend to become more strongly connected (and vice versa)
    • HEBB - “fire together / wire together” via functional and structural changes
    • activity in 1 area is more likely to induce activity in connected areas
  • the biological basis of learning is thought to depend in part upon the ability of neurons to modify their synaptic connections within neural circuits based on experience
  • memory retrieval happens when we re-experience same or similar spatiotemporal patterns in cortical activity
  • cortex is “reorganized” during “learning”:
    • train specific fingers to perform sensory tasks
  • Learning / Memory is the behavioral and psychological consequence of changes in sensory / information processing
    • psychophysical monism
    • neurons are adaptive - can shift or modify response to stimuli based on previous experiences with that stimuli


  • ways to categorize learning and memory
  • stages of "remembering"
  • Nice historical view of memory:
    • primary
    • secondary
  • multistore memory model
    • dual (tri) storage system:
      • sensory
        • capacity
        • duration
        • mechanisms
      • short-term
        • capacity
        • duration
        • mechanisms
      • working memory
        • roles
        • capacity
        • duration
        • mechanisms
        • regions of PFC that control working memory processes
        • central executive and sensory / episodic buffers model
      • incidental learning
      • speed of processing
      • working memory in animal studies


  • encoding memories
    • the distributed cortical engram
    • cognitive model of encoding - schemas
    • auto-associative neural networks
    • things that effect the encoding of memories
    • consolidation in LTM
      • 2 phases
      • role of the hippocampus
        • parahippocampal buffer
        • comparator function
        • recording episodic memories
        • activity during sleep
        • damage leads to lack of consolidation
          • partial vs. total
      • storage "site"
      • hippocampus vs. cortex:
        • capacity
        • plasticity


  • Hippocampus vs. neocortex
    • capacity
    • plasticity speed
    • plasticity “longevity”
  • Interaction of hippocampal networks with cortical networks
  • final repository of a memory
  • LTM organized by “schemas”
  • Initial role of hippocampus
  • memory retrieval
    • characteristics
    • testing effects
    • prospective memory
    • false memories
  • forgetting - possible mechanisms
  • “dissociations” suggest different biological systems
    • double dissociations
    • at least two distinct functional memory systems
      • operate simultaneously and in parallel
      • in each proposed system:
        • a simple yet gradually learned form of memory (e.g., procedural, simple associative)
          • brain areas
        • a more complex but rapidly learned form (e.g., declarative, spatial, explicit, contextual associative)
          • brain areas
    • “Mono-hierarchy” idea
    • model circuit illustrating how sensations may interact with various neural structures to form a memory of episodes and facts
  • Procedural Memory
    • characteristics
    • brain areas
      • motor cortex
      • cerebellum
      • basal ganglia
    • effects of brain damage
  • adaptive control of thought theory


canceled due "swinesque" flu symptoms


  • two neural “learning” systems display different developmental time-courses.
    • developmental dissociation of habituation from sensitization
  • development of sensory systems and simple associative learning mediating by those systems
  • developmental dissociation may result from a caudal-to-rostral sequence of maturation
  • the ability to form configural / relational associations emerges later
    • evidence from studies of younger versus older rats
    • evidence from studies of younger versus older human children
      • facilitation of configural performance in younger children
  • development of the hippocampus in rats and humans
    • correlation with ability to form behavioral configural associations
      • infantile amnesia
    • effects of neonatal brain damage in general
      • effects of neonatal hippocampal damage

Disability accommodation

If you are registered with the Dean's office for disability accommodation, please see me as soon as possible to discuss any course accommodations that may be necessary. If you have a disability and wish accommodation, please visit the Dean’s Office of your school. See the Accommodation for Disability policy.

Academic integrity policy

Don’t cheat.