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Medicinski fakultet Sveučilišta u Zagrebu. Učenje i pamćenje – pregled najvažnijih spoznaja i koncepata. Izborni predmet Učenje i pamćenje. Goran Šimić 10. travnja 2018. K rivulje zaboravljanja.
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Medicinski fakultet Sveučilišta u Zagrebu Učenje i pamćenje – pregled najvažnijih spoznaja i koncepata Izborni predmet Učenje i pamćenje Goran Šimić 10. travnja 2018.
Krivulje zaboravljanja S obzirom da je proces zaboravljanja najbrži tamo gdje je krivulja najstrmija, poslije usvajanja sadržaja prvo se ponavljanje preporučuje napraviti što ranije (najkasnije u prvih 24 sata nakon učenja). Ebbinghaus H. Translated by H.A. Ruger and C.E. Bussenius. New York: Teachers College, Columbia University; 1885. Memory: a contribution to experimental psychology. (Originally in German, Ueber das gedächtnis: Untersuchen zur experimentellen psychologie). Ann. Neurosci. 2013; 20: 155-156.
Klasični model triju skladišta pamćenja 15-30 s Fonološka petlja Vidno-prostorna crtanka Ikoničko (< 1 s) Ehoičko (1-4 s) Haptičko (1-10 s) Upamćivanje Skladištenje Prisjećanje Atkinson RC, Shiffrin RM (1968). "Chapter: Human memory: A proposed system and its control processes". U: Spence KW, Spence JT.The psychology of learning and motivation.
http://www.abacon.com/slavin/t62.html http://www.abacon.com/slavin/t62.html
Što je radno pamćenje? Crystallized systems Fluid systems Working memory has been conceived and defined in three different, slightly discrepant ways: as short-term memory applied to cognitive tasks, as a multi-component system that holds and manipulates information in short-term memory, and as the use of attention to manage short-term memory (Cowan N. What are the differences between LTM, STM, and working memory? Prog Brain Res 2008; 169: 323-338.) Atkinson RC, Shiffrin RM (1968). "Chapter: Human memory: A proposed system and its control processes". U: Spence KW, Spence JT.The psychology of learning and motivation.
salamander turtle man lizard oposum monkey hedgehog
Heteromodalna (transmodalna) asoc. polja omogućuju kogniciju • Građena od homotipnog izokorteksa • Prefront., dorzolat. i orbitofront. cx (BA9 i 10, BA45-47, BA8, 11 i 12), stražnji parijet. cx (BA39-40) i lat. sljepoočni cx (BA20, 21, 37, 38) • Povezujući osjetne inform. s emocijama, nagonima i strategijama djelovanja omogućuju intermedijarno (integrativno) procesiranje (kogniciju), tj. povezuju modalitet-specifične fragmente informacija u cjelovita iskustva, sjećanja i misli (tako se ostvaruje transcedencija instinktivnog ponašanja, te povećava repertoar ponašanja koje je u skladu s kontekstom)
Razlike u duljini trajanja neurogeneze Mus musculus Macaca mulatta Homo s. sapiens
Vrste pamćenja MTL + diencephalon eksplicitno implicitno neocx amygdala Cerebellum simple conditioning neocx neocx striatum
Vrste DUGOTRAJNOG pamćenja DEKLARATIVNO (EKSPLICITNO) - asocijacije istovremenih podražaja - semantičko (referentno) (znam što je to gitara) višestruko je reprezentirano u moždanoj kori i temelji se na plastičnosti asocijativnih područja korteksa - epizodičko (autobiografsko)(sjećam se kad sam kupio svoju prvu gitaru) zavisi isljučivo o djelovanju hipokampalne formacije (činjenice), pa je i vulnerabilnije često sadrži afektivnu (emocionalnu) komponentu (kontekst) koja se pamti odvojeno (“source amnesia” - amnezija izvora, naročito kad je izvor nepouzdan, npr. mediji) REFLEKSIVNO (IMPLICITNO) - ne ovisi o svijesti i pažnji - ne može se verbalizirati, pa ga je teško opisati, npr. kako svirati gitaru - vještine i navike - jednostavno kondicioniranje - neasocijativni oblici učenja (habituacija, senzitizacija)
Funkcija HF 1888. Brown i Schäffer; 1900. Bechterew 1930-1950. Penfield i Rasmussen Bilateral removal of HF: long-lasting inability to form long-term memories (global amnesia) 1953. patient H.M. (Scoville, Milner, Corkin) 1975. patient N.N. (Damasio, Damasio) 1978. patient R.B. (Squire, Zola-Morgan, Amaral) Unilateral HF damage - partial compensation (left: lowered ability to make long-term verbal memories; right: deja vu syndrome, aff. visuo-spatial capabilities)
Kako nastaje epizodičko pamćenje? Duvernoy Niuewenhuys et al.
Uloga NMDA receptora u LTP • NMDA receptor se ponaša kao detektor spregnutog djelovanja (asocijativnost), jer za indukciju LTP zahtijeva: • vezanje za receptorno mjesto (specifičnost) i • prethodnu ili istovremenu depolarizaciju postsinaptičke membrane (da bi se odstranili ioni Mg2+) (kooperativnost) • retrogradni difuzibilni signal (NO) omogućuje potencijaciju susjednih sinapsi
Promjene sinaptičke učinkovitosti HF • 1949. Hebb: hipoteza dvostrukog traga • 1973. Bliss i Lomo: LTP (“vanilija”) • 1991. Llinas: “čokolada” LTP
Dokaz uloge LTP u učenju Injiciranje antagonista NMDA receptora (2-amino-5-fosfonovalerične kiseline): • spriječava nastanak LTP u bloku (lameli) hipokampalnog tkiva, • spriječava učenje na razini ponašanja (Morrisov test pronalaženja platforme uronjene u vodi) - oba učinka razmjerna su dozi (Morris, 1983)
Koliko je vremena potrebno za trajno zapamćivanje? Nakon stjecanja iskustva za konsolidaciju zapamćenog potrebno je najmanje 6-8 tjedana (Squire, Zola-Morgan i Amaral, 1989-1992) HF: 1. Povezuje informacije iz različitih područja Cx za vrijeme učenja, 2. Nakon razdoblja od 6-8 tjedana konsolidira navedene informacije u neokorteks tako da postaju dostupne i neovisno od HF
Fiziološki ritmovi HF i hipoteza učenja u 2 koraka (Buzsaki et al. 1973-2007) 1. Korak: “istraži i iskusi” - theta ritam 2. Korak: “odmori se i konsolidiraj” - oštri valovi (za vrijeme spavanja) - ‘hipokampalna kompresija vremena’ Svojstva CA3/CA1 mreže (autoasoc. matriksa) - učenje iz jednog pokušaja! - generalizacija! - djelomična otpornost na šum! Shema autoasoc. matriksa (Rolls, 2005) Buzsaki, Rolls et al.
Ponavljanje, konsolidacija i spavanje (Buzsaki, Kolb et al.)
How do we forget? • Herman Ebbinghaus, 1885 • subjects memorize a list of meaningless, three letter words • tracked how quickly his subjects forgot the words • became known as the Ebbinghaus or Forgetting Curve Graphic Source: www.york.ac.uk/depts/maths/histstat/people/ebbinghaus.gif
Ebbinghaus Curve: www.psych.purdue.edu/~ben/285su2001/notes/figures/5-forgettingcurve.htm
How do we forget? Forgetting curve would start here if we could remember everything after a lecture However, you have the potential to forget less PLUS remember more if you review immediately after class Forgetting curve would actually start here as we typically remember only about 75% at the end of a lecture – so we have less to remember
Overcoming the Curve Immediately after class 24 hours later 1 week later (or sooner) 1 month later (or sooner) Notice how less is forgotten after each review!!
So Review • 10 min - After class (or in evening) by completing, organizing & comprehending (rewriting, typing) notes (e.g. Cornell right column) • 24 hr – Next day, before Ebbinghaus kicks in, reread notes, condense to main ideas & create questions (e.g. Cornell left column) • 1 week - Before class the following week or earlier, review and self-test your recall (e.g. Cornell, cover right column & self-test from left) • Therefore, when you prepare to study for your midterms and finals, you have already reviewed the material a minimum of 3 times
Kako pospješiti učenje (4) V4 ljudi oruđa životinje • Epicentri u lateralnom temp. i temporopolarnom cx • BA37 i 20 (prednji i srednji dio g. fuziformisa) – percepcija objekata i lica, malo lateralnije, na granici BA19 i BA37 – percepcija Riječi • Vidna integracija je stupnjevit proces: nepoznata lica aktiviraju samo unimodalne areje u i oko g. fuziformisa, poznata lica aktiviraju i transmodalne areje (naročito u g. temp. med.) motorika “GDJE” PUT R V3 V3 V5 R V2 V5 R “ŠTO” PUT kognicija pamćenje • akromatopsija, anomija za boje, akinetopsija • vidna objektna agnozija, čista aleksija (jače lij.) • asocijativna prozopagnozija (jača desno) • vidna amnezija (lijevo), vidna hipoemoc. (desno) L
Sustav zrcalnih neurona (“mirror eurons”) kodira ne samo opaženu radnju, nego i očekivano djelovanje
Kako pospješiti učenje (7) Kolb et al.
Damage to the hippocampal formation interferes with contextual fear associations • Panic disorder, PTSD, obsessive-compulsive disorder, anxiety and fobias are all examples of classical fear conditioning Kolb B, Whishaw I (2008) Fundamentals of human neuropsychology.
a) fear conditioning b) instructed fear c) observational fear learning Three different ways by which humans can learn about the aversive properties of an event. In all of them the activation of amygdala is seen. Le Doux JE (2000) Emotion circuits in the brain. Annu Rev Neurosci 24: 155-184; Gazzaniga et al., 2002.
Auditory fear conditioning Le Doux 1992; Davies 1992; Kandell 2000.
Learned helplessness (naučena bespomoćnost) Escape learning Avoidance learning Seligman M et al.,
- Loss of fear 100% - Oral examination of objects 98% - Hypersexuality 79% - Hypermetamorphosis 77% - Visual agnosia 42% - Bulimia 28%
Concept of “high” and “low” road of emotional processing “Low road” is faster and unconscious; it depends on projections from thalamus to amygdala. “High road” is slower and conscious; it depends on projections from OFC to amygdala. Decreased activity of OFC has been documented in non-premeditated murderers and schizophrenia Le Doux JE (2000) Emotion circuits in the brain. Annu Rev Neurosci 24: 155-184.
…, but there are huge sex differences In recognition test 3 weeks after seeing pictures, they were remembered much better by women (activating signif. fewer structures and the left AMY) than men. Canli et al. (2002) Sex differences in the neural basis of emotional memories. PNAS USA 99: 10789-10794.
Mesolimbic and mesocortical projection from A10 (VTA) An universal brain rewarding system is composed of dopaminergic projections from ventral tegmental area (VTA) to AMY and OFC (the so-called mesolimbic and mesocortical projections), which release dopamine at higher rate during pleasant experiences of eating food, having sex, bonding with a child, etc. (behavioral activation, step 1)
Olds et al. (1961) Heath et al. (1972) Recently, questions have been raised whether DBS shown here might have produced merely “wanting”, rather than actual “liking”=true pleasure Smith et al. (2008)
Normal circuitry When a reward is greater than expected (such as in case of drugs like cocain, amphetamines, opioids, canabinoids, nicotine, partially alcohol), the firing of certain VTA DA neurons increases, which consequently increases the desire or motivation toward the reward in PFC (OFC). It is the PFC (OFC) glutamatergic projections that regulate activation of motor responses to “motivation-related events” through n. accumbens septi and thalamic motor nuclei, while blocking D1 receptors in PFC (OFC) prevents seeking for a drug. Due to change in regulation of motivation circuit and pathological long-term changes in synaptic plasticity of OFC neurons that project to ventral striatum, in the last phase ofdependence and drug abuse such patients are constantly looking for a drug. Drug dependence Arrias-Carrion and Poppel, Acta Neurobiol. Exp., 2007