Auala Neural Path of Stress Integration: Faʻaleagaina o le Faʻamaina o le ava, James P. Herman, Ph.D.

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James P. Herman, Ph.D., o se polofesa i le Matagaluega o Psychiatry and Behavioral Neuroscience, Iunivesite o Cincinnati, Cincinnati, Ohio.

O le fa'atiga o se vaega taua i le atina'eina, tausiga, ma le toe fa'afo'isia o amioga ua fai ma vaisu, e aofia ai le fa'aaogaina o le ava. O lenei tusiga e toe iloilo le tulaga o loʻo i ai nei o tusitusiga e uiga i le tali atu o le popole o le faiʻai, e taulaʻi i le hypothalamic-pituitary-adrenal (HPA) axis. O tali fa'atiga e mafai ona tupu o se tali atu i lu'i po'o fa'amata'u fa'aletino (po'o fa'aletino); fa'ailoga mai le tele o vaega o le fai'ai e tu'uina atu fa'aoga i le paraventricular nucleus (PVN) i totonu o le hypothalamus. Ae ui i lea, e tupu foi tali i faʻalavelave faʻalavelave e vavalo ai faʻamataʻu faʻalavelave (psychogenic stressors). O tali a le Psychogenic e faʻatalanoaina e se faasologa o fesoʻotaʻiga neura i le ala limbic-PVN, faʻatasi ai ma le amygdalar ma le infralimbic cortex circuits e faʻaalia ai le fiafia ma le prelimbic cortex ma le hippocampal neurons e faʻaalia ai le faʻalavelave faʻalavelave. Limbic-PVN feso'ota'iga e fa'asalalauina e le tele o GABAergic neurons i itulagi e pei o le moega o le stria terminalis ma preoptic area. O le faʻalavelave faʻafuaseʻi e aʻafia ai le fausaga ma le gaioiga o le limbic stress circuitry ma maua ai le faʻaleleia o le PVN excitability, e ui lava e le o iloa tonu le masini. O le mea taua, o le faʻaalia o le ava malosi ma le masani e iloa e aʻafia uma ai auala faʻalavelave faʻapitoa ma psychogenic ma e mafai ona fesoʻotaʻi atu i le faʻalavelave faʻalavelave e ala i le faʻalavelaveina o le faʻalavelave faʻafuaseʻi-pei o suiga i amygdalar ma vaega muamua o le fesoʻotaʻiga e pulea le faʻalavelave limbic.

upu autu: Vaisu; 'ava malosi ma isi amioga e sa'ili ai fualaau fa'asaina; le fa'aaogaina o le ava malosi ma le fa'aleagaina; atuatuvale; fa'alavelave; fa'alavelave fa'aletonu masani; tu'ufa'atasiga mamafa; tali physiological i le atuatuvale; tali fa'apopoleina psychogenic; fai'ai; ala neural; ala limbic-paraventricular; feso'ota'iga e pulea le mamafa o le lima; hypothalamic-pituitary- adrenal axis; iloiloga o tusitusiga

O fetuutuunaiga i le feagai ai ma faigata faaletino poʻo le mafaufau e manaʻomia mo le ola, soifua maloloina, ma le manuia o meaola uma. O mea leaga e tutupu, e masani ona taʻua o "faʻalavelave," e amata ai se tali faʻaletino eseese mai le tele o punaoa, e aofia ai le faʻagaoioia o le hypothalamic-pituitary- adrenal (HPA) axis.¹ O le HPA axis e nafa ma le vaega o le glucocorticoid o le faʻalavelave faʻalavelave (faʻataʻitaʻiga, steroid hormone tali; cortisol i tagata, corticosterone i isumu ma isumu). O le fa'alilolilo o le Glucocorticoid e manatu e saofagā i le fetuuna'i fa'apopoleina e ala i le fa'atupuina o suiga umi i le fa'aaliga o kenera e ala i tali fa'ama'i adrenocorticosteroid (ie, mineralocorticoid receptor [MR] ma le glucocorticoid receptor [GR]). O le adrenocorticosteroid receptors e galue e pei o ligand-gated transcription factor (De Kloet et al. 1998) ae mafai foi ona faʻaogaina le faʻaliliuga e ala i le faʻalavelaveina o isi faʻasalalauga faʻasalalauga, e pei o le nuclear factor-kB (NF-kB) ma le activator protein-1 (AP-1). ) (Webster ma Cidlowski 1999). Glucocorticoids e mafai foi ona i ai ni aafiaga vave i le kemisi o le fai'ai ma amioga e ala i masini faailo o le membrane (De Kloet et al. 2008). Glucocorticoids e manatu e saofagā i le faʻamutaina o le amataga o le faʻalavelave faʻalavelave (Keller-Wood ma Dallman 1984) ma auai i le toe faʻaleleia o le homeostasis mo se taimi umi e mafua mai i le tali muamua (Munck et al. 1984).

¹ Mo le faauigaina o lenei ma isi faaupuga fa'apitoa, tagai i le Glossary, itulau 522–524.

Glucocorticoid fa'amamafa tali e mafai ona amataina e fa'alavelave fa'aletino (fa'atusaina tali fa'afoliga) po'o faiga fai'ai e feso'ota'i ai fa'ailoga si'osi'omaga ma i'uga le lelei. O le tali mulimuli lea e taʻua o le "psychogenic" e faʻamoemoe i le natura ma e aofia ai auala faiʻai e nafa ma polokalame puipuia faʻanatura poʻo le manatuaina o mea leaga (Herman et al. 2003). O le mea lea, o le tali a le psychogenic e fesoʻotaʻi ma le poto masani muamua, ma ua mamanuina e saunia ma le malosi le tino e aloese ai mai se taunuuga leaga poʻo le auai i amioga e mafai ona faʻateleina ai le gafatia mo le ola.

O le tele o fa'amaoniga e fa'ailoa mai ai o faiga fa'amamafa e tele sona sao i fa'agaioiga o vaisu, e aofia ai le fa'alagolago i le ava. Mo se faʻataʻitaʻiga, o le faʻaalia i le atuatuvale e mafai ona faʻatupuina le toe faʻafoʻi pe faʻateleina le faʻaaogaina o le ava (Sinha 2007). O gaioiga o le faʻamaʻi / glucocorticoids i luga o le ava malosi e mafai ona fesoʻotaʻi atu i le faʻaogaina o taui / faʻalavelave faʻalavelave, e aofia ai, mo se faʻataʻitaʻiga, faʻaleleia o le tuʻuina atu o le dopamine i totonu o le nucleus accumbens (Sutoo ma Akiyama 2002; Yavich ma Tiihonen 2000) ma le faʻagaioiina o le corticotropin-faʻasaʻo. vaega (CRF) ala (Heilig ma Koob 2007). O le mea moni, o le sootaga i le va o le ava malosi ma le atuatuvale e faigata ona o le aafia i le ava malosi, e pei o le tele o fualaau faasaina, e mafua ai le tuʻuina atu o glucocorticoids pe a faʻaalia ma mafai ai ona faʻavasegaina o se "faʻamalosi" o ituaiga (vaai Allen et al. . 2011).

O lenei tusiga o loʻo iloiloina le faʻatulagaina o neurocircuits e faʻatonutonu ai faʻalavelave faʻalavelave, e taulaʻi i le HPA axis, lea e taua tele i faiga faʻasuesue (silasila Marinelli ma Piazza 2002). O lo'o fa'atalanoaina ai fo'i vaega o feso'ota'iga i le va o le fa'alavelave ma ala o taui, aua e foliga mai e taua tele i le fa'atalanoaina o a'afiaga leaga o le fa'amamafa i le fa'aaogaina o vaila'au ma vaisu.

Circuitry Mediating the Reflexive Stress Response

O le HPA axis e pulea e neurons i totonu o le paraventricular nucleus (PVN) i le hypothalamus (silasila i le ata 1). O nei neu e natia le CRF ma le hormone vasopressin i totonu o le faʻasalalauga, lea e faʻaosoina ai le tuʻuina atu o le adrenocorticotropin hormone (ACTH) mai le pito i luma o le pituitary gland. E fealua'i le ACTH e ala i le ta'amilosaga fa'apitoa e o'o atu ai i le fa'ama'i o le tino, lea e fa'apipi'iina ma fa'asa'oloto ai le glucocorticoids (silasila Herman et al. 2003).

O tali fa'alavelave fa'afuase'i e tupu i taimi o fa'alavelave fa'afuase'i (fa'ata'ita'iga, fa'ama'i, matelaina, fa'amama le vai, po'o le te'i), pe a tatau ona tali atu le fai'ai i se lu'itau tele ile homeostasis e ala ile fa'agaoioia ole HPA axis. O faʻamatalaga faʻamatalaga e fesoʻotaʻi atu i le PVN e ala i neu muamua poʻo le lua, faʻatupuina se faʻagaioiga tuusaʻo o le faʻamalolo CRF (silasila Herman et al. 2003). Mo se faʻataʻitaʻiga, o le toto maualalo e fesoʻotaʻi ma le gau o le toto e tuʻuina atu e ala i neura faʻapitoa i neu faiʻai i le A2 catecholaminergic cell group (Palkovits ma Zaborszky 1977), ona faʻatautaia saʻo lea i le PVN (Cunningham ma Sawchenko 1988) ma faʻavave ai le faʻamalosia o le noradrenergic. CRF neurons (Plotsky et al. 1989).

I le faaopoopo atu i ala neural, o faʻamatalaga e uiga i suiga i le tulaga o le physiological e mafai foi ona tuʻuina atu e ala i mea faʻasalalau e fusifusia i vaega i fafo atu o le toto-faiʻai pa. Mo se faʻataʻitaʻiga, o le faʻateleina o le maualuga o le hormone angiotensin II (faʻailoga o le faʻamalo) e lagona e tali i totonu o le okeni subfornical (lea o loʻo tu i fafo atu o le paʻu o le toto-faiʻai ma faʻatonutonu le paleni o le vai), lea e tuʻuina saʻo le angiotensin II i le PVN CRF neurons, fa'afaigofie le fa'agaoioia o HPA (Plotsky et al. 1988). O nisi fa'aosofiaga fa'apitonu'u, e pei o le fulafula, e maua ai mea e mafai ona fa'ailo i le tele o masini; mo se faʻataʻitaʻiga, o le proinflammatory cytokine interleukin 1-b e foliga mai e faʻagaoioia le axis HPA e ala i alava neura lagona i le nerve vagus; le vaega postrema, lea e i fafo atu o le pa puipui o le toto-faiai; ma sela perivaskular i le itulagi o le vaega o le cell A2 (Ericsson et al. 1997; Lee et al. 1998; Wieczorek and Dunn 2006).

O vaila'au o le fa'aleagaina e mafai fo'i ona maua ai se tali muamua ole corticosterone e ala ile fai'ai ole PVN-projecting pathways. Mo se faʻataʻitaʻiga, o le faʻaalia muamua o le ava malosi e mafua ai le faʻamalolo o le ACTH ma le corticosterone, e ogatasi ma le ava malosi e avea o se faʻamalosi e le faʻamalosia (Allen et al. 2011). Acute HPA axis activation e ala i le ava malosi e ala i le fai'ai noradrenergic faiga (Allen et al. 2011). Ae ui i lea, o le fa'apa'iaina pea o le ava malosi e matua fa'afefeteina ai le fa'agaoioiga o le HPA i le tu'ina o le 'ava malosi (Rivier 1995), e fa'ailoa mai ai, i sina tulaga, e fa'asa'o a'afiaga HPA o le fa'aaogaina o le 'ava malosi i le aluga o taimi.

Circuitry Subserving Anticipatory Stress Responses: Le Limbic Stress-Control Network

Talu ai o "faʻalavelave faʻafuaseʻi" moni o le physiologic e seasea lava, o le tele o tali faʻalavelave o loʻo faʻamoemoeina i le natura, e aofia ai le faʻamatalaina o le faʻamataʻu o le faʻaosofia o le siosiomaga e tusa ai ma le poto masani muamua poʻo polokalama faʻanatura. O tali faʻalavelave faʻafuaseʻi e tele lava ina pulea e limbic forebrain structures, e pei o le hippocampus, medial prefrontal cortex (mPFC), ma amygdala (silasila Ulrich-Lai ma Herman 2009). O nei fausaga uma e maua uma faʻamatalaga faʻapitoa ma e aofia ai i le faʻatonutonuina o lagona, taui, ma lagona.

O faʻamaʻi o le faiʻai ma suʻesuʻega faʻaosofia e faʻaalia ai o le hippocampus e taofia ai le HPA axis. O le fa'aosoina eletise o le hippocampus e fa'aitiitia ai le fa'amalolo o le glucocorticoid i isumu ma tagata. Faʻaleagaina i le hippocampus, poʻo neura o loʻo ave ese mai ai (ie, lateral fornix), e mafua ai le faʻateleina o tali i faʻalavelave faʻalavelave o le mafaufau (faʻataʻitaʻiga, taofiofia) ma faʻaalia o se toe foʻi faaumiumi i le maualuga o le glucocorticoid (mo faʻamatalaga muamua, vaʻai Herman et al. 2003; Jacobson ma Sapolsky 1991). O nisi faʻamatalaga e taʻu mai ai o le hippocampus e faʻalavelaveina foi le gaioiga o le HPA axis, ae e le o vaʻaia lautele (Herman et al. 2003; Jacobson ma Sapolsky 1991). O aʻafiaga o le faʻaleagaina o le hippocampal i luga o le psychogenic HPA axis stress responses e mafai ona faʻaogaina i le ventral subiculum (vSUB), o le vaega autu o le subcortical output o le ventral hippocampus (Herman et al. 2003). O manua tuʻufaʻatasia o le vSUB i isumu e faʻaleleia ai le PVN CRF peptide ma le mRNA faʻamatalaga ma faʻateleina le tuʻuina atu o le corticosterone ma le faʻamalosia o le PVN (e pei ona fuafuaina e ala i le faʻaogaina o le FOS mRNA expression) i le tali atu i le taofiofia (Herman et al. 1998).

O le a'afiaga o le vSUB i le fa'atonuga o le fa'alavelave e fa'apitoa. O manu'a o le vSUB e fa'aumiumi tali atu a le HPA i mea fou ae le afaina ai tali fa'afoliga (fa'ata'ita'iga, i le eter inhalation) (Herman et al. 1998). O nisi faʻamaoniga e taʻu mai ai o le glucocorticoids o loʻo i ai se sao i le hippocampal inhibition of anticipatory tali, e pei o faʻamaʻi e mafai ona poloka faʻalavelave faʻaalia o le HPA axis e le synthetic steroid dexamethasone (Magarinos et al. 1987). E le gata i lea, o isumu ma le muaʻi mafaufau GR faʻamalo, e aofia ai le hippocampus, ua faʻateleina tali i le taofiofia ma le mea fou (ae le o le hypoxia) ma le faʻaleagaina o le dexamethasone le taofiofia o le corticosterone release (Boyle et al. 2005; Furay et al. 2008). Faʻatasi, o faʻamaumauga o loʻo faʻaalia ai o le hippocampus o loʻo faʻapitoa i le faʻatonutonuina o tali i faʻamaʻi faʻalavelave psychogenic, e tusa ai ma lana matafaioi i le gaioiga o le mafaufau ma lagona.

E le pei o le hippocampus, o le amygdala e fesoʻotaʻi ma le fiafia o le HPA axis. Amygdalar stimulation faʻamalosia le faʻamalolo o le glucocorticoid, ae o le tele o manuʻa o le amygdaloid faʻalavelave faʻaitiitia le gaioiga o le HPA (silasila Herman et al. 2003). Ae ui i lea, o loʻo i ai se faʻailoga faʻapitoa faʻapitoa faʻapitoa o galuega faʻalavelave-faʻatasi i totonu ole amygdala. O le totonugalemu o le amygdala (CeA) e matuaʻi tali atu i faʻalavelave faʻalavelave o le homeostatic, e pei o le mumū ma le gau o le toto (Dayas et al. 2001; Sawchenko et al. 2000). Lesions o le CeA attenuate HPA axis tali i nei ituaiga o faʻamalosi ae le o le taofiofia (Dayas et al. 1999; Prewitt and Herman 1997; Xu et al. 1999). I se faʻatusatusaga, o le ogatotonu o le amygdala (MeA) o loʻo faʻaalia ai tali faʻapitoa a le FOS i faʻamalosi, e pei o le taofiofia (Dayas et al. 2001; Sawchenko et al. 2000). O faʻamaʻi o le MeA e faʻaitiitia ai tali a le HPA i le taofiofia ma le malamalama ma le leo faʻamalosi ae le o le tui faʻaogaina o le protein interleukin 1-b poʻo le ether inhalation (Dayas et al. 1999; Feldman et al. 1994). O le mea lea, e foliga mai e mafai ona faʻatulafonoina tali faʻafouga ma faʻamoemoe i se vaega e ala i le amygdaloid circuitry discrete.

O le mPFC e foliga mai o loʻo i ai se sao faʻalavelave i le faʻatonutonuina o faʻalavelave. O vaega uma o le rodent PFC o loʻo faʻamalosia malosi e ala i le faʻalavelave mamafa. Ae ui i lea, o taunuuga o le physiological o le faʻamalosia o le faʻamalosia e foliga mai e eseese i le itulagi. O le vaega muamua o le mPFC (PL) e taua tele i le faʻalavelave faʻalavelave ona o le tele o suʻesuʻega ua faʻaalia ai o le faʻaleagaina o lenei itulagi e faʻaumiina ai tali a le HPA i faʻalavelave faʻafuaseʻi psychogenic (ae le o le homeostatic) (Diorio et al. 1993; Figueiredo et al. 2003; Radley et al. 2006), ae o le faʻamalosi e taofia ai le faʻalavelave faʻalavelave (Jones et al. 2011). O le mPFC e foliga mai o se nofoaga mo faʻamatalaga glucocorticoid o tali a le HPA ona o mea faʻapipiʻi glucocorticoid i le lotoifale e taofia ai le faʻamoemoe (ae le o le faʻaalia) tali i faʻalavelave (Akana et al. 2001; Diorio et al. 1993). I le faʻatusatusaga, o faʻamaʻi e faʻatatau i le tele o le infralimbic PFC (IL) e matuaʻi ese le aʻafiaga o le tino. Faʻaleagaina o le IL faʻaitiitia ai tali tutoʻatasi i faʻamaʻi o le mafaufau (Tavares et al. 2009) ma faʻapipiʻi foi le faʻamalosia o le PVN Fos i le tali atu i le taofiofia (Radley et al. 2006). O le mea lea, o le PL ma le IL e foliga mai o loʻo i ai aʻafiaga faʻafeagai i luga o le faʻalavelave faʻatasi.

Fa'agaoioi le Relay: Limbic-PVN Networks

O le faʻamalosia o le PVN e le hippocampus, prefrontal cortex, ma le amygdala e faʻatapulaʻaina. O le mea lea, o le fa'atonutonuina o galuega a le HPA e ala i nei fausaga e mana'omia ai feso'ota'iga feso'ota'iga (silasila i le ata 2). O suʻesuʻega e suʻe ai faʻataʻitaʻiga mai le tasi vaega o le faiʻai i le isi (faʻataʻitaʻiga, suʻesuʻega suʻesuʻega) e faʻaalia ai le mafai ona maua fesoʻotaʻiga bisynaptic limbic-PVN e fealuaʻi ai le tele o vaega pito i lalo, e aofia ai le moega o le stria terminalis (BNST), dorsomedial hypothalamus. , Medial preoptic area, ma peri-PVN itulagi (e aofia ai le subparaventricular nucleus) (Cullinan et al. 1993; Prewitt ma Herman 1998; Vertes 2004). O suʻesuʻega faʻalua e faʻaalia ai o neura o loʻo ave ese mai le vSUB, MeA, ma le CeA (ie, neura efferent) faʻafesoʻotaʻi saʻo PVN-projecting neurons i nei itulagi, e ogatasi ma fesoʻotaʻiga galue (Cullinan et al. 1993; Prewitt ma Herman 1998) .

O aʻafiaga eseese o le PL ma le IL i luga o faiga faʻalavelave faʻalavelave e mafai ona atagia mai ai lo latou eseesega iloga i sini faʻavae. O le PL o loʻo i ai le tele o faʻataʻitaʻiga e faʻatau ai auala talafeagai, e aofia ai le nucleus accumbens ma le amygdala basolateral, faʻapea foʻi ma le BNST pito i tua, lea e fesoʻotaʻi ma le HPA axis inhibition. I se faʻatusatusaga, o le IL e tele fesoʻotaʻiga faʻatasi ma itulagi o loʻo aʻafia i tulafono faatonutonu tutoʻatasi, e aofia ai le CeA, nucleus of the solitary tract (NTS), anteroventral BNST, ma le dorsomedial hypothalamus (Vertes 2004). O le mea lea, e foliga mai o le aoga o le faʻamalosia o le PFC e manaʻomia ai le tuʻufaʻatasia o le PL ma le IL.

O le faʻamatalaga, mPFC, hippocampal, ma amygdalar efferents e masani ona faʻapipiʻi i itulagi o loʻo tuʻuina atu le γ-aminobutyric acid (GABA) -o loʻo ave faʻataʻitaʻiga i le PVN (silasila i le ata 2). O le mea moni, o le numera tele o PVN-projecting neurons e GABAergic i le phenotype. O faʻasologa o neurons o le vSUB (faʻapea foʻi ma le mPFC) e glutamatergic i le natura, ma faʻaalia ai o nei sela e auai i le transsynaptic inhibition o le PVN pe a uma ona faʻagaoioia e ala i le atuatuvale. I se faʻatusatusaga, o le faʻasologa o neurons o le MeA ma le CeA e sili atu le GABAergic, o loʻo fautua mai ai o le amygdalar excitation o le PVN e faʻatalanoaina e ala i le faʻaleagaina, e aofia ai faʻasologa GABA synapses (Herman et al. 2003).

O le BNST e sili ona fiafia i ai, ona o le mauaina o mea e faʻaogaina mai i le tele o faʻamaʻi faʻapipiʻi faʻapipiʻiina (CeA, MeA, vSUB, IL, ma PL) (Cullinan et al. 1993; Dong et al. 2001; Vertes 2004) . O le faʻamatalaga, o vaega laiti a le BNST e foliga mai e nafa ma le faʻalavelave faʻatasi ma le faʻaosofia o tali faʻalavelave HPA. Mo se faʻataʻitaʻiga, o faʻamaʻi o le itu pito i tua o le BNST e faʻateleina ai le maualuga o le ACTH ma le corticosterone faʻamalolo ma le PVN Fos activation (Choi et al. 2007), e faʻaalia ai se sao i le tuʻufaʻatasia tutotonu o le faʻalavelave faʻalavelave. O faʻamaʻi o le vaega anteroventral o le BNST e faʻaleleia ai foi tali faʻalavelave (Radley et al. 2009). I se faʻatusatusaga, o faʻamaʻi tetele o le BNST pito i luma e faʻaitiitia ai tali faʻamamafa o le HPA (Choi et al. 2007), e ogatasi ma se matafaioi mo lenei itulagi i le faʻalavelave faʻalavelave. O le mea lea, o le matafaioi a le BNST i le faʻalavelave faʻalavelave faʻatasi ma le faʻagaoioia o loʻo tuʻufaʻatasia ma e mafai ona fesoʻotaʻi ma eseesega i le limbic tulimataʻi o vaega taʻitasi o le BNST. Mo se faʻataʻitaʻiga, o le BNST medial pito i tua e maua ai le mamafa mai le vSUB ma le MeA, ae o le vaega o le anteroventral e maua ai faʻamatalaga mai le CeA ma le tele o le IL efferents (Canteras ma Swanson 1992; Cullinan et al. 1993; Dong et al. 2001; Vertes. 2004).

O le vaega pito i luga ole vaeluagalemu ma itu-PVN e matua'i tumu i neu GABAergic ma e foliga mai.

e fa'amuamua le fa'aogaina o le fa'alavelave fa'alavelave (Herman et al. 2003). Neurons i nei itulagi e talitonuina e tuʻuina atu le tonic inhibition i le PVN, lea e mafai ona fetuutuunai e tusa ai ma mea faʻapipiʻi glutamate mai le vSUB (faʻamalosia le faʻaogaina) poʻo le GABAergic mea faʻapitoa mai le MeA (disinhibition). O faʻamaʻi o le medial preoptic nucleus faʻateleina tali faʻamaʻi o le HPA ma poloka tali a le HPA e mafua mai i le medial amygdalar stimulation, e fautua mai ai se matafaioi muamua i le faʻalavelave faʻalavelave (mo faʻamatalaga muamua, vaʻai Herman et al. 2003). O le faʻalavelave faʻapitonuʻu o le faʻailoga o le glutamate i totonu o le peri-PVN e faʻaleleia ai foi le faʻamalosia o tali a le HPA (Ziegler ma Herman 2000), e faʻapea o le axons limbic e muta i lenei itulagi e mafai ona faʻaogaina le PVN activation.

E sili atu le faigata ona faʻamaonia le matafaioi a isi itulagi o le hypothalamic e fesoʻotaʻi ai le limbic efferents i le PVN, e pei o le dorsomedial nucleus (Herman et al. 2003). Mo se faʻataʻitaʻiga, o taunuuga feteʻenaʻi e matauina pe a maeʻa le faʻamaʻi, faʻagaoioia, poʻo le le faʻaaogaina o lenei hypothalamus dorsomedial, atonu ona o le mamafa o le fefiloi o le glutamate ma le GABA neuronal faitau aofaʻi (Herman et al. 2003).

O lo'o tumau pea le su'esu'eina atoatoa o isi fa'amea fa'aonaponei. Mo se faʻataʻitaʻiga, o le raphe nuclei ma le NTS e faʻapipiʻiina le PVN, o loʻo faʻamoemoeina e le limbic structures (e pei o le PL) (silasila i le Vertes 2004) ma o loʻo aʻafia i le faʻalavelave faʻalavelave e le serotonin ma le norepinephrine (Herman et al. 2003), i le faasologa. Ae ui i lea, i le taimi nei, e leai ni suʻesuʻega faʻapitoa e faʻamatalaina bisynaptic limbic-PVN relays i nei itulagi.

Circuitry Subserving Chronic Stress Responses

O le fa'aumiumi po'o le fa'alauteleina o le fa'apopoleina e mafua ai le fa'atupuina o le fa'atonuga umi o le HPA axis, e fa'aalia i le fa'aitiitia o le mamafa o le thymus (fa'atatau i le fa'atuputeleina o le maualuga i GCs); faʻateleina le tele o le tino (faʻatatau i le faʻalauteleina o le ACTH); faateleina le lagona o le tino ile ACTH; fa'afaigofie tali a le HPA i fa'alavelave fou; ma i nisi (ae

e le o mea uma) faʻataʻitaʻiga / tulaga, faʻasaʻoina o le basal GC secretion (silasila Herman et al. 1995; Ulrich-Lai et al. 2006). Suiga i le tuʻuina atu o le hormone peripheral o loʻo faʻatasi ma le faateleina o le PVN CRF ma le vasopressin mRNA (Herman et al. 1995), o loʻo fautua mai ai o le HPA upregulation o loʻo tuʻufaʻatasia. E le gata i lea, o le faʻalavelave masani e faʻateleina ai le glutamatergic ma le noradrenergic terminal e faʻatatau i le PVN CRF neuronal somata ma dendrites, e ogatasi ma le faʻaleleia o le synaptic drive (Flak et al. 2009).

O auala tutotonu o le fa'agaioiina o le axis HPA tumau e le'i fuafuaina. O le matafaioi a le limbic forebrain i le faʻalavelave faʻalavelave e taʻu mai ai o le eseesega o le auai o le PFC, hippocampus, ma amygdala atonu e nafa ma le taʻavale umi. O le faʻamatalaga, o itulagi uma o loʻo faʻaalia le tele o suiga o le neuroplastic faʻaosoina: Dendritic retraction o loʻo manino i le hippocampal ma le mPFC pyramidal neurons, ae o le faʻalauteleina o le dendritic o loʻo matauina i le amygdala (mo faʻamatalaga muamua, vaʻai Ulrich-Lai ma Herman 2009). O nei suʻesuʻega e ogatasi ma le toe tufatufaina atu o le faʻaogaina o le limbic i le HPA excitatory circuits, faʻamalosia le faʻamalosi i luga o le faʻasaina.

O le faʻaleleia o le taʻavale amygdalar ua faʻamoemoe e faia se sao taua i faʻamaʻi faʻamaʻi masani. Mo se faʻataʻitaʻiga, o le faʻalavelave masani e faʻagaoioia ai le faiga o le CeA CRF, lea na faʻatulagaina e avea o se ala faʻatupu faʻalavelave faʻalavelave (Dallman et al. 2003). Ae ui i lea, o le CeA e foliga mai e le manaʻomia mo le atinaʻeina poʻo le tausia o faʻamaʻi faʻamaʻi masani (Solomon et al. 2010). E le gata i lea, o manua o le MeA e le mafai foi ona taofia le faʻalavelave faʻafuaseʻi o le faʻamaʻi o le HPA axis (Solomon et al. 2010). O le mea lea, o le fesoʻotaʻiga lautele i le va o le amygdalar hyperactivity ma le faʻalavelave faʻafuaseʻi-faʻaleagaina o le HPA axis e leʻi faʻamautuina.

O le paraventricular nucleus o le hypothalamus (PVT) e foliga mai o loʻo aofia ai se vaega o le ala masani-faʻalavelave. O manuʻa o le PVT poloka faʻamaʻi faʻamaʻi masani o tali a le HPA i faʻalavelave faʻalavelave (Bhatnagar ma Dallman 1998), faʻaalia ai se matafaioi muamua i le faʻagasologa o le faʻaogaina.

E le gata i lea, o faʻamaʻi PVT faʻalavelaveina le faʻagasologa o le faʻaogaina o le HPA axis habituation i faʻalavelave faʻafuaseʻi (Bhatnagar et al. 2002). Ile tu'ufa'atasia, o fa'amaumauga o lo'o fa'ailoa mai ai o le PVT e tele sona sao i le fa'aogaina o le HPA axis drive i le tulaga o le fa'aumiumi o le fa'alavelave. O le faʻamatalaga, o le PVT ma le limbic forebrain nofoaga e pulea ai faʻalavelave faʻalavelave faʻalavelave e fesoʻotaʻi (vaʻai Vertes ma Hoover 2008), faʻatagaina mo le faʻamaopoopoina o gaioiga faʻamalosi corticolimbic i lenei itulagi. O loʻo faʻatulagaina foʻi le PVT e faʻatautaia faʻamatalaga e uiga i le faʻaauauina o le physiological tulaga, mauaina o mea e faʻaogaina mai neurons orexinergic (lea e faʻatonutonuina le tuʻuina atu o le acetylcholine, serotonin, ma le noradrenaline) o le dorsolateral hypothalamus (lea e taʻalo ai se sao taua i le puleaina o gaioiga faʻafefe) ma le faiʻai aʻe. faiga e a'afia i le pulea tuto'atasi.

O lo'o fa'atulagaina fo'i le BNST e tu'ufa'atasia fa'amatalaga e uiga i fa'alavelave fa'aumi. O manuʻa o le BNST anteroventral e faʻaitiitia ai tali i le faʻalavelave faʻalavelave, ae faʻamalosia le faʻaogaina o le HPA axis e ala i le faʻalavelave masani (Choi et al. 2008). O nei faʻamatalaga o loʻo faʻaalia ai o lenei itulagi o loʻo i ai le tumau-faʻalagolago i matafaioi i le HPA axis control, faʻatasi ai ma le eseese o tagata neural na faʻafaigaluegaina i se taumafaiga e tali atu i le faʻalavelave faʻaumiumi. Ona o fesoʻotaʻiga vavalalata i le va o le BNST ma le mPFC, hippocampus, ma amygdala, e ono mafai ona "toe faʻatulagaina" neu BNST e ala i suiga masani o le faʻalavelave i le gaioiga o le limbic poʻo le faʻaogaina o mamanu.

Si'o Fa'atiga ma le 'Ava

O le aufaitau e masani i tusitusiga o le ava o le a mautinoa lava latou te maua le tele o feso'ota'iga i le va o feso'ota'iga fa'apopoleina o lo'o fa'amatalaina i luga ma le va'aiga o fai'ai e feso'ota'i ma le inuina o le ava. Mo se faʻataʻitaʻiga, o faʻamaumauga tele e lagolagoina ai se matafaioi mo le CeA, BNST, ma le noradrenergic system i le tausiga o le faʻalagolago i le ava (silasila i le Koob 2009), o loʻo fautua mai ai o le faagasologa o vaisu e fesoʻotaʻi ma le faʻagaoioia o le atuatuvale (ma le HPA axis) auala faʻamalosi. O le mea moni, o le faʻaleleia o le CeA / BNST CRF faʻamatalaga e foliga tutusa ma le mea o le a faʻamoemoeina pe a maeʻa le faʻalavelave masani, e oʻo atu ai i le manatu e faʻapea o tulaga le lelei o vaisu (faʻataʻitaʻiga, aloese mai le faʻamavaeina) e fesoʻotaʻi ma le faʻaaogaina o le ava malosi o faʻalavelave faʻalavelave masani (Koob 2009). I le isi itu, o le faʻagaoioia o ala o taui e iloa e matua puipuia ai le faʻalavelave faʻalavelave e ala i le amygdaloid complex, e faʻaalia ai se faiga e mafai ai e aʻafiaga tauia o le ava malosi ona faʻaitiitia ai le atuatuvale (Ulrich-Lai et al. 2010).

O le ava malosi foi e i ai aafiaga loloto i luga o le medial prefrontal cortical neural activity, ma o le faʻaaogaina masani e fesoʻotaʻi ma le faʻaogaina muamua (le lelei le pulea o le manaʻo) i totonu o tagata (silasila Abernathy et al. 2010). O galuega faatino a le mPFC i le CeA ma le BNST ma, a itiiti mai i le tulaga o le vaega muamua, o loʻo i ai se sao taua i le HPA faʻasaina. Faatasi ai ma le mauaina o galuega o loʻo vaʻaia i le amygdalar-BNST circuits, o nei faʻamatalaga o loʻo fautua mai ai o le faʻaaogaina o le ava malosi e mafua ai suiga iloga i luga o le fesoʻotaʻiga o le faʻamaʻiina o le limbic, faʻaituau le tino mo le faʻalavelave faʻalavelave.

I le aotelega, o le pulea lelei o le HPA axis o se mana'oga mo le ola pupuu ma le umi. Talu ai ona o mea autu e pulea ai le gaioiga o le HPA o loʻo taulaʻi i le ava malosi, ma o le ava lava ia e fai ma faʻamataʻu, e le o se mea e ofo ai le corticosteroids, le "pito pisinisi" o le axis, e iai fegalegaleaiga loloto ma le amio ma le physiological tulafono faatonutonu o taumafa. O le felavasa'i ile va ole HPA fa'atonutonu ma vaisu o lo'o fa'ailoa mai ai itu taua e ono fa'atatau mo a'afiaga fa'aleaogaina umi ole ava fa'apea le fa'alagolago lava ia. O le taua o le so'oga fa'amanino o lo'o fa'amamafaina atili e le malosi o le fa'afeagai i le va o le popole i le olaga ma le inu, lea e fa'alavelave ai taumafaiga e fa'avae ma fa'atumauina le fa'amama.

Faʻafetai

O lenei galuega sa lagolagoina e fesoasoani MH–049698, MH–069680, ma MH–069725.

Faailoaina o Tupe

Fai mai le tusitala e leai sana mea tau tupe e tauva ai.

mau faasino

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