Neurobiologia cunostintei

8/18/2019 Neurobiologia cunostintei

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Neurobiologia cunostintei:

Modelele neuroanatomice ale cunostintei:

1. SRAA – implicat in „arousal” – este alcatuit din nucleii reticulari clasici din trunchiul cerebral, darinclud si nucleii colinergici din trunchiul cerebral superior, nucleii noradrenergici, mai ales locuscoeruleus, proiec iaț histaminica din hipotalamusul posterior iș probabil cai ascendente de dopamina si

serotonina din trunchiul cerebral. O mare parte din activitatea eercitat! de aceste c!i este mediata detalamus, care poate "i considerat ca v#r"ul SRAA, precum i unș releu sinaptic critic pentru ma$oritateacanalelor sen%oriale iș intracerebrale &'ononi iș (och, )**+ ))*- 'ononi )*1) )))-/. Rolul SRAAimplic!, de asemenea, coordonarea cu talamusul, care prin colaterale aonale inhibitorii, inhiba"asciculul reticular speci"ic de in"orma iiț catre corte. &0oung iș igott, 1222 )32-/. 'omogra"ia cuemisie de po%itroni &4'/ a demonstrat hipometabolism selectiv talamic iș SRAA 5n timpul somnuluicu unde lente &Ma6uet, )*** 137-/ iș aneste%ie &Al8ire, 9aier iș allon, )*** 3-/.

). Amigdala – neurobiologia a dovedit rolul ei in memoria de lunga durata &;'M/, memoria de lucru& ?-/.Rela ia dintre amigdala si sistemeleț ș țcognitive este bi@directionala, depin%and de lin8@uri "unc ionale corticolimbice. Ast"el, noradrenalina iț șdopamina pot $uca un rol 5n rela ia dintre amigdala i cunoa terea &„cognition”/ superioar! &alleine iț ș ș ș(illcross, )**? 1?-. (im et al, )*11 1*7-/. Apare evidenta nevoia de integrare intr@un modelcuprin%!tor, a bine@cunoscutei implicari a amigdalei 5n emo ie, cu rolul s!u 5n cunoa tereț șsuperioara&helps, )**? 1??-/. Amigdala asigura o "unc ie de vigilen ! generala, care are ca scopț ț

preg!tirea organismului pentru a "ace "a ! unor situa ii di"icile, solicitante, ca% in care r!spunsul vigilentț țal amigdalei poate lua "orma unor semnale neuromodulatorii "a%ice proiectate pe sistemul nervoscognitiv, motor i B sau autonom, 5n timpul unei situa ii ce solicita resurse metabolice suplimentare,ș țcum ar "i, o amenin are "i%ic! sau o activitate mentala complea &Schae"er i CraD, )**E 1+>-/. e de oț ș

parte, amigdala este sensibila la stimuli care posed! o valoare intrinsec! de adaptare &de eemplustimuli emo ionali, sau care generea%a teama etc/. e de alt! parte, amigdala este sensibila la stimuli ațc!ror valoare motivationala este determinat! 5n mod arbitrar de scopul activitatii sau de anumitesolicitari &Richardson, ciudate i =olan, )**E 177-/. rin urmare, acest lucru sugerea%a ca amigdalaș


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trebuie s! primeasc! nu numai intr!ri sen%oriale de ba%!, dar, de asemenea, intr!ri de la nivel 5naltlegate de scopul sarcinii. Amigdala are un model remarcabil de bogat de coneiuni directe i indirecteș cu regiunile subcorticale iș trunchiul cerebral, dar i cuș regiunile neocorticale implicate 5n memoria delucru & 12)-/. =e asemenea, eista teorii ce sustin implicarea proiec iilor ț a"erente din parietal, corteul temporal iș occipital iș sistemul limbic in integrarea de in"orma ii sen%orialeț ,organi%area visuospatial, memoria vi%uala, cuno tin !ș ț iș controlul comportamentului i motiva ie&ș ț Schmahmanniș andDa, 122) 121-/ . Iatura eact! a implic!rii cerebelului 5n procesele cognitive, este p#n! 5n pre%ent, mai

pu in 5n eleas!. Anomalii ale structurii i "unc iei cerebelului au "ost identi"icate neuroimagistic 5n anumiteț ț ș țtulbur!ri psihiatrice &(onars8i et al., )**? 11E-/. Mai mult dec#t at#t, interven ii terapeutice "armacologice iț ș

psihosociale pentru pacien ii cu aceste tulbur!ri au "ost asociate cu modi"ic!ri ale "unc iei cerebelului, suger#ndț țun rol important al acestuia, 5n di"erite procese mentale, care sunt perturbate 5n anumite tulbur!ri psihice &Strata,)*11)*+-/.

E. 'alamus @ neuronii din talamusul central &nucleii intralaminari anterior i posterior, por iunileș ț paralaminare conee ale nucleilor de asociere talamici @ dorsal median, anterior ventral, ventral pulvinar lateral i in"erior/ au speciali%!ri anatomice i "i%iologice speci"ice care spri$in! rolul lor esen ial 5nș ș ț"unc iile generale de aten ie sus inut!, memorie de lucru i preg!tire motorie. Aceste popula ii neuronaleț ț ț ș țsunt interpuse 5ntre sistemul de ecitare &arousal/, care controlea%! nivelul general de activitate talamic!

i cortical!, precum i sistemele de supraveghere "rontale, care controlea%a „shi"t@ul” premotor deș șaten ie i a$ustarea nivelului de vigilen ! i alert! &arsuraman, 1222 1?*-/. Jn paralel, populatiiț ș ț ș

http://www.benbest.com/http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3653219/#ref208http://www.benbest.com/http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3653219/#ref208


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neuronale individuale din cadrul acestor agregate celulare sunt tintite de o varietate de relee sen%orialededicate care au evoluat, pentru a capta rapid aten ia i a redirec iona comportamentul &(rout, el%er iț ș ț ș;oeHD, )**) 117-/. Gmpreuna, aceste speciali%!ri anatomice sugerea%! c! mul i neuroni din talamusulțcentral au rol in comportamentele directionate catre scop i in cuno tin !.ș ș ț 'alamusul central prime teș inerva ieț larga in cadrul sistemului de ecitare &ra"", )**? 1?E-/ cu a"erente neuronale colinergice&(olmac iș Mitro"anis, 1222 113-/ cat iș noradrenergica din locus ceruleus iș a"erente serotoninergicede la ra"eul medial &McFormic8 iș al, 1227 1E3-/. ;e%iuni directe ale talamusului central pot producetulbur!ri ale cuno tin ei la nivel global &arvi%i i =amasio, )**3 1?1-/. =ac! le%iunea este unilaterala,ș ț șse pot produce a"ectari hemispa iale sau stari modi"icate de cunostinta similare cu mania acuta sauțdelirul. ;e%iuni bilaterale restr#nse ale talamusului central pot produce coma acuta, care indic! rolulimportant pe care aceste structuri il $oac! in alerta, sau starea de veghe. Fu toate acestea, spre deosebirede le%iunile trunchiului cerebral care produc com! 5n urma le%iunilor "ocale, recuperarea cuno tin ei inș țle%iunile bilaterale centrale talamice 5n ceea ce prive te comportamentul directionat spre scop iș șabilit! ile de comunicare este "oarte lent i incert &Schi"", )**+ 1++-/.ț ș

>. Fircuitele "rontoparietale:

=aca cuno tin aș ț „"enomenal!” &phenomenal consciousness/ se re"er! la aspectul subiectiv al eperien eiț ,cunostinta „de acces”& access consciousness/ se re"er! la aspectul obiectiv prin controlul direct al eperien eiț

prin ra ionamentț , raportare, si ac iuneț . =istinc ia 5ntre ele &loc8, 1227 )3-/, ar "i relevanta pentru a 5n elegeț ț presupusul rol al %onelor "rontoparietale 5n percep ia con tient!. „ț ș henomenal consciousness” ar putea "i asociata5n principal cu activarea regiunilor sen%oriale, iar „access consciousness” ar putea avea nevoie de implicare din

partea %onelor "rontoparietale. Modelul speci"ic de activare "rontoparietala in timpul percep iei con tiente poateț ș"i di"eren iat mai in detaliu 5n "unc ie de "a%ele i B sau tipul de prelucrare, de eemplu, tran%i ii perceptive "a ! deț ț ș ț ț

percep ie sus inut!. =e asemenea eista teorii care sustin ca suprapunerea activarilor "rontoparietale re"lectaț ț procese cognitive comune, de tip atentional sau memorie de lucru.

?. Forteul pre"rontal – eista teorii contradictorii in ce priveste rolul in cunostinta, totusi cele mai recentesustin c! F $oac! un rol mai important 5n monitori%area con tienta dec#t 5n acumularea automat saușnon@con tient! de dove%i. Sugestia este c! traseul con tient de nivel superior corespunde re elei parieto@ș ș ț

pre"rontal i %onelor corticale superioare asociate &9irstein, )*11 2E-/.ș

7. recuneus @ a "ost sugerat c! activitatea precuneus 5n timpul st!rii con tiente de odihn!ș spri$in! procesarea conceptual! care operea%! pe stocurile interne de in"orma iiț &semnale endogene/, maidegrab! dec#t "unc iile deț Kpercep ieKț &surse de in"orma iiț eterne de creier/. Fu alte cuvinte, acestdomeniu pare s! contribuie la auto@ prelucrarea de „ganduri” pe care oamenii o eperimentea%a 5ntimpul odihnei constiente &Favanna, )**7 33-/. O interac iune 5ntreț precuneus iș corteul pre"rontal a"ost postulata 5n st!ri de cunostinta caracteri%ate de un nivel ridicat de re"lectare a con tiin ei de sine.ș ț &($aer et al ., )**)1*2- Favanna and 'rimble, )**?3)-/.

Modele si teorii neuronale:1. 'eoria dendritelor apicale @ Activitatea sincrona 5n grupurile de dendrite apicale produce unde

electromagnetice &4M/ care pot radia spre eterior, iar 5n ca%ul 5n care acestea sunt su"icient de puternice pentru a a$unge la supra"a a pielii capului pot "i m!surate ca 44C. F#mpul electromagnetic aț"ost propus ca substrat "i%ic al cuno tin ei &Mcadden, )*** 1EE- oc8ett, )*** 1?7-/. Furen iiș ț ț"luctuanti care produc c#mpuri 4M sunt produsi de mai multe surse, printre care: activitatea membraneiaonilor, dendritelor i somei neuronilor din regiunile corticale si subcorticale. Fon"orm acestei teoriișcuno tin a de moment este repre%entata de c#mpul general "ormat din toate aceste domenii 4Mș țindividuale.

). Activitatea I@metDl – = – Asparat@ului

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Apari ia st!rii de cunostinta depinde 5n mod critic de o anumit! categorie de procese computeri%ate,țcare sunt mediate de sDnapse IM=A. Argumentele 5n "avoarea ipote%ei se ba%ea%! pe ideea c!activitatea receptorului IM=A este capabil! s! "orme%e repre%entari 5n creier i toate aneste%iceleșgenerale inhiba activitatea receptorilor IM=A. rin urmare, activitatea receptorului IM=A esteesen ial! pentru cunostinta &Cambrill i colab., )*11 77-/.ț ș

3. Modelul ansamblurilor neuronale:

Ln ansamblu de celule neuronale este de"init ca o structur! di"u%! cuprin%#nd celule din corte iș dience"al,capabil! s! ac ione%eț rapid ca un sistem 5nchis iș sa "acilite%e "unctionarea altor ast"el de sisteme &9ebb, 12E22)-/. Au "ost stabilite ast"el de ansambluri, observand ca 1* de milioane de neuroni isi pot sincroni%a activitatea5n mai putin de )3* ms si s@a postulat ca "iind esen ialeț pentru producerea de st!ri con tiente.ș Jn teorie neuronala&;ibet, 1223 13*-/, cuno tin a este asociata cu activit! ile neuronale care persista pentru o perioad! su"icient deș ț țlung!, cu o durat! minim! de >** ms. Jn sinapsele de ansambluri neuronale, cunoscute sub numele de sinapseMalsburg, puterea sinapselor cre te c#nd eist! o corela ie puternic! 5ntre activitatea pre@ i postsinaptic! &Fric8,ș ț ș12+E E2-/. Aceste con"igura ii tridimensionale de ansambluri de mari dimensiuni din creier, care nu respectațlimitele anatomice conven ionale, se vor corela cu di"erite grade de cunostinta 5n "iecare moment &Creen"ield,ț1227 +?-/. Ln studiu recent spri$ina acest model de asamblare neuronala &(och, )**> 11)- Overgaard iș

Morgensen, )*11 1>7-, ansen i Overgaard, )*11 1*1-/.ș

E. Modelul dialogului talamo@cortical:

'eoria descrie centre pe circuitul thalamocortical care implic! nucleii intralaminari, care con in celule carețconectea%a activitatea trunchiul cerebral &in st!rile generale de veghe i vigilen !/ la di"erite %one aleș țcorteului &ogen, 122> )E-/, distribuite pe scar! larg!. Al ii au propus ca structurile talamice i corticaleț ș&5mpreun! cu rela ia lor cu nucleii reticulati din trunchiul cerebral/ sunt condi ii necesare i su"iciente pentruț ț șeisten a unei stari de cunostinta de veghe, iar proprietile recurente de Kre@intrareK ale circuituluițthalamocortical par a "i esentiale &aars et al, )**3 2-.. Seth et al, )**> 127-/.

>. 'eoria activitatii neuronale cerebrale ba%ale:S@a observat ca i 5n absen a unor sarcini cognitive sau de stimulare sen%orial!, activitatea cerebralaș ț

ba%ala 5n %onele implicate 5n gene%a percep iei con tiente "luctuea%! continuu 5n creierul uman &Agliotiț și Fandidi, )*11 1-/. Mai mult dec#t at#t, datele de cercetare sugerea%a ca auto@con tienti%are iș ș ș

constienti%area eterna nu sunt concurente, ci ar "i mai degrab! 5n competi ie 5n timpul procesului deț percep ie sen%orial!. ;uate 5n ansamblu, aceste date sugerea%! c! "luctua iile in activitatea cerebralaț ț ba%ala sunt susceptibile de a "orma con inutul de K"lu al con tiin eiK noastre 5n curs de des"! urare 5ntr@ț ș ț șun mod decisiv. Fu toate acestea, originea i "i%iologia acestor "luctua ii r!m#ne un mister, i va "aceș ț șobiectul unor studii viitoare &olD et al, )**+ )7-.. olD et al, )*11 )?-/.

'eoria proto@panpsihista a cunostintei:Fon"orm acesteia sistemele con tienteș sunt cele care con inț dou! re eleț care interac ionea%!ț activ, unul

pentru construirea de cuno tin eș ț , i cealalt! pentruș generarea de sentimente. Ioutatea este modul 5n careeste conceput sistemul de sentimente: ca un sistem care "unc ionea%!ț cu modele „Haveli8e”, iar "iecaretip de val corespunde unui sentiment. „Fentrul” care ini ia%!ț sentimentele se propune a "i re eauaț astrogliala &ereira r. iș urlan, )*1* )E?- ereira r, ereira iș urlan, )*1* )E7-/, re"lect#nd stariale 5ntregului corp, 5n procesul de interac iuneț cu lumea. Jn acest sens, studii recente "ac legatura intreundele de calciu din re eauaț de astrocite cu integrarea in"orma iilor ț in creier &Iavarrete et al., )*1))E+-/, starea de con tien !ș ț &'hrane et al., )*1) )E2-/ iș bolile neurologice cu impact asupracunon tin ei,ș ț cum ar "i boala Al%heimer &urman i colabș ., )*1) )>*-/.Re eauaț astrogliala prime te semnale de laș 5ntregul corp prin intermediul neuronilor, "luului de s#ngei lichidulș ce"alorahidian &de eemplu, 5n sistemul vestibular/. Aceast! stare a 5ntregului corp este

comparata dinamic cu in"orma iileț cu privire la eistenta organismului 5n mediul eterior, a a cum esteșrepre%entat de re eauaț neuronal!. Re%ultatul acestei compara iiț este un sentiment de moment careraportea%a in"orma iilor primite laț starea sistemului. Acest sentiment impactea%a intreg organismul, prin

semnale neuro@endocrin@imune @ inclusiv activit! iț ale inimii si sistemului nervos. rin urmare,con tiin aș ț este conceputa ca prelucrarea de in"orma ii despreț corp iș lume, construirea de cuno tin eș ț iș


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investirea lor cu sentimente care in"luen ea%!ț procesarea iș , prin urmare, ac iunileț individului caretr!ieste 5n lume.

Ieuromodulatorii:Sistemul colinergic este unul dintre cele mai importante sisteme de neurotransmitatori la nivelulcreierului i controlea%a activit! ile care depind de aten ia selectiv!, component! esen ial! a cunostintei.ș ț ț ț

=ove%i psiho"armacologice i patologice sus in conceptul de Kcomponent! colinergicaK a cunostintei. eș ț ba%a constatarii ca activarea corticala se mentine in cadrul somnului R4M, in absenta activariineuronale mono@aminergice &serotoninergice sau noradrenergice/ dar in pre%enta descarcarilor constantedin neuronii colinergici ai nucleului ba%al, s@a a$uns la conclu%ia ca sistemul colinergic ascendent estesu"icient pentru mentinerea neocorteului intr@un mod operativ. Se pare insa ca sunt necesare ac iunițcombinate ale AFh i glutamatului 5n acest sens. 'eoriile recente ale rolului AFh includ posibilitatea cașaceasta a"ectea%! procesele discriminatorii creste rata semnal@%gomot, modulea%! e"icien a deț

prelucrare corticala a in"orma iilor sen%oriale i de asociere, controlea%! receptia stimulilor i evaluareaț ș șnivelul lor de semni"ica ie, modi"ic! reac ia corticala 5n ceea ce prive te relevan a i noutatea, iț ț ș ț ș șlimitea%! con inutul "luului con tient. =roguri care antagoni%ea%! receptorii muscarinici inducț șhalucina ii i reduc nivelul de cunostinta, 5n timp ce receptorul nicotinic este implicat 5n mecanismul deț șac iune al aneste%icelor generale &prin inhalare/. Gn bolile degenerative ale creierului, modi"ic!ri alețst!rii de con tien ! sunt asociate cu de"icite regionale 5n sistemul colinergic. Reteaua vasta colinergicaș ț

din trunchiul cerebral care include proiec ii convergente la talamus, pare a "i amplasata strategic pentruța genera i integra constienta. )>1-șrintre di"eritele p!r i ale neuronului , dendritele sunt probabil cei mai buni candidati pentru a "i cheiaț

pentru "unctia cognitiva superioara , deoarece pot integra un num!r mare de intr!ri . Membrananeuronala este site@ul ini ial de r!spuns pentru intr!ri de la alti neuroni , dar ceea ce se a"l! dincolo dețmembrana neuronal! controlea%! nivelul sinaptic de r!spuns prin compararea intr!rilor noi cuin"orma iile stocate 5n memorie . =endritele sunt imbogatite cu microtubuli si proteine asociatețmicrotubulilor &MA/ , al caror rol nu se cunoaste inca pe deplin. Acumularea de dove%i sugerea%a camicrotubulii i MA@urile $oac! un rol critic 5n memorie i con tiin ! , precum i 5n transportulș ș ș ț șneuronal. roteina @ ) asociata microtubulilor & MA) / este o protein! citoscheletala speci"icadendritelor , care , de asemenea, ac ionea%! ca o molecul! de transduc ie a semnalului , mediindț țr!spunsurile chimice interne ca urmare a eliber!rii sinaptice de neurotransmitatori glutamat ișacetilcolina . MA) i microtubulii se leaga 5mpreun! pentru a "orma o matrice care stochea%!ș

memoria. Gn"orma ia stocata 5n matricea de microtubuli este apoi accesata la eliberarea anumitorțneurotransmi !tori , cum ar "i acetilcolin! i glutamat . Acetilcolina controlea%! nivelul de con tien ! ,ț ș ș ț5n principal prin receptorul muscarinic determinand activarea consecutiva a 8ina%elor (F i FaM(GG ,șambele "os"oriland MA) cu rol in memorie . rin activarea de catre neurotransmitatori , microtubulii

pot evalua intr!rile sinaptice actuale 5n contetul activit! ii sinaptice anterioare , i apoi sa creascaț ștransportul de anumite molecule legate de 5nv! are la sinapse . entru aparitia activitatii con tiente esteț șnecesara co@ activare a unui num!r mare de sinapse si transportul 5n mas! de proteine sinaptice esen ialeț, i este posibil ca „6uantum computation” sa $oace un rol 5n diri$area coerenta a acestui transport . Oșteorie anterioara sustine ca acetilcolina "acilitea%! calculele cuantice 5n microtubuli prin "os"orilareaMA). O terorie recenta propune ca modelul de legare a MA) de microtubuli "ormea%! un contur pe

ba%a de gel, care repre%int! in"orma ii stocate de c!tre mecanismul de 5nv! are i o"er! o ba%! "i%ic!ț ț ș pentru aceste in"ormatii. F#nd MA) este "os"orilat, acest contur pe ba%a de gel se etinde propagandin"orma ia longitudinal pe microtubul, i tangen ial, la microtubulii vecini. Jn aceste dou! moduri,ț ș ț

microtubulii activati "i%ic asigura un r!spuns coerent care acoper! mai multe sinapse moduland intr!rinoi 5n contetul in"orma iilor stocate 5n memorie.ț )>)-

Re"erinte:1. Aglioti SM, Fandidi M. Out o" place bodies, out o" bodD selves. Ieuron. )*117*:173–>. ubMed-). Ale8sander G. Mapping the Mind Series. ;ondon:


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251. errD 4,

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