Moputso, dopamine le taolo ea ho ja lijo: moelelo oa botenya (2011)

Litsela tsa Cogn Sci. 2011 Jan; 15 (1): 37-46. doi: 10.1016 / j.tics.2010.11.001. Epub 2010 Nov 24.

Volkow ND, Wang GJ, Baler RD.

mohloli o moholo

Mokhatlo oa Sechaba oa Tlhekefetso ea Lithethefatsi, Mekhatlo ea Sechaba ea Bophelo, Bethesda, MD 20892, USA. [imeile e sirelelitsoe]

inahaneloang

Bokhoni ba ho hanela tšusumetso ea ho ja bo hloka hore ho sebetse hantle ha lipotoloho tsa neuronal tse amehang ka holimo-tlaase ho laola ho hanyetsa likarabo tse hlokomelisitsoeng tse bolellang moputso oa ho ja lijo le takatso ea ho ja lijo. Kelithuto tsa boselamose li bontša hore lihlooho tse ngata li ka 'na tsa e-ba le tšenyo e mpe litseleng tsa dopaminergic tse laolang tsamaiso ea neuronal e amanang le moputso oa tlhokomelo, boemo bo botle le taolo. Hoa tsebahala hore li-neuropeptides tse laolang matla a motlakase (homeostatic tshebetso) ho tswa ho hypothalamus li boetse li etsisa mosebetsi oa lisele tsa dopamine le lihlahisoa tsa tsona libakeng tse amehang mekhoeng e khotsofatsang e tlisoang ke ho ja lijo. Ho boleloa hore sena se ka boela sa e-ba mokhoa oo ka oona ho ja haholo le ho hanyetsa matšoao a homoeostatic ho fokotsa mosebetsi oa lipotoloho tse amanang le ts'ebeliso ea moputso, boemo bo botle le taolo ea ts'oaetso.

Selelekela

Palo ea boraro ea batho ba baholo ba United States ke boima haholo [boima ba 'mele (BMI) ≥30 kg m-2] [1]. Taba ena e na le liphello tse tebileng haholo, hobane botenya bo amahanngoa haholo le mathata a tebileng a meriana (mohlala, lefu la tsoekere, lefu la pelo, sebete sa mafura le kankere e meng) [2]. Ha ho makatse hore ebe litšenyehelo tsa tlhokomelo ea bophelo bo botle feela ka lebaka la ho nona haholo US li hakanngoa li le haufi le US $ 150 [3].

Ha ho pelaelo hore litaba tsa setso le tsa setso li tlatsetsa ho seoa sena. Ka ho khetheha, libaka tse khothalletsang mekhoa e sa jeoang ea lijo (ho fumaneha habonolo ho lijo tse ntseng li sebelisoa haholo le tse sa jekeng) ho lumeloa hore ho na le karolo ea bohlokoa bothateng bo atileng ba ho nona haholo (Websaeteng ea Boima bo Feteletseng le Boima ba Mafura; http://www.cdc.gov/obesity/index.html). Leha ho le joalo, mabaka ka bomong a boetse a thusa ho tseba hore na ke mang ea tla (kapa a ke ke a ba mafolofolo) libakeng tsena. E itšetlehile ka lithuto tsa lefutso, lihlahisoa tsa lefutso li hakanyetsoa ho kenya letsoho pakeng tsa 45% le 85% ea ho fetoha ha BMI [4,5]. Le hoja liphuputso tsa liphatsa tsa lefutso li senoletse ntlha ea liphetoho tse phahametseng batho ba bangata haholo [4], ka kakaretso, botenya bo nahanoa hore bo tlas'a taolo ea polygen [6,7]. Ka sebele, thuto ea morao-rao ea tlhahlobo ea liphatsa tsa lefutso (GWAS) e entsoeng ka 249,796 batho ba tsoang Europe ba khethollang 32 loci e amanang le BMI. Leha ho le joalo, loci ena e hlalosa feela 1.5% ea phapang ea BMI [8]. Ho feta moo, ho hakanngoa hore lithuto tsa GWAS tse nang le lisebelisoa tse khōloanyane li ka tseba ho tseba lintlha tse ling tsa 250 le liphello ho BMI. Leha ho le joalo, esita le ka mefuta e fapaneng e sa tsejoeng, ho hakanngoa hore lipontšo tse tsoang ho tse ling tse tloaelehileng tsa loci li ne li tla ikarabella feela ka 6-11% ea liphatsa tsa lefutso tsa BMI (ho itšetlehile ka ho lekanyelitsoe ho hlokahala ha 40-70%). Tlhaloso e lekanyelitsoeng ea ho fapana ha lipatlisiso tsena tsa liphatsa tsa lefutso e ka 'na ea bonahatsa litšebelisano tse rarahaneng pakeng tsa lits'ebetso tse fapaneng (joalo ka ho khethoa ke liphatsa tsa lefutso) le tsela eo ka eona batho ba amanang le libaka tseo ho tsona lijo li fumanehang haholo, eseng feela mohloli oa phepo, empa hape e le moputso o matla oo ka boeona o khothalletsang ho ja [9].

Hypothalamus [ka li-neuropeptides tse laolang, tse kang leptin, cholecystokinin (CCK), ghrelin, orexin, insulin, neuropeptide Y (NPY), le ka ho utloahala ha limatlafatsi, tse kang glucose, amino acid le mafura a acids] e nkoa e le boko bo ka sehloohong sebaka se laolang ho ja lijo ha se amana le litlhoko tsa caloric le phepo e nepahetseng [10-13]. Haholo-holo, mokokotlo oa khohlopo ka ho kopana ha oona le li-nuclei tse ling tsa hypothalamic le libaka tse ling tsa boko bo feteletseng, tse kenyeletsang nucleus tractus solitarius, li laola lijo tsa homeostatic [12] 'me e ameha ka ho nona haholo [14-16] (Setšoantšo sa 1a, lehlakoreng le letšehali). Leha ho le joalo, bopaki bo ntse bo bokella hore boko bo potoloha ntle le ba laolang tlala le satiety ba kenang lijong tsa lijo le botenyay [17]. Ha e le hantle, limbōpeho tse 'maloa [nucleus accumbens (NAc), amygdala le hippocampus] le libaka tsa cortical boko [orbitofrontal cortex (OFC), cingulate gyrus (ACC) le li-insula] le mekhoa ea neurotransmitter (dopamine, serotonin, opioids le cannabinoids) hammoho le hypothalamus li ameha ka liphello tse putsang tsa lijo [18] (Setšoantšo sa 1a, lehlakoreng le letona). Ka lehlakoreng le leng, tsamaiso ea phepelo ea lijo ka hypothalamus e bonahala e itšetlehile ka moputso le ho susumetsa methapo ea pelo ho fetola boitšoaro ba lijo [19-21].

Setšoantšo sa 1  

Tlhophiso ea ho ja lijo e itšetlehile ka puisano e mengata pakeng tsa moputso o feteletseng le methapo ea methapo ea homeostatic. (A) Setšoantšo sa kemiso sa mokokotlo o pakeng tsa homeostatic (hypothalamus, HYP) le lipotoloho tsa moputso tse laolang ho ja lijo. The ...

Ho itšetlehile ka liphuputso tse entsoeng ke lithuto tsa litšoantšo, mohlala oa ho nona haholo o sa tsoa etsoa tlhahiso ea hore ho ja haholo ho bonahatsa ho se leka-lekane pakeng tsa lipotoloho tse susumetsang boitšoaro (ka lebaka la ho kenella ha bona moputsong le maemo) le lipotoloho tse laolang le ho thibela likarabo tse potlakileng pele [22]. Mokhoa ona o khetholla lipotoloho tse 'nè tse ka sehloohong: (i) moputso-saliency; (ii) motlakase-koloi; (iii) maemo a ho ithuta; le (iv) taolo ea ho thibela-e leng molao oa tsamaiso-tsamaiso ea mosebetsi. Ka ho hlakileng, mohlala ona o boetse oa sebetsa ho lemalla lithethefatsi.

Ibatho ba tlokotsing, ho sebelisoa ha bongata ba lijo tse monate (kapa lithethefatsi tse lemalla) e ka ferekanya ho sebelisana ka tsela e leka-lekaneng pakeng tsa lipotoloho tsena, e leng se hlahisang boleng bo matlafatsang ba lijo (kapa lithethefatsi tse lemalla) le ho fokotseha ha lipotoloho tsa taolo. Ho senyeha ha sena ke phello ea ho ithuta ka boemo bo hlophisitsoeng le ho tsosolosa meeli ea moputso ka mor'a ho sebelisoa ha lijo tse ngata tse phahameng haholo (kapa lithethefatsi tse lemalla) ka batho ba kotsing. Ho senyeha ha marang-rang a marang-rang a holimo-limo a laolang likarabo tsa pele ho moo ho fella ka ho se tsitsisehe le ka ho qeta lijo (kapa ho noa lithethefatsi ho qobella ho lemalla).

Leqephe lena le buisana ka bopaki bo bontsang lipotoloho tsa neural tse amehang holimo-tlaase taolo le ba amehang ka moputso le tšusumetso le ho sebelisana ha bona le matšoao a maiketsetso a laolang lijo tsa homeostatic.

Lijo ke moputso oa tlhaho oa tlhaho le ts'ebeliso ea maemo

Lijo tse ling, haholo-holo tse nang le tsoekere le mafura, ke moputso o matla [23] tse khothalletsang ho ja (esita le ha ho se na tlhokahalo e matla) le ho etsa hore litloaelano li ithute pakeng tsa ts'ebetso le moputso (boemo bo botle). Ka mokhoa oa ho iphetola ha lintho, thepa ena ea lijo tse monate e sebelisetsoa ho ba molemo hobane e netefalitse hore lijo li jeoa ha li fumaneha, ho nolofalletsa matla ho boloka 'mele (joaloka mafura) bakeng sa tlhoko ea nako e tlang metseng eo ho eona mohloli oa lijo o neng o haella le / kapa o sa tšepahaleng. Leha ho le joalo, mekhatlo ea mehleng ea kajeno, moo lijo li fumanehang teng, mokhoa ona o fetohile boikarabelo.

Li-neurotransmitters tse 'maloa, tse akarelletsang dopamine (DA), li-cannabinoids, opioids le serotonin, hammoho le li-neuropetides tse amehang ka taolo ea homeostatic ea lijo, tse kang orexin, leptin le ghrelin, li ameha liphellong tse putsang tsa lijo [24-26]. DA e 'nile ea etsoa lipatlisiso ka ho fetisisa ebile e tsebahala ka ho fetisisa. Ke mohloli oa bohlokoa oa phekolo ea methapo ea meriana ea methapo ea meriana (mehloli ea tlhaho le ea lithethefatsi), eo e sebetsang ka ho khetheha ho latela sepheo sa ventral tegmental (VTA) ho NAc [27]. Litebello tse ling tsa DA li boetse li kenyelelitsoe, ho akarelletsa le dorsal striatum (caudate and putamen), cortical (OFC le ACC) le libaka tsa limbic (hippocampus le amygdala) le lateral hypothalamus. Ha e le hantle, ho batho, ho kenngoa lijo tse monate ho bontšitsoe ho lokolla DA maemong a fokolang ho latela mokhoa oa ho ithabisa o fumanoang ke ho ja lijo [28]. Leha ho le joalo, ts'ebetso ea DA e le moputso e rarahane haholo ho feta ho kopa kholekho ea bohlokoa ea hedonic. Ha ho qala ho pepesehela moputso oa lijo (kapa moputso o sa lebelloang), ho thunngoa ha DA neurons ho VTA ho eketseha ka lebaka la ho eketseha ha DA ho lokolloa ha NAc [29]. Leha ho le joalo, ka ho pheta-pheta khafetsa molemong oa lijo, karabo ea DA e tloaetse ho etsoa 'me e fetisetsoa butle-butle holim'a mohopolo o amanang le moputso oa lijo (mohlala, monko oa lijo), o sebetsanoang e le mokhoa oa ho fumana moputso ho moputso) [30,31]; letšoao la DA le arabelang ka khokahano eo e sebeletsa ho fetisa 'phoso ea ho bolela esale pele ea moputso' [31]. Litho tse ngata tsa glutamatergic li ama li-neurone tsa DA tse tsoang libakeng tse nang le boikutlo bo botle (home insulator kapa li-gustatory cortex), homeostatic (hypothalamus), moputso (NAc), maikutlo (amygdala le hippocampus) le multimodal (OFC bakeng sa sebete) meputso le litlhoko tsa maemo a itseng [32]. Haholo-holo, liphuputso tse tsoang ho amygdala le OFC ho DA neurons le NAc li kenngoa litabeng tsa boemo ba lijo [33]. Ha e le hantle, lipatlisiso tsa litšoantšo li bontšitse hore ha lihlooho tsa banna tse se nang bobebe li ne li botsoa ho thibela takatso ea tsona ea lijo ha li ntse li pepesoa lijo, li fokotseha ts'ebetso ea metabolism ho amygdala le OFC [hammoho le hippocampus (bona le lebokose 1), lihlekehlekeng le lipapaling]; ho fokotseha ha OFC ho ne ho amahanngoa le ho fokotseha ha takatso ea lijo [34].

Lebokose 1. Karolo ea hippocampase ka mekhoa ea ho fepa

Hippocampus hase eona feela e bohareng ba mohopolo, empa e boetse e ameha tabeng ea ho laola boitšoaro ba lijo ka ho ntlafatsa mekhoa e metle (ho kenyeletsa ho hopola hore na motho o ja joang, ho hopola mekhatlo ea boemo bo botle, ho hopola hore na lijo li teng, ho khetholla linaha tsa ho ba le tlala le ho hopola kamoo ho lokisa maemo ana). Ka mohlala, ka litoeba, lisele tse khethileng li-hippocampus li sitoa ho khetholla pakeng tsa tlala le tsa ho ba le bothata [99] 'me, ka likhoto tsa basali, e ile ea fella ka hyperphagia [100]. Ho batho, lipatlisiso tsa ho nahana ka boko ba boletse ho ts'ebetsoa ha hippocampus ka takatso ea lijo, boemo ba tlala, karabo ho litlamorao tsa lijo le ho latsoa lijo [101]. Hippocampus e hlalosa likarolo tse phahameng tsa insulin, ghrelin, glucocorticoids le li-receptor tsa CB1 tsa kannete, e leng se bontšang hore sebaka sena se boetse se laola lijo tsa lijo ka mekhoa e seng ea methe [102,103]. Ho phaella moo, hippocampus e amehile ka ho nona haholo, joalo ka ha ho hlalositsoe lithuto tse bontšang hore ka bobebe empa e se batho ba mahlahahlaha, hippocampus e bontša ho hlekefetsoa ha ho arajoa lijo tse matlafatsang [104].

Litokomane tse tsamaisitsoeng li ka etsa hore li fepe le litoeba tse sated [30] 'me, ho batho, lipatlisiso tsa litšoantšo li bontšitse hore ho pepesehela lijo tse nang le lijo ho etsa hore DA e eketsehe ka statum e amanang le takatso ea ho ja lijo [35]. Ntle le ho ameha ha eona le maemo, DA e boetse e ameha molemong oa ho etsa boitšoaro bo hlokahalang ho fumana le ho ja lijo. Ka sebele, ts'ebetsong ea DA bakeng sa moputso oa lijo e 'nile ea amahanngoa le matla a ho susumetsa kapa ho' batla 'lijo ho fapana le' ho rata 'lijo [36] (lebokose 2), phello e ka 'nang ea kenyelletsa striatum ea marang-rang le mohlomong le NAc [37]. DA e na le karolo ea bohlokoa haholo tabeng ena ea hore litoeba tse sa hlahiseng DA li bolaoa ke tlala ka lebaka la ho hloka matla a ho ja [37]. Ho tsosolosa ts'ebetso ea phekolo ea meriana ea lefapha la phekolo ea lichelete ea lefats'e ea lichelete ka ntle ho bolokolohi ba liphoofolo tsena, empa ho e tsosolosa le NAc ha ho joalo.

Lebokose 2. Ho batla ho bapisa le ho rata: phapang ea bohlokoa

Mekhoa ea meputso ea bokooa e amehang ka ho ja lijo ho khetholla mokhoa o amehang oa ho susumetsa takatso ea lijo, eo ho thoeng ke 'ho batla', ho latela mokhoa o amehang ka lihlahisoa tsa hedonic tsa lijo, tseo ho thoeng ke 'ho rata'36]. Le hoja tsamaiso ea dopamine e hlaselang haholo (le hoja e se feela) e amanang le ho 'batla', tsamaiso ea opioid le cannabinoid haholo (le hoja e se feela) e amanang le lijo 'ho rata'.

Ha e le hantle, lipatlisiso tsa boko ba batho li bontšitse hore ho lokolloa ha dopamine ho bile teng ha batho ba kopana le li-correlate tsa lijo tse nang le litekanyetso tsa bona tsa ho batla lijo [35]. Ka lehlakoreng le leng, ho ts'oara ha li-opioid kapa li-receptor tse nang le li-cannabinoid tse sa feleng ho bonahala li etsa hore ho be le takatso ea lijo ka ho ntlafatsa 'ho rata' lijo (e leng ho lumellana ha eona). Le hoja mekhoa ena e 'meli e fapane, ba etsa kopano ho itekanya mokhoa oa ho itšoara.

The hedonic ('ho rata') thepa ea lijo e bonahala e itšetlehile ka, har'a tse ling, opioid, cannabinoid le GABA neurotransmission [36]. Lihlahisoa tsena tsa 'ho rata' li sebelisoa likarolong tsa moputso ho kenyeletsa hamorao hypothalamus, NAc, ventral pallidum, OFC [9,27,38] le sehlekehlekeng (sebaka sa pele sa ho latsoa bokong) [39].

Pontšo ea li-opioid ka NAc (ka har'a khetla) le ventral pallidum e bonahala e bapisa lijo 'ho rata' [40]. Ka lehlakoreng le leng, ho bonts'oa ha opioid ka li-amygdala tsa basolateral ho kenyelletsoa ho fetisetsa thepa ea lijo, e leng eona e fokotsang bohlokoa ba ho ts'ehetsa lijo le boits'oaro ba ho batla meputso, kahoo e tlatsetsa ho 'ho batla' lijo [41]. Ho thahasellisang ke hore ka litoeba tse 'nileng tsa pepesa lijo tse nang le tsoekere, phephetso ea meriana e nang le naloxone (e nang le lithethefatsi e loantšanang le likokoana-hloko e se nang liphello ho likhoto tsa taolo) e etsa hore lefu la ho tlohela opiate le tšoane le le fumanoang liphoofolong tse' nileng tsa pepesa lithethefatsi tsa opioid [42]. Ho phaella moo, ho pepeseha ha batho kapa liphoofolo tsa laboratori ho tsoekere ho hlahisa karabo ea litšoantšo [43], e leng se bontšang hore tsoekere (le mohlomong lijo tse ling tse ratehang) e na le bokhoni bo tobileng ba ho matlafatsa maqhubu a opioid a sa feleng. Potso ea lipatlisiso e hlahang litabeng tsena ke hore, ho batho, ho ja lijo ho baka lefu la ho itokolla le bonolo le ka etsang hore motho a khutlele morao.

Li-endocannabinoids, haholo-holo ka mokhoa oa ho amohela li-CB1 pontšo (ho fapana le li-receptor tsa CB2), li ameha ka mekhoa ea homeostatic le e putsang ea ho sebelisa lijo le ho sebelisa matla a matla [44-46]. Tlhaloso ea Homeostatic e arolelanoa ka karolo e itseng ka nuclei e tsoetseng pele le e hlahang ka hare ho hypothalamus le ka nucleus ea lekhalo le le leng ka bokong, 'me tsamaiso ea mekhoa ea moputso e arolelanoa karolong e itseng ka liphello tsa NAc, hypothalamus le boko ba maikutlo. Ka hona, tsamaiso ea cannabinoid ke sepheo sa bohlokoa ho ntlafatso ea meriana bakeng sa phekolo ea botenya le lefu la metabolism. Ka mokhoa o ts'oanang, ho feto-fetoha ha lentsoe ka serotonin ea mekhoa ea ho fepa ho amana le moputso le taolo ea homeostatic hape e bile sepheo sa ho ntlafatsa meriana e khahlanong le botenya [47-50].

Ka tsela e tšoanang, ho na le bopaki bo ntseng bo eketseha ba hore phallo ea homeostatic e tsamaisang matla a motlakase, e kang leptin, insulin, orexin, ghrelin le PYY, e boetse e laola boitšoaro bo se nang homeostatic le ho lekanyetsa litšenyehelo tsa lijo [50]. Li-neuropeptides tsena li ka boela tsa ameha ka ho laola tsebo mabapi le ho ja lijo le ka boemo ba ho ba le lijo tse susumetsang [51]. Ka ho khetheha, li ka sebelisana le li-receptor tse nang le li-cognate metseng ea VTA DA neurons, e sa etsoang feela ho NAc, empa le libakeng tsa prefrontal le limbic; ha e le hantle, ba bangata ba bona ba hlalosa li-receptor libakeng tse ka pele le hippocampus le amygdala [50].

Insulin, e leng e 'ngoe ea lihomone tsa bohlokoa tse amanang le taolo ea metabolism ea tsoekere, e' nile ea bontšoa ho hlokomoloha karabelo ea limbic (ho akarelletsa le libaka tsa moputso oa boko) le libaka tsa cortical boko ba motho ho ba le lijo tse matlafatsang. Ka mohlala, ka ho laola bophelo bo botle, insulin e thibelitse ho sebelisoa ha hippocampus, li-cortices tse ka pele le tse bonahalang ha li arabela litšoantšong tsa lijo [52]. Ka lehlakoreng le leng, lihlooho tse sa hanyetsanang le insulin (bakuli ba mofuta oa lefu la tsoekere 2) li ile tsa bontša matla a eketsehileng libakeng tsa mahlakoreng (amygdala, striatum, OFC le insula) ha ba pepeseha lijo ho feta bakuli ba sa tsoekere [53].

Iboko ba motho, hormone leptin e fumanoeng ke adipocyte, e sebetsang ka karolo e le 'ngoe leptin receptors hypothalamus (nucleus e khutsufatsang) ho fokotsa ho ja lijo, e boetse e bontšitsoe ho hlokomoloha karabelo ea libaka tsa litlhoko tsa boko ho e-ba le lijo tse hlahisang lijo. Haholo-holo, bakuli ba nang le bothata ba congenital leptin ba ile ba bontša hore ho na le sepheo sa ho bona lijo tse bonahalang (NAc le caudate), tse neng li amana le lijo tse hlokahalang, esita le ha taba e ne e sa tsoa fepa. Ka lehlakoreng le leng, ts'ebetso ea ho fosahetse ha ea ka ea etsahala ka mor'a beke ea 1 ea phekolo ea leptin (Setšoantšo sa 2a, b). Sena se ne se hlalosoa hore leptin e fokotse likarabo tse khotsofatsang ho lijo [19]. Phuputso e 'ngoe ea fMRI, eo hape e etsoang le bakuli ba nang le bothata ba congenital leptin, e bontšitse hore phekolo ea leptin e fokotsitse ho ts'oaroa ha libaka tse nang le tlala (li-insula, parietal le cortices temporal) empa e ntlafatsa ts'ebetso ea libaka tse amehang ho khethollo ea ho khetholla [prefrontal cortex (PFC)] ha ba pepeseha lijo tse hlahisang lijo [20]. Ka hona, lithuto tsena tse peli li fana ka bopaki ba hore, ka boko ba motho, leptin e tsamaisa mosebetsi oa libaka tsa boko tse amehang feela ka mekhoa ea homeostatic, empa le ka likarabo tse khotsofatsang le ka taolo ea ho thibela.

Setšoantšo sa 2   

Leptin e fokotseha athe ghrelin e eketsa ho finyella ts'ebetso ho lijo tse khothatsang libakeng tsa moputso oa boko. (a, b) Litšoantšo tsa bongoli li bontša libaka tseo leptin e fokotseng ho sebetsa ho tsona (NAc-caudate) litabeng tse peli tse nang le ho haella ha leptin. (B) Histogram bakeng sa karabelo ea ho etsa tlhahiso ...

Harmonone e boetse e bonahala e le ho etsa hore likarolo tsa melemo ea boko li arajoe e le ho susumetsa lijo boko ba motho. Ka mohlala, peptide YY3-36 (PYY), e lokolloang liseleng tsa 'mele ka morao-rao le ho fokotsa ho ja lijo, e ile ea bontšoa ho fetola phetoho ea taolo ea lijo tse kenngoeng ke lipotoloho tsa homeostatic (e leng hypothalamus) ho ea ka taolo ea eona ka lipotoloho tsa moputso ho fetohang tlala ho ea ho satiety . Haholo-holo, ha lipeo tsa PYY tsa mali tsa mafura li ne li phahame (joaloka ha li satiated), ho ts'oaroa ha OFC ka lijo ho etsa hore batho ba je lijo; athe ha mahlaseli a PYY a mali a ne a le tlaase (joalo ka ha lijo li lahlehetsoe) ts'ebetso ea hypothalamic e nepahetseng e ne e boleloa esale pele hore lijo li tla kenngoa [54]. Sena se ne se hlalosoa ho bontša hore PYY e fokotsa likarolo tse khotsofatsang tsa lijo ka ho feto-fetoha ha OFC. Ka lehlakoreng le leng, ghrelin (hormone e fumanoeng ka mpeng e eketsehang naheng ea ho itima lijo 'me e etsa hore lijo li jeoe) e ile ea bontšoa ho eketsa ts'ebetso ho arabela lijo tse hlahisang liphallelo tsa boko ba boko (amygdala, OFC, insula ea ka hare le striatum) mme ts'ebetso ea tsona e ne e le ho amahanngoa le litlaleho tsa tlala (Setšoantšo sa 2c, d). Sena se ne se hlalosoa ho bontša ho ntlafatsoa ha hedonic le likarabo tsa tsusumetso ho tse ling tse amanang le lijo ka ghrelin [55]. Ka kakaretso, liphuputso tsena li boetse li lumellana le ts'ebetso ea boko ba boko bo botle ka lebaka la lijo tse hlahisang batho ba satiated khahlanong le batho ba itima lijo; ho ts'ebetsoa ha meputso ea libaka ho arabela lijo tse fokolang ho fokotseha nakong ea ha ho bapisoa le boemo ba ho itima lijo [15].

Liphuputso tsena li supa ho kopana ho teng pakeng tsa methapo ea methapo ea pelo e laolang moputso le / kapa ts'ehetso le tse laolang matla a metabolism (Setšoantšo sa 1b). Lipontšo tsa poneletso tse laolang lipontšo tsa homeostatic ho fihlela lijong li bonahala li eketsa kutloisiso ea libaka tsa boko ba limbong hore li be le lijo ha li e-na le orexigenic (ghrelin) le ho fokotsa kutloisiso ho ts'ebetsong ha e le anorexigen (leptin le insulin). Ka tsela e ts'oanang, ho utloahala ha moputso oa bokooa libakeng tse fanang ka tšireletso ea lijo nakong ea ho lahleheloa ke lijo ho eketseha, empa ho fokotseha ha nako e ntse e ea. Ka hona, ba potoloho ba homeostatic le moputso ba etsa kopano ho ntšetsa pele boitšoaro bo bobe tlas'a maemo a ho hloka le ho thibela lijo ho ja ka tlas'a maemo a satiety. Ho senyeha ha ho sebelisana pakeng tsa homeostatic le moputso oa potoloho ho ka 'na ha khothalletsa ho ja haholo le ho tlatsetsa ho feteng (Setšoantšo sa 1). Le hoja li-peptide tse ling [tse kang peptide-1 (GLP-1), CKK, bombesin le amylin] li boetse li laola ho ja lijo ka mekhoa ea bona ea ho itšireletsa, liphello tsa bona tsa extrahypothalamic li fumane tlhokomelo e fokolang [12]. Ka hona, ho sa ntse ho e-na le lintho tse ngata tse lokelang ho ithutoa, ho kenyelletsanoa ho sebelisana pakeng tsa mekhoa ea homeostatic le e seng homeostatic e laolang ho ja lijo le ho kenya letsoho ho feteletseng.

Ho senyeha ha moputso le boemo ba lijo ho batho ba fetang boima bo feteletseng le ba boima haholo

Liphuputso tse entsoeng ka mahlakore a mabeli le tsa bongaka li fane ka bopaki ba ho fokotseha ha DA ho bontša libaka tse hlaselang [ho fokotseha ho DAD2 (D2R) receptors le DA release), tse amanang le moputso (NAc) empa hape le mekhoa le litloaelo (dorsal striatum) ka botenya [56-58]. Habohlokoa, ho fokotseha ha D2R e hlaselang e 'nile ea hokahanngoa le ho kenngoa ha lijo ka li-rite tse ngata [59] le ka ho fokotsa tšebetso ea lik'hemik'hale ho OFC le ACC ho batho ba bangata haholo [60] (Setšoantšo sa 3a-c). Ho filoe hore ho se sebetse ho OFC le ACC ho fella ka ho qobelloa [ho hlahlojoa 61], sena e ka 'na ea e-ba mochine oo pontšo ea D2R e fokolang ka ho fetisisa e fanang ka eona e thusang hyperphagia [62]. Ho fokotsa pontšo e amanang le D2R ho boetse ho ka fokotsa ho utloahala ho meputso ea tlhaho, bofokoli boo batho ba boima ba bo khonang ho lefa nakoana ka ho ja haholo [63]. Tlhahiso ena e lumellana le bopaki ba melaho e bontšang hore ho fokotseha ha mesebetsi ea DA ho VTA ho hlahisa ts'ebetso e kholo ea ho sebelisoa ha lijo tse mafura haholo [64].

Setšoantšo sa 3  

Hyperphagia e ka bakoa ke koloi ea ho lefella potoloho e fokolisitsoeng ea moputso (e sebetsoang ka lipotoloho tsa corticostriatal tse laoloang ke dopamine) e kopantsoe le kutloisiso e matla ea phallo (thepa ea hedonic ea lijo e hlophisitsoeng ka karolo e 'ngoe ka…

Ha e le hantle, ha ho bapisoa le batho ba tloaelehileng ba boima, batho ba bangata ba neng ba e-na le litšoantšo tsa lijo tse phahameng tsa calorie (ho susumetsoa ke maemo a bona) ba bontšitse ho eketseha ha neural activation ea libaka tseo e leng karolo ea moputso le lipotoloho tse susumetsang (NAc, dorsal striatum, OFC , ACC, amygdala, hippocampus le insula) [65]. Ka lehlakoreng le leng, ka ho laola boima ba 'mele, ts'ebetsong ea ACC le OFC (libaka tse amehang ka ts'ebetso ea ho etsa mosebetsi oo ho NAc) nakong ea ho hlahisa lijo tse phahameng tsa calorie li fumanoe li sa tsamaisane le BMI ea bona [66]. Sena se bontša phapang e matla pakeng tsa lijo tse jeoang (tse bontšitsoeng karolong e itseng ea BMI) le ho ts'oanela ho sebetsa ha moputso ho libakeng tse phahameng ho lijo tse nang le lik'halori (ho bonahatsoa ha ho etsoa OFC le ACC) ho batho ba tloaelehileng ba boima, ba lahlehileng ka botenya.

Hoa makatsa hore ebe batho ba mahlahahlaha, ha ba bapisoa le batho ba mahlahahlaha, ba ne ba e-ba le ts'ebetso e fokolang ea lipotoloho tsa moputso ho tloha ho sebelisoang ha lijo (moputso oa lijo tse chesang), athe ba ne ba bontša ts'ebetso e kholo ea libaka tsa cortical tse ling tse sebetsang hantle ha li lebeletse ho sebelisoa [67] (Setšoantšo sa 4). Sepheo sa ho qetela se fumanoa se lumellana le thuto e tlalehiloeng e eketsehile ts'ebetso ea motheo ea pheko ea lik'hemik'hale (e leng letšoao la boko bo sebetsang) libakeng tse ling tseo ho tsona ho leng bonolo ho sebetsa hantle, ho kenyeletsa li-insula, ka ho fetisisa ha li bapisoa le lihlooho tse boima [68] (Setšoantšo sa 3d, e). Mosebetsi o ntlafetseng oa libaka tse sebetsang ka katleho o ka etsa hore lihlooho tse matla li fumane lijo ho tse ling tsa li-reinforcers, empa ho fokotsa ts'ebetso ea dopaminergic ka ho sebelisoa ha lijo ho ka etsa hore ho feteletsoe holimo e le mokhoa oa ho lefella lipontšo tsa DA tse fokolang [69].

Setšoantšo sa 4    

Lihlooho tse ngata li na le karabo e fokotsehileng libakeng tse hlaselitsoeng tsa DA ha li fuoa lijo ha li bapisoa le tse tlalehiloeng lihloohong tse boima. (A) Karolo ea Coronal ea ts'ebetso e fokolang ka lehlakoreng le letšehali le le letšehali ka lebaka la ho fumana lebese la lebese ho latela tharollo e sa thabiseng; ...

Liphuputso tsena tsa litšoantšo li lumellana le kutloisiso e matla ea moputso oa potoloho oa potoloho (ho shebella lijo tse phahameng-khalori) tse bolelang esale pele moputso, empa ho fokotseha ho fokolang ho liphello tse khotsofatsang tsa lijo tsa sebele meleng ea dopaminergic ka botenya. Re nahana hore, ho isa tekanyong ea hore ho na le phapang pakeng tsa moputso o lebelletsoeng le ho fana ka palo e sa finyelle tebello ena, sena se tla khothalletsa ho ja lijo tse qobelloang e le teko ea ho finyella tekanyo ea tefo. Le hoja ho hlōleha ha moputso o lebeletsoeng ho fihla ho tsamaea le ho fokotseha ha sele ea DA e chesang liphoofolong tsa laboratori [70], bohlokoa ba boitšoaro ba ho fokotseha ho joalo (ha moputso oa lijo o le monyane ho feta kamoo o neng o lebeletse kateng), ho tsebo ea rona, ha ea etsoa lipatlisiso.

Ka tsela e tšoanang le liphetoho tsena tsa ts'ebetso ho potoloho ea moputso ho lihlooho tse ngata haholo, lipatlisiso tsa litšoantšo li boetse li ngotse ho fokotseha ho tsitsitseng ha reactivity ea hypothalamus ho lipontšo tsa satiety lihloohong tse ngata [71,72].

Bopaki ba ho tsieleha ha tsebo ho batho ba fetang tekanyo le ho feta

Ho na le bopaki bo ntseng bo eketseha ba hore botenya bo amahanngoa le ho se sebetse mesebetsing e itseng ea kutloisiso, e kang mosebetsi o boima, tlhokomelo le mohopolo [73-75]. Ka sebele, bokhoni ba ho thibela litakatso tsa ho ja lijo tse lakatsehang li fapana har'a batho ka bomong 'me e ka' na ea e-ba e 'ngoe ea mabaka a tlatsetsang ho ba kotsing ea ho ja lijo tse ngata [34]. Tšusumetso e mpe ea ho nona haholo ho tsebisoa mohopolo o boetse o bonahala ka ho ata ha tlhokomelo e feteletseng ea ho kula ho matla (ADHD) [76], Lefu la Alzheimer le dementias e 'ngoe [77], lefu la cortic [78] le lefu le tšoeu la taba [79] lihloohong tse ngata. Le hoja boemo ba bongaka bo sebetsanang le ho kula (mohlala, lefu la ho kula, lefu la tsoekere le lefu la tsoekere) le tsejoa hore le tla ama ts'oaetso hampe, hape ho na le bopaki ba hore BMI, ka boeona, e ka senya libaka tse fapaneng tsa tsebo,75].

Ho sa tsotellehe ho se lumellane pakeng tsa lipatlisiso, boitsebiso ba boko ba maikutlo bo boetse bo fane ka bopaki ba liphetoho tse sebetsang le tse sebetsang tse amanang le BMI e mengata ka lintlafatso tse ling tsa bophelo bo botle. Ka mohlala, phuputso ea MRI e entsoeng ho basali ba hōlileng ka ho sebelisa morphometry ea voxel-wise e bontšitse ho se lumellane ho teng pakeng tsa BMI le lihlooho tse bohlooho (ho akarelletsa le libaka tse ka pele), tseo, ho OFC, li neng li amahanngoa le bokooa bo sebetsang [80]. Ho sebelisa positron emission tomography (PET) ho lekanya boko ba tsoekere ea metabolism ho laola bophelo bo botle, ho se lumellane ho sa tšoaneng ho ne ho boetse ho bontšoa pakeng tsa BMI le ts'ebetso ea metsolic a PFC (dorsolateral le OFC) le ACC. Phuputsong ena, mosebetsi oa ts'ebetsong oa PFC o ile oa bolela hore lihlooho li tla sebetsana le litekanyetso tsa mosebetsi oa tsamaiso [81]. Ka mokhoa o ts'oanang, thuto ea NMR e hlollang ea lilemo tse bohareng le taolo ea khale e bontšitse hore BMI e amahanngoa hampe le maemo a N-acetyl-aspartate (letšoao la botšepehi ba neuronal) ka coralx e ka pele le ACC [79,82].

Lipatlisiso tsa ho nahana ka bongoli ho bapisa batho ba bangata le ba omeletseng ba boetse ba tlaleha ho lekanana ha boleng ba bohlooho libakeng tse ka pele (opculum e ka pele le e bohareng ba gyrus) le karolong e ka hare-hare ea gyrus le putamen [83]. Phuputso e 'ngoe, e sa fumaneng phapang pakeng tsa lihlooho tse mahetleng le tse omeletseng, o ile oa tlaleha hore ho na le kamano e ntle pakeng tsa tšoeu e tšoeu ea lihlooho tsa boko le seeta: hip ratio; mokhoa o ileng oa fetoleloa ka thōko ka ho ja lijo [84].

Qetellong, karolo ea DA ho thibela ho thibela ho thibela ho thibela ho thibela ho itšireletsa ho tsebahala hantle le ho ferekanya ha sona ho ka tlatsetsa mathateng a boitšoaro ba ho thibela, joaloka botenya. Khohlano e mpe pakeng tsa BMI le D2R e hlaselang e tlalehiloe ka botlalo [58] hammoho le lihlooho tse feteletseng [85]. Joalokaha ho boletsoe ka holimo, tlhahlobo e tlase ho feta e tloaelehileng ea D2R ho statum ea batho ba mafura haholo e ne e amahanngoa le ts'ebetso ea metabolic ea PFC le ACC [60]. Liphuputso tsena li na le mekhoa e metle ea li-DA e bontšang hore e tlatsetsa ho senyeha ha libaka tsa frontal cortical tse amanang le ho feta boima ba 'mele le botenya. Kutloisiso e molemo ea lits'ebiso tsena ho ka thusa maqheka a ho tataisa ho ntlafatsa, kapa mohlomong le ho khutlisetsa mathata a itseng ka ho khetheha libakeng tse bohlokoa tsa ho nahanisisa.

Ka mohlala, ho lieha ho fokotsa chelete, e leng tšekamelo ea ho fana ka moputso e le mosebetsi oa nako ea nako ea ho fana ka nako, ke e 'ngoe ea liketsahalo tse fuputsoeng ka ho fetisisa tsa lipatlisiso ho amanang le mathata a amanang le ho hloka maikutlo le ho qobelloa. Ho theoleloa ha nakoana ho fuputsoe ka ho fetisisa ka lipatlisiso ho batho ba hlekefetsang lithethefatsi ba khethang meputso e nyenyane-empa-kapele ka ho fetisisa-empa-e liehang [86]. Liphuputso tse 'maloa tse entsoeng ka batho ba mahlomoleng li boetse li bontšitse hore batho bana ba bontša khetho bakeng sa meputso e phahameng, e potlakileng, ho sa tsotellehe monyetla o eketsehileng oa mahlomola a phahameng nakong e tlang [87,88]. Ho feta moo, ho na le kamano e ntle pakeng tsa BMI le ho theoleloa ha hyperbolic, moo melemo e seng mekae ea nakong e tlang e fokolloa ho feta melemo e tlang nakong e tlang, e sa tsoa tlalehoa [89]. Ho theoleloa ha nakoana ho bonahala eka ho itšetlehile ka mosebetsi oa ventral striatum (moo NAc e leng teng) [90,91] le ea PFC, ho akarelletsa le OFC [92], 'me o nahanela maqheka a DA [93].

Hoa thahasellisa hore liso tsa OFC liphoofolong li ka eketsa kapa tsa fokotsa khetho ea meputso e menyenyane ea nakoana ka ho lieha meputso e khōloanyane [94,95]. Ho hlakile hore mokhoa ona o hlakileng oa boitšoaro o ka bontša hore bonyane lits'ebetso tse peli li sebetsoa ka OFC; ngoe ke bonono bo matla, boo motlakase a fumanang bohlokoa ba ho susumetsa maikutlo, 'me e mong o laola litakatso tsa pele ho matla [96]. Ho se sebetse ho OFC ho amahanngoa le matla a sa khoneng ho fetola tšusumetso ea ts'ebeliso ea ts'ebeliso ea motlakase e le mosebetsi oa maemo ao ho oona ho hlahelang (e leng ho fokotsa bohlokoa ba ho khothaletsa lijo ka satiety), e leng se ka etsang hore motho a qobelle lijo [97]. Haeba tšusumetso e matlafatsang haholo (joalo ka lijo le lijo bakeng sa taba e feteletseng) boleng bo matla bo ts'ebelisitsoeng ba motlakase bo tla fella ka sepheo se matlafetseng sa ho se fumana, se ka bonang e le boikemisetso ba ho lieha ho khotsofatsa (joalo ka ho qeta nako ho mela e telele ho reka ice cream).

Leha ho le joalo, ha maemo a fumaneha habonolo, lijo tse ts'oanang li ka etsa hore ho be le boitšoaro bo sa tsitsang (joaloka ho reka le ho ja chokolete e haufi le mofani oa chelete esita leha u sa elelloe ka takatso ea ntho e joalo). Ho se sebetse ho OFC (le ea ACC) ho fokotsa bokhoni ba ho itšunya-tšunya litabeng tsa pele, e leng se fellang ka ho hloka maikutlo le ho fokotsa tekanyo ea theko e fokolang.

Lijo tsa ho nahana

IT e tla hlaha ho tsoa bopaki bo bokeletsoeng bo bontšitsoeng mona, hore karolo e kholo ea batho ba feteletseng ba bonts'a ho se leka-lekaneng pakeng tsa ho utloahala ho matlafatsoa ha moputso oa potoloho o amanang le matla a matla a matla a matla a matla a matla a matla a matla a matla a matla a tsamaiso ea tsamaiso e fokolisang taolo ea tšireletso ka mekhoa e metle ea takatso. Ho sa tsotellehe hore na ho se leka-lekane hona ho bakoa ke kapa ho bakoa ke ho kula ho matla, tšebetso ena e hopotsa ntoa pakeng tsa meputso, maemo a ho potoloha le lipatlisiso tsa potoloho le potoloho e thibelang ho itšireletsa e tlalehiloeng ho lemalla [98].

Tsebo e bokelletsoeng lilemong tse mashome a mabeli tse fetileng tsa liphatsa tsa lefutso, li-neural le tsa tikoloho ea botenya li bontša ka ho hlaka hore bothata bo teng hona joale bo hlahile ho khetholla ha likokoana-hloko tse tsamaisang lijo ka mefuta ea rona le phaello le mefuta e sa tšoaneng ea lijo tse susumetsoang ke rona tsamaiso ea sechaba le moruo. Litaba tse monate ke hore ho utloisisa boits'oaro bo tebileng ba boits'oaro bo bolokang seoa sa botenya bo na le senotlolo sa qeto ea ho qetela (bona le Mabokose 3 'me 4).

Lebokose 3. Lintlha tsa motheo tsa lipatlisiso tse tlang

  • Ho utloisisa hamolemo ho sebelisana ha limolek'hule, lisele le lisele tsa potoloho pakeng tsa mekhoa ea homeostatic le moputso e laolang ho ja lijo.
  • Ho utloisisa karolo ea liphatsa tsa lefutso ho tsamaisa homeostatic le likarabo tsa lichelete bakeng sa lijo.
  • Ho utloisisa habonolo ho ameha ha li-neurotransmitter tse ling, tse kang li-cannabinoids, opioids, glutamate, serotonin le GABA, liphetohong tse tšoarellang tse ntseng li etsahala ka botenya.
  • Ho hlahloba lintlha tsa nts'etsopele ea phekolo ea methapo ea kutlo ea methapo ea lijo (homeostatic le e khotsofatsang) le kutloisiso ea eona ho hlahisoang ke lijo tsa tikoloho.
  • Ho utloisisa liphetoho tsa epigenetic li-circuits tsa neuronal tse amanang le taolo ea homeostatic le e khotsofatsang ea lijo tse kenang bokoeng ba fetal ho latela ho pepesehela lijo le ho hloka lijo nakong ea bokhachane.
  • Ho etsa lipatlisiso tsa li-neuroplastic li-circuits tsa homeostatic le meputso e amanang le ho pepeseha ho sa feleng lijong tse hahang haholo le / kapa lijong tse ngata tse nang le lik'halori.
  • Ho hlahloba kamano pakeng tsa homeostatic le hedonic tshebetso tse laolang lijo le ho ikoetlisa.

Lebokose 4. Kamoso ea lipatlisiso tsa litleliniki

  • Liphuputso tsa ho netefatsa hore na ts'ebetso e kholoanyane ea likarolo tse amanang le melemo e amanang le lijo ho batho ba maholo ka ho fetisisa e tlas'a khatello ea bona ea ho ja kapa ho bonahatsa mokhoa oa boipheliso oa ho ja lijo tse ngata.
  • Ho boleloa hore ho ntlafatsoa ha dopaminergic neurotransmission ho thusa ho ntlafatsa boitšoaro ba ho ja ka ho ntlafatsa le / kapa ho matlafatsa mekhoa ea ho laola likamano tse fanoeng lipakeng ka karolo ea PFC; Leha ho le joalo, lipatlisiso tse tsoelang pele li hlokahalang mekhoeng e sa hlalosoang ka nako eo.
  • Lijo feela ke ka seoelo e leng mokhoa oa ho atleha (e leng ho tsitsitseng) ho lahleheloa ke boima ba 'mele. E ka ba tse rutang ho rarolla hore na: (i) ho ja lijo ho ka baka lefu la ho tlohela ho eketsa kotsi ea ho khutlela morao; le (ii) litekanyo tse fokotsehileng tsa leptin tse amanang le ho ja boima bo bakoang ke lijo-thollo li lebisa ho ts'ebetsong ea potoloho ea moputso le ho ba le mekhoa ea ho batla lijo.
  • Lipatlisiso ho fumana hore na lefu la neurobiology le fokotseha hakae ka takatso ea lijo le tlala ka mor'a ho buuoa ka bongatric.

References

1. Ogden CL, et al. Ho ata ha ho feta tekano le botenya ho United States, 1999 ho 2004. JAMA. 2006;295: 1549-1555. [E fetotsoe]
2. Fleg KM, et al. Ho ata le mekhoa ea botenya ho batho ba baholo ba United States, 1999-2008. JAMA. 2010;303: 235-241. [E fetotsoe]
3. Finkelstein EA, le al. Chelete ea selemo le selemo ea tšebeliso ea bongaka e bakoang ke botenya: ho lefshoa-le tekanyetso e khethehileng ea tšebeletso. Bophelo Aff. 2009;28: w822-w831.
4. Baessler A, et al. Ho kopanya liphatsa tsa lefutso le botsoalle ba liphatsa tsa lefutso tsa hormone secretagogue receptor (ghrelin receptor) ea liphatsa tsa lefutso. Lefu la tsoekere. 2005;54: 259-267. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
5. Silventoinen K, Kaprio J. Genetics ea ho latellana ha setopo sa 'mele ho tloha ho tsoalo ho fihlela bofelong ba lilemo tse bohareng: bopaki bo tsoang ho lifahla le lithuto tsa malapa. Lihlahisoa. Linnete. 2009;2: 196-202. [E fetotsoe]
6. E bua ka E, et al. Litlhahlobo tsa batho ba 249,796 li senola 18 e ncha e amanang le 'mele oa boima ba' mele. Nat. Genet. 2010;42: 937-948. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
7. Thorleifsson G, et al. Mokhatlo oa liphatlalatso ka bophara o fana ka mefuta e sa tšoaneng ea likhetho tse supileng tse latellanang le litekanyetso tsa botenya. Nat. Genet. 2009;41: 18-24. [E fetotsoe]
8. Naukkarinen J, et al. Tšebeliso ea liphatsa tsa lefutso tsa genome-wide ho tsamaisa 'Gray Zone' tsa lithuto tsa GWA li isa liphatseng tsa lefutso tsa botenya ba li-candidate. PLoS Genet. 2010;6 e1000976.
9. Gosnell B, Levine A. Mekhoa ea moputso le ho ja lijo: karolo ea li-opioids. Int. J. Obes. 2009;33 Tlatse. 2: S54, S58.
10. van Vliet-Ostaptchouk JV, le al. Phapang ea liphatsa tsa lefutso litseleng tsa hypothalamic le karolo ea eona ea ho nona haholo. Lihlahisoa. Tšen. 2009;10: 593-609. [E fetotsoe]
11. Blouet C, Schwartz GJ. Hypothalamic nutrient sensing ho laola matla a homeostasis. Behav. Resin ea Boko. 2010;209: 1-12. [E fetotsoe]
12. Coll AP, le al. Ho laola li-hormone tsa lijo. Cell. 2007;129: 251-262. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
13. Dietrich M, Horvath T. Ho fepa matšoao le bokooa ba boko. EUR. J. Neurosci. 2009;30: 1688-1696. [E fetotsoe]
14. Belgardt B, et al. Hormone le glucose pontšo ea POMC le li-neurons tsa AgRP. J. Physiol. 2009;587(Pt 22): 5305-5314. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
15. Goldstone AP. Hypothalamus, li-hormone le tlala: liphetoho boteng ba motho le boloetse. Prog. Resin ea Boko. 2006;153: 57-73. [E fetotsoe]
16. Rolls E. Latsoang, lijo tse fapaneng, le lijo tsa mongobo moputso oa bokooa le botenya. Int. J. Obes. 2005;85: 45-56.
17. Rolls ET. Mesebetsi ea orbitofrontal le ea pele ea lik'hotexte ka ho latsoa, ​​ho khotsofatsa, takatso ea lijo le maikutlo. Acta Physiol. Hung. 2008;95: 131-164. [E fetotsoe]
18. Petrovich GD, et al. Amygdalar le prefrontal tsela ea lateral hypothalamus li ts'oaroa ke sekolo se ithutang se tsosang lijo. J. Neurosci. 2005;25: 8295-8302. [E fetotsoe]
19. Farooqi IS, et al. Leptin e laola libaka tse hlaselang le boitšoaro ba batho. Saense. 2007;317: 1355. [E fetotsoe]
20. Baicy K, et al. Mofuta oa Leptin o fetola karabo ea bokooa ho lijo tsa batho ba baholo ba nang le lefutso la leptin. Proc. Natl. Acad. Sci. USA 2007;104: 18276-18279. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
21. Passamonti L, et al. Boikarabello bo bolela esale pele karabelo ea boko ha ho sheba lijo tse khahlehang: motheo oa neural oa ntlha e kotsi ea ho ja lijo tse matlafatsang. J. Neurosci. 2009;29: 43-51. [E fetotsoe]
22. Volkow ND, et al. Literekeng tsa neuronal tse fetohang li lemalla le botenya: bopaki ba lits'ebeletso tsa mafu. Philos. Trans. R. Soc. London. B. Biol. Sci. 2008;363: 3191-3200. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
23. Lenoir M, le al. Monate o moholo o feta moputso oa cocaine. PLoS One. 2007;2: e698. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
24. Cason AM, et al. Karolo ea orexin / hypocretin ho batla moputso le ho lemalla: ho boleloa ke botenya. Physiol. Behav. 2010;100: 419-428. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
25. Cota D, le al. Li-cannabinoids, li-opioid le boitšoaro ba ho ja: sefahleho sa limolek'hule tsa hedonism? Resin ea Boko. Tšen. 2006;51: 85-107. [E fetotsoe]
26. Atkinson T. Li-peptide tsa li-neuroendocrine tsa methapo ea meriana le liphatsa tsa lefutso 'me li bontša ponong ea takatso ea lijo: ho nahanisisa ka ho nona haholo pharmacotherapy. Lihlahisoa. Tšen. 2008;9: 108-120. [E fetotsoe]
27. Rabo ea Bohlale Rôle ea dopamine ea boko ka lijo le moputso. Philos. Trans. R. Soc. London. B. Biol. Sci. 2006;361: 1149-1158. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
28. DM e nyenyane, le al. Ho lokolloa ha dopamine ho fepa ka dorsal striatum ho kopana le lijo tse monate ho baithaopi ba phetseng hantle. Nako ea mokokotlo. 2003;19: 1709-1715. [E fetotsoe]
29. Norgren R, et al. Moputso oa mekotla le nucleus accumbens. Physiol. Behav. 2006;89: 531-535. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
30. Epstein L, et al. Ho tloaela ho ba moetapele e le ntho e khethollang lijo tsa batho. Psychol. Tšen. 2009;116: 384-407. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
31. Schultz W. Dopamine lipontšo bakeng sa bohlokoa ba moputso le kotsi: dintlha tsa motheo le tsa morao tjena. Behav. Thuto ea Boko. 2010;6: 24. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
32. Geisler S, R. Bohlale bo boleloang ke likarolo tsa glutamaterg ho sebakeng sa ventral tegmental. Rev. Neurosci. 2008;19: 227-244. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
33. Potoloho ea Petrovich G. Forebrain le taolo ea ho fepa ka lithuto tsa thuto. Neurobiol. Ithute. Mem. 2010 Oct 19; [Epub pele ho khatiso]
34. Wang GJ, et al. Bopaki ba phapang pakeng tsa tekano le bokhoni ba ho thibela boko bo sebetsang bo susumelitsoeng ke ho tsosoa ha lijo. Proc. Natl. Acad. Sci. USA 2009;106: 1249-1254. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
35. Volkow ND, et al. 'Phetoho ea li-Nonhedon' ho batho e kenyelletsa dopamine ka dorsal striatum le methylphenidate e eketsa phello ena. Synapse. 2002;44: 175-180. [E fetotsoe]
36. Berridge K. 'Liking' le 'ho batla' lijo tsa moputso: li-substrate tsa boko le boikarabello ba mathata a ho ja. Physiol. Behav. 2009;97: 537-550. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
37. Szczypka MS, le al. Tlhahiso ea Dopamine e hlahisitsoeng ho hlahisa litoeba tse senyehileng tsa dopamine. Neuron. 2001;30: 819-828. [E fetotsoe]
38. Faure A, et al. Mesolimbic dopamine ka takatso le tšabo: ho nolofalletsa ho susumetsoa ke ho senyeha ha glutamate sebakeng sa hau ka nucleus accumbens. J. Neurosci. 2008;28: 7148-7192.
39. Saddoris M, et al. Ho ithuta ka mokhoa o ts'oanang litemoso tsa liphello tsa litloaelo li nolofalletsa ho latsoa-encoding neural ensembles ka cortox e tloaelehileng. J. Neurosci. 2009;29: 15386-15396. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
40. Smith KS, Berridge KC. Potoloho ea li-opioid bakeng sa moputso: ho sebelisana pakeng tsa li-hedonic hotspots tsa nucleus accumbens le ventral pallidum. J. Neurosci. 2007;27: 1594-1605. [E fetotsoe]
41. Wassum KM, et al. Libaka tse fapaneng tsa opioid li khetholla ho lekanngoa le ho khoneha ha liketsahalo tse thabisang. Proc. Natl. Acad. Sci. USA 2009;106: 12512-12517. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
42. Avena NM, et al. Bopaki ba ho lemalla tsoekere: lits'ebeletso tsa boitšoaro le tsa methapo ea metsoako ea methapo ea tsoekere e feteletseng. Neurosci. Biobehav. Tšen. 2008;32: 20-39. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
43. Graillon A, et al. Tsela e fapaneng ea ho arabela li-sucrose tsa li-intraoral, quinine le oli ea poone ho bana ba sa tsoa tsoaloa ba llang. Physiol. Behav. 1997;62: 317-325. [E fetotsoe]
44. Richard D, et al. Boko ba endocannabinoid tsamaiso ea tsamaiso ea matla a motlakase. Molemo o Molemo. Res. Clin. Endocrinol. Metab. 2009;23: 17-32. [E fetotsoe]
45. Di Marzo V, et al. Lenaneo la endocannabinoid e le kamano pakeng tsa litsela tsa homoeostatic le hedonic tse amanang le taolo ea tekanyetso ea matla. Int. J. Obes. 2009;33 Tlatse. 2: S18-S24.
46. Matias I, Di Marzo V. Endocannabinoids le taolo ea matla a tekanyo. Trends Endocrinol. Metab. 2007;18: 27-37. [E fetotsoe]
47. Garfield A, Heisler L. Ts'ebetso ea tšebetso ea serotonergic bakeng sa ho phekola botenya. J. Physiol. 2009;587: 48-60.
48. Halford J, et al. Tlhokomelo ea litlhapi tsa tlhokomelo ea litšebeletso tsa botšepehi ka botenya. Nat. Rev. Endocrinol. 2010;6: 255-269. [E fetotsoe]
49. Lam D, le al. Serotonin tsamaiso ea bokooa ho tsamaisana le ho ja lijo le boima ba 'mele. Pharmacol. Likokoana-hloko. Behav. 2010;97: 84-91. [E fetotsoe]
50. Lattemann D. Endocrine li amahanngoa pakeng tsa lijo le moputso oa caloric homeostasis. Takatso ea lijo. 2008;51: 452-455. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
51. Rosenbaum M, et al. Leptin e fetola liphetoho tse bakoang ke boima ba 'mele boemong ba litsela tsa boipheliso boemong ba litsela tsa boipheliso ba lijo tse bonahalang. J. Clin. Sebelisa chelete. 2008;118: 2583-2591. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
52. Guthoff M, et al. Insulin e tsamaisa mosebetsi o amanang le lijo mohareng o moholo oa methapo. J. Clin. Endocrinol. Metab. 2010;95: 748-755. [E fetotsoe]
53. Chechlacz M, et al. Tsamaiso ea phekolo ea lefu la tsoekere e fetola likarabo litšoantšong tsa lijo libakeng tsa bokooa tse amahanngoa le tšusumetso le maikutlo: thuto ea ho nahana ka litšoantšo tsa matla a khoheli. Diabetologia. 2009;52: 524-533. [E fetotsoe]
54. Batterham RL, et al. PYY ho feto-fetoha ha likarolo tsa cortical le hypothalamic tsa boko bo bolela esale pele boitšoaro ba ho fepa batho. Tlhaho. 2007;450: 106-109. [E fetotsoe]
55. Malik S, et al. Ghrelin e etsa hore boko bo sebetse likarolong tse laolang boitšoaro. Setsi sa Metab. 2008;7: 400-409. [E fetotsoe]
56. Fulton S, et al. Leptin molao oa tsamaiso ea dopamine tsela. Neuron. 2006;51: 811-822. [E fetotsoe]
57. Geiger BM, et al. Ho senyeheloa ha maikutlo a monopolisi a dopamine ho fokolloa ke litekanyetso tsa lithethefatsi. Khopolo-taba. 2009;159: 1193-1199. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
58. Wang GJ, et al. Dopamine ea boko le botenya. Lancet. 2001;357: 354-357. [E fetotsoe]
59. Johnson PM, Kenny PJ. Dopamine D2 receptors ka ho lemalla ho lemalla ho lemala le ho qobella ho ja likhoto tse ngata. Nat. Neurosci. 2010;13: 635-641. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
60. Volkow ND, et al. Li-receptor tse fokolang haholo tsa Dopamine tse amanang le dopamine li amahanngoa le metabolism ea prefrontal lihloohong tse ngata: ho ka khoneha ho tlatsetsa mabaka. Nako ea mokokotlo. 2008;42: 1537-1543. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
61. Fineberg NA, et al. Ho nahana ka boitšoaro bo qobelloang le bo sa tsitsang, ho tloha mehlaleng ea liphoofolo ho ea ho endophenotypes: tlhahlobo ea litlaleho. Neuropsychopharmacology. 2010;35: 591-604. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
62. Davis LM, et al. Tsamaiso ea Bromocriptine e fokotsa hyperphagia le mahlahahlaha 'me ka tsela e fapaneng e ama dopamine D2 receptor le mohoebi a tlama likhoto le litoe tse haelloang ke leptin-receptor ka botenya bo bakoang ke ho ja. Neuroendocrinology. 2009;89: 152-162. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
63. Geiger BM, et al. Bopaki ba ho fokola ha lipalesa tsa dopamine exocytosis ka likhoto tse tloaelehileng. FASEB J. 2008;22: 2740-2746. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
64. Cordeira JW, et al. Ntho e entsoeng ke pelo ea pelo ea neurotrophic e tsamaisa phepo ea hedonic ka ho sebetsa ka mokhoa o hloekileng oa dopamine. J. Neurosci. 2010;30: 2533-2541. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
65. Stoeckel L, et al. Ts'ebetso ea tsamaiso ea meputso e atolositsoeng ho basali ba feteletseng ho arabela litšoantšong tsa lijo tse phahameng tsa lik'halori. Nako ea mokokotlo. 2008;41: 636-647. [E fetotsoe]
66. Killgore W, Yurgelun-Todd D. Boima ba 'mele bo bolela esale pele mosebetsi oa orbitofrontal nakong ea lipontšo tsa litšoantšo tsa lijo tse phahameng. Neuroreport. 2005;31: 859-863. [E fetotsoe]
67. Stice E, et al. Tlhaloso ea moputso o tsoang ho lijo le ho lebella hore lijo li kenelle ho feta botenya: thuto ea ho etsa litšoantšo tsa mahlaseli a motlakase. J. Abnorm. Psychol. 2008;117: 924-935. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
68. Wang G, et al. Mosebetsi o phomotsang oa ho phomola oa cortex ea molomo ka mantsoe a mangata haholoanyane. Neuroreport. 2002;13: 1151-1155. [E fetotsoe]
69. Stice E, et al. Likamano pakeng tsa botenya le ho arabela ka mokhoa o fokolang oa lijo li nkiloe ke TaqIA A1 allele. Saense. 2008;322: 449-452. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
70. Schultz W. Ho itokisa ka dopamine le moputso. Neuron. 2002;36: 241-263. [E fetotsoe]
71. Cornier MA, le al. Litla-morao tsa ho fetela holimo ho karabo ea neuronal ho lijo tse bonahalang tsa lijo tse fokolang le tse fokotsehileng. PLoS One. 2009;4: e6310. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
72. Matsuda M, le al. E fetotse ts'ebetso ea hypothalamic ho arabela ka tsoekere ea tsoekere ho batho ba feteletseng. Lefu la tsoekere. 1999;48: 1801-1806. [E fetotsoe]
73. Bruce-Keller AJ, et al. Botenya le ts'oaetso ea CNS. Biochim. Biophys. Acta. 2009;1792: 395-400. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
74. Bruehl H, et al. Liphetoho tsa ts'ebetso ea tsebo le boikutlo ba boko ba batho ba lilemong tse bohareng le ba seng ba hōlile ba nang le mofuta oa 2 lefu la tsoekere mellitus. Resin ea Boko. 2009;1280: 186-194. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
75. Gunstad J, et al. Palo e phahameng ea 'mele e amahanngoa le ho se sebetse ha molao ho batho ba baholo ba nang le bophelo bo botle. Compr. Psychiatry. 2007;48: 57-61. [E fetotsoe]
76. Cortese S, et al. Mathata a ho felloa ke matla / ho hloka matla a ho kula (ADHD) le botenya: tlhahlobo e hlophisitsoeng ea lingoliloeng. Crit. Rev. Food Sci. Lijo. 2008;48: 524-537. [E fetotsoe]
77. Fotuhi M, et al. Ho fetola litebello mabapi le morao-rao-dementia. Nat. Rev. Neurol. 2009;5: 649-658. [E fetotsoe]
78. Raji CA, et al. Tsela ea botho le botenya. Hum. Brain Mapp. 2010;31: 353-364. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
79. Gazdzinski S, et al. Moelelo oa boima ba 'mele le mahlaseli a matla a khoheli a botšepehi ba boko ba batho ba baholo. Ann. Neurol. 2008;63: 652-657. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
80. Walther K, et al. Ho se tšoane ha boko ba sebōpeho le ts'ebeliso ea likamano tse amanang le boima ba 'mele ho basali ba baholo. Hum. Brain Mapp. 2010;31: 1052-1064. [E fetotsoe]
81. Volkow ND, et al. Kamano e fapaneng pakeng tsa BMI le mosebetsi oa khethollo oa metsolic ho batho ba baholo ba phetseng hantle. Botenya. 2008;17: 60-65. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
82. Gazdzinski S, et al. BMI le botšepehi ba neuronal boemong bo botle ba botsofali ba nang le bokooa: mokhoa oa proton magnetic resonance spectroscopy. Botenya. 2009;18: 743-748. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
83. Pannacciulli N, et al. Bokooa bo sa tloaelehang boteng ba motho: thuto ea voxel-based morphometric study. Nako ea mokokotlo. 2006;31: 1419-1425. [E fetotsoe]
84. Haltia LT, et al. Ho hōla ha taba ea bokooa ka botebo boemong bo botle ba motho le ho hlaphoheloa ha lijo. J. Clin. Endocrinol. Metab. 2007;92: 3278-3284. [E fetotsoe]
85. Haltia LT, et al. Liphello tsa tsoekere e entravenous ka mosebetsi oa dopaminergic boko ba motho Ka vivo. Synapse. 2007;61: 748-756. [E fetotsoe]
86. Bickel WK, et al. Boits'oaro le li-neuroeconomics tsa lithethefatsi tse tsitsitseng lithethefatsi: ts'ebetso ea mahlaseli a khotsofatsang le mekhoa ea ho theola maemo a nakoana. Dithethefatsi Joala. Itšetleha. 2007;90 Tlatse. 1: S85-S91. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
87. Brogan A, et al. Anorexia, bulimia, le botenya: ho fokotsa likarolo tse arolelanoang ho Iowa Gambling Task (IGT) J. Int. Neuropsychol. Soc. 2010: 1-5.
88. Weller RE, le al. Basali ba nang le bokooa ba bontša ho lieha ho fokotseha ho feta basali ba nang le boima bo botle. Takatso ea lijo. 2008;51: 563-569. [E fetotsoe]
89. Ikeda S, et al. Kenyelletso ea li-hyperbolic, pontšo ea pontšo, le index ea 'mele oa' mele. J. Health Econ. 2010;29: 268-284. [E fetotsoe]
90. Cardinal RN. Mekhoa ea Neural e kenyelletsoa ho matlafatsoa le ho tsitsisoa ha maemo. Neural Netw. 2006;19: 1277-1301. [E fetotsoe]
91. Gregorios-Pippas L, et al. Ho fokotsa nako e khutšoanyane ea nakoana ea bohlokoa ba moputso ho batho ba intšang. J. Neurophysiol. 2009;101: 1507-1523. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
92. Bjork JM, et al. Ho fokotsa ho fokotsa likhahla tse tsamaeang le likarolo tse lekanang tsa marang-rang. Biol. Psychiatry. 2009;65: 710-713. [E fetotsoe]
93. Pine A, le al. Dopamine, nako, le tšusumetso ho batho. J. Neurosci. 2010;30: 8888-8896. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
94. Mobini S, et al. Liphello tsa liso tsa orbitofrontal cortex ka kutloisiso ea ho lieha le ho tsitlallela ho matlafatsa. Psychopharmacology. 2002;160: 290-298. [E fetotsoe]
95. Roesch MR, et al. Na ke lokela ho lula kapa ke lokela ho ea? Phetoho ea moputso oa nako-o fokotsoang ka orbitofrontal cortex le lipotoloho tse amanang le boko. Ann. NY Acad. Sci. 2007;1104: 21-34. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
96. Schoenbaum G, et al. Pono e ncha mabapi le karolo ea orbitofrontal cortex ka boitšoaro bo itekanetseng. Nat. Rev. Neurosci. 2009;10: 885-892. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
97. Schilman EA, le al. Karolo ea sebopeho se entsoeng ka ho qobella motho hore a itšoare ka mokhoa o tsitsitseng le o-orbitofrontal-cortex-lekaneng: ho khoneha ho kenya letsoho tsamaiso ea serotonergic. Neuropsychopharmacology. 2010;35: 1026-1039. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
98. Volkow ND, et al. Ho nahana ka karolo ea dopamine ho sebelisa lithethefatsi hampe le ho lemalla lithethefatsi. Neuropharmacology. 2009;56 Tlatse. 1: 3-8. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
99. Davidson T, et al. Menehelo ea hippocampus le li-prefrontal cortex tsa mefuta-futa ho matla le taolo ea boima ba 'mele. Hippocampus. 2009;19: 235-252. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
100. Forloni G, et al. Karolo ea hippocampus ka taolo e itšetlehileng ka thobalano ea boitšoaro ba ho ja: lithuto le ka acid e kainic. Physiol. Behav. 1986;38: 321-326. [E fetotsoe]
101. Haase L, et al. Phetoho ea Cortical ho arabela litakatso tse hloekileng nakong ea maemo a phekolo ea tlala le satiety. Nako ea mokokotlo. 2009;44: 1008-1021. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
102. Massa F, et al. Liphetoho tsamaisong ea hippocampal endocannabinoid ha ho e-na le litoeba tse nang le bothata ba ho ja. J. Neurosci. 2010;30: 6273-6281. [E fetotsoe]
103. ECNay EC. Insulin le ghrelin: li-hormone tsa lipalangoang tse hopolang mohopolo le oa hippocampal. Curr. Opin. Pharmacol. 2007;7: 628-632. [E fetotsoe]
104. Bragulat V, et al. Lihlahisoa tse monate tse amanang le lijo tsa potoloho tsa lipotoloho tsa bokooa nakong ea tlala: phuputso ea motlakase oa FMRI. Botenya. 2010;18: 1566-1571. [E fetotsoe]
105. Benarroch E. Neural e laolang boitšoaro ba ho fepa: kakaretso le likamano tsa bongaka. Neurology. 2010;74: 1643-1650. [E fetotsoe]
106. Olszewski P, et al. Ho hlahlojoa ha marang-rang a fepa li-neuroregulators ho sebelisa Allen Brain Atlas. Neurosci. Biobehav. Tšen. 2008;32: 945-956. [Tlhahiso ea mahala ea PMC] [E fetotsoe]