,Inhoud
Task 1. Working Memory ........................................................................................................................ 6
Behavioral processes ........................................................................................................................... 6
Working memory............................................................................................................................. 7
Cognitve control .................................................................................................................................. 8
Controlled updating of short-term memory buffers ....................................................................... 8
Setting goals and planning .............................................................................................................. 9
Task switching ................................................................................................................................. 9
Stimulus selection and response inhibition .................................................................................. 10
Brain substrates ................................................................................................................................. 10
Frontal lobes and consequences of frontal-lobe damage ............................................................. 10
Frontal brain activity during working-memory tasks .................................................................... 11
Mapping executive processing and working memory onto PFC anatomy ........................................ 13
Maintenance (rehearsal) versus manipulation (cognitive control) ............................................... 13
Visuospatial and phonological-verbal working memory ............................................................... 13
The neural bases of state-based accounts of working memory ................................................... 13
Goal abstraction and frontal-lobe organization ............................................................................ 13
Prefrontal control of long-term declarative memory ................................................................... 14
Clinical perspectives .......................................................................................................................... 14
The prefrontal cortex in schizophrenia ......................................................................................... 14
Attention deficit/hyperactivity disorder (ADHD) .......................................................................... 14
Artikel: Neuron Activity Related to Short-Term Memory (Fuster, M & Garret, E)............................ 15
Internetlink: The End of IQ (and the Dawn of Working Memory)..................................................... 17
Internetlink: How Major Depression Impaired My Cognitive Ability ................................................ 17
Supporting document: encoding, storing, & retrieving memories ................................................... 17
Task 2. Long-term memory and consolidation ...................................................................................... 18
Behavioral processes ......................................................................................................................... 18
Encoding new memories ............................................................................................................... 19
Retrieving existing memories ........................................................................................................ 19
When memory fails ........................................................................................................................... 19
Memory consolidation and reconsolidation ..................................................................................... 19
Brain substrates ................................................................................................................................. 20
Neural networks for semantic memory ........................................................................................ 20
The medial temporal lobes in memory storage ............................................................................ 20
The hippocampus and cortex interact during memory consolidation .......................................... 21
Many kinds of brain damage can impair memory ............................................................................ 22
1
, Damage to the medial diencephalon can also prevent formation of new memories .................. 23
Patients with Korsakoff’s syndrome show damage to midline diencephalic structures and to the
frontal cortex. ................................................................................................................................ 23
Alzheimer’s disease ....................................................................................................................... 24
Memory and aging ........................................................................................................................ 24
Different kinds of learning and memory ....................................................................................... 24
Task 3: Consolidation, plasticity and LTP............................................................................................... 26
The synapse: where neurons connect............................................................................................... 26
Causing changes in neural connections ............................................................................................ 26
Long-term potentiation and long-term depression .......................................................................... 27
New experimental techniques enable rapid progress in research on mechanisms of memory....... 28
Different kinds of neural circuits may underlie memories............................................................ 28
The nervous system may form and store memories in various ways ............................................... 28
Cerebral changes result from training........................................................................................... 29
Training and experience cause the brain to change chemically and anatomically ....................... 29
Learning can produce new synaptic connections ......................................................................... 30
NMDA receptors and AMPA receptors play separate roles in the induction of LTP in the CA1
region............................................................................................................................................. 31
LTP is induces via a cascade of neurochemical steps ........................................................................ 32
Supporting document: cellular consolidation ................................................................................... 34
Cellular consolidation goes in two stages ..................................................................................... 34
The relationship between genes and long-term plasticity ............................................................ 34
Long-term potentiation ................................................................................................................. 35
Can synapses also become weakened?......................................................................................... 35
Supporting document: is the hippocampus more than a tutor? ...................................................... 36
The hippocampus as a spatial map. Preliminary evidence from unit activity in the freely-moving rat
........................................................................................................................................................... 37
Task 4: Classical conditioning ................................................................................................................ 38
Basic concepts of classical conditioning ............................................................................................ 38
Eyeblink conditioning (conditioning of motor reflexes) ............................................................... 39
Refining the basic principles .............................................................................................................. 39
Conditioned compensatory responses .......................................................................................... 39
Extinguishing and old association ................................................................................................. 39
Compound conditioning and overshadowing ............................................................................... 40
Error correction and the modulation of US processing .................................................................... 40
Kamin’s blocking effect ................................................................................................................. 40
The Rescorla-Wagner model of conditioning ................................................................................ 40
2
, Error correction in human category learning ................................................................................ 41
Stimulus attention and the modulation of CS processing ................................................................. 42
Brain substrates ................................................................................................................................. 43
Mammalian conditioning of motor reflexes ................................................................................. 43
Electropfysiological recording in the cerebellum .......................................................................... 44
Cellular perspective ....................................................................................................................... 45
Clinical perspectives .......................................................................................................................... 47
Classical conditioning in tolerance to addictive drugs .................................................................. 47
Reducing medication through classical conditioning .................................................................... 48
Supporting document: explanation details of Purkinje cell depression ........................................... 49
Cerebellum .................................................................................................................................... 49
Anatomy ........................................................................................................................................ 49
Task 5: Emotional learning and memory............................................................................................... 51
What is emotion ................................................................................................................................ 51
Autonomic arousal and the fight-or-flight response ..................................................................... 51
Theorieën van emoties .................................................................................................................. 51
Assessing emotion in nonhuman animals ......................................................................................... 52
Learning emotional responses: focus on fair .................................................................................... 52
Learned helplessness..................................................................................................................... 53
Effect of emotions on memory storage and retrieval ....................................................................... 54
The amygdala: a central processing station for emotions ................................................................ 55
Two pathways for emotional learning in the amygdala ................................................................ 56
The amygdala and episodic memory storage................................................................................ 57
The role of stress hormones .......................................................................................................... 57
Retrievel and reconsolidation ....................................................................................................... 58
Encoding emotional contexts in the hippocampus ........................................................................... 58
Feelings and the frontal lobes ........................................................................................................... 59
Clinical perspectives .......................................................................................................................... 59
Phobias .......................................................................................................................................... 59
Posttraumatic stress disorder ....................................................................................................... 59
Artikel: B-Adrenergic activation and memory for emotional events ................................................ 60
Task 6: Instrumental conditioning ......................................................................................................... 61
The discovery of operant conditioning ............................................................................................. 61
Components of the learned association. .......................................................................................... 62
Discriminative stimuli .................................................................................................................... 62
Response ....................................................................................................................................... 62
3
, Reinforcers .................................................................................................................................... 63
Punishers ....................................................................................................................................... 63
Putting it all together: building the SD->R->O association ................................................................ 64
Timing affects learning .................................................................................................................. 64
Outcomes can be added or subtracted ......................................................................................... 64
Reinforcement need not follow every response ........................................................................... 64
Choice behavior ................................................................................................................................. 65
The dorsal striatum and stimulus-response (SD -> R) learning ......................................................... 66
The orbitofrontal cortex and learning to predict outcomes ............................................................. 67
Mechanisms of reinforcement signaling in the brain ....................................................................... 67
Dopamine: how the brain signals ‘’wanting’’ ................................................................................ 67
Endogenous opioids: how the brain signals liking......................................................................... 69
How do liking and wanting interact?............................................................................................. 69
Clinical perspectives .......................................................................................................................... 69
Drug addiction ............................................................................................................................... 69
Behavioral addiction ...................................................................................................................... 70
Treatments for addiction............................................................................................................... 70
Artikel: Internet Addiction: A Brief Summary of Research and Practice ........................................... 70
Task 7: Skill learning .............................................................................................................................. 72
Encoding new memories ................................................................................................................... 72
More repetition does not guarantee improvement ..................................................................... 72
Time and sequencing of practice matters ..................................................................................... 73
Skill memories are often formed unconsciously ........................................................................... 73
Expertise requires extensive practice............................................................................................ 73
Talent takes time to blossom ........................................................................................................ 74
Retrieving existing memories ............................................................................................................ 74
When memory fails ........................................................................................................................... 75
The basal ganglia and skill learning ................................................................................................... 75
Learning deficits after lesions........................................................................................................ 75
Neural activity during perceptual-motor skill learning ................................................................. 75
Brain activity during cognitive skill learning .................................................................................. 76
Cortical representations of skills ....................................................................................................... 76
Cortical expansion ......................................................................................................................... 77
Are skill memories stored in the cerebral cortex? ........................................................................ 77
The cerebellum and timing................................................................................................................ 77
Parkinson’s disease ........................................................................................................................... 78
4
, Human-machine interfaces: learning to consciously control artificial limbs .................................... 79
Artikel: Where practice makes perfect in texture discrimination: Evidence for primary visual cortex
plasticity ............................................................................................................................................ 79
Task 8: Learning by example ................................................................................................................. 80
Copying what is seen ......................................................................................................................... 80
The social learning theory ............................................................................................................. 80
Studies of true imitation: copying actions..................................................................................... 80
Studies of emulation: copying goals.............................................................................................. 81
Reproducing actions without copying ........................................................................................... 81
Social transmission of information.................................................................................................... 81
Learning through social conformity .............................................................................................. 82
Effects on media on behavior........................................................................................................ 82
Brain substrates ................................................................................................................................. 83
Mirror neurons .................................................................................................................................. 83
Song learning in bird brains ............................................................................................................... 84
Hippocampal encoding of socially transmitted food preferences .................................................... 84
Autism spectrum disorder ................................................................................................................. 84
Imitative deficits after stroke ............................................................................................................ 85
Artikel: Understanding motor events: a neurophysiological study .................................................. 85
5
,Task 1. Working Memory
Behavioral processes
Working memory: het actief behouden en manipuleren van het kortetermijngeheugen.
Cognitive control: de manipulatie en toepassing van het werkgeheugen op plannen, wisselen van
taken, aandacht, stimulus selectie en de inhibitie van ongepaste reflexen.
Er worden vaak drie soorten geheugens onderscheden van elkaar: het sensorische geheugen
(automatische informatie dat snel verdwijnt), het kortetermijngeheugen (blijft zolang de informatie
wordt herhaald, of zolang aandacht besteedt wordt aan de informatie), en het langetermijngeheugen
(informatie kan heel lang of voor eeuwig worden teruggehaald). Hieronder het Model of Memory
(Atkinson-Shirffrin). In dit model is te zien wat de chronologische volgorde is van het geheugen, het
begint dus met de input en kan (als je er iets mee doet in het werkgeheugen) leiden tot een herinnering
in het langetermijngeheugen.
Dit model kent twee transient memories (kortdurende mentale representaties, vaak van maar enkele
seconden): het sensorische- en kortetermijngeheugen. Het sensorische geheugen is alles wat je ziet,
voelt of proeft, dit heeft ongeveer een capaciteit van vier of vijf items.
Short-term memory
Het kortetermijngeheugen (KTG)
is ook gelimiteerd, en tot wel 7
items (-2 & +2). Informatie komt
hier terecht wanneer we
informatie uit het sensorische
geheugen blijven herhalen. Dit
geheugen betreft actieve
content van bewustzijn, is snel
toegankelijk, maar ook weer snel
vergeten. Hiernaast de verschillen tussen het kortetermijngeheugen en het langetermijngeheugen.
Het KTG is ook beperkt tot waar de aandacht op richten. Wanneer we snel even afgeleid zijn, kan
informatie in het KTG verdwijnen. We kunnen meer onthouden door de manier waarop we iets willen
onthouden aan te passen (encoding). Bijvoorbeeld aan de hand van geheugensteuntjes of het
samenvoegen van items in één chunk (chunking).
6
, Working memory
Wanneer informatie in ons KTG gebruikt wordt om informatie een bepaalde tijd te onthouden,
gebruiken we ons werkgeheugen. Dit geheugen werkt als een buffer waarin tijdelijk informatie kan
worden opgeslagen zolang we er actief mee bezig zijn. Een bekend model over het werkgeheugen is
die van Alan Baddeley, het working-memory model. Hieronder een figuur waarin het model
beschreven wordt
Dit model kent twee onafhankelijke buffers van het werkgeheugen: de phonological loop (manipulatie
van auditieve stimuli) & de visuospatial sketchpad (manipulatie van visuele beelden). De central
executive monitort en manipuleert beide buffers en brengt cognitieve controle aan het
werkgeheugen. Deze voegt items toe aan de buffers of verwijdert items waarnaar ze wellicht
doorgestuurd kunnen worden naar het LTG.
In dit model worden twee belangrijke zaken onderscheden. De eerste is het verschil tussen manipulatie
(dit hangt af van de central executive) en behouden (dit vergt herhaling van informatie in de twee
buffers). Ten tweede, informatie wordt los onthouden, de een bevat informatie over geluiden en de
ander informatie over locaties en objecten. Hieronder worden de twee buffers nader besproken
Informatie in de phonological loop blijft meestal maar 2 seconden hangen wanneer er niet herhaald
wordt. Dit zorgt ervoor dat je 7 items ongeveer kan onthouden. Het word-length effect is dat de
meeste mensen denken dat, des te langer woorden zijn die onthouden dienen te worden, hoe
moeilijker het wordt. Dit staat afhankelijk van het visuospatial sketchpad. Dit is bijvoorbeeld gebleken
uit onderstaande onderzoeken.
7
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