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Review
[email protected] www.cogsci.ucsd.edu/~ksweeney/psy260.html
Introduction to Physiological Psychology
n Learning and Memory n Human Communication n Emotion
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What is memory?
n Working Memory: – Limited capacity (7 +/- 2) – Information can be held for several minutes
with rehearsal § (e.g. memory system you use when you have to remember a
phone number but have no place to write it down)
n Long-term Memory: – Very large capacity – Essentially infinite duration
§ e.g. memory system you need when you are reminiscing with friends, or taking a final exam
Forms of Learning
Perceptual Learning
Motor Learning
Stimulus-Response Learning
Relational Learning
Objects
Situations
Form new circuits in the motor system
Form connection between perception and action
Connections between stimuli
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Learning n All forms of learning involve changes in
the ways that neurons communicate.
Stimulus-Response learning n Classical Conditioning
– An unimportant stimulus begins to elicit a similar response as an important one
– It involves an association between two stimuli, one of which is reflexive
n Operant Conditioning (or Instrumental Conditioning)
– A particular stimulus begins to elicit a particular response
– It involves an association between a stimulus and a response
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Classical Conditioning
n Famous example: Pavlov’s dogs – First, present dogs with food and measure
amount of saliva – Then, start ringing a bell just before food is
presented (at first, saliva only occurs at presentation of food)
– In time, salivation occurs in response to the bell
– Conditioning has occurred
Classical Conditioning
n Unconditional Stimulus- dog food n Unconditional Response- salivation n Conditional Stimulus- bell n Conditional Response- salivation
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n Reinforcing stimulus (favorable consequences)
§ Appetitive stimulus that follows a particular behavior and thus makes behavior occur with greater frequency
n Punishing stimulus (unfavorable consequences)
§ Aversive stimulus that follows a particular behavior and thus makes behavior occur more rarely
Instrumental (or Operant) Conditioning
An association between a stimulus and a response
But what has happened in the brain? n Hebb postulated:
– the cellular basis of learning involves strengthening of a synapse that is repeatedly active when the postsynaptic neuron fires
– “neurons that fire together, wire together”
For LTP to occur, the postsynaptic cell must already be depolarized
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NMDA and AMPA n Glutamate binds to NMDA receptors, which controls a
calcium (Ca2+) channel. n So, Ca2+ rushes in, right? NO!
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NMDA and AMPA n At rest, that same calcium channel is ‘guarded’ by a
magnesium ion (Mg2+), so calcium can’t get in through NMDA receptors.
n That Mg2+ ion won’t budge unless cell is depolarized. n But cell can’t depolarize unless Ca2+ can get in, right?
NO!
NMDA and AMPA n If a weak synapse is active by itself, nothing happens… n BUT- if the cell has just fired due to a strong synapse
somewhere else on the cell, a dendritic spike will depolarize the membrane…
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NMDA and AMPA n Depolarization kicks the Mg2+ ion out, and NOW Ca2+
ions can enter the cell. n … and an association between those two synapses is
formed.
We still don’t have LTP! n Ca2+ ions entering the cell bind with the enzyme CaM-
KII n CaM-KII causes more AMPA receptors to to move to
post-synaptic membrane. n More AMPA receptors means it’s easier to depolarize
the cell in the future.
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We still don’t have LTP! n Ca2+ ions entering the cell bind with the enzyme CaM-
KII n CaM-KII causes more AMPA receptors to to move to
post-synaptic membrane. n More AMPA receptors means it’s easier to depolarize
the cell in the future.
n For Ca2+ to enter the cell, NMDA receptors have to be activated by glutamate AND subjected to depolarization simultaneously.
n The fact that both these things must occur together means that NMDA receptors are “coincidence detectors”.
n Thus, they are crucial for LTP.
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Perceptual Learning
n The ventral stream – involved with object
recognition, continues ventrally into the inferior temporal cortex.
n The dorsal stream – involved with perception of
the location of objects, continues dorsally into the posterior parietal cortex.
n The ventral stream is involved with the what of visual perception; the dorsal stream is involved with the where.
Instrumental Conditioning
n Circuits responsible for instrumental conditioning begin in sensory association cortices and end in motor association cortex.
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Instrumental Conditioning
n Two major pathways from sensory to motor association areas: – Direct transcortical connections- involved in
STM, acquisition of episodic memories and of complex behaviors that involve deliberation or instruction (slow and laborious)
– Connections via the basal ganglia and thalamus- which are involved as behaviors become automatic and routine (fast and easy)
H.M.
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What can possibly go wrong? n Anterograde Amnesia:
– Amnesia for events occurring after the precipitating event.
n Retrograde Amnesia: – Amnesia for events occurring before the
precipitating event.
The Medial Temporal Lobe: Crucial in the Declarative Memory System
n Damage to these areas usually results in anterograde amnesia: patients are unable to form new declarative memories.
n Can also result in retrograde amnesia: typically ‘graded’.
n Non-declarative memory is not affected.
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H.M. Effects of Bilateral Medial Temporal Lobectomy
n Minor seizure beginning at age 10, major seizures beginning age 16
n Severe, persistent seizure condition- not controlled with anticonvulsants
n By mid-20’s, condition was so severe he was unable to work
n Surgery at age 27: Bilateral medial temporal lobe resection.
n In HM, the amygdala, entorhinal and perirhinal cortices, and about two-thirds of the hippocampus were removed
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What’s wrong with H.M., and what does it tell us about functions of Hippocampus and MTL?
n What CAN he do? – Intellect is normal – Can remember the past (pre-surgery)
§ He has relatively little retrograde amnesia § His long-term memory is intact
– Can carry on excellent, short conversation § His working memory is intact
– Can learn new skills at a normal rate- and retains those skills over long periods of time § His procedural memory is intact
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What’s wrong with H.M., and what does it tell us about functions of Hippocampus and MTL?
n What CAN’T he do? – Doesn’t retain new semantic or episodic
information
– Can’t form new declarative memories.
What does H.M. tell us about role of Hippocampus and MTL?
n Hippocampus is essential for the formation, but not the storage or retrieval, of long-term declarative memory
n Memory depends on Hippocampus for a short duration
n Hippocampus does not mediate short-term memory
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What does H.M. tell us about role of Hippocampus and MTL?
n STM and LTM are distinctly separate – H.M. is unable to move memories from STM to
LTM, a problem with memory consolidation
n Memory may exist but not be recalled – as when H.M. exhibits a skill he does not know he has learned
Explicit vs. Implicit Memories
n Explicit memories – conscious memories
n Implicit memories – unconscious memories, as when H.M. shows the benefits of prior experience
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Broca’s Area and Patient “Tan”
Lateralization of Function
n For many functions the hemispheres do not differ and where there are differences, these tend to be minimal
n Lateralization of function is statistical, not absolute! – e.g. Right hemisphere has some language
abilities
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Lateralization of Function
Left Hemisphere n “Language”
– Even for deaf people!
n Words, letters
n The details
Right Hemisphere n Emotional Prosody n Music n Spatial ability n Faces, patterns
n The big picture
Language n Language is not a unitary ability
– Production vs. Comprehension
n Production – Requires having something to say, being able
to associate that “thing” with words, and making the mouth move appropriately
n Comprehension – Begins in the auditory system (detection and
analysis of sounds) but there is a difference between recognizing a word and comprehending it
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What can possibly go wrong?
n Aphasia – A difficulty with speech (either production or
comprehension) caused by brain damage rather than, e.g. motor deficits or deafness
What can possibly go wrong? n Broca’s aphasia
– difficulty in language production § Comprehension is normal § Know what they want to say, but can’t say it § “expressive aphasia”, slow laborious speech,
full of disfluencies. § Although words are often mispronounced,
words that are produced are usually meaningful
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What can possibly go wrong?
n Broca’s aphasia – Typically function words are most compromised, with
content words being relatively spared. – Aphasias are a spectrum
What can possibly go wrong?
n Broca’s aphasia: not ONLY a production problem! – Although comprehension is good, it is not normal – Agrammatism is present in production, and
grammatical clues such as word order, tense markers or function words aren’t successfully used in comprehension either.
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What can possibly go wrong?
n Broca’s aphasia: not ONLY a production problem! – Anomia: a difficulty in finding words (in naming
things).
What can possibly go wrong?
n Broca’s aphasia: not ONLY a production problem! – Articulation difficulties: mouth
motor movements are disfluent, so words are often mispronounced
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What else can possibly go wrong?
n Wernicke’s aphasia – Wernicke’s area- difficulty in comprehension;
but production is generally meaningless § Unlike Broca’s Wernicke’s aphasics generally speak
quite fluently, with normal prosody, natural-sounding rhythm and apparently normal grammatical constructions.
§ “jargon aphasia”, natural sounding rhythm and syntax, but output is meaningless (“word salad”)
§ neologisms
Wernicke’s Aphasia
n Difficulty recognizing words n Impaired comprehension (failure to grasp
the meaning of words) n Difficulty converting thoughts into
meaningful words
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Wernicke’s Area n Wernicke’s area is
also implicated in Pure Word Deafness
n Uncompromised recognition of non-speech sounds and intonation.
n Caused by disruption of auditory input to Wernicke’s area, or damage to Wernicke’s area itself
Transcortical sensory aphasia n Wernicke’s aphasics can’t understand
the meaning of words or “translate” their thoughts into meaningful words.
n This seems to be due to trauma to the ‘posterior language area’.
n Damage to just this area often results in transcortical sensory aphasia.
n These patients can recognize words: they can repeat back what you say… but can’t make ‘meaning’.
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Language Areas
Conduction Aphasia
n The fact that transcortical aphasia patients can perform repetition suggests that there is a direct connection between Wernicke’s area and Broca’s area
n This is known as the arcuate fasciculus
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Conduction Aphasia
n Conduction aphasia patients – speak fluently – have pretty good comprehension – Often perform well on repetition tasks, as
long as the sounds have meaning – Often fail at longer repetition tasks,
repeating the gist of a sentence but with different words
The arcuate fasciculus
n A bundle of axons that seems to bring information from Wernicke’s area to Broca’s area about the sounds of words (but not their meanings!)
n Conduction aphasia patients speak fluently, have pretty good comprehension, but fail at repetition tasks… suggesting that the AF is important in STM of words and recently heard speech sounds
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Conduction aphasia
Anomic aphasia
n Speech of anomic aphasics is fluent and grammatical, and their comprehension is fine… but they appear to have difficulty finding the right words.
n Fluent anomia is caused by posterior lesions to the temporal or parietal lobes.
n Patients adopt circumlocutions: alternative ways of saying what they mean
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Next time…