Ts'ebetso ea tsamaiso ea tsamaiso ea dopamine le karabelo ea eona e feto-fetohang le e nang le bothata ba khatello ea kelello (2015)

Pauline Belujon, Anthony A. Grace

DOI: 10.1098 / rspb.2014.2516 E phatlalalitsoe 18 Hlakubele 2015

inahaneloang

Le ha, ho ea ka nalane, tsamaiso ea norepinephrine e hohetse tlhokomelo e ngata lithutong tsa karabelo ea khatello ea maikutlo, sisteme ea dopamine le eona e bile le tšusumetso e tšoanang. Ke nako e telele ho thehiloe hore khatello ea maikutlo e bapala karolo ea bohlokoa ho pathogeneis ea mafu a kelello. Leha ho le joalo, mekhoa ea methapo ea kutlo e arabelang karabelo ea khatello ea maikutlo le phello ea eona ho mafu a mafu a kelello ha e utloisisoe hantle. Sisteme ea dopamine e ka bapala likarolo tse ikhethileng khatellong ea kelello le mafu a kelello. Ho nahanoa hore, leha sisteme ea dopamine (DA) e theha motheo oa mafu a kelello, lefu lena la methapo ea methapo le ka hlaha mehahong e ikamahantseng le tsamaiso ea DA. Tlhahlobo ena e lekola tsebo ea hajoale ea liphetoho tse amanang le khatello ea maikutlo le litaba tsa DA, mme e hlahisa lintlha tsa morao-rao tse shebileng sephetho sa khatello ea maikutlo tsamaisong ea DA le bohlokoa ba eona ho mafu a kelello.

1. Selelekela

Khatello ea maikutlo e hlalosoa ka kakaretso e le tšusumetso efe kapa efe e ka bakang phephetso ho homeostasis ea setho (bakeng sa tlhahlobo, bona [1]). Kahoo, maikutlo a sithabetsang a hlohlelletsa likarabo tse ngata tsa boko ba 'mele tse reretsoeng ho arabela kotsing e ka bang teng. Ka sebele, ts'ebetso ea axisalamic-pituitary-adrenal (HPA) axis, e tsosang karabo ea 'ntoa-kapa-sefofane', ke karabelo e kholo ea neuroendocrine le 'mele ea mmele ha o le kotsing ea ho tsoa maemong a kotsi [2]. Mokokotlo oa HPA ke khokahano ea boko le tsamaiso ea methapo; e na le hypothalamus, e lokollang corticotropin-releasing factor (CRF) le merero ho sethoathoa sa ka ntle, se sephiring sa adrenocorticotropic hormone (ACTH). Sehlahisoa sa pheletso ea ts'ebetso ea axis ea HPA (ke hore, ho lokolloa ha glucocorticoids ke adrenal gland) e sebeletsa ho lemosa setho ka liphetoho tsa tikoloho le 'mele le ho boloka homeostasis [3]. Ka hona, sebopuoa se hloka karabelo ea khatello ea maikutlo ho pholoha maemo a joalo; leha ho le joalo, khatello ea maikutlo le ts'ebetso e sa lekaneng ea adrenocortical e ka ba kotsi hobane e ka nolofatsa maemo a mangata a pathological, joalo ka ts'ebeliso e mpe ea lithethefatsi [4], le khatello e kholo ea maikutlo [5] kapa psychotic [6] mathata. Mokokotlo oa HPA o laoloa ke lihlopha tse fapaneng tsa balekane, mme ka ho khetheha ke likarolo tse amanang le tsamaiso ea limbic tse kang prefrontal cortex (PFC), li-nucleus accumbens (NAc), amygdala le hippocampus [7]. Karabo e 'ngoe ea neuroendocrine ea mantlha maikutlong a sithabetsang ke ts'ebetso ea tsamaiso ea methapo ea kutlo, e hlahisang ho lokolloa ka potlako ha noradrenaline bokong, ka ts'ebetso ea li-neurus tsa locus coeruleus (LC). Mokokotlo oa HPA le Sisteme ea LC-norepinephrine (NE) ke likhokahanyo tse peli tsa bohlokoa tsa boko tse tsamaisanang le khatello ea maikutlo ka mokhoa o hlophisehileng. Leha ho le joalo, ho bonahetse haholoanyane le ho feta hore tsamaiso ea dopamine (DA) e bapala karolo ea bohlokoa karabong ea khatello ea maikutlo, mme haholo karabong ea ts'ebetso ea methapo ea methapo e boneng likhathatso tse ngata tsa mafu a kelello. Sisteme ea DA e bapala karolo e kholo ts'ebetsong ea meputso ea tlhaho le ea maiketsetso. Ka nnete ho bile le tlhahiso ea hore mesolimbic DA e arolelane likarolo tsa mofufutso oa monyaka o khothatsang [8], le hore e sebetsa e le lets'oao la ho ithuta la ho matlafatsa boitšoaro [9]. Boithuto bo bongata bo boetse bo tlalehile hore tokollo ea DA e eketseha ha ho arajoa maikutlo a liphoofolo tse buoang, tse fanang ka maikutlo a hore DA e kanna ea ameha lipapatsong tse susumetsang le tse lebisang tlhokomelo boits'oarong bo boits'oaro bo susumetsang molemong oa tšusumetso e loketseng, ekaba ea lehlasoa kapa la moputso [10,11]. Ho senyeha tsamaisong ea DA ho kentsoe ke mathata a mangata a kelello le methapo ea kutlo, ho kenyeletsoa schizophrenia, lefu la Parkinson, ts'ebeliso e mpe ea lithethefatsi le mathata a maholo a khatello ea maikutlo (MDDs). Leha liphuputso tse ngata li hlalositse ts'ebetso ea mantlha le ea mafu a tsamaiso ea DA bokong, ho senola mokhoa o rarahaneng oa ts'ebetso ea methapo ea kutlo ea DA ho bohlokoa ho fumana mekhoa e lekaneng le e sebetsang ea kalafo ea ho phekola mathata a 'maloa. Ehlile, mathata a mangata a na le monyetla oa hore a ka hlaha meahong e amanang le taolo ea sistimi ea DA. Tlhatlhobo ena e akaretsa tsebo ea hajoale ea ho feto-fetoha ha maemo a khatello ea maikutlo / DA, mme e hlahisa lintlha tsa morao-rao tse shebaneng le phello ea khatello ea maikutlo tsamaisong ea DA le kamano ea eona le mathata a kelello.

2. Khatello ea khatello ea kelello le norepinephrine

Sistimi ea catecholaminergic e hokahantsoe khafetsa le likarabo tsa khatello ea maikutlo. Haholo-holo, Sisteme ea LC-NE e bapala karolo ea bohlokoa taolong ea linaha tsa boitšoaro, ho kenyeletsoa le likarabo tsa khatello ea maikutlo, mme e ka sebeletsa ho matlafatsa boemo ba lihlahisoa tsa bocha e le ho ikamahanya le maemo a phephetso (bakeng sa tlhahlobo, bona [3]). Sisteme ea LC-NE e hlahisoa ke mefuta e mengata ea khatello ea maikutlo e kenyelletsang thibelo [12], footshocks [13] le khatello ea maikutlo sechabeng [14]. Haholo-holo, ka mor'a ho pepesetsoa khatello ea maikutlo, ho na le keketseho ea tšebetso ea li-neuron tsa LC [15], hammoho le keketseho ea phetoho ea NE libakeng tseo morero oa LC neurons o tsoang ho tsona [16]. Lion of the LC e sitoa ho thibela khatello ea kelello ea HPA axis ha e sebetsana le khatello ea maikutlo e telele empa e arabela likarabo tsa li-hormone tsa neuroendocrine ho khatello e matla ea thibelo [17], ho fana ka maikutlo a hore Sisteme ea LC-NE e khothaletsa likarabo tsa mmele ho khatello ea maikutlo. Ho feta moo, ts'ebetso ea li-neuron tsa LC tse nang le CRF e na le litlamorao tse tšoanang le khatello ea maikutlo [18]. Leha ho le joalo, LC ha e hlahelle ka kotloloho ho axis ea HPA empa hape le likarolo tse fapaneng tse amanang le khatello ea maikutlo tse kang amygdala le hippocampus, tseo ka nako e 'ngoe li romelletseng likhakanyo ho axis ea HPA [19]. Subralulum ea hippocampus (vSub), sephetho sa mantlha sa hippocampus, e fumana bolulo bo boholo ho tsoa tsamaisong ea LC-NE [20] hape ea kenella ts'ebetsong ea tlhahisoleseling ea litaba [21]. Kahoo, vSub ke sebopeho sa karabelo ea khatello ea maikutlo, hobane moelelo oa khatello moo o etsahalang o bohlokoa ho tataisa karabelo e hlakileng ea ntho e phelang [22]. Sena se tsamaisana le keketseho ea polelo ea Fos ka hippocampus likoteng tse pepesitsoeng likhatellong tse fapaneng, ho kenyelletsa le khatello ea maikutlo, ho sesa le khatello ea maikutlo [23]. Boithuto bo bonts'itse hore Sisteme ea LC-NE ke sebopeho se sebetsang sa ts'ebetso ea neuron ea vSub, e kenyelletsa inhibition kapa activation ea vSub neurons, e ka bakang khatello ea maikutlo [24]. Ho feta moo, footshocks e etsa hore boholo ba li-neuron ka har'a vSub, bo ikamahanye le likarabo ho khothaletso ea LC ho li-neuron tsa VSub [25]. VSub e hlokomela likarolo tse 'maloa tsa' mele oa sefahleho, joalo ka PFC le amygdala, tseo le tsona li nang le lipakanyo tsa "neurothalamic paraventricular neurons", tse fanang ka maikutlo a hore vSub e na le tšusumetso e matla ea kopaneng ea khatello ea maoto le matsoho [26], e ka amang homeostasis. Amygdala, mme haholo-holo basolateral nucleus (BLA), eo le eona e fumanang projeke e matla ea LC [19], e sebetsa ka mokhoa o ts'oanang nakong ea khatello ea maikutlo [27], mme ts'ebetso ea boiphetetso e teng lipakeng tsa liphetho tsa vSub le BLA [25]. Maemo a NE a eketseha ho BLA nakong ea khatello ea maikutlo [28], le li-neuron tsa BLA ka botsona li arabela mohopolong o khathatsang o bontšitsoeng hore o boetse o kenya li-neuron tsa LC [29]. Matšoenyeho a bolokiloeng kapa a phetoang hangata a hlahisa litlamorao ho hippocampus le BLA, joalo ka dendritic atrophy ho hippocampus, 'me e eketseha ka dendrites le methapo ea methapo ho BLA [30]. Ho na le khokahano e matla pakeng tsa polasetiki ea synaptic le liphetoho tsa morphological masimong [31]. Mohlala, khatello ea maikutlo e matla e etsa hore ho be le keketseho ea nako e telele ea ts'ebeliso ea "adrenergic" e tsamaeang le nako e telele ho BLA [32], e fanang ka maikutlo a hore ho se kopane ha khatello ea kelello ha ho sebetsane le mathata a kelello, ho ka kenyelletsa ho senyeha ha potoloho ena. Joalokaha ho se ho boletsoe, mPFC le eona ke karolo ea bohlokoa ho arabello ea khatello ea maikutlo. E etsoa ka mokhoa o ikhethileng oa khatello ea kelello le ea sechaba [33], mme e tsejoa ho fetolela tšebetso ea neuroendocrine nakong ea khatello ea maikutlo ka modumo ho tsoa tsamaisong ea LC-NE [34]. Ho feta moo, ts'ebeliso ea nako e telele tseleng ea BLA-PFC e thibetsoe ke ho pepesetsoa khatello ea maikutlo e ka se keng ea qojoa [35], hammoho le tšitiso ea polasetiki ea synaptic tseleng ea PFC-BLA [36], ho fana ka maikutlo a hore litšebelisano tsena tsa phetetso le tsona li ka bapala karolo ea bohlokoa karabong ea khatello ea maikutlo.

Le ha ho le joalo, nalane, Sistimi ea NE e amana le likarabo tsa khatello ea maikutlo, lithuto tse 'maloa le tsona li hatelletse tsamaiso ea DA karabong ea khatello.

3. Sisteme ea dopamine

Lithuto tsa Neurochemical li bontšitse hore tsamaiso ea DA e hlahisoa ke ts'usumetso e bolokehileng ea khatello [37]. Khatello ea khafetsa ea khatello ea maikutlo e fetolela karabelo ea mesolimbic DA system ho khatello ea maikutlo, le khatello ea maikutlo e pheta-phetoang joalo ka mohatla-pinch e nolofalletsa ho iphumanela ho itaola ha li-psychostimulants tse kang cocaine le amphetamine (bakeng sa tlhahlobo, bona [37]). Liso tse mpe tsa sebakeng sa "ventral tegmental sebakeng" (VTA) li fetola karabelo e tloaelehileng ea li-hormone ho khatello ea maikutlo ho batho ba baholo, ho bontša hore tsamaiso ea DA e ka ba le tšusumetso ho axis ea HPA [38]. Boemo ba electrophysiological ba sistimi ea DA ea VTA bo ithutile haholo (bakeng sa tlhahlobo, bona [39,40]). Kahoo, Ka vivo liphuputso li bonts'a hore halofo ea li-neuron tsa DA tse ho VTA ha li sebetse ebile ha li thunye ka boomo [41]. Li-neuron tsena li tšoareloa sebakeng se sa phetseng hantle, se sa sebetseng ka tšusumetso ea GABAergic e tsoang ho "ventral pallidum" (VP). Kannete, ho se sebetse ha liphatlalatso tsa pallidal ho lokolla li-neuron ho inhibition ea GABAergic, ho li nolofalletsa ho cha ka mollo hanghang [42]. Ho tse ntseng li thunya ka boithatelo, mefuta e 'meli ea mohlala e bonoa le ho laoloa ke litsamaiso tse fapaneng. Ka hona, li-neurons tsa DA li bonts'a mokhoa o sa tloaelehang, oa ho thula 'moho o le mong (kapa tšebetso ea' tonic '), le mokhoa oa ho thunya o tšabehang (kapa' mosebetsi oa 'phasic' [43,44]. Mokhoa oa phasic o itšetlehile ka ho kenella ha glutamatergic afferent [44], haholo-holo tse hlahang ho pedunculopontine tegmentum (PPTg) [45]. Phasic burst ho shapuoa ho lumeloa hore ke phello e ntle ea sistimi ea DA e fetolang boitšoaro bo lebisitsoeng ho sepheo (bakeng sa ho lekola, bona [39]), le liphetoho tsa phasic ho phatloheng li etsahala ka lebaka la ts'usumetso e nang le boemo, kapa kamora moputso oa mantlha, 'me ba bonts'itsoe ho fana ka karabelo ea phoso ea ho bolela esale pele maemong a tsebahalang [46] le likhoto [47]. Le ha boits'oaro ba koloi bo iketlile bo le sieo 'me ts'ebetso ea methapo e sa sebetse liphoofolong tse fumanehang,' me ho phatloha ka karabelo ea ts'usumetso ha ho le teng, joalo ka ha ho bonts'itsoe nakong ea boroko bo tebileng likatseng [48], thepa e phatlohileng le ho phatloha ha tonic mme tsamaiso ea bona e bapisoa le e bonoang ho liphoofolo tse boitšoaro [49,50]. Ho ke ho hlokomeloe hore litoebeng tse nang le bokhoni ba ho hlaka, ho phatloha ho hoholo ho bonoe ka lebaka la ts'usumetso e bonoang kamora ho hlakoloa ha colliculus e phahameng [51]. Ntle le moo, liphoofolong tse itsoarang, liphoofolo tse iphetotseng li na le thepa e ts'oanang le marokho a hlahisoang ka lebaka la khothalletso [49]. Ntle le moo, lithethefatsi tse thibelang ho thunya ho hoholo, joalo ka bahanyetsi ba NMDA ba kentse VTA [52], hape kenella likarabo tsa ho ithuta [53]. Ka hona, mekoloko e itlhahelang feela e ithutoang liphoofolong tse nang le bana e entsoe ka mokhoa o ts'oanelang le melaoana e phatlohang ka mokhoa o khothatsang liphoofolong tse phelang tse phelang.

Ts'ebetso ea papemaker ea thunya ea tonic, ka lehlakoreng le leng, e fana ka boemo ba tšebetso ba DA [42] e khethollang likhatello tsa DA tsa extrasynaptic. Ts'ebetso ea vSub e baka keketseho ea palo ea li-neuron tse sebetsang tsa DA ka boiketsetso (ke hore, tšebetso ea baahi) empa ha e na phello ho sekhahla sa ho thunya kapa mosebetsi oa ho phatloha [54]. Sena se bontšitsoe hore se etsahala ka tsela ea polysynaptic ho fetela NAc le VP [55]. Ts'ebetso ea phasic ea ho phatloha ho hoholo ha li-neuron tsa DA e ka etsahala feela ho li-neuron tse senyehileng le tse thuntseng hanghang; ka lehlakoreng le leng, "neuronpolarised" ea neuron e bonts'a sekhahla sa NMDA 'me se ke ke sa baka mollo ke NMDA [56]. Ka lebaka leo, vSub e laola palo ea li-neuron tsa DA tse ka sebelisoang ka mokhoa o pharalletseng ke PPTg ka ho beha tokiso ea tonic ea li-neuron tsa DA. Ka hona, PPTg e fana ka 'letšoao', 'me vSub ke' phaello 'ea lets'oao lena. VSub e bapala karolo ea ts'ebetso e itšetlehileng ka maemo mme ts'usumetso ea eona e ka fapana ho latela tikoloho. Kahoo, maemong a kopanetsoeng, tšusumetso e loketseng e tla kenya tšebetsong PPTg, e lumelle karolo e lekanyelitsoeng ea li-neuron tsa DA tse ikemiselitseng ho sebetsa ka vSub ho chesa ka mollo.Setšoantšo sa 1a).

Setšoantšo sa 1. 

Ts'ebetso ea bongata ba dopamine (DA) e laoloa ke meaho e 'meli, "subralulum" ea hippocampus (vSub) le basolateral amygdala (BLA). (a) Tekano lipakeng tsa mabokose ana a neng a ka ba teng ha taba e ntse e le maemong a bobebe e lumella feela karolo e nyane ea li-neuron tsa DA ho chesa hanghang (ka ho le letšehali); maemong ana, tšusumetso e loketseng ea boits'oaro e sebetsang ea ts'ebetso ea pedunculopontine tegmentum (PPTg) e ne e tla lebisa ho phatloha ho hoholo ka karolo e nyane feela ea li-neuron, ho fella ka lets'oao le lenyane la dopamine. (b) Maemong a sithabetsang, vSub e tla etsa hore palo ea li-neurons tsa DA e thunye ka tšohanyetso, e lumelle karabelo e kholo ea dopamine ho tsoa ho khothaletso e loketseng boits'oaro. (c) Kamora ho tlosa khatello ea maikutlo, BLA e nyarosang e tla kenya ts'ebetsong ts'ebetso ea DA, ka tsela eo e lebise ho hloleha ho araba ts'usumetso e ntle.

4. Matšoenyeho le tikoloho

Likarabo tsa khatello ea maikutlo li itšetlehile ka moelelo oa taba. Mohlala, haeba rat e senoloa ke khatello ea maikutlo, joalo ka boemo ba tšabo bo maemong, e tla bonts'a boits'oaro bo ts'oenyehileng ha e khutlisetsoa maemong a ts'oanang a tikoloho [57]. Litlamorao tsa khatello ea maikutlo tsamaisong ea DA li fapana haholo ho latela hore na ke 'mele, maikutlo le tšusumetso e mpe. Mohlala, khothalletso e le 'ngoe e se nang letsoalo, joalo ka thuthuho e hlahisoang liphoofolong tse fumanehang, e bontšitsoe hore e fokotseha ka sekhahla ha sekhahla sa ho thunya ha methapo ea DA [58-60], 'me maemong a mang keketseho [59,61]. Hamorao ho bontšitsoe hore ho fokotseha ho fokotsoang hoa lebelo la ho thunya ha methapo ea DA ho fumaneha boholo karolong ea bongaka ea VTA, empa hore tšibollo ea nakoana e fumaneha karolong ea morao-rao [62]. Ho fapana le hoo, ha ts'usumetso e mpe e fanoa khafetsa ka nako e behiloeng, boemo bo bontšitsoeng ho eketsa maemo a DA libakeng tsa post-synaptic ka ho etsa litoeba [63], keketseho ea nakoana ea tšebetso ea baahi ba li-neuron e khothalletsoa. Ho fapana le tšusumetso e mpe ea maikutlo a hlobaetsang, keketseho ena e ka thibeloa ke ho se sebetse pele ho vSub [62]. Ha khatello ea 'mele e sebelisoa ka kotloloho e tsitsitseng, joalo ka khatello ea maikutlo kapa e thibelang, ho na le keketseho ea ts'ebetso ea neuron ea DA e boetse e khutlisoa ke ho se sebetse ha vSub [62]. Liphuputso tsa Microdialysis li boetse li bonts'itse hore DA e lokolloa seterateng ho arabela khatello e se nang 'nete e kang ho thothomela hoa motlakase le mohatla o pharalletseng [64]. Ho sithabela ho bontšitsoe ho sithabela maikutlo le amphetamine, hobane karabelo ea locomotor ea acute amphetamine, e amanang le ts'ebetso e eketsehileng ea VTA [65] le tokollo e eketsehileng ea DA ho NAc, e ntlafalitsoe [66] ka mokhoa o itšetlehileng ka vSub [62]. Ka hona, boemo bo hatellang ba hyperdopaminergic bo tsamaisoa ke hippocampus, e tsebahalang ka karolo ea eona ho itšetleha ka maemo [21]. Sena se etsa hore sistimi ea DA e behoe maemong a phahameng a arabelang hofeta karabelo molemong oa ts'usumetso ha e fanoa maemong a sebetsang. Ka hona, haeba ts'okelo kapa boemo bo itseng bo hlokang ho falimeha bo le teng, vSub e tla beha sistimi ea DA boemong bo phahameng ba ts'ebetso ea boipheliso, e e fane ka ts'ebeliso e eketsehileng ho fana ka karabelo e nepahetseng khothatsong e kholo (Setšoantšo sa 1b).

Ha khatello ea khatello e bakileng ts'ebetso ea DA e tlosoa kamora moo, sistimi e bonts'a karabo e fapaneng, ketsahalo ea homeostatic e bitsoang ts'ebetso ea mohanyetsi. Ho latela khopolo ena, e hlalositsoeng pele ke R. Solomon ho 1974 [67], maikutlo ke a mabeli a bohanyetsi. Ha maikutlo, a joalo ka tšabo, a fumaneha, a tla baka maikutlo a hanyetsanang, joalo ka phomolo (kapa ho tlohela), kamora nako ea ho lula, e le ho qoba maikutlo a feteletseng kapa ho tlohela boemo bo tšosang. Ka ho tsosoa khafetsa, maikutlo a mohanyetsi a ba matla, a fokolisa boiphihlelo ba maikutlo a pele. Leha ho fanoe ka maikutlo a hore khatello ea maikutlo e matla e baka keketseho e sa khaotseng ea ho lokolloa ha DA ho NAc, liphuputso tsa morao tjena li bonts'itse hore 24 h kamora khatello ea khatello ea kelello, ts'ebetso ea baahi ba DA ho neuron ho VTA e fokotsehile haholo [68]. Ka hona, ts'ebetso ea pele ea li-neurons tsa DA nakong ea khatello ea maikutlo e lateloa ke khatello ea maikutlo ea tšebetso ea DA, ka ho tsamaisana le karabelo ea locomotamine e kenyellelitsoeng ea locomotor [68]. Liketsahalo tse joalo li ka lebelloa ho fumana maikutlo. Phello ena e boetse e hlalositsoe kamora khatello ea maikutlo a batang a sa foleng, a bakang ho fokotseha hoa ts'ebetso ea baahi ba DA ho VTA [69], e bontšang hore khatello ea maikutlo e sa foleng e baka liphetoho tsa nako e telele taolong ea tšebetso ea methapo ea methapo ea mesolimbic DA. Ka hona, ts'ebetso ea pele ea DA e eketseha nakong ea khatello ea maikutlo e rera tsamaiso ea khatello ea maikutlo nakong e tlang ha e tlohela. Mohlala, haeba khatello ea maikutlo e baka tšabo, keketseho ea ts'ebetso ea DA e tla nolofalletsa ho baleha le ho lieha ho fihlella ho ka fokolisa boiphihlelo ba maikutlo a mantlha ha khatello e ka hlaha hape. Leha khopolo ea mohanyetsi e sa le e hlalositsoe ka nako e telele, ho potoloha ha methapo e amanang le tšebetso ea DA e sa tsoa qala ho hlakisoa.

5. The basolateral amygdala

BLA ke sebopeho se nang le matla se nahanang ho hlahisa moelelo oa maikutlo ho susumetso ea tikoloho ka ho hokahanya libaka tse amanang le cortical tse amanang le maikutlo le libaka tse amehang tlhahisong ea likarabo tsa maikutlo [70]. Bohlokoa ba BLA karolong ea maikutlo ea karabelo ea khatello ea maikutlo bo se bo hlakile (ho hlahlojoa, bona [71]). Khatello e sithabetsang, e kang footshock kapa mohatla, e tsosa ts'ebetso ea amygdala [72,73], tse sa foleng le khatello ea maikutlo e matla e eketsa tšebetso ea li-neuron tsa BLA [74]. Ho feta moo, polasetiki ea synaptic ka har'a amygdala [75kapa tseleng ea amygdala-nucleus e bokellanang tseleng [76] e boetse e angoa ke khatello ea maikutlo e fapaneng. Morphologically, chronic [77]78khatello ea maikutlo e phahamisa keketseho e matla masapo a lesapo la mokokotlo le ho kobeha habonolo ha methapo ea kutlo ea BLA. Ntle le moo, khatello ea maikutlo e ka fokotsa ts'ebetso ea li-interneurons tsa GABAergic ho BLA, e fana ka tlhahiso ea ho eketsa khatello ea maikutlo ho susumetsang maikutlo [79]. BLA e boetse e amohela lipeeletso tse matla tse tsoang ho li-neuron tsa LC, tse hlahisoang ke ts'usumetso e hatellang [80,81]. Ntle le moo, ka khatello ea maikutlo e pheta-phetoang, liketso tsa NE ho amygdala li fetoha tse hlasimolohileng ka tlhaho [29].

Ka hona, BLA ke mokhethoa ea matla oa ho buella tšebelisano lipakeng tsa tsamaiso ea DA le karabelo ea ho tlosa khatello maikutlong. Ka sebele, ho thibela modadobolle ea feto-fetohang ea BLA ka ho kenella ha beta-adrenergic antagonist propranolol ho thibela ho fokotseha hoa ts'ebetso ea neuron ea VTA le ts'ehetso ea maikutlo e hlahisoang ke karabelo ea amphetamine locomotor e hlokometsoeng kamora khatello ea thibelo [68]. Li-neurons tsa BLA li romela ho hakanya likarolo tse ngata tse tsejoang ho susumetsa ts'ebetso ea methapo ea VTA DA. Ehlile ho bontšitsoe hore merero ea BLA ho VP [82] le "rostromedial tegmental nucleus [83]. Ka hona, amygdala e ka susumetsa ts'ebetso ea VTA ka marang-rang a rarahaneng. VP ke konokono ea tlhahiso e ka sehloohong e hokahanyang methapo e ka pele ho mokokotlo oa moputso oa makhopho. Boithuto ba morao-rao bo bonts'itse hore phello ea ho thetsoa ha BLA mosebetsing oa DA kamora khatello e matla ea khatello ea maikutlo (CMS) e ka hatelloa ke ho se sebetse VP [84], ho fana ka maikutlo a hore ho fokotseha ha ts'ebetso ea DA nakong ea ho tlosoa ha khatello ea maikutlo ho tsamaisoa ka tsela ea amygdala-pallidal. Ka hona, nakong ea ho ikhula khatello ea maikutlo e tsitsitseng, hyperactivity ea BLA e ne e ka thibela li-neuron tsa DA ka VP, e fokotse tšusumetso ea boits'oaro bo latelang bo amanang le boitšoaroSetšoantšo sa 1c).

Ka hona, "hralocampus" ea "ventral hippocampus" le BLA e fana ka maikutlo a tšoanang, empa litlamorao tse khahlano le tsa VTA DA tsa neuron le ho thunya ha boitšoaro ho amphetamine. Sistimi ea DA e laoloa ka mokhoa o hlakileng ka ho kopantsoe ha maikutlo a ikhethang, a maikutlo le a boits'oaro, 'me tlhahiso ea li-neurons tsa DA e fana ka karabelo e thata lits'ebetsong tsena (haholo-holo NAc) tse laolang boitšoaro bo lebisitsoeng ho sepheo. Ho khahlisang, ho bontšitsoe haufinyane hore infralimbic prefrontal cortex e fana ka taolo ea taelo mabapi le ts'ebetso ea palo ea DA ea li-neuron ho VTA [85ka BLA le hippocampus ea ventral, ho fana ka maikutlo a taolo ea liphetoho tse bakang khatello ea maikutlo mosebetsing oa DA. Ka hona, ho leka-lekana ka har'a potoloho ena ho thusa setho hore se ikamahanye le tikoloho ea sona (Setšoantšo sa 1) hape, ho fokotseha ha tekanyo ena ho ka lebisa karabong e mpe.

6. Karabelo ea khatello ea maikutlo le mathata a kelello

Joalokaha ho hlahlojoa pejana, khatello ea maikutlo e ka baka ts'ebetso e matla ea sistimi ea DA e kopantsoeng ke potoloho ea hippocampal ea potoloho, e lateloe ke karabelo ea ho khaotsa ka ho fokotseha hoa ts'ebetso ea tsamaiso ea DA e tsamaisoang ke amygdala. Leha ho le joalo, haeba karabelo ena e tloaelehileng ea khatello ea maikutlo e nka nako e telele, e ka lebisa litlamorao tsa pathological. Ka sebele, litlamorao tse bohloko tsa khatello ea maikutlo e sa feleng li ka ba maemong a kelello a mangata.

(a) Matšoenyeho le khatello ea maikutlo

Anhedonia, kapa ho lahleheloa ke thahasello kapa thabo ho liketsahalo tse tloaelehileng tse fanang ka moputso, ke letšoao le leholo la mathata a mangata a amanang le neuropsychiatric, ho kenyelletsa MDD. Leha khatello ea maikutlo e kile ea amahanngoa le serotonin ho latela liketso tsa lithethefatsi tse felisang khatello ea maikutlo, lipatlisiso tsa mantlha tsa saense li amahantse anhedonia le pherekano tsamaisong ea DA [86]. Ho joalo ka litoeba, ho ruruha hoa tsamaiso ea DA ho fella ka ho senyeha ha ts'ebetso ea moputso oa boko.87], le DA blockade e nang le li-dopamine D2 receptor antagonists e senya boemo ba moputso bo fanoang [88], ka bobeli li lebisa ho linaha tse kang anhedonic. Ho bakuli ba nang le lefu la Parkinson, tlhaiso-leseling ea litlhare e fana ka maikutlo a hore kantle le tahlehelo ea sistimi ea DA ea nigrostriatal, ho senyeha ha sistimi ea DA ea mesolimbic le eona ea etsahala [89], e ka lebisang ho anhedonia [90]. Ho feta moo, kalafo ea lefu la Parkinson le li-receptor tsa DA tse kang pramipexole kapa l-DOPA e kokobetsa matšoao a anhedonia [87]. Sena se lumellana le mefuta ea liphoofolo e tšehetsang hore anhedonia ke bothata ba lefu la Parkinson [87]. Sisteme ea DA e amahanngoa le ho bolela moputso [9le tse susumetsang [91], 'me ho nahanoa hore DA e ea hlokahala bakeng sa sepheo sa tšusumetso ea ts'usumetso, ho fetola pono ea moputso molemong oa khothatso [10]. Sena se tsamaisana le ho sitisoa ha sepheo sa ho batla liphihlelo tse monate tse hlalositsoeng ho batho ba fumanoeng ba na le MDD [92].

Moetso oa CMS o sa lebelloang, o ntlafalitsoeng ke Paul Willner le basebetsi-mmoho ba li-1980s [93], ke mohlala oa phoofolo e netefalitsoeng ea khatello ea maikutlo ea batho [94]. Moetso ona o na le ho pepesehela khafetsa mefuteng e fapaneng ea khatello ea maikutlo e fapaneng le e sa reroang ka nako e tsitsitseng, e etsa hore ho be le matšoenyeho, ho felloa ke tšepo le anhedonia ka litoeba (bakeng sa tlhahlobo, bona [95]). Ts'ebetso ea li-neuron tsa VTA DA litšoeneng tse senotsoeng ke CMS e ka lopolla likhoto ka mokhoa ona o sithabetsang maikutlo o kang oa khatello ea maikutlo [96], haholoholo ka ha ho lekotsoe ka khetho ea sucrose, e leng sesupo se tsebahalang sa anhedonia ka litoeba [97]. Ho feta moo, thibelo ea li-neurons tsa VTA DA e ka baka mekhoa e mengata ea boits'oaro e kang ea khatello ea maikutlo [96]. Re sebelisa mofuta o le mong, laboratori ea rona e bonts'itse hore likhoto tse pepesitsoeng ho CMS li bonts'a keketseho ea boits'oaro boits'oarong ba boitšoaro bo qobelloang ba ho sesa, le ho fokotseha ho hoholo hoa ts'ebetso ea palo ea baahi ba DA [84]. Phokotso ena e khutlisoa ka ho nolofaletsa BLA kapa VP. Ho feta moo, ts'ebetso ea BLA ka har'a likhoto tse se nang khatello ea maikutlo e fokotsa ts'ebetso ea baahi ba li-neuron tsa DA, tse ts'oanang le tse hlalositsoeng ho litoeba tsa CMS-mmuso o boetse o khutlisoa ka ho thibela VP [84]. Sena se fana ka maikutlo a hore maemong a sithabetsang, ho fokotseha ha tšebetso ea DA ho kopantsoe le tsela ea BLA-VP. Re lokela ho hlokomela hore li-neuron tsa VP li amohela likhakanyo tsa mono- le polysynaptic tse tsoang ho BLA [98], ka hona phello ea li-neuron tsa DA e ka ba e tobileng kapa e kenyelletsang libaka tse pakeng tsa glutamatergic. BLA e amohela lipehelo tse matla tse tsoang ho infralimbic PFC (ilPFC). IlPFC ke leqephe la litoeba tsa tikoloho ea Brodmann (BA) 25, e leng sebaka se tsejoang hore se ka kenngoa bathong ka masoabi a maholo [99] ebile e na le khatello ea maikutlo maemong a sithabetsang [99]. Ka sebele, ho kenella ka mokhoa oa phekolo o sebetsang ho phekola MDD ho batho ho khutlisetsa ts'ebetso ho BA25 [100]. Re fumane hore ho litoeba tsa CMS, inactivation inactivation ea ilPFC e hlahisa ts'ebetso ea disinhibition tsamaisong ea DA, e khutlisetsa ts'ebetso ea bongata ba VTA DA boemong bo ka bapisoang le liphoofolo tse se nang khatello ea maikutlo [101]. Hona joale ho shebiloe ka kakaretso hore MDD ke ts'oaetso ea boemo ba tsamaiso e amang litsela tse kopaneng tse hokahanyang li-kuchagua cortical, subcortical and limbic [102,103]. Ka hona, hyperacaction ea ilPFC / BA25, e bonts'itsoeng e le hyperactive ho MDD ho batho [104] le litoebung tsa CMS, e kenya ts'ilafalo ea ts'ebetso ea methapo ea VTA DA ka tsela ea BLA-VP (Setšoantšo sa 2).

Setšoantšo sa 2. 

Mokhoa o fapaneng oa tsamaiso ea dopamine. (a) Maemong a tloaelehileng, teka-tekano lipakeng tsa ilPFC → BLA → VP le vSub → NAc → potoloho ea VP e lebisa ho halofo ea li-neurons tsa DA li thunya hanghang. (b) Hyperacaction ea ilPFC mathateng a maholo a sithabetsang (MDD) e lebisa ts'ebetsong ea BLA, e ntlafatsa ts'usumetso ea VP sebakeng se phetheselang sa moea, e bakang ho se leka-lekane lipakeng tsa lipotoloho tse peli, 'me sephetho sa eona e bile ho fokotseha ho hoholo ha ho sa thibeloang Ts'ebetso ea sechaba ea DA, e reriloeng ho anhedonia.

Ka hona, ho na le ho leka-lekana lipakeng tsa mekotla e 'meli e arohaneng e sebetsang kapa e amohelang karabelo ea DA ea neuron: ilPFC → BLA → VP e thibela ts'ebetso ea DA [84,101], athe vSub → NAc → VP e buella tsamaiso ea DA [54,55]. Ho bontšitsoe hore khatello ea maikutlo e matla (footshocks) le khafetsa / e tsitsitseng (restent) e kenya letsoho vSub [24,25,62], e bakang keketseho ea tšebetso ea palo ea baahi ba DA [62]. Liphetho tsa khatello ea maikutlo tsamaisong ea DA ka hona li ka amana le mofuta oa eona o boletsoeng esale pele kapa o sa lebelloang [105,106]. CMS ke ts'ebetso e sa lebelloang eo kamehla e lebisang boits'oarong le boikutlong bo joalo ka boits'oaro [107], empa CMS e boletsoeng esale pele joalo ka khatello ea maikutlo ea ho itšireletsa khafetsa ka nako e telele e lebisa ts'ebetsong ea hippocampal, mohlomong ka NE [68], eo ka nako e 'ngoe e sebetsanang le ho thunya ha neuron ea DA. Ho feta moo, liphoofolo li bonts'a sebaka se ikhethileng bakeng sa taba e neng e entsoe pele ka tšisinyeho e lebisitsoeng ho e mong le e mong ea kopantsoeng le ba tšoanang ba sa lebelloang [108]. Ho batho, ho sitoa ho laola ketsahalo e sithabetsang ho kentse khatello ea maikutlo. Ho liphoofolo, khatello ea maikutlo e ke keng ea qojoa e tsebahala e ka etsa hore motho a ithute ho hloka thuso, 'me ke mohlala o ikhethileng oa khatello ea maikutlo. Hoo e ka bang halofo ea liphoofolo tse nang le khatello ea maikutlo e sa laoleheng e ithutile ho hloka thuso (ke hore, tekanyo e fokotsehileng ea ho baleha khatello ea maikutlo e tlang)109]. Halofo e 'ngoe, e sa bontsheng ho hloka thuso, e kanna ea ba le liphetoho tse fapaneng tse ba sirelletsang liphellong tse mpe tsa khatello ea maikutlo e ke keng ea qojoa. Ho bonts'itsoe haufinyane hore likhoto tse bonts'ang ho hloka thuso li na le ho fokotseha hoa ts'ebetso ea baahi ba VTA DA, haholo literekeng tsa kalafo, athe liphoofolo tse sa thuseng letho li na le ts'ebetso ea DA e bapisoang le liphoofolo tse laoloang le ha li fumana palo e ts'oanang ea litšeho [110]. Sena se lumellana le phuputso e fetileng e bonts'ang hore khatello ea maikutlo e sa foleng e fokotsa ts'ebetso ea baahi ba li-neuron litseleng tsa methapo le tse bohareng tsa VTA [111]. Ho feta moo, liphoofolo tse bonts'ang mokhoa oa ho hloka thuso li bonts'a ho fokotseha ha dopamine ka har'a motsoako oa caudate le li-nucleus tsa caudate, tse lumellanang le ts'ebetso ea dopamine e fetisitsoeng ea tsela ea mesolimbic mohlaleng ona [112]. Mokhoa o ithutileng ho hloka thuso, tšitiso ea polasetiki ea synaptic tseleng ea vSub-NAc, e itšetlehileng ka DA [113,114], e fana ka maikutlo a hore ka likhatello tse sa lebelloang, vSub → NAc → potoloho ea VP e bonts'oa, ha e etsoa ka li-stress tse ka bolelang. Ho feta moo, ketamine, mohanyetsi oa NMDA le buka e nkang maikutlo e nkang khatello kapele haholo [115], e bontšitsoe hore e khutlisa phokotso mosebetsing oa sistimi ea DA le ho khutlisetsa LTP tseleng ea vSub-NAc [110], ho lumeloang ka ho kenella ka potlako liprotheine tsa synaptic ho hippocampus [116], le ho khutlisetsa phokotso e bakiloeng ke khatello ea maikutlo mokokotlong [117].

(b) Matšoenyeho le bokhoba ba ho lemalla

Ho na le bopaki bo bongata ba hore mokhatlo o bohlokoa o teng lipakeng tsa khatello ea maikutlo e matla kapa e sa foleng le tšusumetso ea ho sebelisa lintho tse lemalloang ka tlhekefetso (bakeng sa tlhahlobo, bona [4,118]). Keketseho e sa feleng ea mosebetsi oa axis ea HPA e tlalehiloe ka tšebeliso e sa feleng ea li-psychostimulants tse kang cocaine [119], e sebelisang sisteme ea mesolimbic DA. Ka sebele, tsamaiso e matla ea li-psychostimulants tse kang cocaine kapa amphetamine e eketsa mesolimbic DA [120], empa tšebeliso e sa foleng le ho iketla ho matla ha setla-morao ho nyenyefatsa tsela ea DA ea mesolimbic, e lebisang ho fokotseha hoa maemo a DA a basal [121]. Ho batho, lithuto tse inahaneloang li bonts'itse hore ho na le phokotso ea phetisetso ea DA ho "ventral striatum" le "cortex" e tlang pele ho basebelisi ba "cocaine" nakong ea ts'ohanyetso le nako e telele [122], le litšitiso tsa polasetiki ea synaptic ho VTA le NAc li bontšitsoe kamora taolo ea koae e mpe le e sa foleng [123,124]. Ts'ebetso e matla ea li-psychostimulants hape e kenya ts'ebetsong lits'ebetso tsa khatello ea kelello joalo ka axis ea HPA, e hlahisang keketseho ea plasma ACTH le corticosterone [125]. Ho teneha, ho tšoenyeha le khatello ea maikutlo li tloaelehile nakong ea ho itima li-psychostimulants [126]. Boemo bona bo fosahetseng bo hlalositsoe kamora tšebeliso e sa foleng, le tšitiso e tšoanang karabong ea khatello ea maikutlo le lits'ebetso tsa DA (bakeng sa tlhahlobo, bona [127]). Ho latela mohato oa mohanyetsi mohopolo oa tšusumetso [128], maikutlo a matle a hlahisoang ke matla a matlafatsang a lithethefatsi a lateloa ke lits'ebetso tsa sekhahla se kopaneng sa anhedonic se fapaneng le tlhaho le nako e telele ha se bapisoa le maikutlo a pele. Ka hona, kamora ts'ebeliso e sa feleng, setlamo se seng se sebe se lula se le teng nakong ea ho ikhula, se siea motho ea lemaletseng a le maemong a hlobaetsang (bakeng sa ho lekola, bona [129]), e eketsa menyetla ea ho khutla le ka mor'a nako ea nako e telele. Boemo bona bo seng botle bo bonoe feela kamora ts'ebeliso e mpe ea lithethefatsi; lithuto tse 'maloa li lebisitse tlhokomelo molemong oa ho khaotsa ho se sebetse kamora ho entetsoa ka lithethefatsi haholo. Leha ho le joalo, haufinyane tjena ho bonts'itsoe hore ente ea amphetamine e hlohlelletsang ho fokotseha hoa morao hoa ts'ebetso ea palo ea DA ea li-neuron 18 h kamora ho ente, ka phokotso ena e tšoarellang ho fihlela 72 h [130]. Ho feta moo, tsamaiso ea ketamine kapa BLA inactivation e khutlisa ho fokotseha hoa tšebetso ea methapo ea DA [130], ho fana ka tlhahiso ea hore hyperactivity ho ilPFC → BLA → potoloho ea VP, joalo ka ha e bonoa mehlaleng ea MDD, e kanna ea ikarabella bakeng sa boemo bo bobe ba ho tlohela bo eketsang kotsi ea ho khutla le kamora ho entoa ka matla ha psychostimulant. Ka hona, nako ea puso e mpe nakong ea ha e tlohela e ka amana le nako ea ts'ebetso ea ts'ebetso ea DA. Ka sebele, liphetho tsena li fana ka maikutlo a hore tšebeliso e mpe ea lithethefatsi e lokela ho baka kotsi ea ho khutla hape, mohlomong ka lebaka la ho fokotseha ha nako e telele ketsahalong ea methapo ea DA kamora ho khaotsa, athe ts'ebeliso e mpe ea lithethefatsi, e boetse e etsa hore mosebetsi oa neuron oa DA o fokotsehe, Ho itokolla ka bokhuts'oanyane empa mohlomong ho tla khannela motho ea se nang lithethefatsi lithethefatsi hore a nke litekanyetso tse ling tsa lithethefatsi.

Ho utloisisa ka nepo hore na tsamaiso ea lithethefatsi e ka baka boemo bo bobe joang le ho eketsa kotsi ea ho khutla hape ka lithethefatsi le boits'oaro ba ho batla lithethefatsi, ho bohlokoa haholo ho utloisisa litšitiso tsa pathophysiological makaleng a fapaneng a feto-fetohang tsamaisong ea DA, khatello ea kelello le litsela tsa BLA le khatello ea maikutlo. -VSub tsamaiso ea potoloho.

7. Qetello

Sisteme ea DA e ka bapala likarolo tse ikhethileng khatellong ea kelello le mafu a kelello. Ka sebele, khatello ea khatello ea kelello e fokolisang khatello ea kelello sebakeng sa ventral hippocampus e ka kenya tšebetsong tšebetso ea methapo ea VTA DA ka VSub → NAc thibelo ea VP, e lebisang ho phatlalatseng methapo ea methapo ea VTA DA. Leha ho le joalo, kamora ho tlosoa ha khatello ea maikutlo, ho na le phokotso ea nako e telele ea ts'ebetso ea methapo ea DA ka ho ipata ka ilPFC-BLA ea VP, e lebisang ho fihleng ha tšebetso ea methapo ea DA. Ka hona, le ha tsamaiso ea DA e ka theha motheo oa maloetse a mangata a kelello, ts'ebetso ea methapo ea methapo e ka hlaha mehahong e kopaneng e sa sebetseng li-neuron tsa DA. Ka hona, tsepamiso ea maikutlo mabapi le ho khutlisetsa ts'ebetso e tloaelehileng libakeng tsena e kanna ea sebetsa hantle ho feta ho theola tsamaiso ea DA ka kotloloho.

Polelo ea chelete

Mosebetsi ona o tšehelitsoe ke Moputso oa Mofumahatsana oa Tlhahlobo o tsoang NARSAD — The Brain and Behahlangu Research Foundation (PB) le lithuso tsa lithuso tsa litšebeletso tsa bophelo tsa United States tsa United States. MH057440, MH101180 le DA036328 (AAG)

Thahasello e khahlisang

Ngaka Belujon ha e tlalehe lithahasello tsa tlholisano. Ngaka Mohau o amohetse chelete ho Johnson le Johnson, Lundbeck, Pfizer, GSK, Puretech Ventures, Merck, Takeda, Dainippon Sumitomo, Otsuka, Lilly, Roche le Asubio.

  • Re amohetse October 13, 2014.
  • E amohetse ka February 23, 2015.

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