Bothata ba Pulasetiki le Boits'oaro Bocha bo Tsoelang Pele (2011)

J O ka Etsa hore Adad Adccc Psychors. 2011 November; 20 (4): 265-276.

Bryan Kolb, PhD1 'me Robbin Gibb, PhD1
Mohlophisi oa Tlhokomelo: Margaret Clarke, MD le Laura Ghali, PhD
Sehlooho sena se bile teng e boletsoeng ke lihlooho tse ling ho PMC.

inahaneloang

Morero:

Ho lekola melao-motheo e akaretsang ea ntlafatso ea boko, ho khetholla melao-motheo ea tšebetso ea bokopa, le ho tšohla lintlha tse susumetsang nts'etsopele ea bokong le polasetiki.

Mokhoa:

Ho ile ha etsoa tlhahlobo ea lingoliloeng ea lingoliloeng tse ngotsoeng ka letsoho tsa Senyesemane tse mabapi le nts'etsopele ea boko le polasetiki.

Results:

Kholiso ea boko e tsoela pele ka methati e qalang ka ho qala ka neurogeneis le tsoelo-pele ea ho falla ha neural, ho fetoha ha mmele, synaptogenesis, ho faola le ho theha myelin. Ho khetholloa melao-motheo e robeli ea bongoana ea boko. Bopaki ba hore kholo ea kholo ea ts'ebetso ea ts'ebetso ea mokokotlo le ts'ebetso e susumetsoa ke liketsahalo tse fapaneng tsa tikoloho tse kang tšusumetso ea maikutlo, lithethefatsi tse matlafatsang kelello, li-hormone tsa gonadal, likamano tsa motsoali le ngoana, likamano tsa lithaka, khatello ea maikutlo ea morao-rao, mala le lijo.

Qeto:

Kholiso ea kelello e bontša ho fetang feela ho senoloa ha sebopeho sa lefutso empa ho e-na le hoo e bontša motjeko o rarahaneng oa liphatsa tsa lefutso le boiphihlelo bo etsang hore boko bo hlahe. Ho utloisisa motjeko ho fana ka temohisiso khanong e tloaelehileng le e sa tloaelehang.

Keywords: nts'etsopele ea boko, polasetiki ea khoele, ho tsosa tikoloho, epigenetics

Kholiso ea kelello e bontša ho fetang feela ho senoloa ha sebopeho sa lefutso empa ho e-na le hoo e bontša motjeko o rarahaneng oa liphatsa tsa lefutso le boiphihlelo bo etsang hore boko bo hlahe. Boko bo senotsoeng liketsahalong tse fapaneng tsa tikoloho joalo ka tšusumetso ea maikutlo, lithethefatsi, lijo, lihormone kapa khatello ea maikutlo ka hona li ka hlaha ka mekhoa e fapaneng haholo. Morero oa sengoloa sa morao-rao ke ho hlahloba tsela eo boko bo ntseng bo hola bo ka holisoang ka eona ke lintho tse ngata tse pele ho pelehi le kamora 'pelehi. Re qala ka kakaretso ea kholo ea kholo ea kholo ea boko, e lateloe ke tlhahlobo e khuts'oane ea melao-motheo ea bokaholimo ba boko 'me qetellong ho hlahlojoe hore na lintlha li susumetsa nts'etsopele ea boko le boitšoaro ba batho ba baholo joang. Hobane boholo ba seo re se tsebang ka polasetiki ea boko le boitšoaro bo mabapi le nts'etsopele bo tsoa liphuputsong tsa laboratori puisano ea rona e tla tsepamisa maikutlo ho rat empa re tla nka batho ha ho khonahala. Ntle le moo, moqoqo o tla ratoa ho fihlela polokeho ea methapo hobane boholo ba seo re se tsebang ka phetisetso ea kholo ea kholo ea kelello e thehiloe holim'a lithuto tsa tsoekere. Ha ho na lebaka le lekaalo la ho lumela, leha ho le joalo, hore likarolo tse ling tsa boko li ke ke tsa fetoloa ka tsela e ts'oanang.

Nts'etsopele ea Brain

Lilemong tse ling tsa 2000 tse fetileng rafilosofi oa Moroma Seneca o ile a etsa tlhahiso ea hore lehe le emolisitsoeng ke motho e moholo ka mokhoa o fokolang kahoo mosebetsi oa nts'etsopele ke ho hola feela. Mohopolo ona o ne o khahla haholo hoo o neng o lumeloa haholo ho fihlela 19th lekholo. E ile ea totobala qalong ea 20th Lekholong leo kholo ea kholo ea kholo e ileng ea bonts'a methati e mengata eo re ka e bonang e arotsoe ka mekhahlelo e 'meli. Ho liphoofotsoana tse ngata tsa pele li supa tatellano ea liketsahalo ka utero e ka fetoloang ke tikoloho ea bo-mme. Karolo ea bobeli eo e leng pele ho nako ea peleho ho batho, ke nako ea ha khokahano ea kelello e sa ameheng tikoloho feela empa e boetse e bua ka mefuta ea tšebetso ea boko e hlahisoang ke boiphihlelo. Habohlokoa le ho feta, leha ho le joalo, ho se ho amoheloa hore liphetoho tsa epigenetic, tse ka hlalosoang e le liphetoho liphellong tsa nts'etsopele, ho kenyelletsa molao oa polelo ea liphatsa tsa lefutso, li ipapisitse le mekhoa e meng ntle le ea DNA ka boeona (Blumberg, Freeman, & Robinson, 2010). Mohlala, polelo ea liphatsa tsa lefutso e ka fetoloa ke liphihlelo tse ikhethang, 'me sena se ka lebisa ho liphetoho tsa mokhatlo tsamaisong ea methapo.

Methati ea kholo ea bokong

Lethathamo 1 e hlakisa mehato e akaretsang ea kholo ea kholo ea kholo ho liphoofolo tsohle tse anyesang. Lisele tse reretsoeng ho hlahisa tsamaiso ea methapo li qala ho ba teng kamora libeke tse tharo kamora ho ikatisa bathong. Lisele tsena li etsa bokahare ba neural, e leng motsoala oa boko 'me hamorao bo bitsoa sebaka sa subventricular. Lisele tse reretsoeng ho etsa mokokotlo li qala ho arohana ha a le libeke tse ts'eletseng le ka libeke tsa 14 motsoako o shebahala o le motho ka kotloloho, leha o sa qale ho bopa sulci le gyri ho fihlela likhoeli tse ka bang tse supileng. Boholo ba neurogeneis bo felletse ka likhoeli tse hlano, kamora 'ngoe ea bohlokoa e le lisele tsa hippocampus, tse ntseng li tsoela pele ho theha methapo ea kutlo bophelong bohle. Ho na le lisele tse ka bang limilione tse likete tse leshome ho etsa "cortex" ea motho ea motsoako karolong e 'ngoe le e' ngoe ea methapo. Lisele tsena li thehoa ka potlako 'me ho hakanngoa hore ka tlhōrō ea eona, ho na le li-neuron tsa 250,000 tse thehiloeng motsotso. Ho totobetse hore tšebeliso efe kapa efe ea boko ka nako ena e ka ba le litlamorao tse kholo.

Tafole 1. 

Methati ea kholo ea bokong

Hang ha li-neuron li se li thehiloe, li qala ho tsamaea litseleng tse nang le likhoele tse qapiloeng ke lisele tsa radial glial, tse tlohang ho tloha sebakeng sa subventricular ho fihla bokaholimo ba "cortex"Setšoantšo sa 1). Karolo e ka tlas'a lefatše ea lefatše e bonahala e na le 'mapa oa khale oa cortex o rerang esale pele ka lisele tse thehiloeng sebakeng se itseng sa subricricular ho fallela sebakeng se itseng sa cortical. Ha lisele li falla li na le monyetla o sa lekanyetsoang oa tsoelo-pele ea sele empa ha li fihla moo li eang teng tšebelisano ea liphatsa tsa lefutso, ho feto-fetoha ha maemo le tikoloho le tikoloho ea tikoloho li ntse li li matlafaletsa ho khetholla ho mofuta o itseng oa sele. Hang ha lisele li fihla sebakeng sa tsona sa ho qetela li qala ho hola ka: (1) dendrites e ntseng e hola ho fa sebaka se ka holimo sa li-synapses tse nang le lisele tse ling; hape, (2) e holisa li-axons ho lipheo tse nepahetseng tsa ho qala sebopeho sa synapse.

Setšoantšo sa 1. 

Lisele li tloha sebakeng se sa tloaelehang se haufi le moea oa "radial glia" ho ea sebakeng sa tsona sa batho ba baholo.Kolb le Whishaw, 2009).

Ho etsoa ha li-dendrites ho qala ka nako ea tlhaho ho batho empa ho tsoela pele nako e telele kamora ho hlaha. Lits'oaetso tsa masea tse sa tsoa tsoaloa li qala ha mekhoa e ikhethileng e tsoa 'meleng oa sele mme ka lilemo tse peli tse latelang lits'ebetso tsena lia hlakisoa' me ho na le methapo, e leng sebaka seo ho thoeng ke lisosa tsa thabo e kholo. Khōlo ea dendritic e lieha, ka tatellano ea li-micrometer ka letsatsi. Li-Axons li hola ka makhetlo a 1000 ka potlako, e leng hoo e ka bang limilimithara tse le 'ngoe ka letsatsi. Sekhahla sena sa kholo ea phapang se bohlokoa hobane li-axon tse ntseng li hola ka potlako li ka ikopanya le lisele tse qalang pele likarolo tse qalang tsa sele eo li thehoa ka botlalo. Ka lebaka leo, li-axons li ka susumetsa phapang ea dendritic le ho theha mekotla ea likhoerekhoere.

Sebopeho sa Synfall ka har'a cortex ea motho e baka bothata bo boholo, ka kakaretso e fetang ea 100,000 trillion (1014). Palo ena e kholo e ne e ke ke ea khethoa ke lenaneo la liphatsa tsa lefutso, empa ho na le lintlha tse akaretsang tsa khokahano ea neural bokong tse tla rera esale pele. Bongata bo boholo ba li-synapses bo tataisoa ke libaka le mekhoa e fapaneng ea tikoloho. Joalokaha re tla bona, ho qhekelloa ha mefuta e fapaneng le e meng ea matšoao ho ka hlahisa phapang e kholo literekeng tsa likhoerekhoere.

Ka lebaka la ho hloka botsitso palong ea li-neuron tse tla fihla sebakeng sa tsona se nepahetseng le khokahanyo e nepahetseng ea likhokelo tseo e li thehang, bokong bo fetisa li-neuron le khokahanyo nakong ea nts'etsopele, ka tlhōrō ea tlhahiso ea methapo ea kutlo e pakeng tsa selemo se le seng le tse peli, ho latela tikoloho ea cortex. Joalo ka motaki ea etsang seemahale se nang le lejoe la majoe le mokelikeli ho tlosa likotoana tse sa batleheng, boko bo na le tsamaiso e ts'oanang eo ho eona lisele tse sa sebetsoeng le likhokahanyo li tlosoang ke lefu la sele le prunicun. Matšoao a hlalosang kelello bokong e ka ba a mefuta e mengata, ho kenyelletsa le mofuta o mong oa lets'oao la epigenetic, boiphihlelo bo fapaneng, li-hormone tsa gonadal esita le khatello ea maikutlo.

Kameho ea tahlehelo ena ea sele le ho pongoa ha synaptic e ka bonoa ho liphetoho boteng ba cortical ka nako e telele. Ka mantsoe a mang, "cortex" e hlile e ba mosesaane ka mokhoa o hlakileng oa tlhaho ea "caudal-rostral" e qalang ho pota lilemo tse peli ebe e tsoela pele ho fihlela bonyane lilemo tsa 20. Hoa khoneha ho tsamaisa tšilafalo ea cortical ka boitšoaro bo ntlafalitsoeng. Mohlala, liphetho tsa lithuto tsa MRI tsa liphetoho boteng ba cortical li bonts'itse hore ho eketseha ha motor motor ho tsamaisana le ho fokotseha ha letheba la cortical sebakeng sa letsoho la motor cortex e ka letsohong le letšehali (O'Hare & Sowell, 2008). Phapang e le 'ngoe ho moferefere ke molao o betere o bonoa ntlafatsong ea tse ling, empa eseng kaofela, lits'ebetso tsa puo. Kahoo, lithuto tsa MRI li bonts'itse kholo ea karolo e ka tlase e ka tlase ea cortex e kaholimo (hoo e batlang e le sebaka sa Broca) e amahanngoa le ts'ebetso ea ntlafatso ea fono (ke hore, kutloisiso ea melumo ea puo). Kamano ena e ikhethang lipakeng tsa bokima ba cortical le boitšoaro ha e khetholle mesebetsi ea puo ka kakaretso. Mohlala, nts'etsopele ea mantsoe e tsamaellana le botebo bo fokotsehileng ba cortical libakeng tse fapaneng tsa cortical (O'Hare & Sowell, 2008).

Kamano pakeng tsa botenya ba cortical le boits'oaro bo botle e kanna ea ba moelelo bakeng sa phapano ea kholo ea boits'oaro baneng. Mohlala, kholo e liehang ea nts'etsopele ea puo ho bana ba nang le bohlale bo tloaelehileng le motor dexterity (hoo e ka bang 1% ea bana) e ka ba litholoana tsa butle butle ho feta liphetoho tse tloaelehileng ho boteng ba cortical. Lebaka leo e ka bang sena ha le tsejoe.

Karolo ea ho qetela ea kholo ea bokong ke nts'etsopele ea glial ho theha myelin. Tsoalo ea li-astrocyte le oligodendrocyte li qala kamora hore boholo ba neurogeneis bo felletse mme bo tsoele pele bophelo bohle. Le ha li-axon tsa CNS li ka sebetsa pele ho myelination, ts'ebetso e tloaelehileng ea batho ba baholo e fumanoa feela ka mor'a hore myelination e phethe, e leng lilemo tsa 18 tsa lilemo tse libakeng tse kang pele ho pele, parietal ea poster, le anterior temport cortex.

Ka hona, nts'etsopele ea boko e na le liketsahalo tse ngata tse qalang ka ho qala ha methapo le ho fela ka ho theoa hoa myelin. Matla a phallo ea kelello le liphihlelo li tla fapana ka methati e nepahetseng ea kholo ea kholo. Ha rea ​​lokela ho makala, ka mohlala, hore liphihlelo le / kapa lits'oants'o nakong ea mitosis li ka ba le litlamorao tse fapaneng ho feta liketsahalo tse tšoanang nakong ea synaptogenesis kapa hamorao nakong ea ho pongoa. Liphihlelo li sebetsa ka bohlale bo fapaneng haholo maemong a fapaneng a ntlafatso.

Likarolo tse ikhethang tsa nts'etsopele ea boko

Likarolo tse peli tsa nts'etsopele ea boko li bohlokoa haholo ho utloisisa hore na liphihlelo li ka fetola mokhatlo o hlophisitsoeng oa cortical joang. Taba ea mantlha, lisele tse hokahaneng le karolo e tlase ea bophelo ke lisele tsa stem tse lulang li sebetsa bophelo bohle. Lisele tsena tsa stem li ka hlahisa lisele tsa neural kapa glial progenitor tse ka fetelang kahareng e tšoeu kapa e bohlooho, leha e se e le motho e moholo. Lisele tsena li ka lula li arotsoe libakeng tsena nako e telele empa li ka hlahisoa ho hlahisa li-neuron le / kapa glia. Karolo ea lisele tsena ha e utloisisoe hantle hajoale empa e ka ba motheo oa bonyane mofuta o le mong oa neurogeneis ea kamora ho kula, haholoholo kamora ho lemala (mohlala, Gregg, Shingo, & Weiss, 2001; Kolb et al., 2007). Ntle le moo, boko ba 'mele, ho kenyelletsa le bokong ba khale, bo ka hlahisa li-neurons ha li se li le batho ba baholo tse reretsoeng bulb ea olkidory, sebopeho sa hippocampal, mohlomong le libaka tse ling (mohlala, Eriksson et al., 1998; Gould, Tanapat, Hastings, & Shors, 1999; Kempermann & Gage, 1999). Karolo e sebetsang ea lisele tsena e ntse e le phehisano empa karolo ea bona e ka susumetsoa ke lintlha tse ngata ho kenyelletsa boiphihlelo, lithethefatsi, lihormone le ho lemala.

Karolo ea bobeli e ikhethang ke hore li-dendrites le li-spine li bonts'a polasetiki e tsotehang ho arabela boiphihlelo mme e ka theha li-synapses ka lihora le mohlomong esita le metsotso ka mor'a liphihlelo tse ling (mohlala, Greenough & Chang, 1989). Bokantle, sena se ka bonahala se sa lumellane le ts'ebetso ea tlhahiso e kholo ea li-synapses e lateloang ke ho faola ka mokhoa o hlalositsoeng pejana. Taba ea bohlokoa ke hore le ha pruning e le karolo ea bohlokoa ntlafatsong ea boko, boko bo tsoela pele ho theha li-synapses nakong eohle ea bophelo mme ha e le hantle li-synaps tsena lia hlokahala bakeng sa ho ithuta le ho hopola lintlha. Green, e ntšo le Wallace (1987) ba boletse hore ho na le phapang ea mantlha lipakeng tsa lits'ebetso tse laolang ho etsoa ha li-synapses kholisong ea bongoana ea pele le tse nakong ea nts'etsopele ea bongoana le ho ba motho e moholo. Ka ho khetheha, ba pheha khang ea hore mantsoe a khale a boqhetseke ke “liketsahalo” tse lebelletsoeng, tse sebetsang ho li khutlisa. Ba li bitsa li-synapses tsena e le "ba lebelletseng boiphihlelo" 'me ba hlokomela hore li fumanoa ka mokhoa o fapaneng nakong eohle ea kokoana-hloko. Ho fapana le hoo, sebopeho sa synapse sa morao se tsepamisitse maikutlo ho feta libakeng tse amehang ho sebetsanang le boiphihlelo bo ikhethang. Ba re mantsoe ana a hlalosoa e le "a ts'epahallang boiphihlelo." Karolo e 'ngoe e makatsang ea litlamorao tse tlisoang ke boiphihlelo mabapi le li-synapses ke hore ha se feela liphihlelo tse ikhethileng tse lebisang ho sebopeho se ikhethileng sa tatellano empa hape le tahlehelo ea tatellano ea synaptic. Kahoo, liphihlelo li fetola marang-rang a neural ka ho eketsa le ho faola maqhubu. Sena se re lebisa bothateng ba boteng ba polasetiki ea boko.

Melao-kakaretso ea Plasticity ho Brain ba tloaelehileng

Pele re bua ka liphihlelo tse susumetsang bokaholimo ba bokong, re tlameha ho hlahloba ka bokhutšoanyane melao-motheo e 'maloa ea bohlokoa ba polasetiki bokong bo tloaelehileng.

1. Liphetoho bokong li ka bonts'a maemong a mangata a tlhahlobo

Phetoho ea boits'oaro e tlameha ho bakoa ke phetoho e itseng bokong empa ho na le mekhoa e mengata ea ho etsa lipatlisiso ka liphetoho tse joalo. Liphetoho li ka fokotsoa mehatong ea lefats'e ea ts'ebetso ea boko, joalo ka mefuta e fapaneng ea Ka vivo monahano, empa liphetoho tse joalo li suthele hole le lits'ebetso tsa limolek'hule tse li tsamaisang. Liphetoho tsa lefats'e li bonts'a liphetoho tsa synaptic empa liphetoho tse bakoang ke liphetoho tsa limolek'hule tse kang liphetoho tse teng liteisheneng, polelo ea mofuta, joalo-joalo. Bothata ba ho ithuta polasetiki ea boko ke ho khetha lesupa le loketseng hantle potso e botsoang. Liphetoho mekhoeng ea khalsiamo li kanna tsa nepahala bakeng sa ho ithuta liphetoho tsa li-synaptic maemong a ikhethang a ka bang teng mabapi le ho ithuta habonolo empa a sa khone ho utloisisa phapang ea thobalano ts'ebetsong ea puo. Ho ka ba molemo ho feta ho ithuta ka tsona Ka vivo imaging kapa tlhahlobo ea postmortem ea morphology ea sele (mohlala, Jacobs & Scheibel, 1993). Karolo e loketseng e lokela ho lebisoa ho potso ea lipatlisiso e haufi. Boithuto ba ho etsa lipatlisiso maano a ho ntlafatsa ntlafatso ea tšebetsong kamora ho lemala hangata a sebelisa anatomical (cell morphology le kgokahano), physiological (cortical stimulation), le Ka vivo monahano. E 'ngoe le e' ngoe ea mekhahlelo ena e ka hokahana le liphetho tsa boitsoaro lithutong tsa batho le tsa batho athe litekanyetso tsa limolek'hule li ipakile li le thata haholo ho amana le boits'oaro, haholoholo boitšoaro ba kelello.

2. Mehato e fapaneng ea neuronal morphology e fetoha e ikemetseng 'me ka linako tse ling e fapana

Ho bile le tloaelo ho lingoliloeng ho bona liphetoho tse fapaneng tsa neuronal e le ho etsa joalo ka e mong. E 'ngoe ea tse atileng haholo ke ho nahana hore liphetoho tsa boholo ba lesapo la mokokotlo li bonts'a liphetoho ho bolelele ba dendritic le ka mokhoa o fapaneng. Sena ha se fetohe joalo ka ha mehato e 'meli e ka fapana ka boits'oaro' me ka linako tse ling e ka lehlakoreng le fapaneng (mohlala, Tloha, McDonald, & Kolb, 2010; Kolb, Cioe, & Comeau, 2008). Ho feta moo, lisele tse likarolong tse fapaneng tsa cortical, empa ka likholumong tse tšoanang tsa ho iketelletsa, li ka bonts'a likarabo tse fapaneng haholo liphihlelong tse tšoanang (mohlala, Teskey, Monfils, Silasi, & Kolb, 2006).

3. Liphetoho tse itšetlehileng ka boiphihlelo li ba le sepheo

Leha ho na le tšekamelo ea ho nahana ka liphetoho tsa polasetiki ho arabela liphihlelong ka hore li atile hohle bokong, ha se hangata ho etsahalang joalo. Mohlala, lithethefatsi tsa psychoactive li ka hlahisa liphetoho tse kholo tsa boitšoaro mme tsa ba le litlamorao tse mpe ho li-neuron, empa liphetoho tse sa fetoheng tsa polasetiki li shebile ka mokhoa o makatsang mme haholo li tsamaisana le "cortex" ea pele le li-nucleus. Robinson & Kolb, 2004). Ka lebaka leo, bafuputsi ba hloka ho nahana ka hloko mabapi le hore na libaka tse ntle haholo li hokae bakeng sa ho hlokomela liphihlelo tse ikhethang. Ho sitoa ho fumana liphetoho tse amanang le phetoho ea boitšoaro ha se bopaki ba ho ba teng ha liphetoho.

4. Liphetoho tsa polasetiki li itšetlehile ka nako

Mohlomong liphetoho tse kholo ka ho fetisisa mokhatlong oa synaptic li ka bonoa ho arabela ho beha liphoofolo tsa lab libakeng tse rarahaneng (tseo ho thoeng li "ruisitse"). Kahoo, ho na le liphetoho tse pharaletseng ho pholletsa le sensory le motor cortex. Liphetoho tsena li bonahala li tlola molao-motheo oa liphetoho tse itšetlehileng ka boiphihlelo ha li shebane empa ho fetoha ha liphetoho ho ka etsahala ka lebaka la semelo sa lefats'e sa liphihlelo ho kenyeletsa le liphihlelo tse sa bonahaleng joalo ka pono, tactile, auditory, olgicory, motor, le liphihlelo tsa sechaba. Empa liphetoho tsena tsa polasetiki ha se tsa ka mehla ebile li ka fetoha haholo ha nako e ntse e tsamaea.

Mohlala, ha likhoto li behiloe tikolohong e rarahaneng ho na le keketseho e nyane ea bolelele ba dendritic ho cortex ea pele e ka bonoang kamora matsatsi a mane a matlo a rarahaneng empa a nyamela kamora matsatsi a 14. Ka lehlakoreng le leng, ha ho na liphetoho tse totobetseng ho sensor cortex ka mor'a matsatsi a mane empa liphetoho tse hlakileng, ebile li bonahala li le teng ka mora matsatsi a 14 (Woau et al., 2010).

Monyetla oa hore ho na le liphetoho tse fapaneng tse sa foleng le tse liehang ho latela li-neurons tsa "cerebral" li lumellana le lipatlisiso tsa liphatsa tsa lefutso tse bonts'ang hore ho na le mefuta e fapaneng e hlahisitsoeng hantle le e sa fetoheng ho arabela maemong a rarahaneng (mohlala. Rampon et al., 2000). Phapang ea hore na liphetoho tse liehang ho feta le tse tsoelang pele li-network tsa li-neuronal li amana joang le boitšoaro ha li tsejoe.

5. Liphetoho tse itšetlehileng ka boiphihlelo lia kopana

Batho ba na le liphihlelo tsa bophelo bohle ba bona ba qalile ho tsa bo-mme le ho tsoela pele ho fihlela lefung. Liphihlelo tsena lia kopana. Mohlala, re bonts'itse litlelaboreng tsa laboratori hore haeba liphoofolo li pepesetsoa li-psychomotor tse hlasimollang ebang ke bana kapa li se li le batho ba baholo, liphihlelo tsa morao-rao li ba le tšusumetso e kholo (kapa ka linako tse ling e le sieo). Mohlala, ha likhoto li fuoa methylphenidate e le bana kapa li-amphetamine e le batho ba baholo ebe ka nako e 'ngoe li beoa libakeng tse rarahaneng kapa li koetliselitsoe mesebetsi ea ho ithuta, liphetoho tse itšetlehileng ka boiphihlelo li a koaloa. Se makatsang ke hore le ha lithethefatsi li sa bonts'e tšusumetso efe kapa efe e hlakileng e hlakileng ho libaka tsa sensory cortical, ho pepesetsoa pele ho nako ho thibela liphetoho tse lebeletsoeng libakeng tsena (mohlala, Kolb, Gibb, & Gorny, 2003a). Litšebelisano tsena tse amanang le lithethefatsi ha li ea hlokahala. Ha likhoto tsa bakhachane li fuoa khatello ea maikutlo bakeng sa metsotso ea 20 habeli ka letsatsi nakong ea bokhachane bo matla ba neurogeneis baneng ba bona (matsatsi a embryonic 12-18), litloholo tsa bona li bonts'a liphetoho tse amanang le khatello ea lesapo la mokokotlo mokokotlong oa preortal cortex (PFC) empa ha ho litlamorao tse amanang le lithethefatsi (Muhammad & Kolb, ka ho tobetsa a). Ha ho hlake hore na hobaneng ho se na litlamorao tse amanang le lithethefatsi kapa hore na sena se tla bolelang ho lemalla empa se bonts'a hore liphihlelo li kenella liphellong tsa bona bokong.

7. Liphetoho tsa polasetiki li latela lilemo

Ka kakaretso ho nahanoa hore boko bo ntseng bo hola bo tla arabela liphihlelong tse fetang tsa kelello ea motho e moholo kapa senescent. Sena se nepahetse haholo empa ho na le khokahano e 'ngoe ea bohlokoa: ho na le liphetoho tse fapaneng bokong ho arabela se shebahalang e le boiphihlelo bo le bong ka lilemo tse fapaneng. Mohlala, ha ho longoa likhoto tsa batho ba baholo, kapa ba baholo, kapa tsa senescent li beoa tikolohong e rarahaneng, lihlopha tsohle li bonts'itse liphetoho tse kholo empa li ne li fapane ka mokhoa o makatsang. Haholo-holo, athe re ne re lebelletse keketseho ea letsoalo la lesapo la mokokotlo ka lebaka la matlo a rarahaneng, sena e ne e le 'nete ho likhoto tsa batho ba baholo le ba senesi. Likhoto tse kentsoeng tikolohong eo bacha ba bontšitseng a fokotseha letsoai la lesapo la mokokotlo (Kolb et al., 2003a). Ho theoha ho tšoanang ha letsoele la lesapo la mokokotlo ho ile ha fumanoa lithutong tsa morao moo litoeba tse sa tsoa tsoaloa li fuoeng thobalano e bonolo ka borashe bo bonolo bakeng sa metsotso ea 15, makhetlo a mararo ka letsatsi ka matsatsi a leshome a pele a bophelo empa eseng joalo haeba ts'usumetso e se e le motho e moholo (Gibb, Gonzalez, Wagenest, le Kolb, 2010; Kolb le Gibb, 2010). Sebopeho se itšetlehileng ka lilemo tsa phetoho ea synaptic se bohlokoa ka ho hlaka bakeng sa ho utloisisa hore na liphihlelo li fetola kelello joang.

8. Ha se polasetiki eohle e ntle

Le ha mohopolo o akaretsang oa lingoliloeng ke hore liphetoho tsa polasetiki bokong li ts'ehetsa mesebetsi e ntlafalitsoeng ea makoloi le a kelello, liphetoho tsa polasetiki le tsona li ka sitisa boitšoaro le bona. Mohlala o motle ke liphetoho tse bakiloeng ke lithethefatsi tse bonoang ha li araba li-psychomotor stimulants (mohlala, Robinson & Kolb, 2004). Hoa utloahala ho fana ka maikutlo a hore boitšoaro bo bong bo bobe ba lithethefatsi tsa lithethefatsi bo ka bakoa ke liphetoho tse amanang le lithethefatsi ho morphology ea pele ho neuronal. Ho na le mehlala e meng e mengata ea ts'ebetso ea polasetiki ea methapo ho kenyelletsa le bohloko ba pathological (Baranauskas, 2001), karabelo ea methapo ea mafu ho kula (Tlhaselo, Capuron, & Miller, 2006), lefu la sethoathoa (Teskey, 2001), schizophrenia (Ntsu et al., 2004), le 'dementia'Mattson, Duan, Chan, & Guo, 2001).

Le ha ho na le lithuto tse ngata tsa boteng ba methapo ea kelello bokong bo ntseng bo hola, mohlala o totobetseng ke boteng ba bongoana ba bana. Mohlala o mong ke litlamorao tsa khatello ea maikutlo e tebileng ea prenatal, e bonts'itsoeng haholo ho fokotsa ho rarahana ha li-neuron ho cortex ea pele (mohlala, Murmu et al., 2006) 'me ka lehlakoreng le leng ho ka ama mesebetsi e tloaelehileng ea kelello le ea makoloi kholisong le botsofaling (mohlala, Halliwell, 2011). Leha mekhoa e tlisang liphetoho tsena e sa utloisisoe hantle, hoa tsebahala hore khatello ea pelehi ea kamora ho pelehi e ka fetola polelo ea liphatsa tsa lefutso bokong (Weaver et al., 2004; Weaver, Meaney, & Szf, 2006).

Lintlha Tse Sitisang Ntlafatso ea Brain

Ha bafuputsi ba qala ho ithuta liphetoho tse itšetlehileng ka bokong bo ntseng bo eketseha ho li-1950 le li-1960, ho ne ho e-na le khopolo ea tlhaho ea hore liphetoho tsa kholo ea kelello li ne li tla bonahala feela ha li arabela liphetoho tse kholo tsa boiphihlelo, joalo ka ho tsosoa lefifing. Lilemong tse fetileng tsa 20 ho ile ha hlaka hore esita le liphihlelo tse batlang li hloka toka li ka ama kholo ea kholo ea kholo le hore liphihlelo tse ka fetolang nts'etsopele ea boko li kholo haholo ho feta kamoo ho neng ho lumeloa pele (bona Lethathamo 2). Re tla totobatsa tse ling tsa litlamorao tse ithutoang hantle.

Tafole 2. 

Lintho tse susumetsang nts'etsopele ea boko le tšebetso ea tsona

1. Liphihlelo tsa maikutlo le likoloi

Mokhoa o bonolo oa ho thetsa boiphihlelo ho theosa le lilemo ke ho bapisa sebopeho sa boko ho liphoofolo tse lulang sebakeng se tloaelehileng sa mesebetsi ea matsoho ho ea ho liphoofolo tse behiloeng libakeng tse futsanehileng haholo kapa tseo ho thoeng li entsoe tikoloho. Ho holisa liphoofolo libakeng tse hlokang thuso joalo ka lefifi, khutso, kapa ho itšehla thajana ho etsa hore boko bo hōle. Mohlala, malinyane a lintja a holisitsoeng a le mong a bonts'a mefuta e mengata ea boits'oaro bo bobebe, ho kenyelletsa le ho hloka kutloelo-bohloko boiphihlelo bo bohloko (Hebb, 1949). Ka mokhoa o ts'oanang, ho holisa liphoofolo tse fapaneng joalo ka litšoene, likatse le litoeba lefifing ho kena-kenana le nts'etsopele ea sebono. Mohlomong lithuto tsa tsieleho tse tsebahalang ka ho fetisisa ke tsa Weisel and Hubel (1963) ea fepileng leihlo le le leng la li-kittens tse koetsoeng mme hamorao a bontša hore ha leihlo le buloa ho na le tahlehelo e sa feleng ea pono ea sepakapaka (amblyopia) (mohlala, Giffin & Mitchell, 1978). Haufinyane tjena, haufinyane bafuputsi ba ile ba nahana ka ketsahalo e fapaneng le ena, e leng ho fa liphoofolo menyetla ea pono ho bona hore na pono e ka ntlafatsoa. Phuputsong e le 'ngoe e ntle, Prusky et al. (Prusky, Silevera, Tschetter, Alam, & Douglas, 2008) o sebelisitse mofuta o mong oa mokhoa oa ho hlasimolla oo likhoto li kentsoeng tsamaisong ea "optokinetic" moo mela e otlolohileng ea maemo a sa tšoaneng ea sepakapaka e fetisang phoofolo pele. Haeba mahlo a bulehile 'me a shebile molekong o sisinyehang, ho ke ke ha khoneha hore liphoofolo, ho kenyelletsa le batho, ba qobe ho latela mela e tsamaeang, haeba maqhubu a sebaka a le lipakeng tsa pono. Bangoli ba beha liphoofolo ka har'a lisebelisoa tse ka bang libeke tse peli kamora letsatsi la ho bula mahlo (letsatsi la kamora ho phomola la 15). Ha e lekoa bakeng sa acuity ea bonono ha e se e le motho e moholo, liphoofolo li ile tsa bontša hoo e ka bang ntlafatso ea 25% ho acuity ea pono e amanang le liphoofolo ntle le kalafo ea pele. Botle ba thuto ea Prusky ke hore ts'ebetso ea pono e ntlafalitsoeng e ne e sa ipapisa le koetliso e ikhethileng, joalo ka ho ithuta bothata, empa e bile teng ka tlhaho ho arabela ho tlatsetso ea pono e ntlafalitsoeng.

Re lekile ho ntlafatsa boiphihlelo bo bobe ka ho sebelisa mokhoa o qalang oa Schanberg le Tšimo (1987). Lithutong tsena litoeba tsa masea li ile tsa fuoa takatso e matla ea ho li khothatsa ka brashi e nyane bakeng sa metsotsoana ea 15 makhetlo a mararo ka letsatsi bakeng sa matsatsi a 10-15 ho qala ha a hlaha. Ha masea a ithuteloa bongoaneng a bonts'a ts'ebetso ea makoloi e ntlafalitsoeng le ho ithuta ka sepakapaka hammoho le liphetoho mokhatlong o hlophisitsoeng oa synaptic ka har'a cortex ea likhoerekhoere. Kolb le Gibb, 2010). Leha mokhoa o hlakileng oa ts'ebetso ea ts'usumetso ea tactile o sa tsejoe, re bonts'itse hore ho ts'oaroa ha tactile ho lebisa keketseho ea tlhahiso ea neurotrophic factor, ukuaji oa fibroblast factor-2 (FGF-2) ka bobeli letlalong le bokong (Gibb, 2004). FGF-2 e tsejoa e bapala karolo ho nts'etsopele e tloaelehileng ea bokong 'me e ka susumetsa ho hlaphoheloa ho tsoa likotsi tsa boko ba perinatal (mohlala, Tloha, Hastings, & Kolb, 2007). Polelo ea FGF-2 e boetse e eketseha ho arabela mekhoa e fapaneng ea kalafo e kenyelletsang matlo a ntlafalitsoeng le lithethefatsi tsa psychoactive, tseo ka bobeli li susumetsang liphetoho tsa polasetiki bokong (bona ka tlase).

Mokhoa o mong oa ho ntlafatsa lits'oants'o tsa kutlo le makoloi ke ho beha liphoofolo libakeng tse rarahaneng tseo ho tsona ho nang le monyetla oa hore liphoofolo li ikopanye le maikutlo a feto-fetohang a maikutlo le sechaba le ho etsa ketso e fetang ea koloi hangata. Boithuto bo joalo bo thathamisitse liphetoho tse ngata tsa neural tse amanang le mofuta ona oa "ho ruisa." Tsena li kenyelletsa keketseho ea boholo ba bokong, botenya ba cortical, boholo ba neuron, lekala la dendritic, letsoalo la lesapo la mokokotlo, li-synapses ka neuron, lipalo tsa glial le ho rarahana le ho ruruha ha methapo ea methapo. (bap Greenough & Chang, 1989; Siervaag le Greenough, 1987). Boholo ba liphetoho tsena ha boa lokela ho nyahamisoa. Mohlala, lithutong tsa rona tsa litlamorao tsa makhulo a manyane a matlo bakeng sa matsatsi a 60 libakeng tse ntlafalitsoeng, ka boiketlo re bona liphetoho ka bongata ba boima ba boko ka tatellano ea 7-10% (mohlala, Kolb, 1995). Keketseho ena ea boima ba bokong e bonts'a ho eketseha hoa palo ea glia le methapo ea mali, boholo ba neuron soma, li-dendritic element, le li-synapses. Ho ka ba thata ho hakanya palo eohle ea li-synapses tse eketsehileng empa mohlomong e ka tatellano ea 20% ka cortex, e leng phetoho e sa tloaelehang. Habohlokoa, le ha lithuto tse joalo li bonts'a liphetoho tse itšetlehileng ka maemo nakong efe kapa efe, ho na le likhohlano tse sa lebelloang. Taba ea pele, likhoto tsa batho ba baholo lilemong life kapa life li bontša keketseho e kholo ea bolelele ba dendritic le letsoalo la lesapo la mokokotlo ho pholletsa le boholo ba kokoana-hloko ea likhoerekhoere empa likhoto tsa bana li bontša se tšoanang nyollelo ka bolelele ba dendritic empa a fokotseha boteng ba lesapo la mokokotlo. Ka mantsoe a mang, liphoofolo tse nyane li bonts'a phetoho e fapaneng molemong oa phepelo ea li-synapses ka li-neuron tsa pyramidal ha li bapisoa le liphoofolo tse tsofetseng (Kolb et al., 2003a). Taba ea bobeli, ha matamo a bakhachane a ne a beoa tikolohong e thata bakeng sa lihora tse robeli ka letsatsi pele ba ima mme ka nako eohle ea libeke tse tharo, tlhahlobo ea kelello ea bana ba bona ba baholo e bontšitse fokotseha ka bolelele ba dendritic le a nyollelo boteng ba lesapo la mokokotlo. Kahoo, ha ho na phello ea prenatal boiphihlelo empa phello e ne e fapana le boiphihlelo nakong ea bongoana kapa ho ba motho e moholo. Ho makatsang ke hore, liphetoho tsohle tse arabelang matlo a rarahaneng li lebisa tšebetsong ea kelello le tsa makoloi.

Ho na le melaetsa e meraro e hlakileng ho tsoa lithutong tsena. Taba ea pele, liphihlelo tse fapaneng tsa maikutlo le makoloi li ka hlahisa liphetoho tsa polasetiki tse telele bokong. Taba ea bobeli, boiphihlelo bo tšoanang bo ka fetola boko ka tsela e fapaneng ka lilemo tse fapaneng. Taba ea boraro, ha ho na kamano e bonolo pakeng tsa lintlha tsa polasetiki ea synaptic le boitšoaro nakong ea kholo. Taba ke eng, le ha ho le joalo, ke hore liphihlelo tsena tsa pele li na le phello e matla ho tlhophiso ea boko nakong ea kholo le botsofaling.

2. Lithethefatsi tse nang le kelello

Haesale ho tsejoa hore ho pepesetsoa joala kapele ho bohlokoa ka ho fetisisa bakeng sa nts'etsopele ea kelello, empa haufinyane ho sa tsoa bontšoa hore lithethefatsi tse ling tsa psychoactive, ho kenyeletsa le meriana e fanoeng, li ka fetola kholo ea kelello. Robinson le Kolb (2004) e fumane hore ho pepesehela li-psychomotor tse susumetsang botsofaling ho hlahisitse liphetoho tse kholo sebopeho sa lisele ho PFC le nucleus accumbens (NAcc). Ka ho khetheha, ha lithethefatsi tsena (amphetamine, cocaine, nicotine) li hlahisa tlhahiso ea dendritic bolelele le letsoapo la lesapo mokokotlong oa "medial prefrontal cortex" (mPFC) le NAcc, ho bile le phokotso ea mehato ena ho orbital frontal cortex (OFC), kapa maemong a mang , ha ho na phetoho. Kamora moo ba bonts'itse hore hoo e batlang e le sehlopha se seng le se seng sa lithethefatsi tsa psychoactive le sona se hlahisa liphetoho ho PFC, le hore litlamorao li fapana ho ea ka litereke tse peli tse tlang pele. Ka lebaka la hore boko bo ntseng bo hola hangata bo pepesetsoa lithethefatsi tse fanang ka kelello, kapa li utero kapa nakong ea bokhachane kamora ho pepa, re ile ra botsa hore na lithethefatsi tse li ka ba le phello efe ntlafatsong ea bongaka.

Lithuto tsa rona tsa pele li ile tsa sheba litlamorao tsa amphetamine kapa methylphenidate e fanoeng nakong ea bongoana (mohlala, Diaz, Heijtz, Kolb, le Forssberg, 2003). Lithethefatsi ka bobeli li fetotse mokhatlo oa PFC. Liphetoho tse entsoeng ka mokhoa oa dendritic li ne li amahanngoa le boits'oaro bo sa tloaelehang ba litšoantšiso tse sebelisoang ke lithethefatsi, kaha li bonts'itse ts'ebetso e fokotsoang ea papali ha e bapisoa le ba bapalang ba tšoaroang ke letsoai hammoho le ts'ebetso e sa sebetseng hantle boitekong ba ho hopola. Li-stimulants tsa Psychomotor ka hona li bonahala li fetola nts'etsopele ea PFC mme sena se bonahala liketsong tse mpe tsa boitšoaro bo amanang le boits'oaro bo amanang le pele ho bophelo.

Bana le bona ba ka fuoa meriana e fanoeng ke ngaka ka utero kapa ka mora nako. Mefuta e meraro e tloahelehileng ea lithethefatsi ke li-antipsychotic, li-antidepressants le li-anxiolytics. Ka boraro ba tsona li na le litlamorao tse kholo kholisong ea cortical. Frost, Cerceo, Carroll, le Kolb (2009) sekaseka boqapi ba dendritic ho litoeba tsa batho ba baholo ba tšoaroang ke paradigmatic tloaelehileng- (haloperidol) kapa lithethefatsi tsa atpsicalotic atypical (olanzapine) maemong a nts'etsopele a tsamaellanang le nako ea bokhachane (matsatsi a ka morao ho lefu la 3-10) kapa mekhahlelo ea bokhachane le pelehi (matsatsi a ho qetela a 3-20) bathong. Lithethefatsi ka bobeli li hlahisitse phokotso ea bolelele ba dendritic, ho rarahana ha dendritic, le letsoalo la mokokotlo ka har'a cortex ea medial pele le orbital. Phuputsong e ileng ea latela ea ho sebelisa likhoto bangoli ba bontsitse ho senyeha hoa mesebetsi e amanang le neuropsychological e amanang le PFC joalo ka memori e sebetsang.

Karolong e ts'oanang ea lithuto re shebile phello ea ho pepesetsoa bokhachane ho diazepam kapa fluoxetine ho likhoto (Kolb, Gibb, Pearce, & Tanguay, 2008). Ka bobeli lithethefatsi li amme kelello le boits'oaro, empa ka tsela tse fapaneng. Prenatal diazepam e ile ea eketsa bolelele ba dendritic le letsoalo la lesapo la mokokotlo liseleng tsa pyramidal ho parietal cortex mme sena se ne se amana le mesebetsi e ntlafalitsoeng ea makoloi. Ka lehlakoreng le leng, fluoxetine e fokotse mehato ea ho ngola 'me sena se ne se tsamaellana le bofokoli ba thuto ea sepakapaka bo holileng tsebong.

Potso e 'ngoe ke hore na ho pepesehela lithethefatsi tse nang le kelello "ho sa le pele, ho ka fetola bokaholimo ba kelello hamorao bophelong." Pele re ne re bonts'a hore haeba likhoto tsa batho ba baholo li fuoa amphetamine, cocaine, kapa nicotine ebe hamorao e beoa libakeng tse rarahaneng, polasetiki ea neuronal e ile ea koaloa (Hamilton le Kolb, 2005; Kolb, Gorny, Samaha, & Robinson, 2003b). Phuputsong e ileng ea latela ea morao ra fana ka litoeba tsa bana ba banyenyane methylphenidate mme ka nako ea ho ba motho e moholo re beha liphoofolo tsena libakeng tse rarahaneng mme, hape, ra fumana hore ho pepeseha ha lithethefatsi tsa pele ho thibetse liphetoho tse lebelletsoeng ho latela cortex (Comeau & Kolb, 2011). Ntle le moo, thutong e ts'oanang re bonts'itse hore mocha oa methylphenidate o pepesa ts'ebetso ho mesebetsi e amanang le methapo e amanang le ts'ebetso ea pele ho ts'ebetso.

Ka kakaretso, ho pepesetsoa litlhare le lithethefatsi tsa tlhekefetso ho na le tšusumetso e kholo ntlafatsong ea pele le boits'oarong bo amanang le pele. Litlamorao tsena li bonahala e le tsa nako e telele kapa li tšoarella 'me li ka susumetsa polasetiki ea boko ho ba motho e moholo. Litlamorao tse mpe tse sa lebelloang tsa litlhare tse fanoang ke ngaka ho bokeng le boits'oarong ntle ho pelaelo li bohlokoa ho ntlafatso ea bongoana ba motho. Ho hlakile hore ha se mohala o bonolo oa hore na bo-mme baimana ba nang le khatello ea maikutlo e tebileng, psychosis, kapa khatello ea maikutlo ba lokela ho fuoa meriana e fuoeng hore maemo ana a boitšoaro a ka ama tsoelo-pele ea boko leseeng mme haholoholo ho fihlela boiphihlelo ba 'm'a- litšebelisano tsa masea. Patlisiso e fana ka maikutlo a hore meriana e joalo e lokela ho sebelisoa ka tekanyetso e tlase e sebetsang joalo ka ho sebelisoa eseng feela bakeng sa litlamorao tsa bona tsa "ho khutsisa" bo-mme.

3. Li-hormone tsa Gonadal

Phello e totobetseng ea ho pepesetsoa li-hormone tsa gonadal nakong ea kholo ke phapang ea liphatsa tsa lefutso tse qalang pele ho nako. Maemong ana tlhahiso ea testosterone ke banna e lebisa kholong ea botona ba botšehali. Hamorao bophelong, estrogen le testosterone ka bobeli li ama li-receptor libakeng tse ngata tsa 'mele, ho kenyelletsa le bokong. Boithuto ba MRI ba nts'etsopele ea boko ba motho bo bonts'itse phapang e kholo ka tekanyo ea kholo ea kholo ea botona ho tsa botona le botšehali (O'Hare & Sowell, 2008). Ka ho khetheha, palo eohle ea boko e fihla ho asymptote ho basali ho pota lilemo tsa 11 le 15 ho ba batona le ba batšehali ka ho latellana. Empa ho na le taba e fetang ea botona ba botona le botšehali ho feta tekano ea kholo. Ka mohlala, Kolb le Stewart (1991) e bonts'itsoe likhoto hore li-neuron tsa MPFC li na le masimo a maholo a dendritic ho banna le hore li-neuron tsa OFC li ne li na le lisele tse kholo ho basali. Liphapang tsena li ile tsa fela ha liphoofolo li ne li gonadectomised ka nako ea tsoalo. Ka mokhoa o ts'oanang Goldstein et al. (2001) e entse tlhahlobo e felletseng ea bophahamo ba 45 ea libaka tse fapaneng tsa boko ho tsoa ho li-MRI scans tsa batho ba baholo ba phetseng hantle. Ho bile le phapang ea thobalano ka bongata, e lekanang le kakaretso ea khoele, mme sena e ne e le 'nete haholo ho PFC: basali ba ne ba e-na le bophahamo bo boholo ba PFC ea banna empa banna ba ne ba e-na le bonyenyane ba boholo ba OFC. Lesorphism lena la thobalano le amana le maemo a phahameng a tikoloho ea li-steroid tsa thobalano nakong ea bophelo ba pele liphoofolong tsa liphoofolo. Kahoo e hlaha ho batho le ho liphoofolo tse sebetsang hore li-hormone tsa gonadal li fetola kholo ea "cortical". Sena se bohlokoa haholo ha re nahana hore litlamorao tsa liphihlelo tse ling tse kang ho pepesehela matlo a rarahaneng kapa li-psychomotor tse matlafatsang le tsona ke tlhekefetso ea thobalano. Ho bonahala monyetla oa hore liphihlelo tse ling tsa ntlafatso li ka fetoha ka mokhoa o fapaneng matsoalo a basali le a banna, leha lithuto tse fokolang li hlile li entse papiso ena.

4. Kamano ea motsoali le ngoana

Masea a Mamary a tsoaloang a sa le monyane a tobana le phephetso e kholo bophelong ba pele. Ba itshetlehile ho batswadi ba bona mme ba tlameha ho ithuta ho tseba, ho hopola le ho khetha bahlokomeli ba bona. Le ha joale re tseba hore liphoofolo tse nyane (esita le liphoofolo tse anyesang) li ka ithuta ho feta kamoo li neng li ananeloa ka teng (bona tlhahlobo ka Hofer le Sullivan, 2008), ha ho na pelaelo ea hore likamano tsa motsoali le ngoana li bohlokoa ebile li bapala karolo ea bohlokoa kholisong ea boko. Phapang tseleng ea litšebelisano tsa pelehi tsa masea le masea e ka hlahisa litlamorao tsa nako e telele tse tsoelang pele ho ba motho e moholo.Myers, Brunelli, Squire, Shindledecker, & Hofer, 1989). Mohlala, lipatlisiso tsa litoeba li bonts'itse hore nako e sebelisitsoeng ho ikopanya, palo ea ho nyeka ha bo-mme le nako eo bo-'mè ba e qetang ba le maemong a hloahloa a ho phomola a phahameng ka mokhoa o fapaneng ka mefuta e fapaneng ea boits'oaro le boits'oaro. Lilemong tse leshome tse fetileng Meaney le basebetsi mmoho le yena (mohlala Cameron et al., 2005) ba khonne ho bonts'a tšebelisano ena ea methapo ea bokhachane le masea ka mokhoa o hlophisehileng nts'etsopele ea karabelo ea khatello ea maikutlo le maikutlo a fapaneng a boits'oaro le kelello. Liphetoho tsena li hokahane le liphetoho liphatseng tsa "corticosterone cell" tsa hippocampal cell membrane, tseo ka tsona li laoloang ke liphetoho lipuong tsa mofuta oa gene (Weaver et al., 2006).

Litlamorao tsa phapang e teng tlhokomelong ea bo-mme ha li fuoe hippocampus, leha ho le joalo, ebile e ka ba e atile. Ka mohlala, Fenoglio, Chen le Barum (2006) e bonts'itse hore tlhokomelo e ntlafalitsoeng ea bo-mme nakong ea beke ea pele ea bophelo e hlahisitse liphetoho tse tsitsitseng tseleng ea ho bua ka seleng ho hypothalamus le amgydala (bona hape tlhaiso ka Fenoglio, Bruson, & Barum, 2006).

Ha re tsebe lithuto tse tšoanang tse talimang boithuto ba pelehi, 'me bo pele-pele, e le polasetiki ho arabela karohanong ea litšebelisano tsa masea le masea, empa liphetoho tse joalo li bonahala li ka etsahala. Mohlala, re bontšitse, hore karohano ea ka mehla ea bo-mme, ke ts'ebetso e neng e sebelisoa ho eketsa tšebelisano ea masea le masea nakong ea bokhachane. Fenoglio et al. (2006) ho ithuta, ho eketsa bolelele ba dendritic le letsoalo la lesapo la mokokotlo ka bobeli mPFC le OFC litekong tsa batho ba baholo (Muhammad le Kolb, 2011).

5. Likamano tsa lithaka

Kamano ea lithaka e 'nile ea tsejoa ho susumetsa boitšoaro ba batho ba baholo ho tloha ka lithuto tsa Harlow (mohlala, Harlow & Harlow, ka 1965). E 'ngoe ea likamano tse matla tsa lithaka ke ho bapala, e bontšitsoeng e le bohlokoa bakeng sa nts'etsopele ea bokhoni ba batho ba baholo ba sechaba (mohlala, Pellis le Pellis, 2010). Bokapele bo ka pele bo phetha karolo ea bohlokoa boits'oarong ba sebapali. Ho lemala ha lesea ho MPFC le OFC beha melao ea boitšoaro papaling, leha e le ka mekhoa e fapaneng (mohlala, Pellis et al., 2006). Ka lebaka la liphetho tse joalo, re ile ra nahana hore nts'etsopele, le ts'ebetso e ileng ea latela ea litereke tse peli tse tlang pele li ka fetoloa ka mokhoa o fapaneng haeba boitšoaro ba papali bo ne bo laoloa nts'etsopele. Likhoto tsa bana joale li fuoe monyetla oa ho bapala ka likhoto tsa batho ba baholo tsa 1 kapa 3 kapa ka 1 kapa 3 liphoofolo tse ling tsa bacha. Ha ho na mokhoa oa ho bapala le liphoofolo tsa batho ba baholo empa boits'oaro ba ho bapala bo ne bo eketseha le liphoofolo tse nyane tse neng li le teng. Tlhahlobo ea lisele ho PFC e bonts'itse hore li-neurons tsa OFC li arabile ho palo ea lithaka tse teng, eseng hore na ho bapala ho etsahetse kapa che, ha li-neurons tsa MPFC li arabile tekanyo ea papali empa eseng palo ea Conspecifics (Bell, Pellis, le Kolb, 2010). Kamora moo re bonts'itse letotong la lithuto tsa hore maiphihlelo a fapaneng a pele a fetola sebapali sa papali, ho kenyelletsa khatello ea bokhachane, ho tsosa takatso ea bongoana, le ho pepesetsoa ha bacha ka methylphenidate (mohlala, Muhammad, Hossain, Pellis, & Kolb, 2011) hape, maemong 'ohle, ho na le lits'oants'o tsa tlhabollo ea pele. Ho ka ba le thuto ea bohlokoa mona ha re nahana ka maemo ao ho bapala karolo ea motho ho sa tloaelehang, joalo ka autism kapa tlhokomeliso ea khatello ea kelello (ADHD). Litšitiso tsa boitšoaro ba papali li ka ama kholo ea pele le boits'oaro ba batho ba baholo ha morao.

6. Khatello ea maikutlo

Ho na le lingoliloeng tse ngata tse bokelitsoeng lilemong tse fetileng tsa 60 tse bonts'ang litlamorao tsa khatello ea kelello le boits'elo ho batho ba baholo empa ke haufinyane tjena moo ho ananeloang karolo ea khatello ea maikutlo ho masea. Hajoale hoa tsebahala hore khatello ea maikutlo le ea bokhachane e rera motho e mong bakeng sa boitšoaro bo fapaneng ba maladaptive le psychopathologies. Mohlala, khatello ea maikutlo makhopho a bokhachane ke sesosa sa kotsi ea kholo ea kholo ea maikutlo, ADHD, khatello ea maikutlo le bokhoba ba lithethefatsi (Anda et al., 2006; van den Bergh le Marcoen, 2004). Boithuto ba liteko mabapi le liphoofolo tsa lab bo netefalitse liphumano tsena ka sephetho se akaretsang ke hore khatello ea kelello, lithutong le linthong tse seng tsa batho, e hlahisitse lits'oants'o tsa boits'oaro tse joalo ka karabelo e phahameng le e telele ea khatello ea maikutlo, ho sitisoa ho ithuta le ho hopola, ho haella tlhokomelong, ho fetotsoe tlhahlobo boits'oaro, mekhoa e fetotsoeng sechabeng le papaling, le keketseho e atileng ea joala (mohlala, tlhahlobo ea Weinstock, 2008).

Liphetoho tsa polasetiki mokhatlong oa synaptic oa boloi ba liphoofolo tse sithabetseng maikutlo ha li ithutoe hantle, leha ho le joalo, litlamorao li bonahala li amana le lintlha tsa boiphihlelo bo sithabetsang. Ka mohlala, Murmu et al. (2006) ea tlaleha hore khatello ea maikutlo ea bokhachane bo lekaneng bekeng ea boraro ea boimana e bakile ho fokotseha ha letsoalo la mokokotlo le bolelele ba dendritic ho mPFC le OFC ea lifus ea batho ba baholo. Ka papiso, Muhammad & Kolb (2011) e fumane hore khatello ea maikutlo a bokhachane nakong ea beke ea bobeli ea boimana e fokotse khatello ea lesapo la mokokotlo ho mPFC empa ha e na phello ho OFC le ho eketsa letsoalo la mokokotlo ho NAcc ea likhoto tsa batho ba baholo. Tlhahlobo ea bolelele ba dendritic e bontšitse sebopeho se fapaneng kaha ho bile le keketseho ea bolelele ba dendritic ho mPFC le NAcc empa ho fokotseha ha OFC. Ka mokhoa o makatsang, Mychasiuk, Gibb le Kolb (2011) o fumane hore khatello ea maikutlo nakong ea beke ea bobeli ea boikoetliso e ile ea eketsa letsoalo la mokokotlo ho mPFC le OFC ha ho hlahlojoa methapo ea bongoana ho fapana le likhoto tsa batho ba baholo. Ho etsoa liphuputso tsena ho bonts'a hore liphapang tse bang teng nakong ea khatello ea bokhachane le nako eo boko bo hlahlojoang ka eona e fella ka liphetoho tse fapaneng lipolasetiking makaleng a neuronal. Ntho e 'ngoe e hlakileng, leha ho le joalo, ke hore litlamorao tsa khatello ea maikutlo ea boimana li bonahala li fapane le tsa khatello ea kelello ea batho ba baholo. Ka mohlala, Liston et al. (2006) Pele ho bonts'oa hore khatello ea batho ba baholo e lebisitse ho fokotseheng ha makala a dendritic le letsoai la mokokotlo ho mPFC empa keketseho ea OFC.

Re tseba ka phuputso e le 'ngoe feela e shebileng litlamorao tsa khatello ea pelehi ea pelehi (karohano ea bo-mme) holima mokhatlo o hlophisitsoeng ka methapo e meholo ea batho ba baholo. Kahoo, Muhammad & Kolb (2011) o fumane hore karohano ea bo-mme ea basali e eketsa letsoalo la lesapo ho mPFC, OFC, le NAcc litekong tsa batho ba baholo. Ho sa ntseng ho ikemiselitse ho latela kameho ea khatello ea bokhachane kapa ea masea ke hore liphapang tsena tsa phetoho ea synaptic li amana joang le boits'oaro bo morao kapa hore na li-neurons tsa polasetiki li tla sebetsana joang le liphihlelo tse ling tse kang matlo a rarahaneng, papali kapa likamano tsa motsoali oa masea. Lithuto tse joalo li netefalitsoe hore e tla ba mohloli oa lithuto tsa kamoso.

7. Limela tsa kahare

Hang ka mor'a ho hlaha, liphoofolo tse anyesang li lula ka potlako ho mefuta e mengata ea likokoana-hloko tsa tlhaho. Likokoana-hloko tsena li susumetsa nts'etsopele ea mesebetsi e mengata ea 'mele. Mohlala, "gut micobiota" e na le litla-morao tsa ts'ebetso ea ts'ebetso ea sebete (mohlala, Björkholm et al., 2009). Hobane ho na le kamano e tsebahalang lipakeng tsa maloetse a methapo ea methapo ea methapo e kang Autism le schizophrenia le tšoaetso ea pathogen ea microbioal nakong ea nako ea pherekano (mohlala, Finegold et al., 2002; Mittal, Ellman, & Cannon, 2008), Diaz Heijtz et al. (ka khatiso) o ile a ipotsa hore na tšoaetso e joalo e ka fetola kholo ea kelello le boits'oaro. Ba joalo. Bangoli ba ile ba bapisa litekanyetso tsa boits'oaro ba motor le bokong ho litoeba tse tsoetseng pele le ka ntle ho 'mele "gut mircrobiota". Bangoli ba fumane hore libaktheria tsa gut li susumetsa ho ts'oara litsela, neurotransmitter turnover, le tlhahiso ea liprotheine tse amanang le synaptic ho cortex le striatum ho nts'etsapele litoeba 'me liphetoho tsena li ne li amana le liphetoho mesebetsing ea makoloi. Ena ke phello e thabisang hobane e fana ka leseli la tsela eo ts'oaetso e ka fetolang kholo ea kelello le boits'oaro bo latelang ba batho ba baholo.

8. Lijo

Ho na le lingoliloeng tse ngata ka litlamorao tsa liprotheine le / kapa lijo tse thibetsoeng tsa caloric bokong le boits'oarong (mohlala, Lewis, 1990) Empa ha ho tsejoe haholo ka litlamorao tsa lijo tse matlafatsoang kholisong ea boko. Ka kakaretso ho nahanoa hore mmele o fola hantle ha o fuoa phepo e nepahetseng kahoo hoa utloahala ho bolela esale pele hore nts'etsopele ea boko e ka tsamaisoa ke li-vithamine le / kapa litlatsetso tsa liminerale. Tlatsetso ea phepo ea lijo ka lijo nakong ea bongoana e hlahisa mefuta e fapaneng ea phetoho le boits'oaro (Meck & Williams, 2003). Mohlala, tlatsetso ea perineatal choline e lebisa tlhokomelong e ntlafalitsoeng ea sebaka litekong tse fapaneng tsa ho sesa ka sepakapaka (mohlala, Meck & Williams, 2003; Tee, & Mohammadi, 1999) mme e eketsa maemo a kholo ea kholo ea methapo (NGF) ho hippocampus le neocortex (mohlala, Sandstrom, Loy, & Williams, 2002). Halliwell, Tee and Kolb (2011) o ile a etsa lithuto tse tšoanang mme a fumana hore tlatsetso ea choline e eketsa bolelele ba dendritic ho pholletsa le cortex ea mokokotlo le ho hippocampal CA1 pyramidal neurons.

Halliwell (2011) hape o ithutile ka litlamorao tsa ho eketsoa ha tlatsetso ea vithamine / mineral ho lijo tsa likhoto tse lactating. O ile a khetha ho sebelisa tlatsetso ea lijo tse tlalehiloeng ho ntlafatsa mohopolo le mabifi ho batho ba baholo le ba lilemong tsa bocha ba nang le mathata a fapaneng (Leung, Wiens le Kaplan, 2011) le ho fokotsa khalefo, maemo a ts'ebetso le ho tlosoa hoa sechaba ho autism ka keketseho ea boits'oaro (Mehl-Madrona, Leung, Kennedy, Paul, & Kaplan, 2010). Tlhahlobo ea bana ba baholo ba likhoto tse lactating tse fepelitsoeng tlatsetso e tšoanang e fumane keketseho ea bolelele ba dendritic ho li-neuron ho MPFC le parietal cortex empa eseng ho OFC. Ntle le moo, lijo li bile le phepo e ntle ea ho khutlisa litlamorao tsa khatello ea maikutlo ea bokhachane phokotsong ea bolelele ba dendritic ho OFC.

Ho sa ntse ho hlokahala hore re ithute ka litlamorao tsa thibelo ea lijo le tlatsetso ho ntlafatso ea marang-rang a boitšoaro le boitšoaro. Ts'ebetso tsena ka bobeli li fetola kholo ea boko empa joalo ka lintho tse ling tse ngata tse tšohliloeng mona, ha re na setšoantšo se hlakileng sa hore na liphihlelo tsa pele li tla sebetsana joang le liphihlelo tsa morao-rao, joalo ka lithethefatsi tsa kelello, ho fetola kelello le boitšoaro.

Nahanisisa

Kutloisiso ea rona ea mofuta oa kholo ea ts'ebetso e tloaelehileng ea boko e tsoetse pele haholo lilemong tse fetileng tsa 30 empa re sa tsoa qala ho utloisisa lintlha tse ling tse fetolang nts'etsopele ena. Ho utloisisa modumo ena ho tla ba bohlokoa hore re qale ho bula makolopetso a mathata a methapo ea methapo le ho qala liphekolo tsa pele ho thibela kapa ho khutlisa liphetoho tsa methapo ea methapo. Khatello e totobetseng ke hore liphihlelo ha se liketsahalo tse ikhethang empa ha re ntse re phela bophelong, liphihlelo li hokahana ho fetola boitšoaro le kelello, e leng mohato o tsejoang hangata e le metaplasticity.

Ha re ntse re buisana ka liphetoho tse fapa-fapaneng tse bileng teng bokong bo ntseng bo hola re sebelisitse “kelello e ntseng e hola” jwaloka nako e le 'ngoe. Ho hlakile hore ha ho joalo 'me re na le khoao ea hore qetellong re tla fumana hore ho na le lifensetere tse bohlokoa tsa nako eo mohopolo o holang o arabelang ho feta ka linako tse ling. Ntle le moo, ho ka etsahala hore libaka tse fapaneng tsa likomkomere li bonts'a lifensetere tse fapaneng tsa bohlokoa. Ka mohlala, re fumane hore haeba motor cortex e lemeha bongoaneng ho na le sephetho se fosahetseng se amanang le ho lemala ho tšoanang nakong ea bohlankanaNemati le Kolb, 2010). Ka mokhoa o makatsang, leha ho le joalo, morao-rao ke 'nete bakeng sa ho lemala mokokotlong oa pele. Ho khetholla li-windows tse bohlokoa tse itšetlehileng ka batho ba bang e tla ba phephetso lilemong tse leshome tse tlang.

Re tsepamisitse maikutlo mona mehatong ea polasetiki ea synaptic empa re fela re elelloa hore liphetoho tsa polasetiki mokhatlong oa boko li ka ithutoa maemong a mang a mangata. Qetellong mochine oa motheo oa phetoho ea synaptic o tla fumanoa ka polelo ea gene. Ho thata ke hore ho na le monyetla oa hore liphihlelo tse fetolang boitšoaro ka mokhoa o fapaneng li tla amana le liphetoho lipakeng tsa mefuta e makholo kapa makholo. Phephetso ke ho supa liphetoho tse amanang haholo le liphetoho tsa boitšoaro bo shebiloeng.

Liteboho / Likhohlano tsa khahleho

Re lakatsa ho leboha ka bobeli NSERC le CIHR ka tšehetso ea bona ea nako e telele bakeng sa lithuto tse amanang le mosebetsi oa rona tse boletsoeng tlhahlobisong ena. Re boetse re leboha Cathy Carroll, Wendy Comeau, Dawn Danka, Grazyna Gorny, Celeste Halliwell, Richelle Mychasiuk, Arif Muhammad, le Kehe Xie ka menehelo ea bona e mengata lithutong.

References

  • Anda RF, Felitti VJ, Bremner JD, Walker JD, Whitfield C, Perry BD, Giles WH. Litholoana tsa nako e telele tsa tlhekefetso le liphihlelo tse bohloko tse amanang le tsona bongoaneng. Khokahano ea bopaki bo tsoang ho neurobiology le lefu la mafu. Li-Archives tsa Europe tsa Psychiatry le Clinical Neuroscience. 2006; 256: 174-186. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
  • Baranauskas G. plastikiity e kentsoeng ke lesapo la mokokotlo. Ka: Shaw CA, McEachern J, bahlophisi. Ho ea ka khopolo ea Neuroplasticity. Philadelphia, PA: Psychology Press; 2001. maq. 373-386.
  • Bell HC, Pellis SM, Kolb B. Juvenile tsebo ea papali ea lithaka le nts'etsopele ea cortex ea orbitof Pambal le medial prefrontal. Patlisiso ea Bitso ea Boitšoaro. 2010; 207: 7-13. [E fetotsoe]
  • JE e ntšo, Kodish IM, Grossman AW, Klintsova AY, Orlovskaya D, Vostrikov V, Greenough WT. Pathology ea safu ea "V" pyramidal neurons kahara "cortex" ea bakuli ba nang le lefu la kelello. Koranta ea Amerika ea Psychiatry. 2004; 161: 742-744. [E fetotsoe]
  • Blumberg MS, Freeman JH, Robinson SR. Moeli o mocha oa ntlafatso ea boits'oaro bo amanang le boitšoaro. Ka: Blumberg MS, Freeman JH, Robinson SR, bahlophisi. Oxford Handbook of Developmental Behaisheral Neuroscience. New York, NY: Oxford University Press; 2010. maq. 1-6.
  • Björkholm B, Bok CM, Lundin A, R After J, Hibberd ML, Pettersson S. Intestinal micobiota e laola metabolism ea xenobiotic ho sebete. MAHLOMOLA MOTHO. 2009; 4: e6958. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
  • Cameron NM, Champagne FA, Carine P, Fish EW, Ozaki-Kuroda K, Meaney M. Lenaneo la liphapang tsa batho ka bomong mabapi le likarabo tse itšireletso le maano a ho ikatisa ka mokhoa oa phapano ka phapano ea tlhokomelo ea basali. Tlhahlobo ea Neuroscience le Biology. 2005; 29: 843-865. [E fetotsoe]
  • Woau W, Hastings E, Kolb B. Tšusumetso e fapaneng ea ts'ebeliso ea pele le ea morao-rao ea FGF-2 ka mor'a ho lemala ha kalafo ea methapo. Patlisiso ea Bitso ea Boitšoaro. 2007; 180: 18-27. [E fetotsoe]
  • Woau WL, McDonald R, Kolb B. Liphetoho tse entsoeng pele ho li-cortical. Patlisiso ea Bitso ea Boitšoaro. 2010; 214: 91-101. [E fetotsoe]
  • Woau W, Kolb B. Ho pepesetsoa ha bana malapeng a methylphenidate lithutong tsa morao-rao tsa boiphihlelo ka hara motho e moholo. 2011. Buka e ngotsoeng ka letsoho.
  • Diaz Heijtz R, Kolb B, Forssberg H. Na mokhoa oa phekolo oa amphetamine nakong ea bongoaneng o ka fetola sebopeho sa mokhatlo o hlophisitsoeng oa synaptic bokong? Koranta ea Europe ea Neuroscience. 2003; 18: 3394-3399. [E fetotsoe]
  • Diaz Heijtz R, Wang S, Anuar F, Qian Y, Björkholm B, Samuelsson A, Pettersson S. Tloaelehileng ea gut microbiota e hlophisa kholo ea kelello le boitšoaro. Ts'ebetso ea National Academy of Science (USA) (ka khatiso). [Tlhahiso ea mahala ea PMC] [E fetotsoe]
  • Eriksson PS, Perfi lieva E, Björk-Eriksson T, Alborn AM, Nordborg C, Peterson DA, Gage FH. Neurogeneis ho hippocampus ea motho e moholo. Bongaka ba Tlhaho. 1998; 4: 1313-1317. [E fetotsoe]
  • Fenoglio KA, Brunson KL, Baram TZ. Hippocampal neuroplasticity e khothalletsoang ke khatello ea pele ea bophelo: likarolo tsa mosebetsi le tsa limolek'hule. Frontiers ho Neuroendocrinology. 2006; 27: 180-192. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
  • Fenoglio KA, Chen Y, Baram TZ. Neuroplasticity ea hypothalamic-pituitary-adrenal axis esale pele ea bophelo e hloka ho hloa lipakeng tsa libaka tse tsamaisang khatello ea kelello. Tlaleho ea Neuroscience. 2006; 26: 2434-2442. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
  • Finegold SM, Molitoris D, Sefela Y, Liu C, Vaisanen ML, Bolte E, Kaul A. Gastrointestinal microflora lithuto tsa morao-rao tsa ho qala. Maloetse a tšoaetsanoang a Clinical. 2002; 35 (Suppl 1): S6-S16. [E fetotsoe]
  • Frost DO, Cerceo S, Carroll C, Kolb B. Ho pepeseha ha haloperidol kapa Olanzapine ho etsa liphetoho tsa nako e telele tsa mofuta oa dendritic. Synfall. 2009; 64: 191-199. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
  • Phihlelo ea Gibb R. Polao le ho hlaphoheloa ho tsoa likotsi tsa bokong. 2004. Thesis e sa phatlalatsoang ea PhD, Univesithi ea Lethbridge, Canada.
  • Gibb R, Gonzalez CLR, Wegenast W, Kolb B. Tactile stimulation e thusa ho hlaphoheloa kamora ho lemala ha kotlo ea sethoathoa ho likhoto tsa batho ba baholo. Patlisiso ea Bitso ea Boitšoaro. 2010; 214: 102-107. [E fetotsoe]
  • Giffin F, Mitchell DE. Sekhahla sa ho hlaphoheloa pono ka mor'a ho nyahlatsoa ke batho ba le mong nakong e tlang. Tlaleho ea Physiology. 1978; 274: 511-537. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
  • Goldstein JM, Seidman LJ, Horton NJ, Makris N, Kennedy DN, Cainess VS, Tsuang MT. Litorphism tse tloaelehileng tsa botona le botšehali tsa kelello ea motho e moholo tse hlahlojoang ke monahano oa vivo oa magnetic resonance. Cerebral Cortex. 2001; 11: 490-497. [E fetotsoe]
  • Gould E, Tanapat P, Hastings NB, Shors TJ. Neurogeneis ka ho ba motho e moholo: Karolo e ka bang teng ho ithuteng. Mekhoa ea Mahlale a Kelello. 1999; 3: 186-192. [E fetotsoe]
  • Greenough WT, Black JE, Wallace CS. Boiphihlelo le kholo ea bokong. Psychobiology ea Nts'etsopele. 1987; 22: 727-252. [E fetotsoe]
  • Green WT, Chang FF. Plasticity ea sebopeho sa synapse le paterone ho cortex ea cerebral. Ka: Peters A, Jones EG, bahlophisi. Cerebral Cortex, Vol 7. New York, NY: Plenum Press; 1989. maq. 391-440.
  • Gregg CT, Shingo T, Weiss S. Neural stem cell ea 'mele oa maqhubu a limela. Symposium ea Mokhatlo oa Biology ea Tlhahlobo. 2001; 53: 1-19. [E fetotsoe]
  • Halliwell CI. Ho kena lipakeng kalafo ho latela khatello ea maikutlo le pherekano ea bana. 2011. Thesis e sa phatlalatsoang ea PhD, Univesithi ea Lethbridge, Canada.
  • Halliwell C, Tee R, Kolb B. Prenatal choline e ntlafatsa ho hlaphoheloa ho tsoa likotsi tsa sefahleho tsa pele ho litoeu. 2011. Buka e ngotsoeng ka letsoho.
  • Hamilton D, Kolb B. Nicotine, boiphihlelo le polasetiki ea bokong. Neuroscience ea Boitšoaro. 2005; 119: 355-365. [E fetotsoe]
  • Harlow HF, Harlow MK. Lits'ebetso tsa lerato. Ka: Schier A, Harlow HF, Stollnitz F, bahlophisi. Mekhoa ea litloholo tsa batho bao e seng batho. Moq. 2. New York, NY: Phatlalatso ea Litaba; 1965.
  • Hebb HLALE. Mokhatlo oa Boitšoaro. New York, NY: McGraw-Hill; 1949.
  • Hofer MA, Sullivan RM. Ho ea ho neurobiology ea khokahano. Ka: Nelson CA, Luciana M, bahlophisi. Handbook of Developmental Cognitive Neuroscience. Cambridge, MA: Press ea MIT; 2008. maq. 787-806.
  • Jacobs B, Scheibel AB. Tlhahlobo e ngata ea ho lekola ea sebaka sa Wernicke ho batho. I. Lifespan e fetoha. Tlaleho ea Nearology e Bapisang. 1993; 327: 383-396. [E fetotsoe]
  • Kempermann G, Gage FH. Lisele tse ncha tsa methapo bakeng sa boko ba batho ba baholo. Mahlale a Amerika. 1999; 280 (5): 48-53. [E fetotsoe]
  • Kolb B. Brain plasticity le boits'oaro. Mahwah, NJ: Erlbaum; 1995.
  • Kolb B, Cioe J, Comeau W. Phapang e fapaneng ea mesebetsi ea ho ithuta ka makoloi le pono ea pono mabapi le dendritic arborization le lesapo la mokokotlo ka likhoto. Neurobiology ea ho Ithuta le Memori. 2008; 90: 295-300. [E fetotsoe]
  • Kolb B, Gibb R. Tactile e tsosa maikutlo e thusa ho hlaphoheloa ha phetoho le phetisetso ea dendritic kamora marapo a morao-rao a morao-rao a medal kapa a posterior parietal ka litoeba. Patlisiso ea Bitso ea Boitšoaro. 2010; 214: 115-120. [E fetotsoe]
  • Kolb B, Gibb R, Gorny G. Liphetoho tse itšetlehileng ka ts'ebeliso ea dendritic arbor le letsoalo la lesapo la mokokotlo ho neocortex li fapana ka lilemo le thobalano. Neurobiology ea ho Ithuta le Memori. 2003a; 79: 1-10. [E fetotsoe]
  • Kolb B, Gorny G, Li Y, Samaha AN, Robinson TE. Amphetamine kapa cocaine e fokotsa bokhoni ba boiphihlelo ba morao-rao ho ntlafatsa polasetiki ea sebopeho ho neocortex le li-bokellase tsa nucleus. Ts'ebetso ea National Academy of Science (USA) 2003b; 100: 10523-10528. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
  • Kolb B, Gibb R, Pearce S, Tanguay R. Ho pepesetsoa bongaka ba meriana ho phekola ho hlaphoheloa ha litlhare kamora ho tsoa likotsi tsa ts'enyo ea nako ea pele ho likhoto. Mokhatlo oa Neuroscience Abstracts. 2008; 349: 5.
  • Kolb B, Morshead C, Gonzalez C, Kim N, Shingo T, Weiss S. Keketseho-e matlafatsang molumo oa lithane tse ncha tsa cortical le ho hlaphoheloa hoa ts'ebetso kamora ho senyeha ha setla-morao ho litla-morao tsa makoloi. Ralitaba tsa Cerebral flow flow le Metabolism. 2007; 27: 983-997. [E fetotsoe]
  • Kolb B, Stewart J. Liphapang tse amanang le thobalano ho dendritic branching ea lisele tse khubung ea pele ea likhoto. Tlaleho ea Neuroendocrinology. 1991; 3: 95-99. [E fetotsoe]
  • Kolb B, Whishaw IQ. Bohlokoa ba Neuropsychology ea Batho. Khatiso ea 6th. New York, NY: E na le bohlokoa; 2009.
  • Leung BM, Wiens KP, Kaplan BJ. Na tlatsetso ea micrutrient ea bongoana e ntlafatsa tsoelo-pele ea bana kelellong? Tlhahlobo e hlophisehileng. Ho ima ha lesea la BCM. 2011; 11: 1-12. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
  • Lewis PD. Phepo e nepahetseng le nts'etsopele ea bokong. Ka: van Gelder NM, Butterworth RF, Drujan BD, bahlophisi. (Mal) Phepo e Ntle le Boko ba masea. New York, NY: Wiley-Liss; 1990. maq. 89-109.
  • Liston C, Miller MM, Godwater DS, Radley JJ, Rocher AB, Hof PR, McEwen BS. Liphetoho tse bakiloeng ke khatello ea maikutlo ho tsa pele-pele tsa cortical dendritic morphology li bolela esale pele menyetla ea ho senyeha ha maemo a fetohang tlhokomelo ea kelello. Tlaleho ea Neuroscience. 2006; 26: 7870-7874. [E fetotsoe]
  • Palamente ea Mattson, Duan W, Chan SL, Guo Z. Ho: Ho Theory of Neuroplasticity. Shaw CA, McEachern J, bahlophisi. Philadelphia, PA: Psychology Press; 2001. maq. 402-426.
  • Meck WH, Williams CL. Ho ts'oaroa ha metabolism ea choline ka ho fumaneha ha eona nakong ea boimana: Litlamorao bakeng sa ho hopola le ho hlokomela lintho ka hloko ho pholletsa le nako ea bophelo. Tlhahlobo ea Neuroscience le Biology. 2003; 27: 385-399. [E fetotsoe]
  • Mehl-Madrona L, Leung B, Kennedy C, Paul S, Kaplan BJ. Micronutrients khahlanong le taolo e tloaelehileng ea meriana ho autism: Phuputso ea taolo ea tlhaho ea tlhaho. Tlaleho ea Psychopharmacology ea bana. 2010; 20: 95-103. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
  • Mittal VA, Ellman LM, Cannon TD. Tšebelisano ea tikoloho ea tikoloho ea botona le botšehali ho schizophrenia: Karolo ea mathata a thibelo. Bulletin ea Schizophrenia. 2008; 34: 1083-1094. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
  • Muhammad A, Hossain S, Pellis S, Kolb B. Tactile e susumetsang nakong ea nts'etsopele e fumana sensorization ea amphetamine le alter neuronal morphology. Neuroscience ea Boitšoaro. 2011; 125: 161-174. [E fetotsoe]
  • Matšoenyeho a bokhachane a prenatal modulates modulates le methapo ea kutlo ea mokokotlo ntle le ho ama maikutlo a amphetamine. Nuroscience ea Ntlafatso (ka khatiso). [E fetotsoe]
  • Muhammad A, Kolb B. Ho arohana ha 'mele ha bolekane bo fetotsoeng le khatello ea methapo ea methapo ntle le ho susumetsa maikutlo a amphetamine. Patlisiso ea Bitso ea Boitšoaro. 2011; 223: 7-16. [E fetotsoe]
  • Murmu M, Salomon S, Biala Y, Weinstock M, Braun K, Bock J. Ho fetoha ha letsoalo la lesapo la mokokotlo le ho rarahana ha dendritic kahara cortex ea pele ho bana ba bo-mme ba pepesitsoeng khatello ea maikutlo nakong ea kemaro. Koranta ea Europe ea Neuroscience. 2006; 24: 1477-1487. [E fetotsoe]
  • Ho sithabela ha Mychasiuk R, Gibb R, Kolb B. Ho imeloa kelellong ho bakang khatello ea maikutlo ho mpefatsa liphetoho tse bang teng sethaleng le hippocampus ea ho holisa bana. Patlisiso ea Brain. 2011; 1412: 55-62. [E fetotsoe]
  • Myers MM, Brunelli SA, squire JM, Shindledecker R, Hofer MA. Boitšoaro ba bo-'mè ba likhoto tsa SHR kamanong ea bona le khatello ea mali ea bana. Psychobiology ea Nts'etsopele. 1989; 22: 29-53. [E fetotsoe]
  • Nemati F, Kolb B. Kotsi ea motor cortex e na le litlamorao tse fapaneng tsa boits'oaro ho litoeba tsa bongoana le tsa bocha. Neuroscience ea Boitšoaro. 2010; 24: 612-622. [E fetotsoe]
  • O'Hare ED, Sowell ER. Ho etsa liphetoho tsa boqhetseke tse putsoa le tse tšoeu bokong ba motho. Ka: Nelson CA, Luciana M, bahlophisi. Handbook of Developmental Cognitive Neuroscience. Cambridge, MA: Press ea MIT; 2008. maq. 23-38.
  • Pellis SM, Hastings E, Takeshi T, Kamitakahara H, Komorowska J, Tšoarelo ML, Kolb B. Litlamorao tsa tšenyo ea orbital ea kapele ea cortex ho module ea likarabo tse itšireletsang ka likhoto maemong a bapalang le a sa bapalang. Neuroscience ea Boitšoaro. 2006; 120: 72-84. [E fetotsoe]
  • Pellis S, Pellis V. Sebapali se Bapalang. New York, NY: Lingoliloeng tsa Oneworld; 2010.
  • Prusky GT, Silver BD, Tschetter WW, Alam NM, Douglas RM. Pulasetiki e itšetlehileng ka boiphihlelo ho tloha ho buloa mahlo e etsa hore ho be le ntlafatso e tšoarellang ea pono ea "cortex-based" Tlaleho ea Neuroscience. 2008; 28: 9817-9827. [E fetotsoe]
  • Raison C, Capuron L, Miller AH. Cytokines e bina ka bokhabane: Ho ruruha le pathogenesis ea khatello ea maikutlo. Mekhoa ea thuto ea boitšoaro. 2006; 27: 24-31. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
  • Rampon C, Jiang CH, Dong H, Tang YP, Lockart DJ, Schultz PG, Hu Y. Liphello tsa ho ruisa tikoloho ho polelo ea gene bokong. Ts'ebetso ea National Academy of Science (USA) 2000; 97: 12880-12884. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
  • Robinson TE, Kolb B. Tsela ea polasetiki e amanang le lithethefatsi tsa tlhekefetso. Neuropharmacology. 2004; 47 (Suppl 1): 33-46. [E fetotsoe]
  • Sandstrom NJ, Loy R, Williams CL. Tlatsetso ea prenatal choline e eketsa maemo a NGF ho hippocampus le frontort cortex ea bacha le ba baholo. Patlisiso ea Brain. 2002; 947: 9-16. [E fetotsoe]
  • Schanberg SM, Tšimo ea TM. Matšoenyeho a ho nyenyefatsa le ho hlasimolla ho kopaneng ea popo le neonate ea pele ho motho. Kholiso ea bana. 1987; 58: 1431-1447. [E fetotsoe]
  • Sirevaag AM, Greenough WT. Kakaretso e akaretsang ea lipalo-palo ea methapo ea "testaptic" ea likhoto tse fumanehang maemong a rarahaneng, a sechaba le a batho ka bomong. Patlisiso ea Brain. 1987; 441: 386-392. [E fetotsoe]
  • Tees RC, Mohammadi E. Litlamorao tsa tlatsetso ea lijo tsa neonatal choline ho thuto ea mahlale a batho ba baholo le ho ithuta ka kemiso ho memeng. Psychobiology ea Ntlafatso. 1999; 35: 226-240. [E fetotsoe]
  • Teskey GC, Monfils MH, Silasi G, Kolb B. Neocortical kindling e amana le liphetoho tse hanyetsanang morphology ea dendritic ho neocortical layer V le striatum e tsoang ho neocortical layer III. Synfall. 2006; 59: 1-9. [E fetotsoe]
  • Teskey GC. Ho sebelisa kindling ho etsisa liphetoho tsa neuroplastic tse amanang le ho ithuta le ho hopola, mathata a neuropsychiatric, le lefu la sethoathoa. Ka: Shaw CA, McEachern JC, bahlophisi. Ho ea ka khopolo ea neuroplasticity. Philadelphia, PA: Taylor le Francis; 2001. maq. 347-358.
  • van den Bergh BR, Marcoen A. Matšoenyeho a phahameng a bokhachane a amanang le matšoao a amanang le bokhachane bo amana le matšoao a ADHD, mathata a kantle ho naha, le matšoenyeho ho bana ba lilemo li 8- le 9. Kholiso ea bana. 2004; 75: 1085-1097. [E fetotsoe]
  • Weaver ICG, Cervoni N, Champagne FA, D'Alessio AC, Sharma S, Seckl JR, Dymov S, Szf M, Meaney MJ. Lenaneo la Epigenetic ka boits'oaro ba bo-mme. Neuroscience ea Tlhaho. 2004; 7: 847-854. [E fetotsoe]
  • Weaver ICG, Meaney M, Szf M. Litlhokomelo tsa batsoali tlhokomelong ea bo-mme le maikutlong a amanang le ho tšoenyeha ka hara bana ba fetoloang boteng. Ts'ebetso ea National Academy of Science (USA) 2006; 103: 3480-3486. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
  • Weinstock M. Litlamorao tsa boits'oaro ba nako e telele tsa khatello ea maikutlo ea prenatal. Tlhahlobo ea Neuroscience le Biology. 2008; 32: 1073-1086. [E fetotsoe]
  • Wiesel TN, Hubel DH. Likarabo tsa sele e le 'ngoe ka li-cortex tsa striate tsa kittens li amohuoe pono ka leihlo le le leng. Tlaleho ea Neurophysiology. 1963; 26: 1003-1017. [E fetotsoe]