I-Neurodevelopmental trajectories ye-cortex ye-cerebral yomntu (2008)

J Neurosci. 2008 Apr 2;28(14):3586-94. doi: 10.1523/JNEUROSCI.5309-07.2008.

Shaw P1, Kabani NJ, Lerch JP, Eckstrand K, Lenroot R, Gogtay N, UGreenstein D, Udidi L, UEvans A, Rapoport JL, Giedd JN, Ubulumko SP.

Abstract

Ukuqonda umbutho we-cerebral cortex kuhlala kuyingqwalasela ephambili ye-neuroscience. Iimephu zeCortical zithembele ngokukodwa kuvavanyo lwezicubu ze-postmortem ukwakha imephu yesakhiwo, ye-architectonic. Ezi mephu ziye zahlula ngokungaguquguqukiyo phakathi kweendawo ezineeleya ezimbalwa ezibonwayo, ezinepateni engantsokothanga kakhulu yokunyibilika kunye neziswele umaleko wegranular wangaphakathi, kunye nezo zineengqimba ezintsonkothileyo zoyilo lwelaminar. Eyangaphambili ibandakanya iindawo ezininzi ze-agranular limbic, kwaye le yokugqibela ibandakanya i-homotypical and granular area of ​​association kunye ne-sensory cortex. Apha, sinxulumanisa ezi mephu zemveli kunye nedatha yophuhliso esuka kwi-neuroimaging engaphazamisiyo. Utshintsho kubunzima becortical yamiselwa kwi-vivo ukusuka kwi-764 ye-neuroanatomic magnetic resonance imifanekiso efunyenwe ngobude ukusuka kuma-375 edla ngokuphuhla abantwana kunye nabantu abadala abancinci. Sifumana amanqanaba ahlukeneyo okuntsonkotha kokukhula kwecortical kuyo yonke i-cerebrum, ehambelana ngokusondeleyo neemephu zoyilo lwezakhiwo. Imimandla yeCortical enolwakhiwo olulula lwelaminar, kubandakanya uninzi lweendawo zemilenze, ubukhulu becala ibonisa iindlela ezilula zokukhula. Ezi ndawo zichonge ngokucacileyo ii-homologues kubo bonke ubuchopho bezilwanyana ezanyisayo kwaye ke kusenokwenzeka ukuba zavela kwizilwanyana ezanyisayo zakuqala. Ngokwahlukileyo, i-polysensory kunye ne-high-order association iindawo ze-cortex, ezona ndawo zinzima kakhulu ngokubhekiselele kwi-architecture yazo ye-laminar, nazo zineendlela ezinzima kakhulu zophuhliso. Eminye yale mimandla yahlukile, okanye yandiswe ngokumangalisayo kwiiprimates, iboleka ukubaluleka kwendaleko kwizinto ezifunyanisiweyo. Ngaphaya koko, ngokwenza imephu uphawu oluphambili kwezi trajectories zophuhliso (iminyaka yokufikelela incopho yobunzima becortical) sibhala ukuvuthwa okuguquguqukayo, okune-heterochronous kwe-cerebral cortex ngokusebenzisa ulandelelwano lokudlula kwexesha (“imiboniso bhanyabhanya”).

Icandelo langaphambiliIcandelo elilandelayo

intshayelelo

Uninzi lweemephu ze-cerebral cortex yomntu ziye zahlulahlula ngokweempawu ze-histological, ezinje ngokusasazwa kwemizimba yeeseli okanye i-myelin, kwaye, kutsha nje, iziphawuli zeemolekyuli (von Economo kunye noKoskinas, ngo-1925; I-Ongur et al., 2003; Zilles et al., 2004). Ukuthelekiswa kokuhlelwa okufanayo phakathi kweentlobo ezininzi zeentlobo zibonelela ngembono yendaleko, kwaye uhlalutyo olunjalo luchonge iindidi ezimbini ze-cortical. Olunye uhlobo, i-allocortex, lunefomu yamandulo enamaleko amathathu efana kakhulu ne-homologues yayo kwizirhubuluzi. Olunye uhlobo, i-isocortex, alunazo ii-homologues kwaye lunolwakhiwo oluphuma ngaphezulu oluphawulwa ngamacandelo angaphezulu kwesithathu (ngokuqhelekileyo amathandathu) kunye nepateni entsokothileyo yoqikelelo oludibeneyo kunye ne-efferent (Kaas, 1987; Puelles, ngo-2001; UAllman et al., ngo-2002; Striedter, ngo-2005). Phakathi kwe-allocortex kunye ne-isocortex, iindawo ngamanye amaxesha ezibizwa ngokuba yi-"transition cortex" zinezakhiwo eziphakathi. Ukusebenzisa i-neuroimaging yesakhiwo kwi vivo Ukukhula kwengqondo, siye saphonononga ukuba kunokwenzeka ukuba ezi ntlobo zahlukeneyo ze-cortex zibonisa amanqanaba ahlukeneyo okuntsonkotha kwiindlela zokukhula kwabo ebuntwaneni nasekufikiseni.

Sisebenzisa i-neuroanatomy yokubala, sichaze ubukhulu becortical kumanqaku angaphezu kwama-40,000 kuyo yonke i-cerebrum kwiqela labantwana abasempilweni abangama-375 kunye nolutsha olufikisayo. Ubukhulu beCortical bonyulwe njengemetric ethi zombini ibambe uyilo lwekholamu ye-cortex kwaye inovelwano kutshintsho lophuhliso ekuphuhliseni ngokwesiqhelo kunye nabemi bekliniki (Lerch et al., 2005; O'Donnell et al., 2005; UMakris et al., ngo-2006; UShaw et al., 2006a,b; Lu et al., 2007; Sowell et al., 2007).

Uninzi lwabantwana abakwiqela lethu babephindaphinda i-neuroanatomic imaging, kwaye idatha enjalo yobude inokudibaniswa nedatha yecandelo elinqamlezayo ukwenza imodeli yotshintsho lophuhliso, kunye nedatha yelongitudinal inolwazi ngakumbi. Kubunzima becortical, eyona trajectory ilula enokufakwa ukuchaza utshintsho lwayo ngokuhamba kwexesha ngumgca othe tye. Imifuziselo yokukhula entsonkothileyo ngakumbi ibandakanya izigaba ezicacileyo zokukhula kunye nokuncipha kobukhulu becortical: imodeli yequadratic inezigaba ezibini ezinjalo (ngokuqhelekileyo ukonyuka kokuqala okufikelela kwincopho ngaphambi kokuncipha) kwaye imodeli ye-cubic inezithathu. Ubunzima bokukhula bunokwahluka kwi-cortex ye-cerebral, kwaye siye sazama ukuphonononga ukuba ngaba olu tshintsho luhambelana neempawu ze-cytoarchitectural.

Iimpawu ezithathwe kwii-curves zophuhliso, ezifana nobudala bokufikelela kwiindawo ezahlukeneyo zokuguquguquka, zihlala zisetyenziswa njengezalathisi zophuhliso (Tanner et al., 1976; UJolicoeur et al., ngo-1988). Kubunzima becortical, iminyaka apho ubungqingqwa becortical incopho bufikelelwa khona (indawo apho ukonyuka kunika indlela yokuncipha kobukhulu becortical) kunokumiselwa ngamanqaku ecortical nokuba yicubic okanye quadratic (kodwa hayi emgceni) trajectory kwaye, ke, ivela njenge. isalathiso esinokuba luncedo sophuhliso lwecortical. Ke ngoko siphinde savavanya ipateni yokufikelela incopho yobukhulu be-cortical kwi-cerebrum, ukuqinisekisa kunye nokwandisa uqwalaselo lwangaphambili lolandelelwano lwe-heterochronous, kunye neendawo eziphambili zeemvakalelo ezifikelela incopho yobukhulu be-cortical phambi kwe-polymodal, iindawo ezihambelanayo eziphezulu (UGogtay et al., 2004).

Impahla nenkqubo

Abathathi-nxaxheba.

Abantwana abangamakhulu amathathu anamashumi asixhenxe anesihlanu kunye nolutsha, abantwana abasempilweni abangenayo imbali yomntu okanye yentsapho yokuphazamiseka kwengqondo okanye ye-neurological, babenayo yonke imifanekiso ye-764 magnetic resonance. Isifundo ngasinye sigqibe uLuhlu lokuHlola lokuziphatha kwaBantwana njengesixhobo sokuhlola kwaye emva koko senziwa udliwano-ndlebe olucwangcisiweyo lokuxilonga ngugqirha wengqondo yomntwana ukuba akhuphe naluphi na uxilongo lwengqondo okanye lwe-neurological (UGiedd et al., 1996). Ukunikezelwa kwesandla kunqunywe kusetyenziswa i-PANESS (Uviwo lwePhysical and Neurological Examination for Soft Signs), kwaye i-336 (90%) yayiyisandla sasekunene, i-20 (5%) ubukhulu becala ngasekhohlo, kunye ne-19 (5%) i-ambidextrous. I-intelligence quotient (IQ) yayiyi-115 (SD, 13) njengoko imiselwe ukusuka kwiinguqulelo ezifanelekileyo ze-Wechsler Intelligence Scales (UShaw et al., 2006b). Ubume boqoqosho lwentlalo (SES) luchongwe kwi-Hollingshead Scales kunye nenani eliphakathi kwe-40 (SD, 19) (Hollingshead, ngo-1975). Uluhlu lobudala lususela kwi-3.5 ukuya kwi-33 yeminyaka, kwaye ukwabiwa kweminyaka yeskeni kubonisiwe Umzobo 1. Izifundo zaziphuma kwiintsapho ezahlukahlukeneyo ezingama-292; I-196 (52%) yayingamadoda. Uluhlu lobudala lususela kwi-3.5 ukuya kwi-33 yeminyaka. Zonke izifundo zineskeni esinye ubuncinane (iminyaka yobudala ekuskena ngayo okokuqala, iminyaka eyi-12.3; SD, 5.3); I-203 (54.1%) inezikena ezimbini ubuncinane (iminyaka yobudala, i-13.8; SD, 4.6); I-106 (28.3%) yayinezikena ezintathu ubuncinane (zithetha, 15.3; SD, 4.2); kunye ne-57 (15.2%) ine-scan ezine okanye ngaphezulu (ithetha i-18, i-SD 4.5).

Umzobo 1. 

Ukuhanjiswa kweminyaka yobudala bedatha. Ubudala kwiskena ngasinye buboniswa ngedayimani eluhlaza. Kwisifundo ngasinye, iskena sokuqala sisoloko sisesona sisekhohlo; izifundo ezineskeni eziphindaphindiweyo zinomgca othe tye ozotyiweyo odibanisa iminyaka ekuqaleni kokuskena kunye nobudala kwizikena zamva.

I-Neuroimaging.

Imifanekiso ene-T1 ene-contiguous 1.5 mm axial slices kunye ne-2.0 mm ye-coronal slices yafunyanwa kusetyenziswa i-gradient eyonakeleyo ye-three-dimensional ikhunjulwe i-echo kwisimo sokuzinza kwi-1.5-T General Electric (Milwaukee, WI) I-Signa scanner. Iiparamitha zomfanekiso bezimi ngolu hlobo lulandelayo: ixesha le-echo, 5 ms; ixesha lokuphinda, 24 ms; flip angle, 45°; matrix yokufumana, 256 × 192; inani le-excitations, i-1; kunye nebala lokujonga, i-24 cm. Ukubekwa kwentloko kuye kwalungiswa njengoko bekuchaziwe ngaphambili (UGiedd et al., 1999). Kwasetyenziswa iskena sinye kulo lonke uphononongo. Izikena ze-MRI zomthonyama zabhaliswa kwindawo esemgangathweni ye-stereotaxic kusetyenziswa uguqulo lomda kunye nokulungiswa kwezinto zakudala ezingekhoyo (Sled et al., 1998). Imiqulu ebhalisiweyo neyalungiswayo yahlulahlulwe yaba yinto emhlophe, ingwevu, iCSF, kunye nemvelaphi kusetyenziswa i-neural net classifier (Zijdenbos et al., 2002). I-algorithm yokuguqulwa komphezulu isetyenziswe kuqala ehambelana nomphezulu wemiba emhlophe, emva koko yandisa ngaphandle ukuze ifumane i-grey matter-CSF intersection ichaza ubudlelwane obaziwayo phakathi kwe-vertex nganye ye-white matter surface kunye ne-gray matter surface counterpart; Ubukhulu becortical buchazwa njengomgama phakathi kwezi vertices eziqhagamshelweyo (kwaye zilinganiswe ngama-40,962 ezo nkqo) (IMacDonald et al., 2000). I-kernel ye-30-mm-bandwidth blurring kernel yasetyenziswa; obu bukhulu bukhethwe ngokwesiseko sokulinganiswa kwabantu apho olu luhlu lwebhanwidth lwandise amandla eenkcukacha-manani ngelixa kuncitshiswa izinto ezibubuxoki (ULerch noEvans, ngo-2005). Le kernel ivumela ukwenziwa kwendawo ye-anatomical, njengoko i-30 mm isenza luzizi kumphezulu usebenzisa i-diffusion smoothing operator igcina iimpawu ze-cortical topologic kwaye imele i-cortex encinci kakhulu kune-volumetric elinganayo ye-Gaussian blurring kernel.

Ukunyaniseka kwesi singqinisiso esizenzekelayo ngokuchasene noqikelelo lwe-neuroanatomical lwe-cortical ubukhulu be-cortical luye lwaboniswa ngaphambili kwimimandla ye-cortical ekhethiweyo kubantu abadala (Kabani et al., 2001) Siphindaphinda olu phononongo lokuqinisekisa kuluntu lwethu lwabantwana kwimimandla ye-cortical ebandakanyiweyo kwisifundo sokuqala (i-gyri yangaphambili kunye ne-post central, i-gyrus yangaphambili ephezulu, i-gyrus ephezulu yesikhashana, i-cuneus, i-lobule ephezulu ye-parietal kunye ne-supramarginal gyrus) (Kabani et al., 2001). Sikwavavanye imimandla enomdla ngokukodwa kolu phononongo. Le yayiyi-insula, i-orbitofrontal cortex (elinganiswe ngokulinganayo kwi-anterior, posterior, medial, and lateral divisions), kunye nemimandla ye-cortical ephakathi (i-anterior ne-posterior cingulate, i-medial dorsal prefrontal cortex, kunye ne-parahippocampal gyrus). Ukuskena okungamashumi amabini kukhethwe ngokungakhethiyo kwiqela (ukususela kwiminyaka eyi-6 ukuya kwe-15). Kummandla ngamnye wobuchopho i-neuroanatomist (NK) esetyenzisiweyo yokuhlaziya umfanekiso wesoftware (MacDonald, ngo-1996) ukuphawula inqaku elinye okanye ithegi kwi-CSF kunye nomda we-grey emele umphandle we-cortex, kunye nenye indawo yomda we-grey ne-white matter emele indawo yangaphakathi ye-cortex. Umgama phakathi kweethegi ezimbini ubalwa, ukulinganisa i-algorithm esetyenziswe sisixhobo esizenzekelayo. Kwithegi enikiweyo ebekwe yi-neuroanatomist kumphezulu we-cortical yangaphandle, i-vertex ekufutshane kumphezulu we-cortical ekhutshwe ngokuzenzekelayo ichongiwe kwaye ubukhulu bayo be-cortical ehambelana nayo baphawulwa. Imveliso yeendlela kunye neendlela ezizenzekelayo zifaniswa kusetyenziswa imilinganiselo ephindaphindiweyo ye-ANOVA elandelwa ngamabini t iimvavanyo zokuchonga iiyantlukwano zengingqi. Kwakukho umahluko obalulekileyo wohlobo lomlinganiselo kunye noqikelelo oluzenzekelayo lube lukhulu (lithetha, 4.62; SE, 0.06) kunencwadana (ithetha, 4.41; SE, 0.04; F(1,684) = 8.8, p = 0.02). Kwakukho intsebenziswano ebalulekileyo yohlobo lomlinganiselo kunye nommandla (F(35,684) = 2.59, p <0.001) eyaphononongwa ngakumbi. Ngokubanzi, akukho mahluko ubalulekileyo phakathi kweendlela zemanyuwali kunye ne-automated kwi-30 yemimandla ye-36, kunye nokusebenza kakubi okuphawulwe ngokudibeneyo kwi-precentral gyrus, nakwi-gyrus yangasemva esekhohlo, nakwi-gyrus ephakathi engaphambili, i-gyrus rectus kunye ne-cuneus ekhohlo ihemisphere. Ngokucacileyo, inye kuphela kule mimandla ibekwe kwindawo yomdla othile kolu phononongo (i-gyrus rectus yasekhohlo). Kwakungekho lunxulumano phakathi kweminyaka yobudala kunye nomahluko phakathi koqikelelo oluzenzekelayo kunye nolwezandla (r = 0.02, p = 0.53). Ke, akukho bungqina bokuba umahluko phakathi kweemethrikhi zombini unalo naluphi na ucalucalulo olunxulumene nobudala.

Ukumisela i-trajectories yophuhliso kwindawo nganye ye-cortical, uhlalutyo oluxubileyo lwemodeli edibeneyo lukhethwa njengoko luvumela ukufakwa kwemilinganiselo emininzi ngomntu ngamnye, idatha engekhoyo, kunye nezithuba ezingaqhelekanga phakathi kwemilinganiselo, ngaloo ndlela ukwandisa amandla ezibalo (I-Pinheiro kunye neBates, ngo-2000). Ulwahlulo lwethu lweendlela zophuhliso lusekelwe kwinkqubo yokunyula imodeli yokuhla: kwindawo nganye ye-cortical silinganisa ubukhulu be-cortical usebenzisa i-mixed-effects polynomial regression model, ukuvavanywa kwe-cubic, i-quadratic kunye ne-linear age effects. Ukuba isiphumo cubic yobudala yayingabalulekanga kwi p <0.05, yasuswa kwaye sehla kwimodeli ye-quadratic njalo njalo. Ngale ndlela, siye sakwazi ukuhlela uphuhliso lwendawo nganye yecortical njengokucaciswa ngcono yi-cubic, quadratic, okanye umsebenzi womgca wobudala. Sicinga ukuba iimodeli ze-cubic zinzima ngakumbi kune-quadratic, ezithi nazo zibanjwe zinobunzima kuneemodeli zelinear. Isiphumo esingacwangciswanga somntu ngamnye sakhiwe ngaphakathi kwesiphumo esingenamkhethe kusapho ngalunye, ngaloo ndlela kubalelwa kokubini phakathi komntu kunye nokuxhomekeka kusapho. Ngaloo ndlela, kumanqaku e-cortical kunye nemodeli ye-cubic, i kUbunzima becortical ye iumntu ngamnye kwi jusapho lwayo lwaluyimodeli yobunzimaijk = nqamla + dij + b1(ubudala) + β2*(ubudala)**2 + β3*(ubudala)**3 + eijk, apho dij zifakwe kwindlwane yeziphumo ezingacwangciswanga phakathi komntu kunye nangaphakathi kokuxhomekeka kusapho, i-intercept kunye ne-β imigqaliselo ziziphumo ezisisigxina, kwaye eijk imele impazamo eshiyekileyo. Iimodeli ze-quadratic zazingenalo ixesha le-cubic yobudala, kunye neemodeli ezihambelana neminyaka ye-cubic kunye ne-quadratic yobudala. Uhlalutyo lwaluphinda lungena kwi-SES kunye ne-IQ njengama-covariates.

Ubudala obufunyanwe ngayo ubukhulu becortical ubukhulu bubalelwe iimodeli ze-cubic kunye ne-quadratic ukusuka kwi-odolo yokuqala yokuphuma kweegophe ezifakelweyo.

iziphumo

Kuyo yonke i-isocortex esecaleni esecaleni, esecaleni, ekwi-parietal kunye ne-occipital, iindlela zophuhliso ziyi-cubic, kunye nexesha lokukhula komntwana, kulandelwa kukwehla kolutsha kunye nokuzinzisa kokuqina kwecortical xa umntu emdala.Ikhiwane. 2). Ukukhula okuphawulwa ngokunyuka kunye nokuncipha, kodwa ukunqongophala kwesigaba sokuzinzisa kumashumi amathathu eminyaka okuqala obomi (imodeli ye-quadratic) ikhona kuninzi lwe-insula kunye ne-anterior cingulate cortex. I-trajectory yomgca ibonakala kwi-operculum yangasemva kunye ne-frontal operculum, iinxalenye ze-piriform cortex, i-cortex ye-medial temporal, i-subgenual cingulate area, kunye ne-medial occipitotemporal cortex. Iigrafu ezibonisa amanqaku edatha azimeleyo avela kwimimandla emeleyo enecubic, quadratic okanye linear trajectory ibonisiwe Umzobo 3.

Umzobo 2. 

Ubunzima beendlela zophuhliso kuyo yonke i-cortex ye-cerebral. Iimephu zobuchopho zibonisa i-vertices ene cubic (red), quadratic (green) okanye linear (blue) yophuhliso trajectory. Iigrafu zibonisa ipateni yokukhula yecandelo ngalinye kula. Ngokulandelelana kukho i-dorsal, icala lasekunene, i-medial ekhohlo, i-left lateral, kunye ne-right medial iimbono. I-corpus callosum yenziwe mnyama.

Umzobo 3. 

Iigrafu ezibonisa idatha yobukhulu becortical ekrwada ngombala ozuba, kunye nomzila ofakelweyo obekwe ngaphezulu ngepinki. a-c, Imifanekiso emithathu yokuqala ibonisa ngokulandelelana ubukhulu be-cortical kunye ne-trajectory yemimandla emele: i-gyri yangaphambili ephezulu, ene-cubic trajectory (a); inxalenye ye-insula ene-quadratic trajectory, ebonwa kuluhlaza kwi Umzobo 5 (b); isahlulo se-orbitofrontal cortex ene-linear trajectory, ebonwa ngeblue in Umzobo 4 (c).

Sivavanye ukuntsokotha kweendlela zophuhliso malunga nemimandla yecortical yeentlobo ezahlukeneyo ze-cytoarchitecturonic, sisebenzisa iiatlasi ze-histological ukwabela amasimi e-cytoarchitectonic (I-Ongur et al., 2003). Olu hlalutyo lubonakalise ukuhambelana okucacileyo phakathi kweentlobo ezisisiseko ze-cortex kunye nepateni yophuhliso lwe-cortical. I-cortex ye-orbitofrontal ngumzekelo wembalelwano phakathi kweentlobo ze-cortical kunye neendlela zophuhliso (Ikhiwane. 4). Kweyona nxalenye yangaphambili yalo mmandla, i-cubic trajectory ibonakalisa i-homotypical (i-4-layered) isocortex yepali yangaphambili kunye nemimandla ye-orbitofrontal esecaleni. Ngokwahlukileyo, uninzi lwecortex kumphezulu we-orbital ongasemva ilandela iindlela ezilula zokukhula ezine-quadratic kunye nemigca. Lo mmandla unepateni ye-lamination ye-transitional cortex: xa kuthelekiswa ne-homotypical isocortex ineeleya ezimbalwa, ezingaphuhliswanga kakuhle kwaye ayinayo ingxinano ecacileyo yeeseli ze-nonpyramidal zomaleko wesi-XNUMX, umaleko wegranular wangaphakathi.Brockhaus, ngo-1940; IMesulam kunye neMufson, ngo-1982; I-Ongur et al., 2003). Kwindawo engasemva kwalo mmandla, ukukhula komgca kunye ne-quadratic kubonisa i-piriform cortex, indawo ye-allocortical yamandulo egcina i-olfaction.

Umzobo 4. 

A, Ukuntsokotha kweendlela zophuhliso kuyo yonke i-orbitofrontal cortex, eqikelelweyo kwitemplate yengqondo eqhelekileyo. I-cortex yangaphambili kunye ne-lateral orbitofrontal cortex ine-cubic fit (obomvu); i-medial and posterior orbitofrontal cortex ine-quadratic elula (green) kunye ne-linear (blue) trajectories. B, I-trajectories ibekwe phezulu kwimephu ye-cytoarchitectonic yommandla ngo Öngür et al. (2003) ukubonakalisa ukugqithelana phakathi kwemimandla yecytoarchitectonic kunye nokwahlukana kwengingqi kwiitrajectories. C, Umkhondo wecandelo ngalinye.

Nangona Umzobo 4 igxininise kwi-cortex ye-orbitofrontal, imigaqo efanayo ibonwa ngokubanzi, apho ukuguqulwa kwe-isocortex ukuya kwiifom ezilula kwenzeka. Iziphumo ze-medial frontal cortex ziyafana nezo zikwi-orbitofrontal cortex, ngokukhula kwe-cubic ngaphambili, ngakumbi kwi-homotypical cortex ye-medial frontal pole- kunye ne-linear or quadratic trajectories ngasemva ngakumbi kwimimandla ye-dysgranular okanye i-architecture ye-agranular (Ikhiwane. 5, phezulu). Kwi-insula (Ikhiwane. 5, ezantsi), ipateni iyafana kakhulu. I-insula yangaphambili, kunye ne-agranular kunye ne-cortex elayitiweyo engekho mgangathweni, inomgca wophuhliso oluhamba phambili. Ukuhamba ngasemva kwi-insula ye-dysgranular kunye ne-homotypical, kukho okokuqala i-quadratic fit enzima ngakumbi; ngakumbi ngasemva, njengoko i-cortex isiya ikhula i-homotypical, i-trajectory iba cubic. Ngokufanayo kwi-lobe yesikhashana, inxalenye ye-allocortical efana ne-piriform cortex ibonisa ngokugqithiseleyo i-trajectory yomgca. Ngokwahlukileyo, i-isocortex ye-lateral temporal ine-cubic trajectory kunye neendawo zotshintsho ezifana nemimandla ye-entorhinal kunye ne-perirhinal ine-quadratic kunye ne-linear trajectories (Ikhiwane. 2). Ezi ziphumo zishwankathelwa kwi 1 Table. Kukho imimandla ethile ye-cortical apho olu nxulumaniso phakathi kweentlobo ze-cortical kunye ne-trajectories yophuhliso alubambeli, ngokukodwa kwi-occipitotemporal ye-medial kunye neendawo zangaphambili eziphezulu ze-temporal, zombini ziyimimandla ye-isocortical nangona kunjalo inomgca we-linear kunye ne-quadratic trajectory, ngokulandelanayo. Ipateni yeziphumo ezibanjwe xa i-SES kunye ne-IQ zingeniswa njenge-covariates, ngokwahlukileyo okanye kunye.

Umzobo 5. 

Phezulu, Iimbono ezineenkcukacha zetrajectories kwi-cortex yangaphambili ye-medial yangaphambili, apho imimandla ye-isocortical ine-cubic trajectory, kunye neendawo eziguqukayo zine-quadratic trajectory (umzekelo, i-agranular kunye ne-lamine cortex engekho kakuhle yendawo ye-24a kwi-cingulate gyrus) okanye ukwehla komgca. kubukhulu (umzekelo, i-cortex encinci kunye ne-agranular ye-gyrus rectus). Ezantsi, I-insula yasekunene ibonisa ngokuqhubekayo iindlela ezintsonkothileyo ezihambayo: isahlulo esingasemva sinecubic trajectory (ebomvu), umzimba we-insula unokulingana kwe-quadratic (luhlaza) kunye ne-insula yangaphambili inomda womgca (oluhlaza okwesibhakabhaka). Ipateni efanayo ibambe i-insula yasekhohlo.

Ithebula 1. 

Iiodolo ezahlukeneyo zetrajectories zinikwa kunye nemimandla yecortical ehambelanayo kunye nohlobo olusezantsi lwecortical.

Siye sagqiba ngokulandelayo iminyaka apho ubungqingqwa becortical bencochoyi bufunyenwe kuwo onke amanqaku ane cubic okanye iquadratic trajectory, kusetyenziswa ulandelelwano lokuqala oluphuma kwigophe elifakelweyo kwindawo nganye. Ubudala bencopho ayinakumiselwa amanqaku anendlela yomgca. Iziphumo zinikezelwe njengolandelelwano oluguquguqukayo oluphelelwa lixesha (imiboniso bhanyabhanya eyongezelelweyo 1, 2, ekhoyo www.jneurosci.org njengemathiriyeli eyongezelelweyo), "isele" ithathwe kwiimuvi (Ikhiwane. 6), kunye nobudala obuqikelelwayo bobunzima be-cortical ye-peak ye-56 yemimandla yobuchopho (njengoko kuchazwe ngesixhobo sokwahlula i-ANIMAL).

Umzobo 6. 

Ubudala bokufikelela incopho yobukhulu becortical kwi-cerebral cortex. Ubukhulu bencopho bunokuqikelelwa kuphela imimandla ene-cubic okanye i-quadratic trajectory kwaye ingeyiyo imimandla enotshintsho lomda (oboniswe ngomthunzi omnyama obomvu). Utshintsho lubonakaliswe ngamandla kwiiMovie ezongezelelweyo 1 kunye ne-2, ekhoyo www.jneurosci.org njengemathiriyeli eyongezelelweyo.

Ukushwankathela iziphumo, ngaphakathi kwe-isocortex, i-sensory yokuqala kunye neendawo zeemoto zihlala zifikelela incopho yazo yobukhulu be-cortical phambi kweendawo eziziisekondari ezikufutshane, naphambi kwezinye iindawo zombutho we-polymodal. Kwingqondo yangasemva, indawo yokuqala yokufikelela kwincopho yobukhulu bayo yi-somatic sensory cortex (∼7 iminyaka), ilandelwa zizibonda ze-occipital, equlethe uninzi lwendawo yokubona ephambili (∼7 iminyaka ngasekhohlo, kunye ne-8 iminyaka. ngakwesokudla) kwaye emva koko i-cortex ye-parieto-occipital eseleyo, kunye nemimandla ye-polymodal (efana ne-posterior parietal cortex) ifikelela kwi-peak thickness later (∼9-10 iminyaka). Kwi-cortex engaphambili, i-primary motor cortex ifikelela incopho ye-cortical thickness kuselithuba (∼9 iminyaka), ilandelwa ziindawo zemoto ezongezelelweyo (~iminyaka eli-10) kunye nobuninzi bepali engaphambili (∼10 iminyaka). Iindawo eziphakamileyo ze-cortical, ezifana ne-dorsolateral prefrontal cortex kunye ne-cingulate cortex, zifikelela kwi-peak thickness last (~10.5 iminyaka). Kwiimbono zangaphakathi, iipali ze-occipital kunye nezangaphambili zifikelela kwincopho yobukhulu kwangoko, kwaye emva koko i-centripetal wave itshayelwa ukusuka kwezi ndawo, kunye ne-medial prefrontal kunye ne-cingulate cortex efikelela kwincopho yobukhulu bokugqibela. Kukwakho ne-dorsal ephawulweyo kwinkqubela ye-ventral yophuhliso. Iziphumo ezineenkcukacha zommandla ngamnye wobuchopho zinikiwe 2 Table.

Ithebula 2. 

Ubudala obuqikelelwayo bencopho yobukhulu becortical bunikwa imimandla yobuchopho engama-56

ingxoxo

Ukulungelelaniswa kweentlobo ze-cortical kunye neendlela zophuhliso

Olu phononongo lubonisa ulungelelwaniso oluqinileyo phakathi kwendlela yophuhliso kunye neentlobo zecortical eziboniswe kwiimephu zesiqhelo zecytoarchitectonic, ukunika ezi mephu zamandulo ukubaluleka kophuhliso. Uphononongo luxhasa kwaye lwandisa umsebenzi wangaphambili (UGogtay et al., 2004; Sowell et al., 2004; O'Donnell et al., 2005) ngokubandakanywa kobungakanani besampulu enkulu kakhulu evumele ukubonwa kweziphumo zomgangatho ophezulu wobudala.

Olunye uphononongo lwexesha elide lophuhliso oluqhelekileyo luxhasa ezinye zeziphumo ezikhoyo. Ngokomzekelo, ukukhula okulula komgca esikuxelayo kwindawo enye ye-allocortex, indawo ye-piriform, nayo ifunyenwe ngaphambili kwi-hippocampus (UGogtay et al., 2006). Asikwazanga ukulinganisa i-hippocampus ngokuthe ngqo kwisifundo esikhoyo, kodwa uGogtay kunye noogxa bafumanisa ukuba umkhondo wokutshintsha umthamo we-allocortical hippocampus wawumgca. Ukugqwesa kwe-isocortical thinning ebusheni kuqinisekiswa kwizifundo kusetyenziswa ezinye iindlela ze-cortical morphometric ezifana nobuninzi begrey, ebonisa ukuhambelana kwezi milinganiselo zahlukeneyo zeempawu ze-cortical (UGogtay et al., 2004; Sowell et al., 2004).

Imodeli esetyenzisiweyo apha isebenza kuphela kuluhlu lweminyaka yobudala kwaye ayinakuchazwa. Umzekelo, ukuba i-cubic trajectory yandiswe ngaphaya koluhlu lobudala, oko kuya kuthetha ukuba kukho ukonyuka kobunzima becortical xa umntu emdala (ukuqala malunga nobudala obungama-∼25), nto leyo engenakuthenjwa ngokwebhayoloji okanye exhaswa yidatha ekhoyo kolu luhlu lobudala. (Sowell et al., 2007). Endaweni yoko, iminyaka apho inqanaba le-cortical thinning liyekayo (inqaku lesibini lokuguquguquka kwi-cubic curve) liqondwa ngcono njengamanqaku okutshintshela kwimilinganiselo yecortical ezinzileyo yobudala. Iindawo ezine-cubic trajectories zifikelela kweli nqanaba lokuguquguquka ngokukhawuleza kunezo zineegophe ezizii-quadratic, kwaye ngale ndlela zinokucingelwa njengokukhula okukhawulezayo.

Imiba yendlela

Kubalulekile ukuqwalasela ukuba nokwenzeka kwe-artifact yendlela yokwenza igalelo kwiziphumo. Oku kunokuvela, umzekelo, njengoko ukwakhiwa ngokutsha komphezulu wecortical, okusisiseko sobuchule obuzenzekelayo, kunokuba nzima ngakumbi kwimimandla yeallocortex kunye necortex yenguqu, kwaye ukwanda okubangelwa yimpazamo yokulinganisa kunokufihla iipateni zokukhula ezintsonkothileyo (cubic). . Ziliqela izinto ezenza oku kungenzeki. Ubunyani bomlinganiselo wecortical ubukhulu obugwetywe ngokuchasene noqikelelo olwenziwa ngesandla aluzange luhluke ngokucwangcisiweyo ngohlobo lwecortex. Uqikelelo lobungakanani becortical oluzenzekelayo kwimimandla ye-allocortical ye-orbitofrontal kunye ne-medial cortex yayisemthethweni njengemilinganiselo yemimandla ye-isocortical. Ukongeza, i-algorithm esiyisebenzisileyo kunye nezinto eziphuma kuyo inokukhupha ngokuchanekileyo indawo ye-cortical yengqondo "yephantom", ibone ukucutha okufanisiweyo kwe-cortex kwaye ibambe iipateni ze-neuropathologically zokuqhubeka kwesifo.IMacDonald et al., 2000; ULerch noEvans, ngo-2005; Lerch et al., 2005; ULee et al., 2006). Okokugqibela, uphononongo lwethu luzuza kubungakanani besampulu enkulu kunye nenani eliphezulu ledatha elindelekileyo, izinto ezibonelele ukubonwa kweziphumo zelinear kunye ne-curvilinear yobudala ekukhuleni kwecortical eziye zombini ngokwezibalo kwaye, sixoxa ngokwebhayoloji, ibalulekile.

Iziphumo zokusingqongileyo kunye nemfuzo kwiindlela zokukhula

Ubume beziganeko zeselula ezixhasa utshintsho lwe-cortical ebantwini aluzange lusekwe. Eminye imiba yokuqala yophuhliso lwecortical, njengokuvela kunye nesisombululo se-subplate njengoko i-neuroblasts ifuduka isuka kwi-neuroepithelium iye kwiindawo zazo ezivuthiweyo ze-laminar.Kostovic kunye noRakic, ngo-1990; Kostovic et al., 2002) misela ukutyunjwa kwe-cerebral utero kunye ne-perinatal, kodwa iwela ngaphandle kwefestile yeminyaka esiyifundileyo. Izifundo kwizilwanyana ezingengomntu zibonisa ukuba imilinganiselo ye-cortical ngexesha elibalulekileyo lokuphuhliswa kwemisebenzi yokuqonda inokubonisa ukubunjwa okuxhomekeke kumava kwi-architecture ye-cortical columns kunye ne-dendritic spine kunye ne-axonal remodeling (Chklovskii et al., 2004; Mataga et al., 2004; IHensch, 2005; Sur and Rubenstein, 2005). Ezo ziganeko ze-morphological zinokuba negalelo kwisigaba sobuntwana sokunyuka kobunzima becortical, eyenzeka kwimimandla ene-cubic okanye i-quadratic trajectory. Inqanaba le-cortical thinning elawula ukufikisa kunokubonisa ukupheliswa okuxhomekeke kusetyenziso lwee-synapses (IHuttenlocher kunye ne-Dabholkar, 1997) ezinokucokisa iisekethe ze-neural, kubandakanywa nezo zixhasa izakhono zokuqonda (IHensch, 2004; UKnutudsen, 2004). Iziganeko ezenzeka kumdibaniso phakathi kwezinto ezimhlophe kunye negrey, ezifana nokunyuka kwe-myelin kwi-peripheral cortical neuropil ebuntwaneni nasekufikiseni, kunokuchaphazela ubukhulu be-cortical.Yakovlev kunye neLecours, ngo-1967; Sowell et al., 2004).

Le ngxelo yeziganeko ezinokwenzeka zeselula igxininisa indima yamava njengesithinteli esinye se-cortical architecture. Uvavanyo lwethu lwemiba yokusingqongileyo lwalukhawulelwe kwimeko yentlalo noqoqosho yomntwana kwaye ukungena kulo mlinganiselo njenge-covariate akuzange kutshintshe iphethini yeziphumo. Nangona kunjalo kuya kuba nomdla ukuvavanya impembelelo yezinye izinto eziphambili, ngakumbi usapho kunye nemeko yesikolo kuphuhliso lwecortical. Umahluko womntu ngamnye kubukrelekrele uchaphazela ubukhulu be-cortical kunye nophuhliso lwayo (Narr et al., 2006; UShaw et al., 2006b). Nangona kunjalo, iziphumo zethu eziphambili ezibanjelwe xa i-IQ yayingeniswa njenge-covariate, ethetha ukuba nangona ubukrelekrele bunokuchaphazela ezinye iipropathi zeendlela zokukhula kwe-cortical, njengesantya kunye neminyaka yokufikelela kwincopho yobukhulu be-cortical, ayinampembelelo kwikhonkco elisisiseko phakathi kobunzima bobunzima. i-cytoarchitecture kunye nobunzima bendlela yophuhliso.

Imiba yemfuza nayo ibalulekile ekumiseleni i-cortical architecture (UThompson et al., 2001; I-Lenroot et al., 2007). Iipolymorphisms eziqhelekileyo ezifana ne icatechol-O-methyltransferase Val158Met i-polymorphism, i-polymorphism eyodwa ye-nucleotide kwi-regulator ye-G-protein ebonisa i-4 gene, kunye nommandla okhuthazayo we-polymorphism ye-serotonin transporter gene (5-HTTLPRzonke zifunyenwe zinefuthe elithile kumthamo wecortical, ubukhulu okanye ubunzima (UBrown noHariri, ngo-2006; Meyer-Lindenberg et al., 2006; Zinkstok et al., 2006; IBuckholtz et al., 2007; Taylor et al., 2007). Eyona nto inomdla kakhulu ziimfuza ezinegalelo ekukhuleni kwe-cortical kunye nobunzima kwaye zibonakala ziphantsi kokhetho olufanelekileyo kwi-primate evolution, ngakumbi kwimigca ekhokelela kubantu banamhlanje (UGilbert et al., ngo-2005). Ezi ziquka i ASPM (engaqhelekanga spindle-like, microcephaly ezinxulumene) kunye MCPH1 (microcephaly, primary autosomal recessive) genes (Evans et al., 2004a,b). Kuya kuba ngumdla ukufumanisa ukuba ngaba ukwahluka kwimbonakalo yecortical yengingqi yofuzo olunjalo luhambelana nazo zombini iintlobo zecortical kunye neemephu zendlela yophuhliso.

Iingqwalasela ezisebenzayo

Ukuqwalaselwa nzulu kwezi patheni zophuhliso kunye nobudlelwane babo obunokwenzeka kuphuhliso lwengqondo luhlala lusemsebenzi wexesha elizayo, kodwa amanqaku ambalwa angenziwa. Ngokomzekelo, i-posterior medial orbitofrontal areas ziye zadibaniswa kunye ne-limbic system kunye nokulawulwa kwenkqubo ye-nervous autonomic, kwaye ibonisa umgca womgca. Ezi ndawo zicingelwa ukuba zibeke iliso kwiziphumo ezinxulumene nokuziphatha, ngakumbi isohlwayo okanye umvuzo (Imiqulu, i2004; Kennerley et al., 2006), imisebenzi yokuqonda ibaluleke kakhulu kangangokuba akunakwenzeka ukuba ibe nophuhliso olude. Ngokwahlukileyo, imimandla ye-isocortical ihlala ixhasa imisebenzi yengqondo enzima ngakumbi, ebonisa i-gradients ezicacileyo zophuhliso, ezibonakaliswa ngophuhliso olukhawulezayo ngexesha elinzima. Singaqikelela kuphela malunga nobudlelwane obunokwenzeka phakathi kwamaxesha abalulekileyo ophuhliso lwezakhono zoluntu kunye neendlela zophuhliso ezichazwe apha. Ukuchazwa kwamaxesha abalulekileyo kuphuhliso lwezakhono zoluntu kuntsonkothile, kodwa ubuntwana kade lixesha lophuhliso olukhawulezayo lwezakhono zokulawula, inkumbulo yokusebenza kunye nokuguquguquka kwengqondo, ixesha lobudala elihambelana nokunyuka kobunzima becortical kwi-lateral prefrontal cortex. (UChelune noBaer, ​​ngo-1986; Idayimani, i-2002; Huizinga et al., 2006; UJacob et al., 2007). Ngokwahlukileyo, ixesha elibalulekileyo lemisebenzi ethile ebonakalayo (efana ne-letter acuity kunye nokufunyanwa kwentshukumo yehlabathi) kuqikelelwe njengokuphela komntwana ophakathi (∼ ubudala 6 okanye 7) (Lewis kunye noMaurer, ngo-2005) kwaye, ngokunjalo, ixesha lokunyuka kwe-cortical thickness kwi-cortex ebonakalayo iphinda iphele malunga neli xesha (malunga ne-7-8 yobudala). Olu nxulumano phakathi kobude bamaxesha athile abalulekileyo kunye nesigaba sokunyuka kobungqingqwa becortical ngokuqinisekileyo asiyondawo yonke. Inqunyelwe ngokuyimfuneko kubukho beenkqubo (ezixhaswe yimimandla ye-cortical efanayo) kunye namaxesha amaninzi abalulekileyo, ngalinye linefestile yesikhashana eyahlukileyo, njengoko kwenzeka kwiinkqubo ezithile zoluvo. (UHarrison et al., 2005; Levi, ngo-2005; Lewis kunye noMaurer, ngo-2005). Le ngxoxo igxininise kumaxesha abalulekileyo kwaye akufanele ithathwe njengokugxotha ukubaluleka kokuqhubeka nokucocwa kwezakhono ezininzi zokuqonda ngexesha lesigaba sokufikisa se-cortical thinning (I-Luna et al., 2004; Luciana et al., 2005).

isiphelo

Iziphumo ezichazwe apha zixhasa ingcamango yokuba umbutho we-cerebral cortex unokuqondwa ngokoluhlu lwamakhonkco agxininisekileyo, kunye ne-isocortex (ene-cubic trajectory) kumbindi wayo, i-allocortex (ebonisa ubukhulu becala ukukhula komgca) kwi-periphery. , kunye neendawo zotshintsho (ezinomxube we-quadratic kunye ne-linear trajectories) phakathi. I-isocortex kulo mzekelo ayikho kuphela kwi-core ye-cortex ye-cerebral kule ngqiqo, kodwa iphinda ivele kamva ekuziphendukeleni kwemvelo kunommandla we-piriform (i-allocortex ye-lateral) kunye ne-hippocampus (i-allocortex ye-medial), kunye neendawo ezincinci ezongezelelweyo ze-allocortical. Ngaloo ndlela, ngokusebenzisa kwi vivo I-neuroanatomic imaging sibonisa ukuba uphuhliso lwe-cortical izibuko zombini i-cytoarchitecture kunye nembali ye-cerebral cortex.

Imihlathi

  • Ifunyenwe ngoNovemba 30, 2007.
  • Uhlaziyo lufunyenwe ngoFebruwari 7, 2008.
  • Wamkelwe ngoFebruwari 26, 2008.

  • Lo msebenzi wawuxhaswa yiNkqubo yoPhando lwe-Intramural yeZiko lezeMpilo leSizwe. Sibulela bonke abathathi-nxaxheba kuphononongo kunye neentsapho zabo.

  • Ababhali bavakalisa ukuba akukho mfuno yemali ekhuphisanayo.

  • Imbalelwano kufuneka ibhekiswe kuPhilip Shaw, iSebe leNgqondo yoMntwana, iGumbi 3N202, iSakhiwo se-10, iZiko leZiko, iZiko leSizwe leMpilo yengqondo, iBethesda, MD 20892. [imeyile ikhuselwe]

Ucaphulo

  1. ↵
    1. Allman J,
    2. Haeem A,
    3. Watson K

    (2002) Iingcali ezimbini ze-phylogenetic kwingqondo yomntu. Neuroscientist 8: 335-346.

  2. ↵
    1. Brockhaus H

    (1940) Die cyto-und myleoarchitcktonik des crotex clastralis und des clastrum beim menschen. J Psychol Neurol 49: 249-348.

  3. ↵
    1. Brown SM,
    2. Hariri AR

    (2006) Izifundo ze-Neuroimaging ze-serotonin gene polymorphisms: ukuhlola ukusebenzisana kofuzo, ingqondo kunye nokuziphatha. Yamkela uBehav Neurosci 6: 44-52.

  4. ↵
    1. Buckholtz JW,
    2. Meyer-Lindenberg A,
    3. Honea RA,
    4. Straub RE,
    5. Pezawas L,
    6. Egan MF,
    7. Vakkalanka R,
    8. Kolachana B,
    9. Verchinski BA,
    10. Sust S,
    11. UMatay VS,
    12. Weinberger DR,
    13. Callicott JH

    (2007) Ukwahluka kwe-allelic kwi-RGS4 kuchaphazela unxibelelwano olusebenzayo kunye nolwakhiwo kwingqondo yomntu. J Neurosci 27: 1584-1593.

  5. ↵
    1. Chelune GJ,
    2. Baer RA

    (1986) Imimiselo yoPhuhliso yovavanyo lwe-Wisconsin Card Sorting. J Clin Exp Neuropsychol 8: 219-228.

  6. ↵
    1. Chklovskii DB,
    2. UMel BW,
    3. Svoboda K

    (2004) I-Cortical rewiring kunye nokugcinwa kolwazi. indalo 431: 782-788.

  7. ↵
    1. Idayimani A

    (2002) kwi-Principles of frontal lobe function, Uphuhliso oluqhelekileyo lwe-prefrontal cortex ukususela ekuzalweni ukuya ebudaleni obuncinane: imisebenzi yokuqonda, i-anatomy kunye ne-biochemistry, i-eds Stuss DT, i-Knight RT (i-Oxford UP, i-New York), i-pp 466-503.

  8. ↵
    1. Evans PD,
    2. UAnderson JR,
    3. Vallender EJ,
    4. Choi SS,
    5. Lahn BT

    (2004a) Ukwakha ngokutsha imbali ye-evolution ye-microcephalin, i-gene elawula ubungakanani bobuchopho bomntu. Hum Mol Genet 13: 1139-1145.

  9. ↵
    1. Evans PD,
    2. UAnderson JR,
    3. Vallender EJ,
    4. UGilbert SL,
    5. UMalcom CM,
    6. Dorus S,
    7. Lahn BT

    (2004b) I-adaptive evolution ye-ASPM, eyona nto ibalulekileyo yobungakanani becortical yecerebral ebantwini. Hum Mol Genet 13: 489-494.

  10. ↵
    1. UGiedd JN,
    2. Snell JW,
    3. Lange N,
    4. Rajapakse JC,
    5. Casey BJ,
    6. Kozuch PL,
    7. Vaituzis AC,
    8. Vauss YC,
    9. Hamburger SD,
    10. Kaysen D,
    11. Rapoport JL

    (1996) Ubungakanani bomfanekiso wemagnethi wokukhula kwengqondo yomntu: iminyaka eyi-4-18. Cereb Cortex 6: 551-560.

  11. ↵
    1. UGiedd JN,
    2. Blumenthal J,
    3. UJeffries HAYI,
    4. Castellanos FX,
    5. Liu H,
    6. Zijdenbos A,
    7. Paus T,
    8. Evans AC,
    9. Rapoport JL

    (1999) Ukuphuhliswa kwengqondo ngexesha lobuntwana kunye nokufikisa: isifundo se-MRI sobude. Nat Neurosci 2: 861-863.

  12. ↵
    1. UGilbert SL,
    2. Dobyns WB,
    3. Lahn BT

    (2005) Amakhonkco emfuza phakathi kophuhliso lobuchopho kunye nokuvela kwengqondo. Nat Rev Genet 6: 581-590.

  13. ↵
    1. Gogtay N,
    2. UGiedd JN,
    3. Lusk L,
    4. Hayashi KM,
    5. Greenstein D,
    6. Vaituzis AC,
    7. Nugent TF III.,
    8. Herman DH,
    9. Udidi lwe-LS,
    10. AW,
    11. Rapoport JL,
    12. Thompson PM

    (2004) Imephu eguqukayo yophuhliso lwecortical yomntu ngexesha lobuntwana ukuya ebuntwini obukhulu. IProc Natl Acad Sci USA 101: 8174-8179.

  14. ↵
    1. Gogtay N,
    2. Nugent TF III.,
    3. Herman DH,
    4. Ordonez A,
    5. Greenstein D,
    6. Hayashi KM,
    7. Udidi L,
    8. AW,
    9. UGiedd JN,
    10. Rapoport JL,
    11. Thompson PM

    (2006) Imephu eguqukayo yophuhliso lwesiqhelo lomntu lwehippocampal. Hippocampus 16: 664-672.

  15. ↵
    1. Harrison RV,
    2. Gordon KA,
    3. INtaba iRJ

    (2005) Ngaba kukho ixesha elibalulekileyo lokufakelwa kwe-cochlear kubantwana abazizithulu abazalwa nabo? Uhlalutyo lokuva kunye nentetho yokuqonda ukusebenza emva kokufakelwa. Dev Psychobiol 46: 252-261.

  16. ↵
    1. Hensch TK

    (2004) Ummiselo wexesha elibalulekileyo. Annu Rev Neurosci 27: 549-579.

  17. ↵
    1. Hensch TK

    (2005) Ixesha elibalulekileyo leplastiki kwiisekethe zecortical yendawo. Nat Rev Neurosci 6: 877-888.

  18. ↵
    1. Hollingshead AB

    (1975) Isalathiso sezinto ezine kwisimo sentlalo (Yale UP, New Haven, CT).

  19. ↵
    1. Huizinga M,
    2. Dolan CV,
    3. van der Molen MW

    (2006) Utshintsho olunxulumene nobudala kumsebenzi olawulayo: iindlela zophuhliso kunye nohlalutyo oluguquguqukayo olufihlakeleyo. I-Neuropsychologia 44: 2017-2036.

  20. ↵
    1. Huttenlocher PR,
    2. Dabholkar AS

    (1997) Umahluko wengingqi kwi-synaptogenesis kwi-cortex ye-cerebral cortex. J Comp Neurol 387: 167-178.

  21. ↵
    1. Jacobs R,
    2. UHarvey AS,
    3. UAnderson V

    (2007) Umsebenzi olawulayo olandela izilonda zelobe engaphambili: impembelelo yexesha lesilonda kwisiphumo. Cortex 43: 792-805.

  22. ↵
    1. Jolicoeur P,
    2. Pontier J,
    3. Pernin MO,
    4. Sempe M

    (1988) Igophe lobomi elingena-symptotic kubude bomntu. iibhayometrikh 44: 995-1003.

  23. ↵
    1. Kaas JH

    (1987) Umbutho we-neocortex kwizilwanyana ezanyisayo: iimpembelelo kwiithiyori zokusebenza kwengqondo. Annu Rev Psychol 38: 129-151.

  24. ↵
    1. Kabani N,
    2. Le Goualher G,
    3. MacDonald D,
    4. Evans AC

    (2001) Umlinganiselo wobunzima be-cortical usebenzisa i-algorithm ye-3-D ezenzekelayo: isifundo sokuqinisekisa. NeuroImage 13: 375-380.

  25. ↵
    1. Kennerley SW,
    2. Walton ME,
    3. Behrens TE,
    4. Buckley MJ,
    5. Rushworth MF

    (2006) Ukwenza izigqibo ezifanelekileyo kunye ne-anterior cingulate cortex. Nat Neurosci 9: 940-947.

  26. ↵
    1. Knudsen EI

    (2004) Amaxesha anovakalelo kuphuhliso lwengqondo kunye nokuziphatha. J Cogn Neurosci 16: 1412-1425.

  27. ↵
    1. Kostovic mna,
    2. URaki P

    (1990) Imbali yophuhliso yezowuni ye-subplate yesikhashana kwi-cortex ebonakalayo kunye ne-somatosensory ye-macaque monkey kunye nengqondo yomntu. J Comp Neurol 297: 441-470.

  28. ↵
    1. Kostovic mna,
    2. Judas M,
    3. Rados M,
    4. Hrabac P

    (2002) Umbutho weLaminar wecerebrum yomntwana otyhilwe ziimpawu ze-histochemical kunye ne-imaging resonance magnetic. Cereb Cortex 12: 536-544.

  29. ↵
    1. Lee JK,
    2. Lee JM,
    3. UKim JS,
    4. UKim I,
    5. Evans AC,
    6. UKim SI

    (2006) Inoveli yobuninzi bokuqinisekiswa kwe-cortical surface reconstruction algorithms usebenzisa i-MRI phantom. NeuroImage 31: 572-584.

  30. ↵
    1. Lenroot RK,
    2. Schmitt JE,
    3. Ordaz SJ,
    4. Wallace GL,
    5. UNeale MC,
    6. Lerch JP,
    7. Kendler KS,
    8. Evans AC,
    9. Giedd JN

    (2007) Umahluko kwiimpembelelo zofuzo kunye nokusingqongileyo kwi-cortex ye-cerebral yomntu ehambelana nophuhliso ngexesha lobuntwana kunye nokufikisa. Hum Brain Mapp, kushicilelo.

  31. ↵
    1. Lerch JP,
    2. Evans AC

    (2005) Uhlalutyo lobungqimba lweCortical luhlolwe ngohlalutyo lwamandla kunye nokulinganisa abemi. NeuroImage 24: 163-173.

  32. ↵
    1. Lerch JP,
    2. UPruessner JC,
    3. Zijdenbos A,
    4. Hampel H,
    5. Teipel SJ,
    6. Evans AC

    (2005) Ukwehla okugxininise kobunzima becortical kwisifo se-Alzheimer esichongwe yi-computational neuroanatomy. Cereb Cortex 15: 995-1001.

  33. ↵
    1. Levi DM

    (2005) Ukufunda ngengqondo kubantu abadala abane-amblyopia: ukuphononongwa kwakhona kwamaxesha abalulekileyo kumbono womntu. Dev Psychobiol 46: 222-232.

  34. ↵
    1. Lewis TL,
    2. UMaurer D

    (2005) Amaxesha amaninzi anovakalelo kuphuhliso lokubonwa kwabantu: ubungqina obuvela kubantwana abangaboniyo. Dev Psychobiol 46: 163-183.

  35. ↵
    1. Lu LH,
    2. ULeonard CM,
    3. Thompson PM,
    4. Kan E,
    5. UJolley J,
    6. Wamkelekile SE,
    7. AW,
    8. Sowell ER

    (2007) Utshintsho oluqhelekileyo lophuhliso kwi-gray yangaphambili engaphantsi ihambelana nokuphuculwa kwe-phonological processing: uhlalutyo lwe-MRI longitudinal. Cereb Cortex 17: 1092-1099.

  36. ↵
    1. Luciana M,
    2. Conklin HM,
    3. Hooper CJ,
    4. Yarger RS

    (2005) Ukuphuhliswa kwememori engasebenziyo kunye neenkqubo zokulawula ulawulo kulutsha. Isixhobo somntwana 76: 697-712.

  37. ↵
    1. Luna B,
    2. Garver KE,
    3. Urban TA,
    4. Lazar NA,
    5. Sweeney JA

    (2004) Ukuvuthwa kweenkqubo zokuqonda ukusuka ebuntwaneni ukuya ebudaleni. Uphuhliso lwabantwana 75: 1357-1372.

  38. ↵
    1. UMacDonald D

    (1996) i-MNI-display (i-McConnell Brain Imaging Centre, i-Montreal Neurological Institute, eMontreal).

  39. ↵
    1. MacDonald D,
    2. Kabani N,
    3. Avis D,
    4. Evans AC

    (2000) Ukukhutshwa kwe-3-D ngokuzenzekelayo kwendawo yangaphakathi nangaphandle ye-cortex ye-cerebral evela kwi-MRI. NeuroImage 12: 340-356.

  40. ↵
    1. UMakris N,
    2. Biederman J,
    3. Valera EM,
    4. Bush G,
    5. Kaiser J,
    6. Kennedy DN,
    7. Caviness VS,
    8. uFarone SV,
    9. Seidman LJ

    (2006) I-Cortical thinning yengqwalasela kunye nothungelwano lomsebenzi olawulayo kubantu abadala abanengxaki yokungahoywa / ukuphazamiseka kwengqondo. Cereb Cortex 17: 1364-1375.

  41. ↵
    1. Mataga N,
    2. Mizuguchi Y,
    3. Hensch TK

    (2004) Ukuthenwa okuxhomekeke kumava e-dendritic spines kwi-cortex ebonakalayo yi-activator ye-plasminogen yezicubu. Neuron 44: 1031-1041.

  42. ↵
    1. Mesulam MM,
    2. Mufson EJ

    (1982) Insula yenkawu yehlabathi endala. I. I-Architectonics kwi-insulo-orbito-temporal component ye-paralimbic brain. J Comp Neurol 212: 1-22.

  43. ↵
    1. Meyer-Lindenberg A,
    2. Nichols T,
    3. UCallicott JH,
    4. Ding J,
    5. Kolachana B,
    6. Buckholtz J,
    7. UMatay VS,
    8. Egan M,
    9. Weinberger DR

    (2006) Impembelelo yenguqu enzima yofuzo kwi-COMT ekusebenzeni kwengqondo yomntu. Mol Psychiatry 11: 867-877.

  44. ↵
    1. Narr KL,
    2. Woods RP,
    3. Thompson PM,
    4. Szeszko P,
    5. URobinson D,
    6. Dimcheva T,
    7. UGurbani M,
    8. AW,
    9. Kwi-RM

    (2006) Ubudlelwane phakathi kwe-IQ kunye ne-cortical grey yengingqi ubukhulu kubantu abadala abanempilo. Cereb Cortex 17: 2163-2171.

  45. ↵
    1. O'Donnell S,
    2. Noseworthy MD,
    3. Levine B,
    4. UDennis M

    (2005) Ubukhulu beCortical yendawo ye-frontopolar ngokuqhelekileyo ekuphuhliseni abantwana kunye nabafikisayo. NeuroImage 24: 948-954.

  46. ↵
    1. UOngur D,
    2. Isikhitshane AT,
    3. Ixabiso JL

    (2003) Ulwahlulo lwe-Architectonic ye-orbital yomntu kunye ne-medial prefrontal cortex. J Comp Neurol 460: 425-449.

  47. ↵
    1. UPinheiro JC,
    2. Bates DM

    (2000) Iimodeli ezixubeneyo kwi-S kunye ne-S-PLUS (Springer, New York).

  48. ↵
    1. Puelles L

    (2001) Iingcinga ngophuhliso, ulwakhiwo kunye nokuvela kwe-mammalian kunye ne-avian telencephalic pallium. I-Philos Trans Trans R Soc Lond B Biol Sci 356: 1583-1598.

  49. ↵
    1. Imiqulu ET

    (2004) Imisebenzi ye-orbitofrontal cortex. Ukwahlula ingqondo 55: 11-29.

  50. ↵
    1. Shaw P,
    2. Lerch J,
    3. Greenstein D,
    4. Ubukhali W,
    5. Udidi L,
    6. Evans A,
    7. UGiedd J,
    8. Castellanos FX,
    9. URapoport J

    (2006a) Imephu yobude be-cortical ubukhulu kunye nesiphumo seklinikhi kubantwana kunye nolutsha olunenkathalo-intsilelo/yokuphazamiseka kokusebenza ngamandla. Arch Gen Psychiatry 63: 540-549.

  51. ↵
    1. Shaw P,
    2. Greenstein D,
    3. Lerch J,
    4. Udidi L,
    5. Lenroot R,
    6. Gogtay N,
    7. Evans A,
    8. Rapoport J,
    9. UJiedd J

    (2006b) Isakhono sobukrelekrele kunye nophuhliso lwecortical kubantwana nakwishumi elivisayo. indalo 440: 676-679.

  52. ↵
    1. Isileyi JG,
    2. Zijdenbos AP,
    3. Evans AC

    (1998) Indlela ye-nonparametric yokulungiswa ngokuzenzekelayo kwe-intensity nonuniformity kwidatha ye-MRI. IEEE Trans Med Imaging 17: 87-97.

  53. ↵
    1. Sowell ER,
    2. Thompson PM,
    3. ULeonard CM,
    4. Wamkelekile SE,
    5. Kan E,
    6. Toga AW

    (2004) Imephu yobude be-cortical ubukhulu kunye nokukhula kwengqondo kubantwana abaqhelekileyo. J Neurosci 24: 8223-8231.

  54. ↵
    1. Sowell ER,
    2. Peterson BS,
    3. Kan E,
    4. Woods RP,
    5. Yoshii J,
    6. Bansal R,
    7. Xu D,
    8. Zhu H,
    9. Thompson PM,
    10. Toga AW

    (2007) Umahluko ngokwesondo kubukhulu be-cortical imephu yabantu abasempilweni abayi-176 phakathi kwe-7 kunye ne-87 yeminyaka ubudala. Cereb Cortex 17: 1550-1560.

  55. ↵
    1. Striedter GF

    (2005) Imigaqo yokuguquka kwengqondo (Sinauer, Sunderland, MA).

  56. ↵
    1. Sur M,
    2. Rubenstein JL

    (2005) Ipateni kunye neplastiki ye-cerebral cortex. inzululwazi 310: 805-810.

  57. ↵
    1. Tanner JM,
    2. I-Whitehouse RH,
    3. Marubini E,
    4. Khangela kwakhona i-LF

    (1976) Ukukhula kokufikisa kwamakhwenkwe namantombazana kuphononongo lokukhula kweHarpenden. UAnn Hum Biol 3: 109-126.

  58. ↵
    1. Taylor WD,
    2. Zuchner S,
    3. Payne ME,
    4. I-Messer DF,
    5. UDoty TJ,
    6. MacFall JR,
    7. Beyer JL,
    8. Krishnan KRR

    (2007) I-COMT Val158Met polymorphism kunye ne-lobe morphometry yesikhashana kubantu abadala abanempilo. Psychiatry Res 155: 173-177.

  59. ↵
    1. Thompson PM,
    2. I-Cannon TD,
    3. Narr KL,
    4. van Erp T,
    5. Poutanen VP,
    6. Huttunen M,
    7. Lonnqvist J,
    8. Standertskjold-Nordenstam CG,
    9. Kaprio J,
    10. Khaledy M,
    11. I-Dail R,
    12. Zoumalan CI,
    13. Toga AW

    (2001) Iimpembelelo zemfuzo kwisakhiwo sobuchopho. I-Nature Neuroscience 4: 1253-1258.

  60. ↵
    1. von Economo C,
    2. Koskinas GN

    (1925) Die dytoarchitektonik der hirnrinde des erwachsenen menschen (Springer, Berlin).

  61. ↵
    1. Yakovlev PI,
    2. Izifundo AR

    (1967) kuphuhliso lweNgingqi yobuchopho ebomini bokuqala, Imijikelezo ye-myelinogenetic yokuvuthwa kommandla wengqondo, ed Minokowski A (Blackwell Scientific, Oxford).

  62. ↵
    1. Zijdenbos AP,
    2. Forghani R,
    3. Evans AC

    (2002) Uhlalutyo oluzenzekelayo "lombhobho" lwedatha ye-3-D ye-MRI yezilingo zeklinikhi: isicelo kwi-multiple sclerosis. IEEE Trans Med Imaging 21: 1280-1291.

  63. ↵
    1. Zilles K,
    2. Palomero-Gallagher N,
    3. Schleicher A

    (2004) I-Transmitter receptors kunye ne-anatomy esebenzayo ye-cortex ye-cerebral. J Anat 205: 417-432.

  64. ↵
    1. Zinkstok J,
    2. Schmitz N,
    3. van Amelsvoort T,
    4. de Win M,
    5. i-van den Brink W,
    6. Baas F,
    7. Linzen D

    (2006) I-COMT val158met polymorphism kunye ne-brain morphometry kubantu abadala abasempilweni. Neurosci Lett 405: 34-39.

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