Ho lemalla joala le adrenal cortex (2013)

E phatlalalitsoe marang-rang pele ho 2013, doi: 10.1530 / EC-13-0028 Endocr Connect vol. 2 che. 3 R1-R14 

  1. Caroline H Brennan

+ Boikarabello ba Mongoli

  1. Sekolo sa Saense ea Likokoana-hloko le Lik'hemik'hale, Mofumahali oa Univesithi ea London ea London, London E1 4NS, UK
  2. Litaba li lokela ho romelloa ho GP Vinson Email: [imeile e sirelelitsoe]

inahaneloang

Bopaki bo bonts'ang bo bonts'a hore li-axis tsa "hypophyseal-pituitary-adrenal (HPA)" li-axis le corticosteroids li kentse letsoho ts'ebetsong ea ho lemalla li-agents tse fapaneng, mme adrenal cortex e na le karolo ea bohlokoa. Ka kakaretso, methapo ea methapo ea cortisol (kapa corticosterone ka litoeba kapa litoeba) e eketseha ho tlosoeng hoa lithethefatsi ka mokhoa o bonts'ang kamano le tloaelo le matšoao a hlakileng ho monna le ho liphoofolo tse lekang. Maemo a Corticosteroid a khutlela ho boleng bo tloaelehileng ho qala ts'ebeliso ea lithethefatsi. Tšebelisano e ka bang teng lipakeng tsa li-corticotrophin tse lokollang li-hormone (CRH) le lihlahisoa tsa proopiomelanocortin (POMC) le HPA ea systemic, hape le tsamaiso ea lehae ea CRH-POMC ka har'a gland ea adrenal ka boeona, e rarahane. Leha ho le joalo, bopaki bo eketseha bo fana ka maikutlo a hore tsohle li ka kopitsoa le hore lihlahisoa tsa POMC bokong le bokong li sebelisana le HPA e tsamaeang le mali ka kotloloho kapa ka kotloloho. Corticosteroids ka boeona e tsejoa e ama maikutlo haholo mme e ka ba lekhoba la eona. Ntle le moo, ho na le monyetla oa hore lithuto tse lemalloang li khutlele maemong a amanang le phetoho mosebetsing oa HPA, joalo ka khatello ea maikutlo kapa ka linako tse fapaneng tsa letsatsi. Boithuto ba morao-rao bo fana ka bopaki bo kholisang ba hore karolo ea bohlokoa ea matšoao a mangata a tlatsetsang e hlahisoa ka kotloloho ke ts'ebetso ea lekunutu ea adrenal cortex le liketso tsa corticosteroids. Ho feta moo, phapang ea thobalano le ts'ebeliso ea lithethefatsi e kanna ea hlahisoa ke ts'ebetso ea adrenocortical: bathong, banna ba ka sirelletsoa ka secretion e phahameng ea DHEA (le DHEAS), le litekong, basali ba ka ts'oaroa habonolo ka lebaka la secretion e phahameng ea corticosterone.

Keywords

Selelekela

Morero oa tlhahlobo ena ke ho bonts'a karolo ea bohlokoa ea adrenal cortex le ts'ebelisong le ho fana ka maikutlo a hore phapang ea thobalano ts'ebetsong ea adrenocortical e ka kenya letsoho phanong ea thobalano le tšebeliso ea lithethefatsi. Moo ho hlakileng, thobalano ea liphoofolo tse lekiloeng kapa ea lithuto tsa batho lithutong tse bontšitsoeng, le ha ho le joalo maemong a mangata ho ne ho sa hatisoe liphapang tsa ho kopanela liphate.

Ho na le nalane e telele ea ho kopanya temallo le adrenal. Ehlile e ne e le hantle pele lihormone tsa adrenocortical li ne li khetholloa ka hore chefo ea morphine e amana le tšoelesa ea adrenal. Kahoo, Lewis (1) le Mackay & Mackay (2) e bonts'itse hore adrenalectomy e eketsa kutloelo-bohloko ea morphine litekong tsa basali, le kalafo e sa foleng e nang le morphine ho banna kapa methadone ho tsa bong bo hlahisa tlhahiso ea adrenocortical hypertrophy (3, 4). Ka lebaka leo, ho bile le thahasello liketsong tsa lihormone tsa adrenal joalo ka baemeli ba ka khonehang ho lemalla ho tloha nakong ea ho sibolloa ha bona. Phekolo ka cortisone (kalafo ea kalafo ea corticosteroid ea khetho ka nako eo) e ile ea sebelisoa kapele taolong ea matšoao a tsoang ho meperidine le morphine ho banna (5), kamoo ho bonahalang ka teng e bile le litlamorao tse ntle, ha Lovell o ne a amahanya botaoa ba joala le bokhoba ba lithethefatsi le hypoadrenocorticism (6).

Boithuto bo hlophisehileng haholoanyane bo ile ba theola li-corticosteroids hammoho le 'lipheko' tse ling tsa lipale tsa ho tlohela, le Fraser & Isbell (7) e bile ba pele ba ho fana ka maikutlo a hore, ha e le hantle matšoao a ho ikhula (ho morphine) ho banna a ne a amahanngoa le eosinopaenia, mohato o neng o sebelisoa ka nako eo ho bonts'a maemo a phahameng a ho potoloha ha corticosteroids (8). Eosinophil e baleha ka mokhoa o potlakileng ha morphine e tsosolosoa. Bangoli bana ba boetse ba fumane hore kalafo ka cortisone kapa ACTH e khutsufalitse nako ea nts'etsopele ea matšoao a bang teng ho banna, ka hona, bona ka bo bona ba ka nkoa e le sesosa. (7, 9, 10, 11). Ka sebele, kalafo e sa foleng e nang le corticosteroid ka boeona e ka lebisa matšoao a ho tlohela hamorao (12).

Kahoo ho na le lipotso tsa mantlha mabapi le karolo ea corticosteroids ho ho lemalla. Na ts'ebetso e tlase ea adrenocortical ts'ebetsong ea morphine e tsitsitseng, mme ho phahama ha eona ha tsamaiso e khaotsa, sesosa kapa phello ea likarabo tse tlatsetsang? Na drive ea lithethefatsi tse lemalloang e hlile e ka emisa drive ho ea tlase cortisol, ka tatellano ea eona? Kapa na secretion e atolositsoeng ea li-corticosteroid ka ho tlohella lithethefatsi e mpa e le karabelo ea khatello ea maikutlo? Mona re pheha khang ea hore adrenal cortex e na le karolo ea bohlokoa ho fumaneng bokhoba ba tahi le ho e sirelletsa khahlanong le eona.

Hypophyseal-pituitary-adrenal axis bokong le ho lemalla

Mabapi le bokhoba ba ho lemalla, ho lebisitsoe tlhokomelo e matla haholo ho likarolo tsa hypophyseal-pituitary-adrenal (HPA) bokong ho feta systemic (ke hore, e tsamaeang le mali) axis ea HPA. Likarolo tsohle li teng bokong, 'me, ho ipapisitse le tšitiso ea maikutlo hore adrenal ka boeona e bohlokoa ho bokhoba, ho bohlokoa ho hlakola kamano lipakeng tsa boko le tšebetso ea HPA. Karolo ena e hlahloba bopaki ba tšebetso ea boko HPA ho bokhoba ba eona 'me e bonts'a hore ha e ikemetse,' me ts'ebetso ea eona e laoloa haholo ke 'me e hokahane le systemic HPA.

Corticotrophin e lokollang hormone

Corticotrophin e lokollang hormone (CRH) e hlahisoa likarolong tse fapaneng tsa boko (13). Taba ea mantlha, CRH e fana ka litlamorao tsa tsamaiso ea eona kamora ho lokolloa ha eona bohareng ka lipampitšana tsa neuronal tse simolohang ho "patventricular nucleus (PVN)" ea hypothalamus. CRH e isoa corticotrophs ea anterior pituitary ka tsamaiso ea methapo ea methapo ebe e tsosa sephiri sa ACTH. ACTH le eona e tsamaisoa ho potoloha ka kakaretso 'me e ntlafatsa secretion ea corticosteroids ka har'a cortex ea adrenal.

Ntle le moo, CRH, li-receptors tsa eona CRHR1 le CRHR2, hammoho le protheine e kopanyang ea CRH (CRH-BP), e fetolang liketso tsa CRH, e fumanoa libakeng tse ling tsa boko, moo ho nahanoang hore CRH e sebetsa haholo e le neurotransmitter. Liwebosaete tsena li kenyelletsa "cerebrocortex", limbic system, hippocampus, amygdala, locus coeruleus, bulb ea olapyory le cerebellum (14, 15, 16, 17, 18, 19, 20). Leha ho kenya letsoho ha CRH e matla ea tlatsetso ka tlhekefetso e ka ikemela ho HPA (18, 20), ka sebele ho na le mekhoa eo ka eona e kenyang taelo ea ho kopanya li-hypothalamic CRH (Feie. 1).

Setšoantšo sa 1 

Sheba tlhahiso e kholoanyane: 

Setšoantšo sa 1 

Mokokotlo oa HPA e holisitsoeng. Ho tloha (20, 49, 80, 82, 192, 193) mme o bone mongolo. BNST, mookotaba oa stria terminalis; PFC, cortex ea pele-pele; PVN, nucleus ea patricricular; VTA, sebaka sa tegumental sa ventral (se amanang le likarabo tsa moputso); CRH, corticotrophin e lokollang hormone; POMC, proopiomelanocortin; +, e susumetsang; -, inhibitory. Metsu e tiileng e bonts'a molao o netefalitsoeng, le metsu e nang le matheba e bonts'a liketso tse reriloeng. CRH e bolokiloeng e bontšoa ka litlhaku tse putsoa, ​​'me libaka tsa CRH le POMC li tšoaetsoa ka bofubelu le botala ka ho latellana: mona, metsu e bontša litsela tse laoloang ntle ho pelaelo empa li ka kenyelletsa liketso tsa CRH le POMC peptides. Kameho e thibelang ea li-peptide tsa neural POMC ho PVN CRH e khahla haholo, mme, ha e bapisoa le lits'ebetso tse ling, e ka fana ka tlhahiso ea mokhoa o mobe oa ho fana ka maikutlo; leha ho le joalo, ho na le bopaki bo fokolang ba karabelo ea CRH ea POMC bokong. Sebakeng seo, taolo ea neural POMC ke multifactorial (mohlala (65, 67), mme sena se amana haholo le karolo ea eona ho tekatekano ea matla le phepo e nepahetseng, bona sengoloa. Leha ho le joalo ho na le bopaki bo bongata bo bonts'ang karabo ea li-glucocorticoids ka polelo ea CRH libakeng tse 'maloa tsa boko. Haholo, sena se fosahetse, ntle le ho amygdala, sebaka sa bohlokoa ka ho lemalla (19), moo e nepahetseng.

 

Ka bokong, CRH e tlama ho mefuta ea li-receptor ka bobeli, CRHR1 le CRHR2. Ntle le CRH ka boeona, li-receptor tsena ka bobeli li tlama ligands tsa lelapa la urotensin. Li-receptors tsena tse peli li buisana ka likarabo tse fapaneng; CRHR1 agonists e hlahisa likarabo tse amanang le khatello ea maikutlo moo CRHR2 e kanna ea ba le phello e fokolang, ha e batla e baka khatello ea lijo ka bongata (21, 22, 23, 24).

Ho hlile ho na le bopaki bo hlakileng bakeng sa karolo ea CRH ho ho lemalla (18, 25), hape haholo-holo ka ho khutlisetsoa morao, empa data ha e lule e lumellana kamehla. Mohlala, cocaine e hlohlelletsa axis ea HPA ka mochine o kopanetsoeng oa hypothalamic / CRH-hare ho likhoto tsa banna (26, 27), leha e le hore sena ha se amanang haufi le corticosterone hangata (28), bobeli Crh Ngoliso ea mRNA le corticosterone e potolohang e ntse e eketseha ka ho eketsehileng ka ho khaotsa koae (29). Ho fapana le moo, ho ts'oaroa ha heroine kapa joala bo ts'oaroang ka ho ts'oaroa ho latela ka ho hlakileng ho latela CRH, empa eseng ho corticosterone, ho latela bangoli ba bang (30, 31, 32). Leha ho le joalo, ts'ebetso ea adrenal e ea hlokahala nakong ea boits'oaro ba koae bakeng sa ho khutlisetsoa ho ts'oaroang ke ts'ebeliso ea ts'oaetso ea CRH hore e etsahale (33). Moemeli oa liketso tsa CRH, CRH-BP, joale e hlaha e le ntlha e tlatsetsang, leha e sa ithutoe haholo lefapheng la bokhoba ba lithethefatsi. (34, 35). Leha secretion ka bobeli corticosterone le ACTH li eketsoa ke ho pepesetsoa joala bo hlobaetsang, li thibeloa ho pepeseha nako e telele (36, 37). CRH kapa cortisol ha e kenelle ho khutlisetsong ha koae ho litšoene tsa squirrel (38).

Mabapi le morphine le li-opioid ka ho khetheha, ho hlakile hore ho tsepamisa maikutlo ha corticosteroid ho ka ba le phello ea thibelo ea opioid ea secretion ea CRH, ho sebetsa ka li-receptor tsa μ- le κ-mofuta oa opioid ho hypothalamus ea banna. (39, 40, 41). Ka ho batho, li-opioids ka ho toba li thibela secretion ea CRH le axis ea HPA, e etsang hore cortisol e fokotsehe. Litekong tsa banna, phello ke biphasic, ka ntlafatso ea pele ea CRH (le HPA) e lateloang ke thibelo kamora matsatsi a 'maloa a kalafo (41, 42); likarabo tse joalo li angoa ke khatello ea maikutlo ho likhoto tsa banna (43). Ho joalo, bopaki bo fana ka maikutlo a hore mekhoa ea opioidergic bonyane e ka nyatsa ka boomo litla-morao tsa boits'oaro ba CRH ho likhoto tsa banna (44) hape le keketseho ea secretion ea CRH tlasa maemo a khatello. Sena se kanna sa se be joalo maemong a mang joalo ka ts'ebetso e eketsehang ea HPA ho liphoofolo tse adrenalectomised (45). Ts'ebetso ena habeli ho likhoto e ka ba hobane li-opioid li na le litlamorao tse fapaneng mefuteng e fapaneng ea lisele: ehlile li thibela secretion ea CRH e khothalletsoang ke li-neurotransmitters (46). Ho kenyelletsoa ho hoholo ha li-opioid bokhobeng ba joala ho batho (47) e boetse e bontšitsoe hore e tla sebelisoa ka ntle ho lits'ebetso tsa HPA (48).

Ho na le phapang e hlakileng lipakeng tsa liketso tsa lithethefatsi tse fapaneng tse lemalloang Crh sengoloa sa mRNA ho hypothalamus, 'me leha joala bo sebetsa ka kotloloho ho PVN, lithethefatsi tse ling, ho kenyeletsa cocaine, nikotine le cannabinoids, li kenya activate Crh transcript libakeng tse ling tsa boko (49). Ts'ebetso ea adrenocortical e kanna ea e-ba boima, ho etsa mohlala, ho khutlisetsoa ha lithethefatsi tsa koae ho litoeba tsa banna (33). Nako ea ho pepesa le eona e bohlokoa; ho pepeseha kapele ho ka ama likarabo tse latelang (50)le likotong tsa banna, ho pepesetsoa mocha oa moro oa joala kapele ho fetoha motho e moholo ea latelang Crh Karolo ea phetisetso ea joala bo bongata (51).

Ntshetsopele ea bahanyetsi ba itseng ba CRHR1 e fane ka leseli le eketsehileng. CRHR1 blockade e thibela ho nooa ho eketsehileng ka har'a litoeba tsa banna tse tloaetseng ho ja haholo (52), hape, hammoho le lithuto tse ling tse sebelisitsoeng Crh1 Ho kenoe ke liphoofolo, ho bontšitsoe hore CRHR1 ea ho ts'oasa litsela ke ea bohlokoa bakeng sa ho susumetsa bokhoba ba tahi ho litoeba tsa banna. (53); polelo e tloaelehileng ea neuroadaptations e susumetsoang ke ho pepesehela khafetsa lithethefatsi tse lemalloang ke karabelo e tsitsitseng ea boitšoaro mabapi le thepa ea bona e matlafatsang. Bangoli bana ba boetse ba bonts'a hore ho fumana le ho susumetsa maikutlo ho laoloa ka mokhoa o fapaneng. Ho fumana ho kenyelletsa axis ea HPA mme ho thijoa ke glucocorticoid blocker mifepristone hammoho le CRHR1 blockade, athe kutloisiso ha e sebetse ke mifepristone. Moruti et al a. (53) etsa tlhahiso ea hore sena se bonts'e tsela e hokahaneng le hypHR1-e kopaneng ea sensitization. Litlamorao tse fapaneng li ile tsa bonoa likarabo tsa methamphetamine (MA), moo maikutlo a boitšoaro a lekantsoeng joalo ka ha a eketsa ts'ebetso ea indomotor e sa sebetseng ho eona Crh1 ho kokota kapa ke mohanyetsi oa CP 154 526 ho litoeba tsa DBA / 2J, athe ho hlakoloa Crh2 attenuated MA-e susumetsoa ke boits'oaro ba boits'oaro. Mona, ho ile ha khothalletsoa ketso ea li-urocortin tsa mehleng ea khale, e tsepamisitsoeng mohahong o ka tlase ho naha le bohareng ba amygdala (54).

Proopiomelanocortin

Proopiomelanocortin (POMC) e fana ka, ho ACTH le α-melanocyte e susumetsang hormone (α-MSH), likarolo tse ling tsa HPA axis, mme moelelong ona, sebaka sa eona sa mantlha sa polelo le ts'ebetso ke sebaka sa pele le (ho litoeba) sa. pars intermedia. POMC e boetse e hlahisoa libakeng tsa bokong, haholoholo ka ho kholoa ho tsoa ho arcuate nucleus ea hypothalamus le ho tloha ho "nucleus" trususitarius ea "brainstem" (55, 56, 57). Karolo ea eona ea mantlha bokong ke moloko oa α-MSH, o nkang karolo taolong ea tšebeliso ea lijo le ho hlahiseng β-endorphin, taolo ea bohloko. α-MSH e sebetsa ka makhetlo a mabeli letotong la melanocortin receptor (MCR), MC3R le MC4R, 'me ea morao-rao e ka boela ea laola likarolo tsa tlhokomeliso ea bohloko (25, 58).

Polelo le ts'ebetso ea POMC e fana ka maikutlo a hore le ha lihlahisoa tsa ACTH le tse ling tsa POMC tse kang β-endorphin li ka fumanoa libakeng tse se nang hypothalamic tsa bokong kapa mokelikeli oa "cerebrospinal fluid" (59, 60), tse ling li ka fetisetsoa bokong li tsoa maling (60, 61). Ho tloha nts'etsopele ea pele, sehlahisoa se ka sehloohong sa POMC se amanang le adrenocortical e amanang le adrenocortical bokong ke α-MSH (62), mohlomong e amanang le kabo ea prohormone e fetolang PC1 le PC2 (63, 64). Ho fihlela joale, sepheo sa mantlha se shebaneng le sena ke karolo ea α-MSH e nang leptin, ghrelin le agouti protheine ea taolo ea phepelo ea lijo le botsitso (56, 62, 65, 66, 67, 68).

Ntle le karolo ea eona ea matla a ho leka-lekana, α-MSH e boetse e bapala karolo ea physiology ea ho lemalla, 'me MC4R, joalo ka li-receptors tsa CRH, e arabela morphine (69, 70, 71), le litlamorao tsa boits'oaro ba morphine kapa cocaine bo hlophisoa ka thibelo ea MC4R e khethiloeng (72, 73). Ho ekelletsa moo, kalafo ea joala e matla e fokotsitse polelo ea α-MSH libakeng tsa hypothalamic le libakeng tse ling tsa boko liteketeng, empa kalafo e sa foleng e ile ea e ntlafatsa. (74).

Ho joalo, ts'ebetso ea POMC mabapi le bokhoba ba tahi e ke ke ea nkoa e le mesebetsi e amanang le HPA feela. Ho hlahisoa ha β-endorphin ho lebisa ka kotloloho ho litlamorao lits'ebetsong tsa ho lemalla. Ketso ea eona ea mantlha e kopantsoe ke μ-receptors joalo ka li-opiates morphine, heroin le methadone, 'me bathong, li-opiates tsa endo native le tsona li sitoa ho thibela ts'ebetso ea HPA, leha e le hore e susumetsa ebile ha e thibele litheko (49, 75).

Ho sa kang ha hlaka ho fihlela joale ke hore na poleloana ea 'HPA axis' ka 'nete e ka eketsoa ho likarolo tsena bokong. Ka mantsoe a mang, ha ho hlake hore na, mohlala, CRH e seng hypothalamic e baka tšebeliso, ho sebetsa kapa ho lokolla POMC bokong, empa libaka tse fapaneng tsa polelo ea likarolo tsena li ka fana ka maikutlo a hore ha e (Feie. 1). Ka mokhoa o ts'oanang, ha ho na bopaki ba hore lihlahisoa tsa boko CRH kapa POMC li na le tšebelisano leha e le efe le adrenal cortex le secretion ea glucocorticoids, ntle le ka hypothalamus. Ho fapana le hoo, ka linako tse ling ho 'nile ha nahanoa hore ha ba etse joalo ka mohlala (53)). Leha ho le joalo, ho senyeha ha neural glucocorticoid receptor (GR), ho kenyelletsa le PVN, e ntlafatsa litholoana tsa matšoenyeho hape e etsa hore mosebetsi oa HPA o phahamele ho litoeba tsa banna. (76), e tsamaellanang le tahlehelo ea glucocorticoid inhibition ea CRH (20, 77). Ho fapana le hoo, ho kokota ka ho khetheha ha GR, e sa kenyeng PVN, ho eketsa boitšoaro ba ho tšoenyeha empa ho na le phello e ts'oanang ea ho fokotsa thibelo ea glucocorticoid ea CRH ho litoeba tsa banna. (77). Ho hlakile ho tsoa thutong ena hore HPA e laoloa karolo e itseng ke tšohanyetso ea GR-Mediated inhibition. Ka hona, se hlokang ho hlakoloa ke bohlokoa ba likarolo tsa boko tsa CRH / POMC ka mokhoa o fapaneng le oa HPA ea systemic, le hore na lits'ebetso tse ikemetseng li hlile li lemaletse joang.

Tšebelisano pakeng tsa CRH ea boko le α-MSH

Le ha ts'ebetso e ka sehloohong e amoheloang ea α-MSH bokong, taolo ea phepelo ea lijo le phepo e ntle e bonahala e sa amane haholo le CRH, haele hantle ho na le bopaki bo bongata ba crosstalk pakeng tsa bona. Ehlile, joalo ka ts'ebetso ea HPA ea methapo, li-neurones tse sebetsanang le POMC li hlahisoa ke khatello ea maikutlo 'me li bapala karolo ea karabelo e latelang ea boitšoaro literekeng tsa banna (78, 79). Ntle le moo, li-peptide tsa neuronal POMC tse hlahisitsoeng li laola hypothalamic CRH ka hona secretion ea ACTH ho litoeba tsa banna le basali (80). Ho feta moo, α-MSH ea hlasimolla Crh transcript ho PVN ea likhoto tsa banna (81, 82), leha, joalo ka γ-MSH, e boetse e thibela tšebetso ea HPA ea li-interleukin-1β, mohlomong e le ka har'a li-MCR tse bohareng. (83). Hore mokokotlo o kopanyang boko le HPA ea systemic e felletse ho fanoa ke ho fumana hore li-glucocorticoids li ntlafatsa MC4R ho supa mola oa selefounu ea hypothalamic neuronal (84). Ka hona re ka bolela esale pele hore ho na le axis ea HPA e atolositsoeng eo likarolo tse tšoanang, CRH, lihlahisoa tsa POMC le corticosteroids joalo ka ts'ebetsong ea li-classical, le tsona li kenang bokong.Feie. 1) e nang le litlamorao tse khethehileng ho maikutlo le boits'oaro. Lits'ebetso tsena tse peli, boko le somatic, li sebelisana ho isa tekanyong ea hore na eng kapa eng e susumetsang ts'ebetso ea systemic, "khatello" ka bophara le nako, le tsona li tlameha ho ba le litlamorao ho maikutlo le boitšoaro.

Li -teroids bokong

Sebopeho sa meralo le tšebetso ea li-neurotheroids li pharalletse haholo ho theha lekala la endocrinology (kapa bonyane paracrinology) ka tokelo ea eona. Boholo bo entsoe ka mokhoa oa lehae, leha ho le joalo hangata bo hloka likarolo tse tsoang mehloling eo eseng ea tlhaho. Li-Oestrogens li hlahella har'a tsena 'me li hlahisoa ke lintho tse aromatase hippocampus, ho nahanoang hore ho latela C e hlahisoang sebakeng sa heno.19 li-substrates tsa steroid (85). Li na le likarolo tsa polasetiki ea neural (86) le neuroprotection (85, 87, 88) le ho laola ts'ebetso ea li-agents tse ling tse sebetsang, ho kenyeletsa le neuroprogesterone, eo le eona e hlophiselitsoeng sebakeng seo (89). Ho na le liphapang tse amanang le ho kopanela liphate likarabo tsa neural ho estrogen (90, 91, 92). Ketso ea estrogen bokong e arolelanoa ka li-receptor tsa "estrogen" tsa khale le β hape ka li-receptor tsa membrane metabotropic glutamate (93, 94). Li-steroids tse nang le 'mele tse sebetsang hantle haholo NLi-receptor tsa -methyl-d-aspartate kapa gamma-aminobutyric acid (GABA) li kenyelletsa adrenal androgen DHEA, eo ka ha DHEAS conjugate e leng steroid e ngata ka ho fetisisa plasma ea motho (95, 96, 97, 98). DHEA ha e patiloe ke rat adalal cortex: boteng ba eona le ts'ebetso bokong li bonts'a mofuta oa eona oa lehae (99). DHEA le imaenolone, ka bobeli Δ5, 3β-hydrosteroids, le tsona ke li-agioists tsa opioid sigma, athe progesterone, e nang le Δ4, 3-e ngoe tokiso, ke mohanyetsi (100). Ka liketso tsa bona tsa agmaist tsa sigma-1, ho qhekella ka DHEA kapa bokhachane ba bokhachane bo nang le mofuta oa cocaine o behiloeng sebakeng se ratoang ke litla-morao tsa CPE ho litoeba. (100) empa e bontsa boitšoaro bo batlang koae (101). Ho bakuli, DHEA le DHEAS li amahanngoa le liketso tse molemo tsa ho khaotsa koae (102, 103), mme ts'ebeliso ea tsamaiso ea DHEA ho thusa ho tlosa opioid e ithutile, ka liphetho tse fapaneng (104, 105).

Li-neurosteroids tse ling tse tsejoang li kenyelletsa 3cy-hydroxy-5α-pregnantan-20-one (tertrahydroprogesterone, allopregnanolone, THP) le 3cy, 21-dihydroxy-5cy-pregnantan-20-one (tetrahydrodeoxy le theCCCC, LIHCCCC le lCCCC, liTDCCCC, liXPUMX-one (tetrahydrodeoxy le lysCDCC, tse tsoang ho li-ubongo le li-XCUMUMELENG. progesterone le deoxycorticosterone (106, 107). Ba na le mesebetsi e khathatsang, e khahlanong le ho tsitsipanya le e susumetsang 'me ba tsebahala hore ba phahame ka tsela e tšoanang ka plasma le bokong ho arabela ethanol ka likhoto. (106, 108). Ntle le moo, axis ea HPA e tlas'a thibelo ea tonic GABA maemong a hypothalamic (75). Habohlokoa le ho feta, tlhahiso bokong ba bobeli ba THP le THDOC e ipapisitse le li-steroids tsa pele ho tlhaho tsa adrenal (106).

Li-corticosteroid ka botsona li na le litlamorao tsa methapo, 'me likelello tsa kelello tsa corticosterone ka sebele li na le bohlokoa bo amanang le boits'oaro ba lithethefatsi ho likhoto tsa banna (109), 'me u bone ka tlase. Leha ho le joalo, bohlokoa ba sebopeho sa boko ba lehae ba corticosteroids ha bo hlake. Ehlile, li-enzyme tsohle tse hlokahalang tsa tsela ea corticosteroid biosynthetic e tsoang cholesterol e teng, haholo hippocampus, hammoho le proteni ea STAR (110, 111, 112), empa boemo ba tsona ba tlhahiso bo ka ba bo tlase ha bo bapisoa le likhahla tse tšelang mali - boko, 'me ha ho nahanoe hore li hlahisoa bokong ho isa tekanyong e kholo. (113, 114). Ho makatsang ke hore, ka li-neurrateoids tse tsejoang, li-corticosteroids li ka oela sehlopheng sa tsona ka botsona li itšetlehileng ka mohloli o tsoang hole: adrenal cortex.

Karolo ea adrenal cortex

Corticosteroids le mongobo

Ka ho hlakileng, karolo ea corticosteroids bokhoba ba tahi e ke ke ea utloisisoa ntle le ho bua ka sebopeho sa likarolo tsa kelello le boits'oaro ba liketso tsa corticosteroids ka boeona. Hoo e batlang e le li-corticosteroids li qalile ho tsebahala, matla a bona a makatsang a ho hlahisa thabo le khatello ea maikutlo ho batho li tsebahala, leha li sa utloisisehe hantle (115, 116). Liphetoho tsa maikutlo ke karolo ea kalafo ea corticosteroid e sa foleng, ka euphoria e bonolo ka nako e khuts'oane mme e eketseha le ho teba ha matšoao a amanang le khatello ea maikutlo, kapa esita le psychosis ka nako e telele, 'me tsena li etsahala hangata ho basali. (116, 117, 118, 119, 120), leha ho bile le phapang e kholo ea tlhaiso lithutong tse fapaneng. Ho feta moo, boemo ba cortisol ka bobeli le karabelo ho ACTH li phahame haholo maemong a sithabetsang kapa a khatello ea maikutlo (121), le liteko tsa liphoofolo li bonts'a hore tsena ka bobeli li ka amana le secretion e phahameng ea CRH (29). Ho 'nile ha fanoa ka maikutlo a hore li-corticosteroids li ka ba le karolo ho mafu a kelello a amanang le dopamine (122), hape ho nahanoa hore likarolo tse ling tsa boits'oaro ho liphoofolo le batho li ka hlaha ka lebaka la liphetoho tsa sebopeho kapa tse ling bokong tse ka 'nang tsa baka li-corticosteroids, kapa bonyane tsa tsamaisa (114, 123, 124). Phokotso ea maemo a potoloho ea corticosteroid, hammoho le li-indices tse ling, le tsona li ka sebelisoa e le letšoao bakeng sa karabelo kalafong ea ho tšoenyeha (125, 126). Ho phatlalalitsoe hore khatello ea maikutlo ha e le hantle e bonts'a ho nyahama ha GR, ho fana ka maikutlo a lefutso la glucocorticoid ho hypothalamus, ka hona a eketsa ts'ebetso ea HPA. Mohlaleng ona, ketso e le 'ngoe ea li-antidepressants ke ho khutlisetsa tšebetsong tšebetso ea mongolo oa GR (125), ntle le ketso ea bona ea ho khutlisa hape monoamine, empa mohlomong e kenyelletsa taolo ea ho felisoa ha steroid seleng ka ts'ebetso ea ho fetisa ea lithethefatsi P-glycoprotein membrane (127, 128). Ka kopanelo, lithuto tsena li fana ka maikutlo a hore liphetoho tse fetohileng tsa corticosteroid-evoke li ka amana le karabelo ea boitšoaro ho lemallo.

Corticosteroids le bokhoba ba tahi

Leha kamano ea pejana pakeng tsa adrenal cortex le bokhoba ba eona e nkiloe haholo ho tsoa ho bopaki bo potolohileng, hona joale ho na le datha tse bonts'ang khokahano e tobileng ea ho baka. Ho tsoa phihlelong ea bona le bakuli ba fumanang kalafo e sa foleng ea steroid, bangoli ba bang ba ikemiselitse ho ho bitsa corticosteroids e le lithethefatsi tsa ho lemalla (129, 130, 131, 132, 133, 134), leha boholo ba bopaki ba pejana bo thehiloe litlalehong tsa linyeoe ka bomong. Liphetho tsena li fana ka maikutlo a kamano e haufi pakeng tsa corticosteroids le bokhoba ba maikutlo, mohopolo o hlahisoang haholo ke lithuto tsa morao tjena. Tsamaiso ea joala e etsa hore secretion ea ACTH e sebetse ka mokhoa o joalo (106). Ho banna ba tloaetseng ho tsuba ba tsubang haholo empa eseng koae, "plasma ACTH" le cortisol li etsahala kamora metsotso ea ho tsuba. (135). Bopaki bo eketsehileng bakeng sa liketso tsa bohlokoa tsa cortisol e phahameng bo fanoa ke setsoalle sa eona le thuto e sa tsitsang le mohopolo ho banna le basali ba lemaletseng koae. (136), le ha maemo a phahameng a basal cortisol a amahanngoa le ts'ebetso e ntlafalitsoeng ea mohopolo ho taolo e phetseng hantle. Litlamorao tsena tse hopoloang li bonts'a karohano ea karabelo ea cortisol e nang le sefahleho sa U; maemong a tlase, cortisol e eketsehang e molemo ho likarabo tsa kelello tsa hippocampal, empa maemong a phahameng, ha ho joalo (137). Tekanyo ea cortisolaemia le khatello ea maikutlo tse amanang le khatello ea maikutlo, e hokahanngoa le ho eketseha ha positivity ka mor'a amphetamine ho banna le basali. (138).

Ho feta moo, bopaki bo bongata ba liteko bo tšehetsa mohopolo oa kakaretso (bona Lethathamo 1). Likhoto tsa banna le tsona li itlhomme corticosterone ka tsela e bonts'ang ho its'epaha (139, 140). Kahoo, de Jong et al a. (141) e fumane hore sensacization ea cocaine e kenyellelitseng maikutlo ho litoeba tse tšehali tsa adrenalectomised e khutlisitsoe ka phetiso ea li-adrenaline le corticosterone, 'me boitšoaro ba cocaine- kapa joala bo tahang litoeba tsa basali bo thibetsoe ha ho e-na le inhibitor ea GR. (142). Ntle le moo, haeba tšebeliso ea corticosteroid e koetsoe, boitšoaro ba cocaine le bona bo khutla ho latela lingoli tse ling (143). Ba bang ba fumana phetoho eo hore corticosterone e thusa ho khutla, le ha dexamethasone e sa etsa joalo, ho fana ka maikutlo a ho nka karolo ha mineralocorticoid receptor (NR3C2, MR) (144). Litlamorao tse joalo, joalo ka tse lithethefatsi tsa antipsychotic, li ka arolelanoa ka sisteme ea mesolimbic dopaminergic (145, 146). Hoa makatsa hore likarabo tse itšetlehileng ka dopamine ho morphine li hloka li-receptor tsa glucocorticoid (147).

Ho liphoofolo tse lekang, bopaki bo hlakileng ba karolo ea bohlokoa ea corticosteroids tlatsetsong ea ho lemalla bo tsoa lipatlisisong tsa morao-rao litlamorao tsa GR ho ea pele le polelo. Boloetse bo ikhethang ba GR ho litoeba bo fokotsehile ho itaola ha koae, ha corticosterone e khutlisitsoe (148). Ts'usumetso e khethehileng ea GR ho dopaminocepts empa eseng dopamine neurones e fokotse ho ikarola ha koae (149), athe pherekano ea GR ka mefuta e 'ngoe le e' ngoe e fumana k'hok'heine ea koae, e se na phello ea boitšoaro bo khothalletsoang ke morphine (150). CPP e kenyellelitsoeng ke morphine e itšetlehile ka hippocampal le nucleus accumbens GR (151). Ho litoeba tse tona, ho ba le khatello e feteletseng ea 'mele e tlisoang ke nama ho etsa hore maikutlo a batho ba bangata a eketsehe ka koae hammoho le matšoenyeho (152).

Ho boetse ho na le bopaki ba karolo ea bohlokoahali ea GR lithutong tsa li-polymorphisms tsa GR bathong, tse senotseng mokhatlo oa linyeoe tse itseng ka qaleho ea tlhekefetso ea joala ho bacha ba basali. (153). Lintlha tsena le tse ling tsa liteko tseo hona joale li hokahanyang boitšoaro le matšoao a corticosteroids, haholo ha a arabela cocaine, li akaretsoa ho Lethathamo 1.

Lethathamo 1 

Glucocorticoids le bokhoba ba lithethefatsi. Bopaki bohle bo tobileng ba tlhahlobo bakeng sa karolo ea bohlokoa ea li-glucocorticoids bo fumanoe liphoofolong tsa liteko, joalo ka ha ho bonts'itsoe mona. Bopaki bo tsoang ho mofuta oa motho ha bo na molemonyana ebile bo potile empa bo bonahala bo tšehetsa qeto e akaretsang ea hore li-glucocorticoids, tse laoloang ke axis ea HPA e atolositsoeng, tlasa likarolo tsa bohlokoa tsa ho lemalla.

 

Phapang pakeng tsa thobalano

Monyetla oa phapang ea thobalano karabong ea lithethefatsi tsa tlhekefetso ea boko CRH, POMC, li-neurotheroids le axis ea HPA ha o so buisoe kae kapa kae ka har'a lingoliloeng tse hlahlojoang mona. Ka linako tse ling, thobalano la liphoofolo tse lekiloeng le sebelisoang ha le fanoe, leha hona ho le joalo. Mohopolo ke hore lithuto li etsoa khafetsa ho liphoofolo tsa bong bo le bong - likhoto tsa banna li sebelisoa khafetsa - ho fokotsa phapang. Leha ho le joalo phapang ea thobalano le ts'ebeliso ea lithethefatsi e hlakile ebile bopaki bo pharaletseng bo sekasekiloe lithutong tsa batho le ho liphoofolo tse lekang. Kahoo, basali ba kotsing ea ho lemalla ebile ba kotsing e kholo ea ho oela hape ho feta banna (154, 155), 'me likhoto tsa basali li pepeseha habonolo ho feta likhoto tsa banna. Bopaki bo bongata bo hokahanya sena le li-hormone tsa gonadal (156).

Leha ho le joalo ho na le lebaka le letle la ho hakanya hore lihormone tsa adrenocortical le tsona lia ameha. Ka bobeli batho le likhoto li na le liphapang tsa ho kopanela liphate mesebetsing ea adrenocortical, mme leha li fapana ka sebopeho, ka bobeli li ka tlatsetsa karohanong ea thobalano le ho lemalla.

Ho batho, phapang e potolohang cortisol ho banna le basali e mahareng haholo, leha ho ka ba le phapang ea karabelo ho ACTH (96, 157, 158). Leha ho le joalo, sehlahisoa se seholo sa tšoelesa ke ha e le hantle DHEA, e sa patiloeng feela joalo ka steroid ea mahala, empa hape, mme haholoholo, joalo ka sulphate, DHEAS. Likhahla tsa Plasma tsa DHEA le DHEAS ho banna ba baholo li ka ba 12 nM le 10 μM ka tatellano, ha li bapisoa le 8 nM le <7 μM ho basali, maemo a fokotseha ka lilemo empa liphapang tsa thobalano lia bolokoa (96, 159, 160, 161).

Taba ke hore DHEA e bonts'itsoe e le ts'ireletso khahlanong le lithethefatsi tsa bokhoba ba lithethefatsi, joalo ka ha ho boletsoe pele. Bopaki bo tsoang mokelikeli oa "cerebrospinal fluid" bo bontša hore adrenal DHEA, esita le DHEAS, li ka fihla bokong ka bongata ba bohlokoa (162), leha e le hore na sena se amana joang le lipalo tse bokeletsoeng ka bokong li ke ke tsa hlahlojoa. Leha ho sa tlaleheloa liphapang tsa botona le botšehali mokelikeling oa 'mele, ho ntse ho utloahala hore banna ba fumana tšireletso ea DHEA ho lithethefatsi tse lemalloang ho feta basali (154, 162).

Litekong, boemo bo fapane, 'me ha ho na secretion ea bohlokoa ea DHEA. Leha ho le joalo, ho na le phapang e kholo ho secretion le ho potoloha ho tsepamisitsoeng ha corticosterone (glucocorticoid e ka sehloohong ho rat); basali ba baholo ba adrenals ba batla ba lekana le banna ka makhetlo a mabeli; 'me tlhahiso ea corticosterone e kholo ka ho lekana (163, 164, 165, 166). Leha ho le joalo ka ha ho boletsoe pejana, DHEA e hlophisitsoe bokong ba rat, ha ho na phapang ea thobalano, 'me likelello tsa kelello li tšoana ho ba batona le ba batšehali. (167). Ka hona, ho rat, ho ka utloahala hore maikutlo a phahamisitsoeng a lithethefatsi tse lemalloang ho basali a amahanngoa le maemo a potolohileng a corticosterone.

Boiketsetso, bokhoba le oache

Haeba ke adrenal gland ka boeona e bohlokoa bakeng sa lits'ebetso tse lemalloang tsa HPA, lintlha tse ling tse bohlokoa ho hlahiseng likarabo tsa adrenocortical li ka lebelloa ho sebetsana le ho lemalla. Ho susumetso ea 'mele e khothalletsang adrenal cortex, khatello ea maikutlo ke eona e hlahelletseng ka ho fetisisa ebile e loketse. Leha ho le joalo, molaoli ea matla oa adrenal cortex ke eena nako.

Khatello eo, leha e hlalosoa, e thusa ho lemalla bakuli le mehlala ea liphoofolo e utloisisoa hantle (168, 169, 170, 171, 172). Hoa thahasellisa ho tseba hore nako ea nako le eona e na le litlamorao ho litakatso le boits'oaro bo bobebe, leha lingoliloeng tsena ka kakaretso li sa bue ka HPA, empa li tsepamisitsoe ho pineal le melatonin bokong ba litoeba tsa banna. (173), kapa, haholoholo, ka mefuta ea li-wat. Nako ea PER1 le sensitivity ea cocaine li amahanngoa le likhoto tsa banna le litoeba tsa mefuta e fapaneng (174), ho khutlisetsoa ha lithethefatsi ho ka hatelloa ke lifoto tsa linako ho litoeba tsa banna (175), le mefuta ea li-wat tsa mefuta-futa li amahanngoa le sensitization ea cocaine ho Drosophila (176) joalo ka ho lemalla litoeba (ho kopanela liphate ha ho fanoe) (177) le bathong, ho latela bangoli ba bang (178, 179, 180, 181) empa hase bohle (182). Bakeng sa banna, tšebeliso ea joala ka nako e fetang ea lihora tsa 26 ha e-so ka e ama melatonin kapa phapang ea curnisol secretory diurnal (183, 184).

Autonomy ea adrenal

Karolo e ngoe ea ts'ebetso ea adrenocortical e sa tsotelleng ho nahanoa, kamanong le bokhoba kapa eng kapa eng, ke hore mekhoa e teng eo sephiri sa glucocorticoid se hlahang se laoloa ke karolo ea susumetso ea lehae. CRH ea tsebahala har'a tsena. Kamano pakeng tsa mesebetsi ea hypothalamic CRH le CRH e entsoeng sebakeng sa heno adrenal ha e sa bonahala. Hore boteng ba adrenal ea mefuta e fapaneng e ka patela CRH ho tloha ho medulla ho bontšitsoe ts'usumetso ea splanchnic nerve, joalo ka tšusumetso e tobileng ea CRH ho secretion ea corticosteroid (185, 186, 187, 188). CR7 ea adrenal e fapana joang le lithethefatsi? Sena ke sehlooho sa nako e tlang.

fihlela qeto e

Ho na le mokhoa o hlakileng kamanong ea ts'ebetso ea HPA ho nts'etsopele ea mekhoa e lemalloang ho arabela lithethefatsi tse fapaneng haholo. Ke eng eo kaofela ba tšoanang ka eona? Na ho na le tsela e kopanyang eo maemong a mangata e lebisang ho seo ka nako e 'ngoe e ka' nang ea bonahala e le temallo ho adrenal cortex le secretion ea glucocorticoids?

Ntlha e ngoe e ntse e hlakile: CRH le POMC libakeng tse fapaneng tsa boko li na le likhokahanyo tse hlakileng tsa ts'ebetso le HPA ea khale (Feie. 1), 'moho, ba ka bapala karolo e tšoanang phetatsong e tlatselletsang boits'ebetsong bo lematsang. Li kanna tsa nkuoa boemong ba bokhoba ba ho lemalla e le HPA e atolositsoeng, eo pheletso, 'me ea bohlokoa, motsoako ke adrenal cortex ka boeona.

Bopaki ba bohlokoa ba bohlokoa ba adrenal cortex le glucocorticoids boits'oarong le matšoao ho tloheng hoa lithethefatsi le ho khutlisetsoa ha bo bonahala bo phethehile. Taolo ea kalafo ea secretion ea glucocorticoid kapa thibelo ea ketso ea glucocorticoid ho receptor ea eona e ka ba tsoelo-pele ea bohlokoa ea bokamoso (148, 189) ka tsela efe kapa efe e le pono e lerootho ea kalafo (48, 189, 190, 191).

Phatlalatso ea phaello

Bangoli ba phatlalatsa hore ha ho na khohlano ea lithahasello tse ka nkoang e le leeme le hlokang leeme la tlhahlobo.

dithuso tsa ditjhelete

Patlisiso ena ha e a ka ea fumana thuso e khethehileng ho tsoa ho lefapha lefe kapa lefe la lichelete setjhabeng, khoebong kapa bakeng sa phaello.

  • E amohetse 24 May 2013
  • E amohetse 31 May 2013
  • E fumaneha inthaneteng e le 31 e amohetsoeng ea Preprint 2013

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