Tsela ea Neural ea khatello ea kelello Botsoako: Bohlokoa ba Tlhekefetso ea Joala, James P. Herman, Ph.D.

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James P. Herman, Ph.D. ea bophelo bo botle., ke moprofesa Lefapheng la Psychiatry and Behavioral Neuroscience, Univesithi ea Cincinnati, Cincinnati, Ohio.

Khatello ea maikutlo ke karolo ea bohlokoa ntlafatsong, tokisetsong le ho khutliseng boits'oaro bo lemaletseng, ho kenyelletsa ts'ebeliso ea joala. Sengoloa sena se lekola boemo ba hajoale ba lingoliloeng tse mabapi le karabelo ea khatello ea kelello, e tsepamisang maikutlo ho hypothalamic-pituitary- adrenal (HPA) axis. Likarabo tsa khatello ea maikutlo li ka hlaha e le karabelo ho phephetso kapa tšabo; lipontšo tse tsoang likarolong tse ngata tsa bokong li romela ho kenella ka har'a patricricular nucleus (PVN) ka har'a hypothalamus. Leha ho le joalo, likarabo li boetse li hlaha ho khatello ea maikutlo e bolelang lintho tse ka behang kotsing (khatello ea kelello). Likarabo tsa Psychogenic li kopantsoe le letoto la likhokahano tsa lisele tsa methapo tseleng ea limbic-PVN, li-circuits tsa amygdalar le infralimbic cortex li bonts'ang li-excation le prelimbic cortex le hippocampal neurons tse bontšang khatello ea maikutlo. Linkic-PVN likhokahano li fetisoa ke li-neuron tsa GABAergic hangata libakeng tse kang mookotaba oa sebaka sa stria terminalis le sebaka sa preoptic. Matšoenyeho a sa feleng a ama sebopeho le tšebetso ea khatello ea methapo ea maoto le ho baka pherekano e ntlafalitsoeng ea PVN, leha mohato o nepahetseng o sa tsejoe. Habohlokoa, ho pepeseha joala bo sa foleng le bo sa foleng ho tsejoa ho ama mekhoa le khatello ea maikutlo mme li ka hokahana le khatello ea maikutlo ka ho hlahisa khatello ea maikutlo joalo ka phetoho ea amygdalar le likarolo tsa pele tsa netweke ea taolo ea khatello ea methapo.

mantsoe a sehlooho: Tlatsetso; joala le litloaelo tse ling tsa ho batla lithethefatsi; tšebeliso e mpe ea joala le tlhekefetso; khatello ea maikutlo; khatello ea maikutlo; khatello ea maikutlo e sa foleng; ho kopanya khatello ea maikutlo; karabelo ea mmele ho khatello ea maikutlo; likarabo tsa khatello ea kelello; boko; litsela tsa neural; limbic-paraventricular tsela; likhoele tsa taolo ea khatello ea maoto; hypothalamic-pituitary- adrenal axis; kakaretso ea lingoliloeng

Ho feto-fetoha le maemo ha ho le tsietsing ea 'mele kapa ea kelello ho hlokahala bakeng sa ho pholoha, bophelo bo botle le boiketlo ba lintho tsohle. Liketsahalo tse mpe, tse atisang ho hlalosoa e le "khatello ea maikutlo", li qala karabelo e fapaneng ea 'mele ho tsoa mehloling e mengata, ho kenyelletsa ts'ebetso ea hypothalamic-pituitary- adrenal (HPA) axis.¹ Mokokotlo oa HPA o ikarabella bakeng sa karolo ea glucocorticoid ea karabelo ea khatello ea maikutlo (ke hore, karabelo ea lihormone tsa steroid; cortisol ho batho, corticosterone ho litoeba le likhoto). Secretion ea Glucocorticoid ho nahanoa hore e kenya letsoho ho feto-fetoha ha khatello ea maikutlo ka ho baka liphetoho tsa nako e telele ho polelo ea gene ka li-receptor tsa crenate adrenocorticosteroid (ke hore, mineralocorticoid receptor [MR] le glucocorticoid receptor [GR]. Li-receptors tsa adrenocorticosteroid li sebetsa e le lintlha tsa ligand-gated transcript (De Kloet et al. 1998) empa e ka boela ea fetola mengolo ka ho kena-kenana le balaoli ba bang ba ho ngola, joalo ka nuclear factor-kB (NF-kB) le activator protein-1 (AP-1 ) (Webster le Cidlowski 1999). Glucocorticoids le eona e ka ba le litlamorao tse potlakileng k'hemistri ea boko le boits'oaro ka methapo ea matsoho e sa sebetseng (De Kloet et al. 2008). Li-Glucocorticoids ho nahanoa hore li kenya letsoho ho feliseng karabelo ea pele ea khatello ea maikutlo (Keller-Wood le Dallman 1984) le ho nka karolo ho khutliseng nako e telele ea homeostasis e bakiloeng ke karabelo ea pele (Munck et al. 1984).

¹ Bakeng sa litlhaloso tsa mantsoe ana le a mang a mahlale, bona Glossary, maq. 522-524.

Likarabo tsa khatello ea maikutlo tsa Glucocorticoid li ka qalisoa ke litšebeletso tsa 'mele (tse bontšang likarabo tse hlakileng) kapa lits'ebetso tsa boko tse hokahanyang likhakanyo tsa tikoloho le litlamorao tse mpe. Karabo ea morao-rao e bitsoang "psychogenic" e lebelletse tlholeho mme e kenyelletsa litsela tsa boko tse ikarabellang bakeng sa mananeo a ts'ireletso ea kahare kapa ho hopola liketsahalo tse khathollang (Herman et al. 2003). Ka hona, karabelo ea psychogenic e amana le boiphihlelo ba pele, mme e etselitsoe ho matlafatsa mmele ka sepheo sa ho qoba sephetho se sebe kapa ho etsa boits'oaro bo ka ntlafatsang monyetla oa ho pholoha.

Bopaki bo bongata bo bontša hore lits'ebetso tsa khatello ea maikutlo li bapala karolo e kholo lits'ebetsong tse lemalloang, ho kenyelletsa ho lemalla joala. Mohlala, ho pepeseha khatello ea maikutlo ho ka fokotsa ho khutla kapa ho eketsa tšebeliso ea joala (Sinha 2007). Ketso ea khatello ea maikutlo / glucocorticoids ho joala e ka hokahanngoa le phetisetso ea moputso / khatello ea khatello ea maikutlo, ho kenyelletsa, mohlala, ntlafatso ea tokollo ea dopamine ka har'a li-nucleus accumbens (Sutoo le Akiyama 2002; Yavich le Tiihonen 2000) le ts'ebetso ea ts'ireletso ea corticotropin e bohareng. factor (CRF) pathways (Heilig le Koob 2007). Haholo-holo, khokahano lipakeng tsa tahi ea joala le khatello ea maikutlo li thatafatsoa ke taba ea hore ho pepesetsoa joala, joalo ka lithethefatsi tse ngata tsa tlhekefetso, ho baka ho lokolloa hoa li-glucocorticoids mme ka hona ho ka khethoa e le "khatello ea maikutlo" ea mefuta ea bona (bona Allen et al . 2011).

Sengoloa sena se lekola mokhatlo o hlophisitsoeng oa li-neurocircuits tse laolang likarabo tsa khatello ea maikutlo, o tsepamisa maikutlo ho axPA ea HPA, e amanang haholo le lits'ebetso tsa ho kenya letsoho (bona Marinelli le Piazza 2002). E boetse e bua ka likarolo tsa karohano lipakeng tsa khatello ea maikutlo le litsela tsa moputso, kaha tsena li bohlokoa molemong oa ho loants'a litlamorao tsa khatello ea tlhekefetso ea lithethefatsi le ho lemalla joala.

Ho Potoloha Ho Arabela Karabo ea Khatello ea Khatello ea Kelello

Mokelikeli oa HPA o laoloa ke li-neurons ka har'a "solarricular nucleus" (PVN) ho hypothalamus (bona setšoantšo 1). Li-neurons tsena tse bolotsana tsa CRF le vasopressin ea lihormone ho potoloha ho potoloha, e leng hona li baka ho ntšoa ha hormone ea adrenocorticotropin (ACTH) ho tsoa mokokotlong oa anterior pituitary. ACTH e tsamaea ka potoloho ea systemic ho fihla ho adrenal cortex, moo li-glucocorticoids li entsoeng le ho lokolloa (bona Herman et al. 2003).

Likarabo tse ntlafatsang tsa khatello ea maikutlo li etsahala nakong ea maemo a tšohanyetso (mohlala, ts'oaetso, tlala, ho felloa ke metsi kapa ho tsitsinyeha), ha boko bo tlameha ho arabela phephetsong e kholo ea homeostasis ka ho tsamaisa axis ea HPA. Tlhahisoleseling ea sensens e fetisetsoa ho PVN ke li-neuron tsa pele kapa tsa bobeli, e hlahisang ts'ebetso e tobileng ea tokollo ea CRF (bona Herman et al. 2003). Mohlala, khatello e tlase ea mali e amanang le tahlehelo ea mali e fetisoa ka methapo ea kutlo ho li-neurology tsa methapo ea methapo sehlopheng sa lisele tsa A2 catecholaminergic cell group (Palkovits le Zaborszky 1977), eo ka nako eo e kenang ka kotlolloho ho PVN (Cunningham le Sawchenko 1988) 'me e potlakise tšebetso ea Noradrenergic activation of Li-neurons tsa CRF (Plotsky et al. 1989).

Ntle le litselana tsa neural, tlhaiso-leseling ka liphetoho tsa mmele le eona e ka fetisoa ka lintho tse potolohang tse amanang le libaka tse kantle ho mokoallo oa mali le bokong. Mohlala, keketseho ea pherekano ho li-angiotensin II (ho bontša ho felloa ke metsi 'meleng) e utloahala ke li-receptors tse ka hare ho setho sa subfornical (se ka ntle ho methapo ea mali le tsamaiso ea mokelikeli oa mali), se romellang ka kotlolloho li-angiotensin II ka kotloloho ho li-neuron tsa PVN CRF, ho thusa ho kenya letsoho ha HPA (Plotsky et al. 1988). Lintho tse ling tse fokolisang, tse kang ho ruruha, li hlahisa lintho tse ka supang ka mekhoa e mengata; mohlala, proinflammatory cytokine interleukin 1-b e bonahala e kenya tšebetsong axis ea HPA ka methapo ea kutlo ea methapo methapong ea methapo ea kutlo; sebaka sa postrema, se kantle ho tšitiso ea mali le boko; le lisele tse potolohileng tikolohong ea sehlopha sa lisele tsa A2 (Nokia et al. 1997; Lee et al. 1998; Wieczorek le Dunn 2006).

Lithethefatsi tsa tlhekefetso le tsona li ka hlahisa karabelo ea pele ea corticosterone ka mekhoa ea projeke ea brainstem PVN. Mohlala, ho pepesetsoa joala ho etsa hore ho nkoe ACTH le corticosterone, e tsamaellanang le joala bo etsang joalo ka tšusumetso e se nang boikaketsi (Allen et al. 2011). Acation HPA axis activation ke joala e kenella pakeng tsa brainstem noradrenergic system (Allen et al. 2011). Leha ho le joalo, ho pepesetsoa joala ho sa feleng ho etsa hore ts'ebetso ea HPA e sebetse haholo ho pepesehang joala (Rivier 1995), e fanang ka maikutlo a hore, ho isa bohōleng bo itseng, e lebise litlamorao tse matla tsa ts'ebeliso ea lino tse tahang ka HPA.

Likarabo tsa ho Pholosa Likhatello tsa Khatello ea Kelello:

Hobane "maemo a tšohanyetso" a 'nete "ha a na seoelo, likarabo tse ngata tsa khatello ea maikutlo li lebelletse tlholeho, li kenyelletsa ho hlalosoa ha ts'okelo ea ts'usumetso ea tikoloho mabapi le boiphihlelo bo fetileng kapa mananeo a tlhaho. Likarabo tsa khatello ea maikutlo tse lebelletsoeng li laoloa haholo ke meetso ea li-limbic forebrain, joalo ka hippocampus, medial prefrontal cortex (mPFC), le amygdala (bona Ulrich-Lai le Herman 2009). Likaho tsena kaofela li fumana leseli le sebetsanang le maikutlo a sebetsang mme le karolo ea taolo ea maikutlo, moputso le maikutlo.

Lithuto tsa brain le lithuto tse susumetsang li bonts'a hore hippocampus e thibela axis ea HPA. Matla a motlakase a hippocampus a fokotseha ho lokolloa ha glucocorticoid ho likhoto le batho. Tšenyo ea hippocampus, kapa methapo e tsamaeang le eona (ke hore, lateral fornix), e baka likarabo tse fetelletseng tsa khatello ea kelello (mohlala, ho ithiba) le ho iponahatsa e le khutlelo ea nako e telele mehatong ea basel ea glucocorticoid (bakeng sa lipontšo tsa mantlha, bona Herman et al. . 2003; Jacobson le Sapolsky 1991). Boitsebiso bo bong bo bontša hore hippocampus e boetse e thibela tšebetso ea axis ea HPA, empa sena ha se bonoe hohle (Herman et al. 2003; Jacobson le Sapolsky 1991). Litlamorao tsa tšenyo ea hippocampal ho likarabo tsa khatello ea kelello tsa HPA axis li ka hlahisoa sebakeng sa subralulum (vSUB), tlhahiso e ka sehloohong ea "subral" hippocampus (Herman et al. 2003). Matsoalo a senotsoeng a vSUB ho likhoto a ntlafatsa peptide ea PVN CRF le polelo ea mRNA le ho eketsa tokollo ea corticosterone le ts'ebetso ea PVN (joalo ka ha ho laoloa ke ho kenyelletsa polelo ea FOS mRNA) ha ho arabeloa thibelo (Herman et al. 1998).

Kameho ea vSUB ho melawana ea khatello ea maikutlo e hlakile. Manonyeletso a vSUB a lelefatsa likarabo tsa HPA axis empa li sa ame likarabo tse hlakileng (mohlala, ho inhalation) (Herman et al. 1998). Bopaki bo bong bo fana ka maikutlo a hore li-glucocorticoids li bapala karolo ea thibelo ea hippocampal ea likarabo tse lebelletsoeng, joalo ka ha liso li ka thibela thibelo ea maikutlo ea HPA axis ka syntheteng ea steroid dexamethasone (Magarinos et al. 1987). Ntle le moo, litoeba tse nang le li-forebrain GR Deletions, ho kenyelletsa le hippocampus, li fetisitse likarabo tsa thibelo le botaki (empa eseng hypoxia) le khatello e mpe ea dexamethasone ea tokollo ea corticosterone (Boyle et al. 2005; Furay et al. 2008). Ka bobeli, tlhaiso-leseling e bonts'a hore hippocampus e sebetsa ka kotloloho taolo ea likarabo ho khatello ea kelello, ho boloka karolo ea eona ts'ebetsong ea kelello le maikutlo.

Ho fapana le hippocampus, amygdala e amahanngoa le ho hlohlona ha axis ea HPA. Ho hlohlelletsoa ha Amygdalar ho ntšetsa pele ho lokolloa ha glucocorticoid, athe liso tse kholo tsa amygdaloid tata li fokotsa mosebetsi oa axis oa HPA (bona Herman et al. 2003). Leha ho le joalo, ho na le mofuta o ikhethang o ikhethang oa mesebetsi e amanang le khatello ea maikutlo ka har'a amygdala. Mookotaba oa bohareng ba amygdala (CeA) o arabela haholo ho khatello ea maikutlo ea lapeng, joalo ka ho ruruha le ho lahleheloa ke mali (Dayas et al. 2001; Sawchenko et al. 2000). Lesions ea CeA e lumellana le likarabo tsa axPA tsa HPA ho mefuta ena ea lintho tse susumetsang empa eseng ho ithiba (Dayas et al. 1999; Prewitt le Herman 1997; Xu et al. 1999). Ho fapana le hoo, medial nucleus ea amygdala (MeA) e bonts'a likarabo tse ikhethileng tsa FOS ho susumetsa, joalo ka ho thibela (Dayas et al. 2001; Sawchenko et al. 2000). Lesions ea MeA e fokotsa likarabo tsa axis tsa HPA ho thibela le ho ts'oara le ho ts'oaroa ha molumo empa eseng ho kenngoa ka mokhoa o hlophisehileng oa protheine ea interleukin 1-b kapa ether inhalation (Dayas et al. 1999; Feldman et al. 1994). Ka hona, ho bonahala eka likarabo tse hlakileng le tse lebelletseng li ka laoloa ka karolo ke discrete amygdaloid circry.

MPFC e bonahala e na le karolo e rarahaneng taolong ea khatello ea maikutlo. Likarohano tsohle tsa PFC e makatsang li hlahisoa ka matla ke khatello ea maikutlo e matla. Leha ho le joalo, litlamorao tsa 'mele tsa ts'ebetso ea khatello ea maikutlo li bonahala li fapana ka libaka. Karohano ea prelimbic ea mPFC (PL) e bohlokoa molemong oa khatello ea maikutlo hobane liphuputso tse ngata li bontšitse hore tšenyo tikolohong ena e eketsa likarabelo tsa axel tsa HPA ho li-stress tsa pherekano e mpe (empa e se homeostatic) (Diorio et al. 1993; Figueiredo et al. 2003; Radley et al. 2006), athe tlhoko ea ts'usumetso e thibela likarabo tsa khatello ea maikutlo (Jones et al. 2011). MPFC e bonahala e le sebaka sa tlhahiso ea glucocorticoid ea likarabo tsa HPA hobane li-glucocorticoid tse kenngoeng ka hare li thibela likarabo tse lebelletsoeng (empa eseng tse hlakileng) likarabo tsa khatello ea maikutlo (Akana et al. 2001; Diorio et al. 1993). Ka lehlakoreng le leng, liso tse lebisitsoeng ho "ventral infralimbic PFC" (IL) li na le phello e fapaneng ea 'mele. Tšenyo ea IL e fokotsa likarabo tsa boikemelo ba khatello ea maikutlo (tavares et al. 2009) hape e amohela ts'ebetso ea PVN Fos ho arabela thibelo (Radley et al. 2006). Kahoo, PL le IL e bonahala e na le litlamorao tse hanyetsanang mabapi le khatello ea maikutlo.

Ho sebetsana le phetisetso: Limbic-PVN Networks

Ho hlohlelletsa PVN ke hippocampus, preortal cortex, le amygdala ha ho na moeli. Ka hona, taolo ea tlhahiso ea li-axis tsa HPA ka meaho ena e hloka li-synapses tsa mahareng (bona setšoantšo 2). Boithuto bo fumanehang ho tloha karolong e 'ngoe ea boko ho ea ho e' ngoe (ke hore, lithuto tsa ho lekola tšupa-tsela) li senola monyetla oa likhakanyo tsa li-bisynaptic limbic-PVN tse parolang libaka tse 'maloa tsa subcortical, ho kenyelletsa le mookotaba oa bethe ea stria terminalis (BNST), dorsomedial hypothalamus , sebaka sa medial preoptic, le sebaka sa peri-PVN (ho kenyelletsa subparaventricular nucleus) (Cullinan et al. 1993; Prewitt le Herman 1998; Vertes 2004). Boithuto bo habeli bo bonts'a hore methapo e tsamaisang litšusumetso tse hole le vSUB, MeA, le CeA (ke hore, methapo ea kutlo) ikopanya ka kotloloho le li-neuron tsa projeke tsa PVN libakeng tsena, tse tsamaellanang le likhokahanyo tse sebetsang (Cullinan et al. 1993; Prewitt le Herman 1998) .

Litlamorao tse fapaneng tsa PL le IL lits'ebetsong tsa khatello ea maikutlo li ka bonts'a phokotso ea tsona tse hlakileng lipakaneng tsa subcortical. PL e na le lipakanyo tse kholo tsa ho tsamaisa meputso e amanang le moputso, ho kenyelletsa le li-bokono tsa li-nucleus le li-amygdala tsa basolateral, hammoho le posterior BNST, e hokahaneng le thibelo ea axis ea HPA. Ka lehlakoreng le leng, IL e na le likhokahano tse matla le libaka tse kenyellelitsoeng ho tsamaiso ea boemeli, ho kenyelletsa le CeA, senotlolo sa pampitšana e le mong (NTS), anteroventral BNST, le dorsomedial hypothalamus (Vertes 2004). Ka hona, ho ka etsahala hore phello e matla ea ts'ebetso ea khatello ea khatello ea kelello ea PFC e hloka ho ikopanya ho tlisoang ke PL le IL ntle.

Ehlile, mPFC, hippocampal, le amygdalar efferents li tloaetse ho tsepamisoa libakeng tse romellang merero ea γ-aminobutyric acid (GABA) ho PVN (bona setšoantšo 2). Ho joalo, palo e kholo ea li-neuron tse kenang kahare ho PVN tse sebetsanang le li-GABAergic li phenotype. Li-neurons tsa projeke tsa vSUB (hammoho le mPFC) li benya ka tlhaho, ka hona li fana ka maikutlo a hore lisele tsena li kenella phetisong ea transsynaptic ea PVN e latelang ts'ebetsong ka khatello ea maikutlo. Ka lehlakoreng le leng, li-neurons tsa projeke tsa MeA le CeA ke GABAergic, ho fana ka maikutlo a hore boithabiso ba amygdalar ba PVN bo kopantsoe le disinhibition, ho kenyelletsa li-synapses tsa GABA tse latellanang (Herman et al. 2003).

BNST e khahla haholo, hobane e amohela litlatsetso tse tsoang metsong eohle e meholo ea khatello ea maikutlo (CeA, MeA, vSUB, IL, le PL) (Cullinan et al. 1993; Dong et al. 2001; Vertes 2004) . Bonneteng, likarolo tse fapaneng tsa BNST li bonahala li ikarabella bakeng sa ho thibela khatello ea maikutlo le khatello ea kelello ea HPA. Mohlala, liso tsa sebaka se ka morao sa medial ea BNST li eketsa boholo ba ACTH le tokollo ea corticosterone le ts'ebetso ea PVN Fos (Choi et al. 2007), e bolelang karolo ea karolo e bohareng ea ho kopanya khatello ea maikutlo. Lesions ea karolo ea anteroventral ea BNST e boetse e ntlafatsa likarabo tsa khatello ea maikutlo (Radley et al. 2009). Ka lehlakoreng le leng, liso tse kholo tsa anterior BNST li fokotsa likarabo tsa khatello ea maikutlo tsa HPA (Choi et al. 2007), e tsamaellanang le karolo ea sebaka sena khatellong ea khatello. Kahoo, karolo ea BNST ho khatello ea maikutlo le ts'ebetso ea khatello ea methapo e hlophisitsoe ka bongata mme e ka amahanngoa le liphapang tsa ho ts'oaroa ha letsoho la setho sa BNST. Mohlala, posterior medial BNST e fumana bolulo bo boima ho tsoa ho vSUB le MeA, athe sebaka sa anteroventral se fumana ho tsoa ho CeA le boholo ba li-efferents tsa IL (Canteras le Swanson 1992; Cullinan et al. 1993; Dong et al. 2001; Vertes 2004).

Sebaka sa medial preoptic le libaka tsa peri-PVN li na le batho ba bangata haholo ba nang le li-neurons tsa GABAergic mme li shebahala

ho theola khatello ea khatello ea kelello (Herman et al. 2003). Li-Neurons libakeng tsena ho lumeloa hore li fana ka thibelo ea tonic ho PVN, e ka fetoloang ho latela liphetho tsa glutamate tse tsoang ho VSUB (tlhahiso e ntlafalitsoeng) kapa likenyelletso tsa GABAergic haholo-holo ho tsoa ho MeA (disinhibition). Lesions ea medial preoptic nucleus e eketsa likarabo tsa khatello ea kelello ea HPA le ho thibela likarabo tsa axel tsa HPA tse hlahisoang ke ho hlohlelletsa medial amygdalar, a fana ka maikutlo a karolo ea mantlha ea khatello ea maikutlo (bakeng sa liphatlalatso tsa mantlha, bona Herman et al. 2003). Thibelo ea lehae ea ho tšoaetsoa ha glutamate sebakeng sa peri-PVN e boetse e ntlafatsa likarabo tsa khatello ea kelello ea HPA (Ziegler le Herman 2000), e fana ka maikutlo a hore li-axon tsa maoto tse emeng sebakeng sena li ka fetola ts'ebetso ea PVN.

Ho thata haholoanyane ho supa karolo ea libaka tse ling tsa hypothalamic tse hokahanyang li-activic le PVN, tse kang dorsomedial nucleus (Herman et al. 2003). Mohlala, sephetho se hanyetsanang se bonoa ka mor'a lesion, activation, kapa inactivation ea dorsomedial hypothalamus, mohlomong ka lebaka la ho kopanya ho matla ha glutamate le GABA neuronal popul (Herman et al. 2003).

Liphetoho tse ling tse ka 'nang tsa hlaha li ntse li lokela ho hlahlojoa ka botlalo. Mohlala, li-raphe nuclei le NTS tse kenang kahare ho PVN, li shebiloe ke likarolo tsa li-limbic (joalo ka PL) (bona Vertes 2004) mme li kenella ho khatello ea maikutlo ka serotonin le norepinephrine (Herman et al. 2003), ka ho latellana. Leha ho le joalo, hajoale, ha ho na lipatlisiso tsa anatomical tse hlalosang li-bacynaptic limbic-PVN tse fetelang libakeng tsena.

Potoloho e Pholosa Likarabo tsa khatello ea maikutlo e sa foleng

Ho pepesetsoa nako e telele kapa ho ateloa ke khatello ea maikutlo ho baka ho ts'oaroa ha bolelele ba HPA ka nako e telele, ho khetholloang ka boima ba "thymus" (bo hlahisitsoeng ke maemo a phahameng a phahameng a li-GC); saese e eketsehileng ea adrenal (e hlahisoang ke ho eketseha ha ACTH); kutloisiso e eketsehileng ea adrenal ho ACTH; e nolofalletsa likarabo tsa axPA tsa HPA ho li-stress tsa lipale; le ho tse ling (empa

eseng kaofela) paradigms / maemo, secretion e phahameng ea basal GC (bona Herman et al. 1995; Ulrich-Lai et al. 2006). Liphetoho tsa ho lokolloa ha li-hormone tsa peripheral li tsamaisana le PVN CRF le vasopressin mRNA (Herman et al. 1995), e fanang ka maikutlo a hore ho phahamisa HPA ho bohareng. Ntle le moo, khatello ea maikutlo e sa foleng e eketsa glutamatergic le noradrenergic terminal e tšoarisang PVN CRF neuronal somata le dendrites, e tsamaellanang le drive e ntlafatsang ea synaptic drive (Flak et al. 2009).

Mehato e bohareng ea ts'ebetso ea axis e sa foleng ea HPA ha e e-so tsekoe. Karolo ea karolo ea methapo ea maoto ho laoleng khatello ea maikutlo e bontša hore ho nka karolo ha PFC, hippocampus, le amygdala ho ikarabella bakeng sa ho khanna nako e telele. Ho joalo, libaka tsohle li bonts'a liphetoho tse kholo tsa khatello ea maikutlo tse bakiloeng ke li-neuroplastic: Phetoho ea dendritic e bonahala ho li-neurons tsa hippocampal le mPFC, athe katoloso ea dendritic e bonoa ho amygdala (bakeng sa liphatlalatso tsa mantlha, bona Ulrich-Lai le Herman 2009). Lithuto tsena li tsamaisana le phetisetso ea lipalo tsa maoto le li-circuits tsa HPA tse khahlisang, 'me li rata ho khahloa ke litšitiso.

Amygdalar drive e ntlafalitsoeng e khothalletsoa ho bapala karolo e kholo ho lefu la khatello ea maikutlo le sa foleng. Mohlala, khatello ea maikutlo e sa foleng e kenya ts'ebetsong tsamaiso ea CeA CRF, e reriloeng e le tsela e sa foleng ea khatello ea maikutlo-e hiriloeng (Dallman et al. 2003). Leha ho le joalo, CeA ha e bonahale e hlokoa bakeng sa nts'etsopele kapa ho lokisoa ha matšoao a khatello ea maikutlo (Solomon et al. 2010). Ntle le moo, likotsi tsa MeA li boetse li sitoa ho thibela khatello ea maikutlo ea mokokotlo oa HPA (Solomon et al. 2010). Kahoo, khokahano e akaretsang lipakeng tsa khatello ea maikutlo ea amygdalar le khatello ea maikutlo e sa tsitsitseng ea HPA ha e e-s'o tiisoe.

Ho bonahala eka motsoako oa moea o ka sehloohong oa "hypothalamus" ("PVT)" e le karolo ea tsela ea khatello ea maikutlo e sa foleng. Mefuta ea PVT e thibela khatello ea maikutlo e sa foleng ea likarabo tsa HPA axis ho likarabo tsa maikutlo (Bhatnagar le Dallman 1998), ho fana ka maikutlo a karolo ea mantlha ts'ebetsong ea thupelo.

Ntle le moo, marang-rang a PVT a senya ts'ebetso ea ho lula ha axis ea HPA ho bahatelli ba phehellang (Bhatnagar et al. 2002). Ha e kopantsoe hammoho, tlhahiso-leseling e bontša hore PVT e bapala karolo e kholo ho loantseng HPA axis drive maemong a ho pepesetsoa khatello ea maikutlo nako e telele. Ho joalo, libaka tsa PVT le li-limbic forebrain tse laolang likarabo tsa khatello ea maikutlo li matla li hokahane (bona Vertes le Hoover 2008), li lumella ho hokahana ha tlhahiso ea khatello ea corticolimbic sebakeng sena. PVT e boetse e maemong a ho lekola tlhahisoleseling mabapi le boemo bo ntseng bo tsoela pele ba 'mele, e amohela litlatsetso tse tsoang ho orexinergic neurons (e laolang tokollo ea acetylcholine, serotonin, le noradrenaline) ea dorsolateral hypothalamus (e phethang karolo ea bohlokoa taolong ea lits'ebetso tsa lefutso) le tsamaiso ea kelello e ntseng e nyoloha. litsamaiso tse kenyelletsang taolo ea bochaba.

BNST e boetse e boemong ba ho kopanya tlhahisoleseling mabapi le khatello ea maikutlo e sa foleng. Metsoako ea anteroventral BNST e lumellana le likarabo tsa khatello ea maikutlo e matla, empa ts'ebetso e tataisang ea axel ea HPA ka khatello ea maikutlo e sa feleng (Choi et al. 2008). Lintlha tsena li fana ka maikutlo a hore setereke sena se na le karolo e itšetlehileng ka taolo ea axis ea HPA, mme ho na le batho ba fapaneng ba fapaneng ba neano ba lekiloeng ho leka ho arabela ho pepesoa ha khatello ea maikutlo nako e telele. Kaha ho na le kamano e haufi pakeng tsa BNST le mPFC, hippocampus, le amygdala, ho ka etsahala hore li-neuron tsa BNST li "hlophisoe" ke khatello ea maikutlo e sa feleng- e bakileng liphetoho tsa ts'ebetso ea litho tsa maoto kapa mekhoa ea bolulo.

Khatello ea Kelello le joala

Babali ba tloaetseng lingoliloeng tsa joala ha ho pelaelo hore ba tla ba le phapano e ngata pakeng tsa khatello ea maikutlo e hlalositsoeng ka holimo le ho potoloha ha boko ho amanang le joala. Mohlala, datha tse hlokoang li ts'ehetsa karolo ea CeA, BNST, le lits'ebetso tsa naradrenergic ts'ebetsong ea ts'ebetso ea ts'ebeliso ea ts'ebeliso ea lino tse tahang (bona Koob 2009), e fana ka maikutlo a hore ts'ebetso ea ts'ebetso ea tlhekefetso e hokahane le ts'ebetso ea khatello ea maikutlo (le HPA axis) litseleng tse thabisang. Ho joalo, polelo e ntlafalitsoeng ea CeA / BNST CRF e ts'oana le se neng se ka lebelloa kamora khatello ea maikutlo e sa feleng, e lebisang ho khopolo-taba ea hore linaha tse lemaletseng hampe (mohlala, ho qoba ho tlohela) li hokahane le ho hira motho ea tahiloeng oa li-circuits tsa khatello ea maikutlo (Koob 2009). Ka lehlakoreng le leng, ts'ebetso ea mekhoa ea moputso e tsebahala ka ho kopanya khatello ea kelello haholo ka mochine oa amygdaloid, ho fana ka tlhahiso ea hore na litlamorao tsa joala li ka fokotsa khatello ea kelello joang (Ulrich-Lai et al. 2010).

Tšebeliso ea joala e boetse e na le litlamorao tse matla tšebetsong ea 'mele ea methapo pele ho kalafo ea methapo,' me tšebeliso e sa foleng e amahanngoa le preofunal hypofunction (taolo e mpe ea tšusumetso) bathong (bona Abernathy et al. 2010). MPFC e ts'ehetsa CeA le BNST, mme bonyane tikolohong ea prelimbic, e bapala karolo e hlahelletseng thibelong ea HPA. Ha e kopantsoe le ho ba le ts'ebetso e bonoang liserekising tsa amygdalar-BNST, liphuputso tsena li fana ka maikutlo a hore tšebeliso e mpe ea joala bo sa foleng e baka phetoho lipakeng tsa khokahano ea taolo ea khatello ea methapo, e leeme ho setho sa khatello ea maikutlo.

Ka kakaretso, taolo e lekaneng ea axPA ea HPA ke tlhokahalo ea ho pholoha ha nakoana le halelele. Ka lebaka la hore likarolo tsa bohlokoa tsa taolo ea HPA ea axis li lebisitsoe ho joala, le hore joala ka bo bona bo beha kotsing, ha ho makatse hore li-corticosteroids, "khoebo ea ho fela" ea axis, li na le tšebelisano e matla le taolo ea boits'oaro le ea 'mele. Phapang e pakeng tsa taolo ea HPA ea taolo le bokhoba e bonts'a lintlha tsa bohlokoa tseo e ka bang liphofu tsa litlamorao tsa tlhekefetso ea nako e telele le tšebeliso ea boits'oaro. Bohlokoa ba ho potoloha ho potoloha bo hatisoa hape ke kamano e matla ea ho khutlisa lipakeng tsa khatello ea bophelo le ho noa, e thatafatsang boiteko ba ho theha le ho boloka tahi.

Liteboho

Mosebetsi ona o tšehelitsoe ke lithuso MH-049698, MH-069680, le MH-069725.

Phatlalatso ea Lichelete

Sengoli se phatlalatsa hore ha a na tlholisano ea lichelete ea tlholisano.

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