Ho fokotseha ho thibela ha neuronal le ho tsamaisana le cortex ea bacha nakong ea boits'oaro (2011)

J Neurosci. 2011 Jan 26;31(4):1471-8.
 

mohloli o moholo

Lefapha la Neuroscience, Univesithi ea Pittsburgh, Pittsburgh, PA 15260, USA.

inahaneloang

Nako ea bocha ke nako ea bokhoni ba ho ba le kelello le ho ba tlokotsing ea mafu a mangata a kelello a kelello le ts'ebeliso ea lithethefatsi. Ho na le temohisiso e ntseng e eketseha ea hore ts'ebetso kapa ts'ebetso ea meriana ka nako ena e ka ba bohlokoa bakeng sa thibelo ea mafu ho batho ba kotsing. Ka hona, re tlameha ho fumana kutlwisiso e tebileng ea hore na boko ba bongoana bo etsa liketsahalo tse ikhethang mabapi le boitšoaro bo susumetsoang joang. Ho etsa sena, re rekota ts'ebetso e le 'ngoe le likarolo tsa tšimo ea lehae kahara orbitofrontal cortex ea likhoto ha ba ntse ba etsa mosebetsi o bobebe o khannoang ke moputso. Bacha ba kenyelelitse likarolo tsa mantlha tsa mosebetsi ka tsela e fapaneng le ea batho ba baholo, ho bontša hore ts'ebetso ea methapo ea methapo ea liketsahalo tse ikhethang e fapana ka lihlopha tsa lilemo tse peli. Ho kenyelletsoa hoa tšimo ea masimo a lehae, ho fapana ha nako ea sekhahla, le karabelo e lekanyelitsoeng ea likarabo tse thibelang maikutlo le tse nyakallisang li fapana ka mokhoa o ikhethileng. Ketsahalo e akaretsang ea bongoana ea phasic e ne e sa thibeloe haholo ebile e fapana le boholo ba mosebetsi. Cortical inhibition e bohlokoa bakeng sa puisano e sebetsang pakeng tsa lihlopha tsa neuronal, mme taolo e fokotsang ea ts'ebetso ea cortical e kentsoe ho schizophrenia le mathata a mang. Kahoo, liphetho tsena li fana ka maikutlo a hore likarabelo tse fokotsang tsa karabo ea methapo ea methapo ea bongo ho liketsahalo tse ikhethang e ka ba mokhoa o thata bakeng sa a mang a bofokoli bo eketsehileng ba nako ena.


Keywords: Bonyatsi, OFC, electrophysiology, schizophrenia, khatello ea maikutlo, bokhoba ba tahi

Selelekela

Bocha ke nako ea phetoho ha motho a phethela phetoho ea 'mele le ea kelello ho ba motho e moholo.Arnett, 1999). E boetse e nkuoa e le nako ea ho ba tlokotsing ha e ntse e tsamaellana le ho qalella ha matšoao bakeng sa mathata a mangata a mantlha a kelello, ho kenyeletsa le mathata a ho feto-fetoha ha maikutlo, lefu la kelello le tšebeliso e mpe ea lithethefatsi (Volkmar, 1996; Pine, 2002; Johnston et al., 2008). Lilemong tsa morao tjena, lithuto tsa batho ba lilemong tsa bocha le mehlala ea liphoofolo li hlalositse liphetoho tse amanang le lilemo mohlaleng oa sebopeho sa lisele tsa cellular le tsa molek'hule litlamorao tsa litlamorao tsa meriana le lithethefatsi tse fapaneng (Spear le Brake, 1983; Spear, 2000; Adriani et al., 2004; Brenhouse et al., 2008; Paus, 2010). Phapang ea boitšoaro e amanang le lilemo le eona e se e hlahlobisitsoe 'me hangata e shebane le eona, le ha boitšoaro ba bongoana bo batla bo tšoana hantle le ba batho ba baholo maemong a mangata ka liphetoho tse fokolang feela tsa bokhoni ba ho etsa liqeto ho tloha lilemong tsa bocha ho ea pele (Spear, 2000; Doremus-Fitzwater et al., 2009; Figner et al., 2009; Cauffman et al., 2010). Leha ho le joalo, bacha ba ka etsa liketsahalo tse ikhethang ho batho ba baholo. Mohlala, phuputso ea morao-rao e hlokometse mocha ea lilemong tsa bocha ho feta polelo ea protheine ea c-fos e kholo ho dorsal striatum le li-nucleus ka mor'a ho pepesetsoa monko o amanang le moputso (Friemel et al. 2010). Phapang ea litekanyo tsa ts'ebetso ea tlhaho ea bongoaneng pele ho cortex (PFC) le ho hokahanya le tsona li hlalositsoe (Ernst et al., 2006; Galvan et al., 2006; Liston et al., 2006; Geier et al., 2009; Uhlhaas et al., 2009a). Leha ho le joalo, ho fokolang ho tsejoa ka sebopeho se nepahetseng sa karohano ena e amanang le lilemo sethong sa neuronal.

Ho bapisa ka kotloloho ts'ebetso e matla ea li-cortical neurons ho bacha le ea batho ba baholo, re tlaleha mosebetsi o le mong le monyetla oa tšimo ea lehae (LFP) ho tsoa ho orbitof Pambal cortex (OFC) ea likhoto ha ba ntse ba etsa boitšoaro bo susumetsoang ke moputso. OFC e ne e shebane ka lebaka la karolo ea eona ea mantlha ts'ebetsong tebello ea boleng le bopaki ba pejana ba nts'etsopele ea eona e tlase ho bacha.Schultz et al., 2000; Galvan et al., 2006; Schoenbaum et al., 2009). Mosebetsi o amanang le boits'oaro o kenyelletsa ho nka karolo ho latela mokhatlo o ithutileng diphetho tsa thuto (Sturman et al., 2010), eo e leng moaho oa mantlha oa boits'oaro bo thata. Bonolo ba mosebetsi ona o ile oa lumella mehato ea boitšoaro hore e ts'oane haholo lipakeng tsa lihlopha. Ka hona re ka leka tlhahlobo ea maikutlo hore le ka ts'ebetso e ts'oanang ea ts'ebetso, mocha oa OFC o kenyeletsa tlhahisoleseling e amanang le mosebetsi ka tsela e fapaneng le ea batho ba baholo. Ho tšoaea liphapang tsa mantlha tsa tšebetso ea neural-le ho etsa joalo maemong a methapo-ho bohlokoa bakeng sa ho khetholla lits'ebetso tsa nts'etsopele tse ka amanang le likotsi tsa neuropsychiatric tsa bohlankana, le moralo oa ka moso oa maano a ho kenella ho thibela le ho phekola mathata a joalo.

Lisebelisoa le Mekhoa

Lihlooho

Adolescent (matsatsi a ho qetela a P28-42; n = 8) le motho e moholo (P70 +; n = 4) ho ne ho sebelisoa likhoto tsa banna tsa Sprague-Dawley (Harlan, Frederick MD). Juvenile (P21) le likhoto tsa batho ba baholo ba ile ba amoheloa beke e le 'ngoe pele ba buuoa. Lintho li ile tsa bolokoa ka har'a vivarium e laoloang ke maemo a leholimo tlasa maemo a leholimo a 12 h (mabone a 7 pm), lib lib ho fihlella chow le metsi pele ho thupelo. Lits'ebetso tsohle tsa ts'ebeliso ea liphoofolo li amohetsoe ke Univesithi ea Pittsburgh Tlhokomelo ea Liphoofolo le Ts'ebeliso.

Mekhoa ea ho buoa le ho buoa ka motlakase

Likhoto li ile tsa etsa liopereishene tse ngata tsa motlakase joaloka ho hlalositsoe pele (Totah et al., 2009). Ka bokhutšoanyane, li-arrider tsa microelectrode (NB Labs, Denison, TX), e nang le lithapo tse robeli tsa tšepe tsa tšepe tsa Teflon tse hlophisitsoeng ka sebopeho sa 2 x 4, li kentsoe ho OFC. Batho ba baholo ba ne ba kenngoa ka makhetlo a mabeli 2.8 ho 3.8 mm ka anterior ho bregma, 3.1 ho 3.5 mm lateral ho bregma, le 4.5 mm ventral ho ea dura holimo. Bacha (P28-29) ba ile ba kenngoa ntle le mabaka (ka lebaka la meeli ea boholo) 2.8 ho 3.2 mm anterior ho bregma, 2.8 ho 3.2 mm hamorao le bregma, le 4.0 mm ventral ho ea dura. Nakong ea lirekoto, hlooho ea motlakase oa tšebeliso ea 'moeli-e kopantseng mohala oa lebokose le boima (NB Labs) e ne e hokahane le commutator (NB Labs) e lumellang likhoto hore li tsamaee ka bolokolohi ka har'a lebokose la liteko. Ts'ebetso e rekotiloeng ea unit e le 'ngoe e ntlafalitsoe ho phaello ea 100 × le analog band-pass e tlhotliloeng ho 300 - 8000 Hz; Li-LPF li ne li entsoe ka sekepe ho 0.7 - 170 Hz. Ts'ebetso ea unit e le 'ngoe e ne e ngotsoe ho 40 kHz mme LFPs e ngotsoe ho 40 kHz mme e theotsoe ho 1 kHz ka software ea Recorder (Plexon). Ts'ebetso ea unit e le 'ngoe e ne e hloekisoa ka tekanyo e phahameng ho 300 Hz mme LFPs e ne e se e le sefepi se tlase ho 125 Hz. Mats'oao a liketsahalo tsa boits'oaro ho tsoa lebokoseng la opereishene a romeloa ho Recorder ho tšoaea liketsahalo tse khahlisang. Diyuniti tse le 'ngoe li ile tsa qheleloa ka thōko ho Offline Sorter (Plexon) li sebelisa mokhoa oa ho kopanya libuka tsa litaelo le tse ikemetseng tsa ho itlhophisa joalokaha ho hlalositsoe pele (Homayoun le Moghaddam, 2008; Totah et al., 2009).

boitshwaro ja

Likhoto tsa batho ba baholo le ba lilemong tsa bocha li ile tsa lekoa ka har'a lisebelisoa tsa mabokose tse sebetsang (Coulborn Instrsets, Allentown, PA) tse nang le khanya ea ntlo, makasine ea pellet e ka tsamaisang li-pellets tsa lijo (dextrose e tiisitsoeng, 45 mg; Bio-serv, Frenchtown, NJ) ho e jareteng ea lijo, le likoti tse tharo tse hulang nko tse khabisitsoeng ka kotloloho leboteng le shebaneng le jareteng ea lijo. Kamora matsatsi a 5-6 a ho hlaphoheloa bocha, liphoofolo li ile tsa thibeloa lijo hanyane ka hanyane, tsa lula ka har'a lisebelisoa tsa tlhahlobo ea boits'oaro, 'me tsa qala ho ikoetlisetsa mosebetsing oa boits'ebetso, o kileng oa tsebahala (Sturman et al., 2010). Ka bokhutšoanyane, likhoto li ithutile ho kenella ka sekoting se boneselitsoeng nko bakeng sa ho matlafatsa lijo. Liteko li ile tsa qala ka ho qala ha lebone la cue kahare ho lesoba le tlohang mpeng. Ha lekhalo le kenella ka sekoting seo, lebone le ile la tima hang-hang 'me la tlisoa ka peterole e le' ngoe ka sejelong sa lijo, se neng se bonesoa ka nako eo. Ho kenella ka har'a senoelo sa lijo ho amohela pellet ho tima lebone la lijo mme ho bakile nako ea liteko ea 5-sSetšoantšo sa 1a). Seboka se seng le se seng se ile sa emisoa kamora liteko tsa 100 kapa ho feta ha 30 min. Mosebetsi o fetileng o bonts'a hore mosebetsi ona o ka ithuta kapele ho bacha le batho ba baholo, ka ts'ebetso e kholo haholo ka letsatsi la boraro la thupelo (Sturman et al., 2010). Mehato ea ts'ebetso ea mantlha e kenyelletsa palo ea liteko tse felletseng nakong ea thupelo e ngoe le e 'ngoe, latency ho tloha ho cue ho ea pokellong ea lipina, le latency ho tloha ho kenella ho kenang ho kenngoeng ha lijo. Mehato ea Age × e pheta-phetoang ANOVA e entsoe mehatong eohle ea sephetho ho SPSS (alpha = 0.05). Maemong ohle moo khopolo ea bonyollo e hatakeloang, litokiso tse tlamelletsoeng ka tlase li ile tsa sebelisoa bakeng sa phetoho e kholo ea maemo a boiketlo.

Setšoantšo sa 1

Meetso ea li-electrode, paradigm ea mosebetsi, le ts'ebetso ea boits'oaro. A) Lits'oaetso tsa bongoana le tsa batho ba baholo li ile tsa koetlisetsoa papaling e bonolo ea ho ithuta eo ho eona li neng li hokahanya mokopu o tsoang ka selomong (hole). ...

Histology

Kamora ho etsa tlhahlobo, likhoto li ne li sa anngoe ka chloral hydrate (400 mg / kg ip) mme hajoale 200 μA e ile ea fetisoa ka ho rekota li-electrode bakeng sa 5 s ho tšoaea li-electrode tip placements. Liphoofolo li entsoe ka letsoai le 10% buffered formalin. Li-brains li ne li tlosoa ebe li kenngoa 10% formalin. Masapo a mokokotlo a ne a arotsoe ka lehare, a koaheloa ka cresyl violet, 'me a hloelletsoa ka methapo ea microscope. Ho beoa ka motlakase ka motlakase ho ile ha hlahlojoa tlasa mochini o bobebe o bobebe. Ke likhoto tse behiloeng ka nepo ka har'a OFC (Setšoantšo sa 1b) li kenyellelitsoe bohlahlobisisong ba electrophysiological.

Tlhahlobo ea Electrophysiology

Lintlha tsa electrophysiological li ile tsa hlahlojoa ka lingoloa tse ngotsoeng ka moetlo, tsa bolaoa ho Matlab (Mathworks, Natick, MA), hammoho le lebokose la lisebelisoa la chronux (Chronux.org) bakeng sa tlhahlobo ea LFP le mesebetsi ea ho thunya ka sekhahla ka mosa e hlahisitsoeng ke Churchland le basebetsi-mmoho (http://www.stanford.edu/∼shenoy/GroupCodePacks.htm) (Churchland et al., 2010). Ka kakaretso, mesebetsi ea neural e ne e notletsoe nako bakeng sa liketsahalo tse itseng: teko ea ho qala ha teko, karabelo ea nko ea pompo, le ho kenella ka sekoleng sa lijo. Mekete ea RF LFP e ne e notletsoe nako bakeng sa liketsahalo tsena, 'me liteko tse nang le liemahale tsa ho taka ha lia ka tsa qheleloa ka thoko pele ho ts'ebetso. Mohlala tekanyetso ea lilemong tsa bocha ea teko e le 'ngoe le batho ba baholo ba ts'oarang LFP nakong ea matlafatso e hlahisoa (Setšoantšo se tlatsetsang 1). Tekanyetso ea motlakase e lekiloeng ea sehlooho ka seng metsotsong e mengata ho potoloha ketsahalo e 'ngoe le e' ngoe ea mosebetsi e baliloe ke FFT. Sena se entsoe ho sebelisoa li-taper tse 13 tse etellang pele, sehlahisoa sa bandwidth sa 7, le 1 s spanning-window (ka mehato ea 250 ms). Meeli ena, ha e bapisoa le e meng eo re neng re e hlahlobile e lumelletse tharollo ea maqhubu a hoo e ka bang 2 Hz, eo hangata e neng e lumella libini tsa makhetlo a mangata sehlopheng ka seng sa thahasello. Ho sebelisitsoe mokhoa o mongata ka ho fetesisa hobane o ntlafatsa likhakanyo tsa spectrogram ha o sebetsana le tlhaiso-leseling e sa feleng ea linakoMitra le Pesaran, 1999), leha ho sebelisoa li-tepe tsa 1, 3, le 9 ho lebisitse ho li-programme tse ts'oanang haholo. Mong le e mong oa mokelikeli (moqolo) o mong le o mong o neng o le matla ka bongata o ne o le Z-lintlha maemong a tloaelehileng ho motlakase o tloaelehileng nakong ea motheo oa basesisi (fensetere ea 2-s e qalang 3 s pele ho molatsoana). Tšoantšiso ea matla a tloaelehileng e ile ea lekanyetsoa bacha le batho ba baholo.

Tekanyo ea nako ea ketsahalo ea nako ea ketsahalo e entsoe bakeng sa yuniti ka ngoe ho lifensetere tse potolohileng liketsahalo tsa mosebetsi. Sekhahla sa teko ea sefapano sa liteko tsa yuniti ka 'ngoe e ne e le lintlha tsa Z tse tloaelehileng ho ea nakong ea tsona ea mantlha. Liyuniti li ile tsa aroloa ka "ho kengoa tšebetsong" kapa ho "thibeloa" ka har'a lifensetere tse khahlisang ho latela hore na ketso ea bona e tloaelehileng e na le lijana tse tharo tse latellanang tsa 50 ms tse nang le Z ≥ 2 kapa Z ≤ -2, ka ho latellana. Mekhoa ena e netefalitsoe ho sebelisoa tlhahlobo e sa sebeliseng parametric bootstrap ka nako ea motheo ea yuniti ka ngoe. Bakeng sa yuniti e ngoe le e ngoe, fensetere ea mantlha e ile ea etsoa sampole ka linako tse ling ka makhetlo a 10,000. Karolo ea lifensetere tsa 2-s tseo ts'ebetso ea tsona e ntlafalitsoeng e fihlelletseng litekanyetso tsa bohlokoa ke tekanyo ea sekhahla se lebelletsoeng sa tšoaetso ea bohata bakeng sa yuniti eo nakong efe kapa efe ea 2-s. Sena se lebisitse ho sekhahla se lebelletsoeng sa tšoaetso ea bohata sa alpha = 0.0034 bakeng sa lihlopha tsohle tsa bocha le alpha = 0.0038 bakeng sa lihlopha tsohle tsa batho ba baholo. Li-alphas tsena tse tlase li bonts'a hore lihlopha tsa bohata li ka ba seoelo ho lekana hore li se ke tsa ama lipalo-palo tsa lipalo pakeng tsa bacha le batho ba baholo. Ho bapisa nako ea likarabo tsa yuniti tlhahlobo ea lihlopha e entsoe ka fensetere e tsamaeang ho potoloha liketsahalo tsa mosebetsi (boholo ba fensetere bo tsamaeang 500 ms ka mehato ea 250 ms). Bakeng sa lifensetere tsa nako tse nang le thahasello e khethehileng bakeng sa papiso ea lipalo-palo tse amanang le lilemo (mohlala, ho li-1 kamora 'cue), ho ile ha etsoa litlhahlobo tsa Chi-square tse kenyeletsang palo ea li-unit tsa batho ba baholo le ba lilemong tsa bocha tse kentsoeng tšebetsong le tse sa sebetseng. Liteko tsa bohlokoa tsa Chi-square li ile tsa lateloa ke papiso ea post-hoc ea boholo ba sehlopha ka seng (mohlala, likarolo tse thibetsoeng lipakeng tsa bacha le batho ba baholo) ba sebelisa Z-teko ea likarolo tse peli (Lethathamo 1). Mosebetsi oa nakong e fetileng le karohano ena ea boits'oaro e bonts'itse hore bacha le baholo ba etsa karabo ea bohlokoa ka mokhoa o tsitsitseng ka nako ea 3 (Sturman et al., 2010). Ka hona, ntle le moo ho boletsoeng ka hosele, litlhahlobo tsa electrophysiological li hlahisoa bakeng sa liboka tsa 3-6, moo mokhatlo oa sephetho sa liketso o ithutiloeng hantle ke lihlopha ka bobeli. Mona le libakeng tse ling, khopolo-taba e fosahetseng e ile ea lahloa ha p <0.05.

Lethathamo 1

Ho bapisoa ha ts'ebetso ea unit ea batho ba lilemong tsa bocha le ba baholo ka lifensetere tse khethiloeng tse bolokiloeng nakong ea liketsahalo. Windows e khahlisang e koetsoe ke cue, ala poke (poke) kapa e kenella ka sejelong sa lijo (FT). Karolo ea bocha (Adol) le motho e moholo ...

Litlhahlobo tsa phapang ea litekanyetso tsa ho thunngoa li ile tsa balelloa joalo ka mabaka a Fano (spike count variance / mean) li sebelisa fensetere ea 80 ms e tsamaeang ka mehato ea 50 ms. Bakeng sa yuniti e 'ngoe le e' ngoe, phapang ea palo ea linokoane le bolele ba spike li ne li baloa ka nako e ngoe le e ngoe. Ho thellisa mohopolong o amanang le ho fapana le meelelo ea likarolo tsohle ho ne ho reriloe mohatong o mong le o mong oa fensetere, ho fana ka mokhoa oa nako oa Fano mabapi le liketsahalo tsa mosebetsi. Ho hlahloba hore na liphetoho tse bonoang ho Fano li felletse ka nako (le liphapang tse amanang le lilemo tsa Fano) li bile teng ka lebaka la liphetoho mokhoeng oa ho thunya ho fapana le phapang, re sebelisitse mokhoa o ts'oanang oa Churchland le basebetsi-mmoho (Churchland et al., 2010). Thutong ea pele re ile ra etsa lintho tse tšoanang ka thoko bakeng sa bana ba lilemong tsa bocha le ba baholo. Mokhoa ona o ne o thusa ho fana ka sekhahla sa ho chechisa moelelo o mong le o mong nako le nako, ka ho hleka ka mokhoa o sa reroang le ka makhetlo. Likhakanyo tsa lintlha tsa Fano bakeng sa ntlha ka 'ngoe ea nako li ne li thehiloe ka karolelano ea li-iterations tsa 10 tsa ts'ebetso ena. Ts'ebetso ena e netefalitsoe e le mokhoa o sebetsang oa ho qoba lintho tsa khale ka lebaka la liphetoho tsa sekhahla sa ho thunya (Churchland et al., 2010). Ntle le sena, ho ile ha etsoa tlhahlobo e ikhethileng e amanang le bolele, eo ho eona ho buuoang ka boholo ba nako ea ho thunngoa historing nakong ea sehlopha sa lilemo (joalo ka holimo) le lipakeng tsa lihlopha tsa lilemo. Ho shebella lintlha tse tšoanang tsa Fano tse tala le tse sa reng letho li ka tiisa hore lithuto tsa nako le phapang e amanang le lilemo Fano factor li bonts'a phapang ea nako ea sepheo 'me e ne e se feela litsebi tsa ho se tšoane ka moelelo oa ho chechisa sekhahla. Bacha ba Fano le batho ba baholo ba ne ba bapisoa ka lipalo ho latela liteko tsa kakaretso ho Matlab.

Results

boitshwaro ja

Nakong ea ts'ebetso ea boits'oaro, bacha ba kenella ka sekoting se koahetsoeng ka bobebe hore ba fuoe matla a ho khothaletsa lijo (Setšoantšo sa 1a). Ha ho na phapang e kholo e hlokometsoeng lipakeng tsa bacha le batho ba baholo ka palo eohle ea liteko F (1,1) = 1.3, p = 0.28, latency ho tloha qalo ea qalo ho ea ho karabelo ea methapo F (1,1) = 0.34, p = 0.57, kapa latency ho tsoa karabelo ea bohlasoa mabapi le ho khutlisetsoa ha lijo pellet F (1,1) = 1.2, p = 0.31. Mosebetsi ona o ne o etsoa ka mehla le ka botlalo ke liphoofolo tse holileng le tse lilemong tsa bocha ke sehlopha sa boraro sa thupelo (Setšoantšo sa 1c).

Bohlokoa ba Tšimo ea Lehae

Ho rekota hoa LFPs ka motlakase, mohato o nahanoang ho bonts'a tšebetso ea barekisi ba libaka, o senotse mekhoa e ts'oanang bakeng sa bacha le batho ba baholo ka mosebetsi o mongata, o nang le phapang e kholo matleng a ponahalo hang hoba kamor'a lijo tse kenang sekoleng hore ba fuoe matla.Setšoantšo sa 2a). Ka nako eo, batho ba baholo ba bonts'a matla a alpha (8-12 Hz) le beta (13-30 Hz). Theta (4-7 Hz) le matla a fokolang a gamma (31-75 Hz) li ne li tšoana lipakeng tsa lihlopha, ha bacha ba ne ba e-na le matla a maholo a "gamma" (76-100 Hz) ho feta batho ba baholo (Setšoantšo sa 2b).

Setšoantšo sa 2

Bacha le batho ba baholo ba li-LCP tsa LCP nakong ea lithuto 3-6. A) Sekhahla sa matla a LFP bakeng sa bacha le batho ba baholo lifesetereng tse potolohileng liketsahalo tsa bohlokoa li ile tsa fetoloa nako ea motheo (3 ho 1 ea bobeli pele ho cue qala) khafetsa e ngoe le e ngoe. Nako ea nako ea tloaelo ...

Tlhahlobo ea Fano Factor

Phapang e amanang le lilemo e ile ea bonoa ho fapana ha litekanyetso tsa ho thunya ho amanang le liketsahalo tse itseng. Karolo ea Fano, e leng moelelo oa kamano lipakeng tsa phapang ea palo ea spike le palo ea palo ea boiketsetso (Churchland et al. 2010), e ne e lekantsoe ho lekola phapang ea nako ea spike ho feta liteko (Setšoantšo sa 3). Bacha (likhoto tse 8 li-unit tse 265) li ne li na le lintlha tsa Fano tse kholo ho feta batho ba baholo (likhoto tse 4 li-unit tsa 184) nakong ea liboka tsa 3-6 (litšoantšiso tse entsoeng ka liteko tsa limaraka) nakong ea 2 ea motheo Z = 6.90, p <0.01, ka Fensetere ea 1-sec hang hang kamora 'qalo ea liteko Z = 5.48, p <0.01, ka fensetere ea 1 s e shebaneng le karabelo ea liletsa Z = 3.12, p <0.01, le motsotsoaneng o le mong o lebisang katolosong ea matlafatso Z = 3.77 , p <0.01 (Setšoantšo sa 3). Hobane lipalo tsa Fano factor li itšetlehile ka boholo ba fensetere le mohato re fapanyetsitseng likarolo tsena ka nako ho potoloha mohopolo oa lipina ho bonts'a hore ha boholo le ho hlaka ha lipalo li ameha, nako le nako le maemo a amanang le lilemo a lula a le teng (Setšoantšo se tlatsetsang 2) Re sebelisitse mokhoa o ts'oanang o bolelang (Churchland et al. 2010) ho ts'oara sekhahla sa ho thunya ho bolelang nako e batlang e feta nako e telele e le hore liphetoho tse bokhutšoanyane ba nakoana tsa ho thunya li se ke tsa fifatsa tlhaloso ea rona ea Fano factor joalo ka tekanyo ea phapang (Setšoantšo se tlatsetsang 3a). Ka mokhoa o ts'oanang re lekantse kabo ea litekanyo tsa ho thunya lipakeng tsa lihlopha tsa lilemo (Setšoantšo sa tlatsetso 3b). Mabaka a Raw Fano a ne a ts'oana haholo le a kopantsoeng ka mokhoa o tšoanang, ho netefatsa hore nako e bontsitsoeng ea Fano e bonts'a phapang ea nako ea spike ho sa tsotelehe hore e bolela matla a ho chechisa sekhahla. Taba e 'ngoe ho sena e ne e le kamora ho khutlisetsoa ka matla, ka nako eo batho ba baholo ba neng ba bontša lintlha tse kholo tsa Fano (Setšoantšo sa 3). Phapang ena e ne e bakiloe bonyane ka karolo ea liphetoho litekanyetsong tsa ho thunya, hobane ho ne ho se na phapang e kholo ea lipalo linthong tse amanang le Fano nakong eo (Setšoantšo se tlatsetsang 3). Liphumano tsena li bonts'a hore liketsahalo tse ikhethang li lebisa phokotsong ea phapang ea nako ea liiketsetso bakeng sa bacha le batho ba baholo, le hore ho khahlisang, nako ea bocha ea spike ea bongoaneng hangata e fapana haholo ho feta ea batho ba baholo nakong eohle ea mosebetsi. Phokotso ea Fano factor e tsamaisoa ke Stimulus ho nahanoa hore ke thepa e akaretsang ea boqapi ba cortical (Churchland et al. 2010). Kahoo, lintlha tse phahameng tsa Fano li ka fana ka maikutlo a hore tšekamelo ea nako ea maiketsetso ha e laolehe haholo ho OFC ea lilemong tsa bocha ha e bapisoa le batho ba baholo.

Setšoantšo sa 3

Tlhahlobo ea Fano e bapisa ho fetoha le maemo ha ba lilemong tsa bocha le ho tsubela batho ba baholo. Karolo ea Fano ke letsoalo la phapang ea palo-ea litekanyetso le palo ea palo ea batho ba sa tšepahaleng. Ho sebelisa fensetere e thellang, khakanyo ena ea ho feto-fetoha ha maemo e ile ea ngoloa ka nako ea nako ...

Mosebetsi oa Yuniti

Tlhahlobo ea tšebetso ea neural unit e le 'ngoe nakong ea mosebetsi e senotse liphapang tse ikhethang tse ikhethang tsa liketsahalo pakeng tsa bacha le batho ba baholo. Nakong ea thuto 1, pele ho ho ithuta mekhatlo ea sephetho, tšebetso e ile ea fetoha hanyane ho ea liketsahalong tsa sehlopha ka seng. Ha mosebetsi o se o ithutile hantle (lithuto tsa 3-6), leha ho le joalo, liketsahalo tsa mosebetsi li ile tsa hlahisa mekhoa e sa fetoheng ea tšebetso ea neural (Setšoantšo sa 4). Litefiso tsa ho hlasela tsa mantlha tse tloaelehileng tsa yuniti e ngoe le e 'ngoe e notletsoeng liketsahalo tsa mosebetsi li bonts'oa Setšoantšo sa 5a, e bonts'a bophara le boholo ba ts'ebetso ea phasic neural. Ho batho ba baholo (likhoto tsa 4, likarolo tsa 184), empa eseng bacha (likhoto tsa 8, likarolo tsa 265), ketsahalo e tloaelehileng e fokotsehile ho cue le pele ho karabelo ea mochini (Setšoantšo sa 5b). Kamora karabelo, ts'ebetso e tloaelehileng ea palo ea lihlopha ka bobeli e ile ea theoha, 'me bacha ba eketseha ho feta batho ba baholo. Ka nako eohle ea ts'ebetso ea matlafalo ea baahi e eketsehile, 'me batho ba baholo ba hola ka pele le ka tekanyo e tlase ho feta ea bacha. Ts'ebetso e kholo ea bocha e fihlile ka nako ea ho kenella lijong; ka nako eo ha karolelano mesebetsi ea batho ba baholo e ne e le tlase haholo. Leha palo e fokola haholo ka palo ho fihlela sephetho se matla, likarolo tsa bocha (n = 8) le batho ba baholo (n = 5) li-interneurons tse potlakileng tsa li-spiking li bonts'itse mokhoa o tšoanang oa ts'ebetso ho pota liketsahalo tse khahlisang joalo ka palo ea batho ka kakaretso ea lihlopha nakong ea mananeo 3 - 6 (Setšoantšo se tlatsetsang 4) .Ho bapisoa le karolo ea liketsahalo tsa thabo le tse thibeloang ke liketsahalo tsa mokete (Setšoantšo sa 5c) ka kakaretso e senotsoe likarabo tse fokotsitsoeng tsa likarabo le likarabo tse ts'oanang kapa tse ntlafatsang tsa bosholu. Tlalehong ea 1 s ho latela mohlala, batho ba baholo ba ne ba e-na le karolo e kholo ea likarolo tse thibelitsoeng ho feta bacha ba nang le li-unit tse ka bapisoang (Lethathamo 1). Kamora karabelo ea bohlokoa, ha bacha le batho ba baholo ba ne ba fokotseha ka mokhoa o tšoanang mesebetsing ea baahi, tekano e ts'oanang ea likarolo tse kentsoeng le tse thibelitsoeng e ile ea bonoa. Tlhahlobo ea likarolo tse tsamaeang ea fensetere, e sebelisitsoeng ho bona nako ea ho haroa hoa neural, e bonts'itse hore ho potoloha karabo, lisebelisoa tse thibeloang batho ba baholo li ile tsa thibeloa pejana 'me tsa ba le nako e telele ho feta lilemong tsa bocha.Setšoantšo sa 5c). Sena se netefatsoa ke ho lekola boholo ba liuniti tse thibetsoeng ka nako windows 0.5 s pele le 1 - 1.5 s kamora karabelo ea seletsa (Lethathamo 1). Le ha likarolo tsa batho ba baholo le tsona li bonahala li hiroa pele ho tsa bocha, liphapang tsena e ne e se tsa bohlokoa lipalo. Boholo ba diyuniti bo arotsoe ka mokhoa o ts'oaroang le o thibetsoeng bo fapane haholo ho potoloha le matlafatso, ka batho ba baholo ba nang le litekanyo tse kholo tsa li-unit tse thibetsoeng le bacha ba nang le likarolo tse kholo tsa li-unit tse butsoeng. Ka 0.5-1 s kamora ho matlafatsoa, ​​ho ne ho se na phapang e amanang le lilemo sehlopheng sa lihlopha. Liphuputso tsena li bonts'a hore leha likarolo tse ts'oanang tsa lilemong tsa bocha le tsa batho ba baholo li ka ts'oaroa kapa tsa thijoa ka linako tse fapaneng (mohlala, poke ea lisebelisoa), boholo ba mosebetsi bacha ba ne ba na le likarolo tse nyane tsa likarolo tse thibetsoeng.

Setšoantšo sa 4

Karolelano ea basekete-e tloaelehileng ea boemo ba ho thunya + 1 sem (shading) bakeng sa likarolo tsohle tsa batho ba baholo le ba lilemong tsa bocha, tse notletsoeng nako ho etsa liketsahalo nakong ea e 'ngoe le e' ngoe ea lithuto tse tšeletseng. Tekanyo e akaretsang ea ho thunya e akaretsang bakeng sa bohle ba lilemong tsa bocha e ne e le 4.66Hz mme likarolo tsohle tsa batho ba baholo e ne e le 5.18Hz. ...
Setšoantšo sa 5

Phasic OFC ea baahi le ts'ebetso ea yuniti e le 'ngoe nakong ea liboka 3-6. A) Merero ea mocheso e emela sekhahla se tloaelehileng sa ho thunya mocha ea lilemong tsa bocha (n = 265; libaka tse holimo) le motho e moholo (n = 184; likarolo tse tlase) yuniti. Mokha o mong le o mong ke ts'ebetso ea motho ka mong ...

Puisano

Maemong kaofela a palo ea baahi le a yuniti e le 'ngoe, OFC ea lilemong tsa bocha e sebetsana le boitšoaro bo susumetsoang ka mokhoa o fapaneng le oa batho ba baholo, ka phapang e totobetseng e sa boleloe haholo phokotso ea bohlankana mesebetsing ea neural nakong ea moputso le liketsahalo tse ling tse ikhethang. Bacha ba boetse ba bontšitse phapang e kholo ea ho boloka nako ea tšebeliso ea mantlha mosebetsing o mongata. Nakong ea matlafatso, ho kenyelletsa phokotso e fokolang ea ts'ebetso, ho ne ho na le karolo e kholo ea likarolo tsa bongoana tse ileng tsa eketsa ts'ebetso ea bona, hammoho le phapang ea matla a alpha, beta le gamma LFP ha e bapisoa le batho ba baholo. Ho bohlokoa hore liphapang tsena tse amanang le neano tse amanang le lilemo li ile tsa bonoa leha ts'ebetso ea mosebetsi e ne e ts'oana, ho bonts'ang hore phapang e joalo ha e bonts'e pherekano ea boitšoaro feela (Schlaggar et al., 2002; Yurgelun-Todd, 2007). Le ha ho eketsoa lithuto tse ling e ne e le ho senola liphapang tsa boitšoaro nakong ea thupelo ea pele, bacha le baholo ba ile ba etsa mosebetsi ka tekanyo e kholo ho tloha sehlopheng sa boraro ho ea pele. Boithuto ba rona ba electrophysiology bo shebana haholo le li-program tsena tsa kamoso, ha mokhatlo oa sephetho sa ts'ebetso o ithutile hantle ke lihlopha ka bobeli. Re khethile mosebetsi oa boits'oaro oo, leha o le bonolo ho lekana hore o ka ithutoa ka nako e khuts'oane ea nako ea bohlano ea bocha, o ka nkuoa e le sebopeho sa motheo sa boits'oaro bo rarahaneng ba ho susumetsa. Kahoo, liphumano tsena li bonts'a hore le ha bacha ba etsa boitšoaro bo tšoanang le ba batho ba baholo, ho kenyelletsa litaba tsa liketsahalo tsa bohlasoa le ts'ebetso e hlakileng ea ts'ebetso (joalo ka ha ho amana le ho fapana ha nako ea spike) ka ho fapana.

Li-neurons tsa lilemong tsa bocha li ne li tloaetse ho ba le ts'ebetso e fokotsehileng ho feta ea batho ba baholo nakong ea liketsahalo tsa bohlokoa tse kang ho qala ha nyeoe, pele ho karabelo ea 'mino, le pele le nakong ea moputso. Phapang e joalo e amanang le lilemo e ka ba ka lebaka la thibelo e nyane ea OFC ea neuronal ka linako tsena. Neuronal inhibition e bapala karolo ea bohlokoa ho hokahanyang mesebetsi ea oscillatory (Fries et al., 2007; Cardin et al., 2009; Sohal et al., 2009), ho laola nako e nepahetseng ea nako ea spike, le ho ntlafatsa ts'ebetso ea puisano ea neuronal (Buzsaki le Chrobak, 1995). Ho feto-fetoha ha maikutlo joalo ka ha ho lekantsoe le EEG le LFP, ke ho feto-fetoha ha maemo le maemo ho thabo ea thabo, eo ho nahanoang hore e bonts'a tšebelisano ea thepa ea methapo ea cellular le potoloho.Buzsaki le Draguhn, 2004), e hlophisang nako ea tlhahiso ea seiketsehi hantle (Meketso, 2005). Synchrony ea oscillations e ka fana ka sebaka sa puisano ea lihlopha tsa neuronal (Meketso, 2005), hape e ka ba karolo ea mantlha ea tlamahano e hlakileng le lits'ebetso tse ling (Uhlhaas et al., 2009b). Mehato ea methapo ea kutlo ea methapo ea methapo e lumellanang le ho sebetsa ha kelello maemong a mangata (Basar et al., 2000; Hutcheon le Yarom, 2000) mme li fokotsoe libakeng tse 'maloa tsa pathological, joalo ka schizophrenia (Uhlhaas le Sebini, 2010). Uhlhaas le basebetsi mmoho ba fumane phapang litabeng tse amanang le mosebetsi tsa EEG pakeng tsa bacha le batho ba baholo (Uhlhaas et al., 2009a). Tumellanong le liphumano tsena, re fumane keketseho e nyane ho matla a alpha le beta ho OFC ea bacha nakong ea matlafatso. Li-frequency band tsena ho nahanoa hore li bohlokoa bakeng sa puisano ea neural ka libaka tse telele (Pfurtscheller et al., 2000; Brovelli et al., 2004; Klimesch et al., 2007), e ka se sebetseng hantle bacheng. Tlhaloso ena e lumellana le lithuto tse bonts'ang hore khokahano e sebetsang ea phetoho e fetoha ho ba sebakeng sa habo ho fetisoa haholo ka nts'etsopele (Fair et al., 2009; Somerville le Casey, 2010).

Re bone hape le liphapang tse amanang le lilemo tsa ho fofa ka sekhahla sa ho thunya ho pholletsa le liteko, tse lekiloeng ho sebelisoa tlhahlobo ea Fano factor. Mosebetsi oa morao-rao o bontšitse hore libakeng tse ngata tsa cortical, ts'ebetso ea spiking ea neuron e ts'oaroa ke boitšoaro bo susumetsang kapa ba ts'ebetso, joalo ka ha bo bonts'itsoe ke lintlha tse fokolisitsoeng tsa Fano (Churchland et al., 2010). Kannete re hlokometse hore ho OFC, boits'oaro ba tšebetsong, mokhoa oa moputso / tebello, le matlafatso (ho batho ba baholo) ho lebisitse ho fokotseng tekanyo ea tekano ea ho thunya ha sekhahla. Phokotso e kholo ka ho fetohang e etsahetse joalo ka likhoto li ile tsa arabela karabo ea bohlokoa le nakong ea tlhatlhobo pele. Tekanyo e kholo ea ho thunya e ne e ka lebelloa haeba nako ea ts'ebetso ea phasic neural e ne e sa laoloe ka thata, joalo ka ha ho le joalo ho OFC ea bacha. Bacha ba ne ba na le mabaka a maholo a Fano ho feta batho ba baholo ka lebaka la mosebetsi o mongata, ntle le nako ea 1-s hang ka mor'a ho kenella lijong. Liphetho tsena li bonts'a hore bacha ba na le phapang e batsi haholo ea ho thunya, e leng se ka supang ts'ebetso e fokotsang ts'ebelisong ea neural. Ka mantsoe a mang, lintlha tsa Fano tse kholo li bontša hore li-neuron tsa lilemong tsa bocha li kenella liketsahalong tse tšoanang le maemo, ho tloha tekong ho ea tekong, e ka bolelang palo e tlase ea lerata ho khoutu e lumellanang le ea batho ba baholo. Sena se tsamaisana le ho fumana hore menyetla e amanang le liketsahalo tsa bana le bacha e na le litekanyetso tse tlase tsa lerata ho feta batho ba baholo, e ka bang ka lebaka la "ho se tsitse ha motho ka mong" libakeng tsa boko tse hlahisang mats'oao ana (Segalowitz et al., 2010). Joalo ka ha ts'ebetso ea neural e le bohlokoa bakeng sa ho kenyelletsa oscillation, marang-rang a inhibitory a fana ka nako e nepahetseng ea ho chechisa lisele tse ka sehloohong (Buzsaki le Chrobak, 1995). Kahoo, khokahano e kanna ea ba teng lipakeng tsa tloaelo ea ho ba lilemo tsa bocha ho bonts'a thibelo e fokolang ea phasic liketsahalong tse ikhethang, le phapang e kholo ea litekanyetso tsa bongoana. Re tlameha ho bontša tlhokomeliso, leha ho le joalo, hore khokahano e joalo ha e na ho bua ka kotloloho, hobane nako ea phapang e kholo ka ho fetisisa ea Fano e ne e se nako ea liphapang tse kholo ho phasic inhibition.

Liphetoho tse kholo tsa neurodevelopmental li etsahala lilemong tsa bocha. Ho na le phokotso ea taba e putsoa le ho eketsoa hoa litaba tse tšoeu nakong ena (Benes et al., 1994; Paus et al., 1999; Paus et al., 2001; Sowell et al., 2001; Sowell et al., 2002; Sowell et al., 2003; Gogtay et al., 2004). Li-receptor tsa li-neuromodulators tse 'maloa tse kang dopamine li hlahisoa maemong a phahameng ho bacha ho feta ho batho ba baholo ho PFC le basal ganglia (Gelbard et al., 1989; Lidow le Rakic, 1992; Teicher et al., 1995; Tarazi et al., 1999; Tarazi le Bal)arini, 2000). Ho litoeba tse sa sebetseng, ts'ebetso ea tlhaho e ikatileng ea li-dopamine neurons e kholo ho bacha ho feta bana ba banyenyane kapa ba baholo (McCutcheon le Marinelli, 2009). Ka li-cortical slices activating the dopamine D2 receptor agonist e ba teng feela ke bongoana kapa ho ba motho ea moholo lilemong, eo ka nako eo ho fetoha ha tšohanyetso ho bonoang (Tseng le O'Donnell, 2007). Polelo ea li-receptor tsa NMDA mabapi le li-neurons tse potlakileng (FS) le tsona li fetoha haholo ho PFC ea bacha. Boholo ba li-interneurons tsa FS tsa lilemong tsa bocha ha li bontše maqhubu a pakeng tsa NMDA receptor-Mediated. Lisele tseo tse nang le tsona li bonts'a tekanyo e fokotsoang hole haholo ea NMDA: AMPA (Wang le Gao, 2009). Lithuto tsena li bonts'a phapang ea mantlha mohahong le boikoetlisong ba 'mele ba bongoana le li-transmit tse amanang le boits'oaro bo susumetsang le ho ba kotsing ea kelello. Boithuto ba hona joale, boo tsebong ea rona e leng bona ba pele ba ho sebelisa rekoto ea motlakase oa kantle ho tsoelo-pele, liphoofolo tse lilemong tsa bocha li ntšetsa pele tšebetso ea liphumano tsena tsa seleng le tsa limolek'hule ka ho bonts'a hore mosebetsi o amanang le neural o fapane haholo ho bacha nakong ea ts'ebetso ea semmuso. liketsahalo.

Lithuto tsa fMRI tsa batho li fumane hore ts'ebetso ea bongoana e fana ka moputso le tebello ea moputso ka mokhoa o fapaneng ho feta batho ba baholo ka lebelo le leholo la tikoloho (Ernst et al., 2005; Galvan et al., 2006; Geier et al., 2009; Van Leijenhorst et al., 2009). Litlhaloso tsa hajoale bakeng sa bofokoli bo itseng ba boits'oaro ba lilemong tsa bocha li kenyelletsa taba ea hore PFC e "ntlafalitsoe" ho latela ts'ebetso ea eona le / kapa ho hokahana ha eona ka ts'ebetso le ho feto-fetoha ha likarolo tsa subcortical (Ernst et al., 2006; Casey et al., 2008; Steinberg, 2008). Boithuto ba hona joale bo fumana hore phapang ea nts'etsopele e bonahala esita le nakong ea boits'oaro bo khothalletsoang ke moputso, mme e bonts'oa maemong a yuniti e le 'ngoe ka tekanyo e fokolisitsoeng ea mosebetsi o fokotsoang oa neural ho bacha ba OFC ho isa ho boholo, empa ha se liketsahalo tsohle tse ikhethang. Le ha ho hlokahala mosebetsi oa nako e tlang ho theha khokahano e joalo, phapang e le 'ngoe ea likarolo tse le' ngoe molemong oa likarabo tse thibelitsoeng e kanna ea ba mohloli oa phapang e bang teng lilemong tsa bocha e shebaneng le matla a sa hlasimoloheng le ho fapana ha nako ea spike. Ka lebaka la bohlokoa ba thibelo ho laola nako e nepahetseng ea li-spikes, ho kenya li-oscillations, mme ka tsela eo ho tsamaisa puisano e sebetsang ea lihlopha tsa neuronal (Buzsaki le Chrobak, 1995; Fries et al., 2007), thibelo e fokotsang ea bongoana ea PFC e tsamaisana le ho shebella liphapang tse kholo ts'ebetsong ea cortical e bonoang thutong ena le tse ling. Leha ho le joalo, tšekamelo ea bacha ea ho ba le phokotso e tlase mesebetsing ea unit ho potoloha liketsahalo tse ikhethang li ka hlaha ka lebaka la phokotso e tlase mesebetsing ea litumelo tse khahlisang hammoho le thibelo e fokotsang.

Mosebetsi o fetotsoeng oa cortical inhibitory o ka susumetsa boits'oaro bo litšila (Chudasama et al., 2003; Narayanan le Laubach, 2006) mme e amana le linaha tse ling tsa pathological (Chamberlain et al., 2005; Lewis et al., 2005; Behrens le Sejnowski, 2009; Lewis, 2009). Mohlala, batho ba nang le lefu la schizophrenia ba fokotse polelo ea GAD67 mRNA, enzyme e kenyang karolo ea motsoako oa inhibitory neurotransmitter GABA (Akbarian et al., 1995). Bakuli ba Schizophrenia le bona ba fokotse li-cartridge tsa GABA membrane (GAT-1) -mmunoreactive axon cartridges ho PFC (Woo et al., 1998). Sena ke sa bohlokoa bo boholo lipatlisisong ho bacha, joalo ka li-cartridge tsa GAT-1 immunoreactive (tseo hape li lokelang ho hlaha ho parvalbumin) pejana feela nakong ea bocha ebe li fokotseha haholo nakong ea bocha.Cruz et al., 2003), nako e tloaelehileng ea ho qala ha schizophrenia. Mosebetsi oa nako e tlang oa ho hlahloba mohloli o nepahetseng oa ts'ebetso ea phasic e amanang le lilemo nakong ea kholo e tloaelehileng e ka sebetsa ka kotloloho ho pathophysiology le matšoao a nako ea mafu a kelello a hlahang nakong ea bocha.

Boitsebiso bo Eketsehileng

Supp1

lumela hore baa fokola

Mosebetsi ona o tšehelitsoe ke National Institute of Mental Health, Pittsburgh Life Sciences Greenhouse, le Andrew Mellon Foundation bakeng sa botsoalle ba predoctoral (DAS). Re leboha Jesse Wood le Yunbok Kim ka lipuisano tse bohlale, le Churchland le basebetsi-mmoho (Churchland et al., 2010) bakeng sa ho etsa hore mesebetsi ea Matlab e fetohe.

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