Filozofski fakultet u Zagrebu Odsjek za fonetiku Predmet: Neurofonetika prof. dr. sc. Damir Horga
NEUROFONETIKA (2008.) RAZVOJ SPOZNAJA O MOZGU O nervnom sistemu saznavalo se sa sporoš ću koja se može objasniti jedino mnogobrojnim teško ćama s kojima se ta vrsta istraživanja oduvijek sukobljavala. (F.J. Gall, 1825).
1882. godine kolekcionar Edwin Smith kupuje u Luksoru papirus. 1930-ih James Breasted (Sveučilište u Chicagu) dešifrira papirus: 17 stupaca rasprava o kirurgiji mozak se prvi puta spominje pod svojim imenom papirus je iz 17. stoljeća pr. Kr., ali to je vjerojatno kopija iz oko 3 000 godine pr. Kr. 48 slučajeva povreda glave i vrata svaki slučaj ima: naslov, ispitivanje, dijagnozu, liječenje slučaj 6 – ako se otkine dio lubanje vide se «bore slične onima što se stvaraju na rastaljenom bakru slučaj 8 – povreda u lubanji uzrokuje skretanje očnih jabučica i bolesnik hoda vukući stopala slučaj 22 – «ako pregledaš čovjeka kojemu je probijena sljepoočnica... kada ga pozoveš, ne odgovara, izgubio je moć govora». slučaj 31 – poslije povrede vratnih kralježaka, bolesnik je neosjetljiv u obje ruke i obje noge, falus mu je u erekciji, a mokri i ejakulira i ne znajući» sažeti, jasni opisi, liječnik ne pribjegava čaroliji nego kaže: «Nema izlječenja». • • • •
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to je prvi dokument koji spominje upravljačku funkciju mozga nad organima koji su udaljeni od glave.
Ipak, to ne znači da je bila shvaćena uloga mozga u intelektualnim i emocionalnim funkcijama. Srce se smatra izvorom života (Egipat, Mezopotamija, Židovi, Gr čka). U Gr čkoj se od 7. do 5. stoljeća pr Kr. traži jedinstveno tumačenje univerzuma i čovjeka. Materija i duša se ne dijele. Voda, zrak, vatra i zemlja, a s Demokritom i atomi, grade i svijet i čovjeka pa, izgleda, i njegove misli.
Demokrit: Misli i osjećaji imaju materijalnu podlogu i zavise od finih, glatkih i okruglih atoma. To su psihički atomi rasprostrti po cijelom tijelu. Mozak je čuvar misli ili uma, srce je «kraljica i hraniteljica srdžbe», a ognjište želja nalazi se u jetri. Dvije su značajke Demokritova učenja: 1. razlikuje nekoliko intelektualnih i emocionalnih funkcija i razmješta ih u različite dijelove tijela (misli su u mozgu) 2. mozak preko atoma uspostavlja veze s organima i vanjskim svijetom (ideja živčanog sustava). Hipokrat (Periklovo doba, 4. stoljeće pr. Kr.) proučava povrede mozga i zna da izazivaju motoričke smetnje zna za kontralateralnost zna za epilepsiju zna za mentalne i neurološke bolesti i pripisuje im moždano podrijetlo «Mozak je sličan nekakvoj žlijezdi... isto je tako bijel i trošan». •
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Platon:
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To je cefalocentrična teza: središte misli je u čovjekovu mozgu.
Aristotel: Srce je središte osjećaja, strasti i uma. Mozak je sastavljen od vode i zemlje i ima ulogu hladionika organizma – snižava temperaturu krvi opskrbljene hranom i donosi san. Aristotel ne poznaje živce, ali poznaje krvožilni sustav, a on konvergira prema srcu. Zaključuje da je srce središnji organ upravljanja. Mozak nije osjetljiv na mehaničko draženje, a srce jest. Životinje bez kralježnice nemaju mozak. Ipak, kaže da duša nikad ne misli bez slika, one su kopije stvarnosti koje do duše dolaze preko osjetnih organa. Zahvaljujući tim slikama, um može računati i razmišljati o budu ćnosti. Ipak, gr čka medicina ostaje encefalocentrična (Epikur). To se prenosi u Aleksandriju pa se u 3. stoljeću pr. Kr. (Herofil i Erazistrat) otkrivaju nove spoznaje o mozgu. Izvode se sekcije na živim ljudima. Herofil je rasjekao više tisuća ljudskih tijela: veliki mozak, mali mozak, kralježnička moždina mozak ima komore površina je naborana živci izlaze iz mozga i kralježničke moždine razlikuju se motorički i senzorički živci čovjekov mozak je naboraniji od mozga životinja • • • • • •
Galen (2. stoljeće) uspijeva odbaciti Aristotelove teze. Uvodi fiziologiju mozga. Izvodi pokuse. Odvaja komore od moždane supstance koja je slična živcima. Otkriva tri šupljine: prednju (podijeljenu na dvije), srednju i stražnju. Ako životinji povrijedi zadnju komoru, ne može se kretati. Mozak je bitan u upravljanju tijelom i intelektualnim funkcijama. Ali Aristotelovo kardiocentri čno gledište ostaje i dalje vladajuće do 18. stoljeća. Postavlja se pitanje odnosa duše i tijela.
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Razumska funkcija ima imaginaciju, razum i pamćenje. Galen tim funkcijama dodjeljuje razli čita mjesta u mozgu. Kasnije, u 4. i 5. stoljeću, Nemezije iz Emesa i sv. Augustin smještaju ih u komore: imaginacija (prednja), razum (srednja), pamćenje (stražnja).
Srednji vijek i skolastika zaboravljaju na spoznaje o mozgu. Renesansa: interes za tijelo – seciranje životinja i leševa. Leonardo da Vinci radi voštani odljev komora i crta vijuge mozga. Descartes smatra da je čovjek stroj. Organe pokreće esprits animaux koji kao zrak u orguljama dolazi do zra čnica. Ali on je dualist i smatra da je duša besmrtna i nematerijalna. Jedinstvo duše ostvaruje se u pinealnoj žlijezdi koja je u mozgu jedinstvena i u kojoj je veza duha i tijela. To je kartezijansko stajalište. Willis (17. stoljeće) proučava subkortikalne strukture (talamus, korpus kalozum). Razlikuje sivu tvar koja stvara esprits animaux i bijelu tvar preko koje se on prenosi u organe i omogućuje pokretanje i senzoriku. Ostaje na temeljima dualizma i Descartesa, ali misli da je spoj duše i tijela u izbrazdanim tijelima. Gassandi (17. stoljeće) priznaje da i životinje imaju pam ćenje, um i druge intelektualne funkcije. Ne određuje točno mjesto spajanje duše i tijela. U 18. stoljeću počinje se odbacivati Descartesov dualizam i vraća se gr čkim atomistima.
Frenologija - javlja se početkom 19. stoljeća Franz Gall smatra da kod čovjeka postoji više urođenih moralnih i intelektualnih sposobnosti. Navodi popis 27 sposobnosti: s posobnosti: nagon razmnožavanja ljubav prema djeci (materinstvo) • •
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osjećaj za mjesto i odnose u prostoru ali i oholost, želja za autoritetom, slavoljublje, metafizički duh, pjesnička obdarenost, odanost
Kako napraviti mapu mozga? Reljef lubanja ponavlja reljef korteksa. To je kranioskopija. Gall skuplja lubanje zločinaca i biste slavnih ljudi. Za govor je pogodio mjesto. Gall je protjeran iz Beča, ni u Parizu nije prihvaćen. Nastavlja lokalistička nastojanja, ali sjedišta sposobnosti traži u korteksu. Otvara mogućnosti eksperimentiranja na životinjama. Flourens provodi eksperimente odstranjujući dijelove korteksa (mali mozak – nema koordinacije pokreta, stražnja komora – disanje). Ipak, zaključuje da funkcije ne nestaju potpuno, da je kora jedinstvena i zato se u njoj može smjestiti duša. Ali njegovi su eksperimenti: 1. zahvaćali i subkortikalne s ubkortikalne strukture 2. bili izvođeni na pticama i nižim kralježnjacima kod kojih je diferencijacija funkcija manja nego kod čovjeka 3. promatranje funkcija nije bilo strogo praćeno. Polovinom 19. stoljeća omeđuju se dijelovi korteksa Silvijusovom i Rolandovom brazdom. Ograničavaju se režnjevi.
Bouillaud (1825) promatra oštećenja mozga i njihov utjecaj na govor. To je početak anatomske patologije govora koja će prerasti u neuropsihologiju. neuropsihologiju. Piše: «Gubitak govora ovisi čas o gubitku pamćenja riječi, čas o gubitku mišićnih pokreta iz kojih se to pamćenje sastoji». Ti nalazi Bouillauda ne uspijevaju se nametnuti znanstvenoj javnosti. Broca (18. travnja 1861.) u Pariškom antropološkom društvu iznosi slučaj pacijenta Leborgnea (21 godinu patio od afazije) čiju je autopsiju obavio večer prije toga. Lokalizira funkciju za govor u tre ćoj vijuzi čeonog režnja lijeve hemisfere. Još 8 pacijenata – kod svih, osim jednoga, centar za govor je u lijevoj hemisferi. Motorika govora.
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Unitaristički (holistički) pogledi smatraju da mozak ipak funkcionira kao cjelina.
ORGANIZACIJA ŽIVČANOG SUSTAVA 1.1. Odnos veličine mozga i funkcija Veličina mozga je važna za njegovu funkciju. a) ontogenetski - novorođenče - 11 mjeseci - 3 godine - odrasla osoba
400 grama 850 grama 1110 grama 1450 grama
b) starenje Nakon 30-te godine mozak se «suši» i do 75-te izgubi prosječno 100 grama c) filogenetski - čovjekoliki majmun - čovjek s Jave - pekinški čovjek
400 grama 800 grama 1100 grama
Veličina mozga nije bitna za njegovu funkciju a) neandertalac i kromanjonski čovjek imali su veći mozak od današnjeg čovjeka - zbog procesa domestifikacije, specijalizacije, ovisnosti o drugim članovima zajednice - domaće, pripitomljene životinje imaju manji mozak od divljih životinja b) osobe koje imaju veće tijelo obično imaju i veći mozak (muškarci – žene) c) nadareni nemaju veći mozak od prosjeka - veličina mozga varira od 1100 do 2000 grama d) delfini imaju relativno ve ći mozak od čovjeka Dakle, težina nije presudna!
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Važna je struktura i organizacija. Broj neurona (180 milijarda), broj neurona koji obrađuju obavijesti (50 milijarda), međusobna povezanost neurona (15 000 fizičkih veza s drugim neuronima). Znanje o anatomskim strukturama mozga prva je stepenica u razumijevanju njegovih funkcija općenito, a to znači i jezičnih i govornih funkcija.
1.2. Podjela živčanog sustava a) CNS, SŽS - kralježnica, lubanjske kosti kralježnička moždina i mozak b) Periferni živčani sustav – izvan tih kostiju Somatski (veza s vanjskom okolinom): eferentni i aferentni živci Vegetativni (autonomni) regulacija mehanizama unutar organizma: aferetni i eferentni živci Autonomni živčani sustav: eferentni živci: simpatički (lumbalni i torakalni) - angažiraju zalihe energije - uzbuđenost parasimpatički (iz mozga i sakralni) - konzerviraju energiju - psihološka opuštenost Svaki organ prima podražaje obiju vrsta, a reakcija ovisi o relativnom odnosu simpatičkog i parasimpatičkog djelovanja. Simpatički živci prekapčaju se daleko od ciljnog organa. Parasimpatički živci prekapčaju se blizu ciljnog organa. Živci izlaze iz kraljžni čke moždine
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Pia mater – meka/tanka ovojnica, uz živčano tkivo Cerebrospinalna tekućina Subarahnoidni prostor Centralni kanal kralježničke moždine Moždane komore (2 lateralne, treća i četvrta) Ako se odstrani dio likvora, pacijenti osjećaju bol pri najmanjem pokretu. Likvor se proizvodi u koroidnim pleksusima. Hidrocefalus – vodena glava; proširenje zidova komora.
1.4. Krvno-moždana barijera Krvne žile u mozgu imaju gusto zbijene stijenke te, za razliku od drugih žila u tijelu, ne propuštaju štetne molekule u okolno moždano tkivo. Neke se velike molekule propuštaju (glukoza, hormoni u određenim dijelovima mozga).
1.5. Kralježnička moždina ( medulla medulla spinalis ) U presjeku ima dva dijela: - unutarnji sivi u obliku slova H (siva tvar, tijela neurona) - vanjski bijeli (bijela tvar, mijelizirani aksoni) Stražnji dorsalni rogovi (senzorički) Prednji ventralni rogovi (motorički) 31 par spinalnih, moždinskih živaca pred kralježničkom se moždinom dijeli na prednji ventralni korijen i stražnji dorsalni korijen.
1.6. Pet glavnih dijelova mozga
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srednji mozak stražnji mozak telencephalon (krajnji mozak) diencephalon (međumozak) mesencephalon (srednji mozak) metencephalon myelencephalon
encephalon – «unutar glave»
- moždane hemisfere (telencephalon) moždano deblo - mesencephalon - metencephalon - myelencephalon 1. Produljena moždina (myelencephalon, medulla oblongata) - putovi između mozga i ostatka tijela - retikularna formacija – reticulum - mrežica 100 jezgara - od stražnje granice produljene moždine do prednje granice srednjeg mozga - retikularni aktivacijski sustav - spavanje, pažnja, pokret, održavanje tonusa mišića, sr čani, cirkulatorni i dišni refleksi 2. Metencephalon
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- Periakvaduktalna siva tvar – oko akvadukta koji spaja treću i četvrtu komoru (analgetski učinak) - Substantia nigra (crna tvar) – komponente senzomotoričkog sustava - Nucleus ruber (crvena jezgra) – komponente senzomotoričkog sustava 4. Međumozak ( diencephalon diencephalon) - talamus – parna struktura na vrhu moždanog debla - po jedan režan sa svake strane treće moždane komore, spaja ih massa intermedia
- na površini su bijeli aksoni - brojni parovi jezgara, to su releji koji impulse iz periferije prenose u korteks - lateralno koljenasto tijelo ( corpus geniculatum laterale) – vid - medijalno koljenasto tijelo (corpus geniculatum mediale) – sluh - ventroposterolateralna jezgra – senzomotorika - hipotalamus – regulacija potreba - otpuštanje hormona iz hipofize (pituitarna žlijezda – šmrkava žlijezda) - optička hijazma – križanje vidnog živca (ipsilateralno – kontralateralno dekusacija) - mamilarna tijela telencephalon) 5. Krajnji mozak ( telencephalon
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kora velikog mozga (cortex cerebri) naborana sulcus (brazda) – vijuge ( gyri gyri) fisura (pukotina) – sulcus fissura longitudinalis moždane komisure, corpus callosum sulcus centralis fissura lateralis
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- hipokampus (morski konjić) – medijalni rub korteksa i zaokreče u temporalni režanj
Limbički sustav i bazalni gangliji Subkortikalni prostor čine aksoni i grupe jezgara. Limbički sustav (rubni – oko talamusa) - instinkti, bijeg, obrana, hranjenje, seksualno ponašanje - amigdala (bademasti oblik) - gyrus cinguli - fornix (svod) - septum Bazalni gangliji (voljna motorika – Parkinsonova bolest) Amigdala Nucleus caudatus (repat) – putamen = striatum (pruga) Globus pallidus (blijed)
STANICE U ŽIVČANOM SUSTAVU Živčane stanice ili neuroni stanice su specijalizirane za primanje, provođenje i prijenos elektrokemijskih signala.
Građa živčanih stanica - multipolarne – s više izdanaka koji se odvajaju od tijela neurona - bipolarne – s dva izdanaka - unipolarne – s jednim izdankom - interneuroni – s kratkim aksonima ili bez aksona; integriraju aktivnost neke neuralne strukture i nije im zadaća da prenose impuls jezgre ili nukleusi – nosioci DNK ili nakupine naku pine tijela neurona u CNS-u, a
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Neuroanatomske tehnike Golgijeva metoda 1870 Nisslova metoda 1880 Elektronska mikroskopija Bojenje mijelina – boji mijelin, ali se ne mogu pratiti pojedini aksoni a ksoni Neuroanatomske tehnike – prati akson
Električna aktivnost neurona Potencijal u mirovanju - negativni ioni: An (organski), Cl - pozitivni ioni: Na, K Membrana je izolator: 1. An ioni ostaju u intracelularnom prostoru 2. polupropusna je za Na i K ione i lakše propušta K ione 3. membrana ima Na-K pumpu koja omogućuje da se intracelularni Na zamijeni ekstracelularnim K. Na se koncentrira s vanjske strane membrane Rezultat tih triju procesa je da u unutrašnjosti membrane ima 350 puta više An iona i 20 puta više K iona nego na vanjskom dijelu membrane, 9 puta više Cl
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Budući da do izjednačavanja potencijala ne dolazi, pretpostavlja se da postoji Na-K pumpa koja dozvoljava zamjenu 2 Na iona za 3 K iona. Ioni Cl slobodno prolaze. Unutrašnjost membrane tako postaje 70 mV (stvarno 55 do 90 mV) negativnija od vanjštine. To je potencijal u mirovanju.
Stimulacija membrane Mijenja se njezina propusnost. Depolarizacija – unutrašnjost postaje manje negativna ili čak pozitivna u odnosu na potencijal u mirovanju zbog većeg ulaska Na iona. Hiperpolarizacija – unutrašnjost postaje još negativnija zbog većeg izlaska K iona ili većeg ulaska Cl iona.
Akcijski potencijal Kod depolarizacija od -50 mV membrana postaje propusna za ione Na (ulaze) i K (izlaze). Akcijski potencijal +50 mV.
Provođenje impulsa Impuls teče uzduž aksona. Brzina 1 do 100 m/sek.
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- inhibitori
PRINCIPI MOŽDANE ASIMETRIJE Od druge polovine 19. stoljeća i Broce, poznato je da je lijeva hemisfera dominantna za jezične i govorne funkcije. Početkom 20. stoljeća, Liepmann je, uspoređujući pacijente s lezijama lijeve i desne hemisfere, pokazao da u kontroli pokreta lijeva hemisfera ima funkcije koje nisu specifične za desnu. Polovinom 20. stoljeća počinju se otkrivati vizualno-spacijalne vizualno-spacijalne sposobnosti desne hemisfere.
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1. ANATOMSKA ASIMETRIJA U LJUDSKOM MOZGU 1860. Hughlings-Jackson, Hughlings-Jackson, Gratioler - izbrazdanost lijeve lijeve hemisfere razvija se brže nego desne. 1960. Geschwind i Levitsky – otkrivaju razlike u veličini sljepoočne površine ( planum planum temporale ). Osam glavnih anatomskih razlika: 1. Desna hemisfera je malo veća i teža. Lijeva hemisfera ima veću specifičnu težinu. 2. Razlika u građi čeonog ražnja Temporalna površina ( planum planum temporale ) veća s lijeve strane; Heschlov girus veći s desne strane. 3. Razlike u sljepoočnom režnju projiciraju se i na talamus. Lateralni posteriorni nukleus veći s lijeve strane – planum planum temprale;
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7. Desna je dulja prema natrag, lijeva prema naprijed. To se odnosi i na lateralne komisure. 8. Razlike na razini neuronske organizacije; razgranatost dendrita je različita.
1.1. ASIMETRIJA KOD NEUROLOŠKIH PACIJENATA 1. Lateralizirane lezije – dvostruka disocijacija Povrede lijeve – jezik Povrede desne – glazba, prostorni odnosi, pjevanje, sviranje instrumenata. 2. Komisurotomija – razdvojeni mozak - presijecanje žuljevitog tkiva (korpus kalozum) – 200 milijuna živčanih vlakana. - radi se kod pacijenata koji pate od epilepsije (1940). Tahistoskopski eksperiment.
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Dikaptičko prepoznavanje predmeta (lijeva ruka) i slova (desna ruka). Prostorni zadatak i sekvencijalni: stimuliranje prstiju (lijeva ruka bolja u prostornom, a desna u sekvencijalnom zadatku). 4. Motorički sustav a) neposredno promatranje - gestikulacija uz verbalne zadatke i govor – desna ruka aktivnija; - slaganje – slagalica: neutralno, verbalno, spacijalno – za verbalni zadatak desna ruka aktivnija, za neverbalni lijeva. - pokreti usana u govoru – desna strana lica aktivnija b) interferencijski zadaci - držanje ravnoteže štapa na kažiprstu - sviranje različitih melodija istovremeno lijevom i
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različitim aspektima (multikanalno procesiranje) - hemisfere se međusobno inhibiraju; ako je dominantna hemisfera oštećena, funkcija se može razviti i u nedominantnoj - informacijsko procesiranje određeno je usmjerenom pažnjom hemisfera. 3. Preferirani kognitivni način Lijevohemisferalni Lijevohemisferalni način Desnohemisferalni način VARIJACIJE CEREBRALNE ASIMETRIJE Postoje li individualne razlike u cerebralnoj asimetriji?
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Dakle, lateralizacija govora je važnija od dominantnosti ruke. - silazni piramidalni put je za 80% ispitanika (uzorak od 300) deblji u desnu ruku (kontralateralni i ipsilateralni) i prelazi na suprotnu stranu na višoj razini produžene kralježničke moždine, nego silazni put u lijevu ruku - isti postotak dobiven je i za ljevoruke ispitanike (82%; za 9 od 11 ispitanika) Dakle, važnija je opet lateralizacija nego dominantnost ruke. - u osoba s dominantnom lijevom rukom ili kod ambidekstra, žuljevito tkivo je za 11% deblje nego u desnorukih. Mogući razlog: veći broj aksona, deblji aksoni ili bolja mijelinizacija. Ako je razlog broj aksona, onda ih ima oko 25 milijuna više. To
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a) Teorija Peloponeskog rata ili teorija štita i mača (muškarci – oružje; žene – djeca) b) Teorija desnorukog svijeta (moguće je odgojiti «desnonoge» ili «ljevonoge» miševe). Dijete živi u desnorukom svijetu i razvija desnorukost. c) Teorija mozgovnog deficita (genetska predodre đenost za desnorukost) – stresni porod može uzrokovati mozgovno oštećenje i ljevorukost. 2. Anatomske teorije a) teorija veće razvijenosti lijeve hemisfere b) Morganova teorija dominantnosti lijeve strane tijela (srce, jajnici, temporalni režanj, lubanja kod primata).
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Cerebralna organizacija u ljevorukih osoba - vjeruje se da su kognitivne funkcije kod ljevaka više bilateralno organizirane - brži oporavak od lezija: Rasmussen i Milner: Wada test za ljevake 70% LH, 15% DH i 15% BL - Kimura: 520 unilateralnih afazija, kod ljevaka je očekivani postotak prema Wada testu - razlike nasljednih i nenasljednih ljevaka - kod mentalno retardiranih osoba češća je ljevorukost zbog prebacivanja funkcija iz oštećene u neoštećenu hemisferu. SPOLNE RAZLIKE U CEREBRALNOJ ASIMETRIJI Muškarci i žene su različiti. Koja je biološka osnova tih razlika?
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- razlika počinje od 2-3 godine - razlog je vjerojatno djelovanje muškog hormona androgena. (Slika razlika verbalnih, spacijalnih sposobnosti, precrtavanja i prisjećanja). - muškarci su bolji u šahu i glazbi (komponiranje i dirigiranje); glazba ima elemente prostornosti (visoki ton – niski ton), utjecaj
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prijenos spacijalnih funkcija; - 25% žena imat će razvijene spacijalne sposobnosti (Slika utjecaja recesivnog gena na prijenos spacijalnih sposobnosti). 4. Brzina sazrijevanja - djevojčice sazrijevaju brže nego dječaci. Što je sazrijevanje sporije, razvija se veći stupanj lateralizacije. Možda je genetski utjecaj upravo u brzini sazrijevanja pa iz toga
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- Bouillard Bouillard (1825) - hemisfere imaju razli različite funkcije za složene pokrete (pisanje, govor) - Dax (1836) – opis pacijenata s lezijama u frontalnom dijelu lijeve hemisfere - Broca (1861) – anatomski dokaz da je govor/jezik u lijevoj hemisferi - Wernicke (1875) – različite zone za produkciju i percepciju
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Na temelju rezultata stimulacija može se zaključiti: 1. Ne potvr đuju strogi lokalistički model (slične efekte ima stimulacija prednjih i stražnjih govornih podru č ja). 2. Stimulacije izvan klasičnih zona utječu na govor. 3. Stimulacijom govornih zona utječe se i na motoriku, kratkotrajno pamćenje i čitanje. 4. Ako se odstrane dijelovi iza Wernickeova područ ja s kojima su povezani govorni i vizualni centri, ne nastaje trajna afazija.
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Wernicke-Geschwindov Wernicke-Geschwindov model: 1. Broca – motorički programi 2. Wernicke – pretvara auditivni ulaz u značenjske jedinice 3. arcuata fasciculus - veza Broce i Wernickea 4. predcentralna i postcentralna zona lica – motorika i senzorika govora 5. angularni girus – vizualne forme jezičnih jedinica i pretvaranje u
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iako primarno slušno područ je funkcionira, osoba kao da ne čuje
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b) osoba može glatko govoriti, ali brka fonemski sastav rije či
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- Benson (1967) – radioskopskom metodom
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Trusted by over 1 million members
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Trusted by over 1 million members
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Lieberman je, na temelju oblika lubanje i položaja grkljana, također