Tšusumetso e ntle ea Lihlahisoa tsa Hyper-Caloric Plastiki ea Neuronal (2017)

Jean-Pascal Morin,^1,2 Luis F. Rodríguez-Durán,^1,3 Kioko Guzmán-Ramos,¹ Claudia Perez-Cruz,⁴ Guillaume Ferreira,^5,6 Sofia Diaz-Cintra,⁷ 'me Gustavo Pacheco-López^1,8,*

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Neural plasticity ke tšobotsi ea bohlokoa le e bohlokoa ea ts'ebetso ea methapo e lumellang liphoofolo "ho ikhaola" ho ikamahanya le tikoloho ea tsona nako eohle ea bophelo ba tsona. Ts'ebetso e itšetlehileng ka ts'ebetso ea methapo ea kutlo ka har'a tsamaiso ea methapo e bohareng ke mofuta oa polasetiki ea neural e etsang motheo oa ho ithuta le ho hopola, hammoho le boits'oaro bo tšoarellang, bo khothalletsang tikoloho, bo joalo le tlhekefetso ea lithethefatsi le ho tšenyo ea nako e matla ea hyper-caloric (PHc). lijo. Lichabeng tsa bophirimela, bongata ba lijo tsa PHc li hlahisitse keketseho e kholo ea tlatsetso ea boima ba 'mele bo feteletseng / botenya le mathata a amanang le ona. Mabapi le hona, ho fanoa ka maikutlo a hore ho atoloha ha botumo ho ka bakoa ke bonyane karolo ea boits'oaro ba boitšoaro ho batho ba anngoeng ke tšusumetso ea tšebeliso e sa feleng ea lijo tsa PHc; Bangoli ba bang ba bile ba lebisa tlhokomelo ho tšoanang ho teng lipakeng tsa ho ja ho tlotlisa le ho lemalla lithethefatsi. Ts'ebeliso e mpe ea nako e telele ea likarolo tse itseng tsa phepo e boetse e hokahane le bokuli bo sa khaotseng ba neuroimmune bo ka 'nang ba etsa hore batho ka bomong ba be maemong a neurodegenerative a kang lefu la Alzheimer's. Sehloohong sena sa tlhahlobo, re bua ka bopaki ba morao-rao bo bonts'ang kamoo ts'ebeliso ea lijo tsa PHc e ka bakang polasetiki ea neural e fetolang likoloi tsa nakoana tse kenang hore e be boits'oaro bo qobellang. Re boetse re buisana ka mekhoa e ka morao ea ho sebelisoa ha lijo tsa PHc ka nako e telele ho ka fetolang ts'ebetso ea boko le ho lebisa ho ho senyeha hoa kelello, ho tsepamisa maikutlo ho bongoaneng, bongoaneng le bohlolong e le methati ea methapo ea methapo ea kutlo ho hlabollo ea tikoloho. Kamora nako, re hlakisa moralo oa sechaba oa ho sebelisa tikoloho e lumellehang ea tikoloho.

Plano ea Neural le Behaviors ba Tlatsehang

E 'ngoe ea litšobotsi tse ikhethang tsa ts'ebetso ea methapo ke bokhoni ba eona ba ho fetola sebopeho sa bona le ts'ebetso ea bona ho latela boiphihlelo, ka tsela ena e lumellang motho ka mong ho ikenya ka har'a "enjena" ho iketsetsa "litekesi" tsa tikoloho. Ntho e makatsang ea neural plasticity e tsejoa e tlasa ho ithuta, ho kopanya le ntlafatso ea boitšoaro bo feto-fetohang le bo bobe (Abbott le Nelson, 2000; Citri le Malenka, 2008; Sehgal et al., 2013). Boemong ba synaptic, liphetoho tse itšetlehileng ka tšebetso tsa matla kapa ho sebetsa ha phetisetso ea phetisetso ea synaptic li bopa thepa ea likaroloana tsa potoloho ea neural. Ho feto-fetoha ha maemo le ho rarahana ha liphatlalatso tsa neural ho etsoa ho khoneha ka mefuta e mengata ea li-cellular tsa methapo ea methapo (Nelson le Turrigiano, 2008). Tsena li kenyelletsa polasetiki ea mofuta oa Bahebrania, joalo ka ts'ehetso ea nako e telele (LTP) le khatello ea maikutlo ea nako e telele (LTD), hammoho le ho hleka-morao hoa homeostatic synaptic le metaplasticity (Pérez-Otaño le Ehlers, 2005).

Boithuto bo bong bo supile hore nts'etsopele ea boits'oaro bo bobebe e arolelana likarolo tse tloaelehileng le mehlala ea ho ithuta ea setso (Setšoantšo (Figure1; 1; Jones le Bonci, 2005). Mohlala, li-receptors block tsa N-methyl-D-aspartate (NMDA), tse thibelang ka nepo LTP le LTD libakeng tse ngata tsa boko (Malenka le Bear, 2004), hape e thibela mekhoa e mengata ea tloaelo e amanang le ts'ebeliso ea lithethefatsi, joalo ka khetho ea sebaka, boemo ba boits'oaro le boits'oaro (Mameli le Lüscher, 2011). Ntle le moo, ho oela hape ho hlahisoang ke ho pepesehela lits'ebetsong tse amanang le ts'ebeliso ea lithethefatsi ke bothata bo boholo ba kliniki bo thusang ho phehella ha lithethefatsi, mme mehato ea eona e ka tlase e nahanoa hore e itšetlehile bonyane ka karolo ea ketsahalo ea ho phethela ha mohlala sebakeng sa hippocampal CA3, e leng lets'oao la ho khutlisa mohopolo oa maemo (Kauer le Malenka, 2007; Kesner et al., 2016). Ka lehlakoreng le leng, tekanyo ea synaptic ea α-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid (AMPA) -receptors expression expression in the nucleus accumbens (NAcc) neurons e hlokometsoe ka ponahalo ea lithethefatsi (Letsatsi le letsatsi Wolf, 2009; Tang le Dani, 2009; Reimers et al., 2014). Ntle le moo, tsamaiso e le 'ngoe ea koae e kenya metaplasticity sebakeng sa ventral tegmental (VTA) ka ho eketseha ha synaptic non-GluA2 e nang le li-receptors tsa AMPA hammoho le NR2B e nang le li-receptors tsa NMDA, e tlatsetsa ho matlafatsong ea ho pepeseha ho eketsehileng, hape le ho theola monyetla oa ho feta. liketsahalo tsa polasetiki tse VTA — NAcc tseleng (Creed le Lüscher, 2013). Phehisano e kholo hape ke mohopolo oa hore batho ba ka nts'etsapele "lijo-ho itšetleha ka lijo" ka ho ithuta le ho etsa tloaelo, le hore botenya bo ka bonoa, bonyane maemong a mang, e le ponahatso ea "ho lemalla lijo" (Volkow and Wise) , 2005; Blumenthal le Khauta, 2010; Volkow et al., 2013a; García-García et al., 2014; Karol et al., 2015). Leha lijo, ho fapana le lithethefatsi tsa tlhekefetso, li hlokahala bakeng sa bophelo ba setho, ho itšepa lijo tsa PHc ho batho le mehlala ea liphoofolo ho fana ka litšobotsi tse tlisoang ke ho lemalla lithethefatsi. (Figure1) .1). Tsena li kenyelletsa ts'ebetso ea mesolimbic dopaminergic system (Blackburn et al., 1986; Hernandez le Hoebel, 1988), ts'ebetso ea likarolo tse tšoanang tsa boko (Robinson et al., 2016), hammoho le matšoao a fetang joalo ka ponahalo ea mamello, boits'oaro bo khothalletsang (Johnson le Kenny, 2010; Rossetti et al., 2014) le matšoao a ho khaohang mabapi le lijo tsa PHc tse 'nileng tsa bonoa kamehla ho batho ba batenya haholo (Iemolo et al., 2012; García-García et al., 2014). Ntlheng ena, ho na le ho tšoana ho hongata lipakeng tsa boitšoaro ba batho ba batenya le mekhoa ea tlhahlobo ea lintho tse itšetlehileng ka Buka ea ho hlahloba le ea Statistical ea Mathata a kelello (DSM -IV, -5). Mohlala, mekhoa ka bobeli e bontša matšoao a: mamello; ho tlosa; Lintho tse nkuoeng ka bongata bo bongata kapa nako e telele ho feta kamoo ho reriloeng; boiteko bo sa atleheng ba ho laola tšebeliso; nako e ngata e sebelisitsoeng ho fumana, ho sebelisa, kapa ho hlaphoheloa ts'ebelisong ea ntho ena; ho hlokomolohuoa ha mesebetsi ea boithabiso, ea mosebetsi kapa ea boithabiso; le tšebeliso e tsoelang pele le ha o na le bothata bo sa khaotseng ba 'mele kapa ba kelello bo bakiloeng kapa bo hlahisoang ke ntho ena (Davis et al., 2011). Kamora mokhoa ona le sepheo sa ho theha sesebelisoa se ts'epahalang sa ho lekola bokhoba ba lijo, litekanyetso tsa DSM-IV tsa ts'ebeliso ea lisebelisoa li fetotsoe ho theha Yale Food Addiction Scale (YFAS, Gearhardt et al., 2009, 2016).

Ho feta moo, ho bohlokoa ho amohela hore metsoako e hloekisitsoeng le e kenelletseng e sebelisetsoang ho hlahisa lijo tsa PHc e tšoana le tlhahiso ea lithethefatsi tse lemalloang tse ntlafatsang cocaine e tsoang makhapetleng a coca kapa heroin e tsoang lipapong (Ifland et al., 2009). Ho sa ntse ho na le ngangisano ea mahlale 'me ha ho tumellano e fihletsoeng mabapi le boholo ba tlhekefetso ea tšebeliso ea lijo mabapi le ho hlalosa botenya (Carter et al., 2016), leha ho le joalo ho hlakile hore hajoale lijo tsa PHc, joalo ka lithethefatsi tse lemalloang, li ka hlahisa liphetoho tse matla morerong oa moputso oa boko tseo re sa kang ra iphetola ho tsona, tse lebisang ho feteng tekano le ho nona. Ho ts'ehetsa pono ena, bopaki ba morao-rao bo bonts'a hore ts'usumetso ea lijo tse lemalloang, joalo ka lithethefatsi, e kanna ea itšetleha ka tekanyo ea ho amoheloa ha eona le metabolism; Lijo tse tlalehiloeng e le litlamorao tse ngata li chesoa ka potlako ebile lia ananeloa (Schulte et al., 2015; Criscitelli le Avena, 2016) hape li na le moputso o moholo kaha re tla bua ka karolo e latelang.

Mofuta oa Moputso o Natefisitsoeng oa Moputso

Ntle le potoloho ea homeostatic e jang lijo tse tlase (e hlahlojoe ho Morton et al., 2014), ho ja lijo ho laoloa ka matla ke lipontšo tsa hedonic kapa meputso, tseo hangata li ka fetisang litsela tsa homeostatic nakong ea matla a mangata ka ho eketsa takatso ea ho ja lijo tse matlafatsoang (Lutter le Nestler, 2009). Tlhahiso ea lijo tse matlafatsoang e etsa hore ho nkoe dopamine ka matla ho NAcc, e thehiloe ho projeke ea VTA, ho kenya letsoho boleng bo khothatsang le bo khotsofatsang ba lijo (Setšoantšo. (Figure2B) .2B). Ka bomalimabe, ts'ebetso ea tsela ena nakong ea lijo e amana le tahlehelo ea taolo ea lijo tsa batho ba bang (Stoeckel et al., 2008).

Karolo ea hedonic ea litheko tsa lijo e ka aroloa le ho feta moputsong le moputsong oa kamora nakoana. Karolo e ka fokotsoang habonolo e ka tlisoa kaha liphoofolo tse anyesang li na le khetho ea kahare bakeng sa litharollo tse natefeloang ka kotlolloho ho feta tse ikemetseng, 'me likhoto li ithuta ho khetha tharollo e nchafalitsoeng ka holim'a metsi hang ha e amoheloa e bolokehile (Bermúdez-Rattoni, 2004; Yarmolinsky et al., 2009; Drewnowski et al., 2012). Ts'ebeliso ea sucralose, e tsoekere e seng caloric ea maiketsetso, e etsa hore keketseho ea NAcc dopamine e lokolloe ka litekanyo tse bapisoang le sucrose (de Araujo et al., 2008). Leha ho le joalo, tatso e monate feela, e ikemetseng ho thepa ea eona e nang le phepo e sa atlehe ho hlahisa "meroalo e metle" e kopantsoeng, e itšetlehileng ka tšebeliso ea "meputso" e ikemetseng ea multisensory, e sa kenyeng monate feela le poso Moputso o sa tsotelleng, empa hape le lithahasello tse lebelletseng le tse mpe tsa litabatabelo (de Araujo, 2011).

Ho nahanisisa ka moputso oa kamoso ea morao-rao ho nahanoa ho bapala karolo ea mantlha phetolelong ea mekhoa ea ho ja (Antoni et al., 2016). Ha e le hantle, bopaki ba morao-rao bo bontšitse hore litoeba li ka ithuta ho khetholla lijo e le moputso o thehiloeng feela ho se nang le khalori ea tsona, ntle le tatso ea tsona. Mohlala, ageus trpm5^{- / -} litoeba, le hoja qalong li ne li hloleha ho khetholla pakeng tsa metsi le tharollo ea sucrose, hamorao li theha khetho ea sucrose e sa amaneng le mefuta ea hlaha (de Araujo et al., 2008; Simone et al., 2008;; Domingos et al., 2011). Matšoao a pele le a tsoang kamor'a ho tsoa ka mpeng a ka fetolang tšebetso ea dopaminergic, ke ka lebaka leo ho nahanoang hore o na le seabo sa tsoekere se ikemetseng ("Araujo et al., '). 2012). Ho joalo, bopaki ba morao-rao bo bonts'itse hore leptin ea li-hormone e kena-kenane le bokhoni ba ho tala ho thunya ho sa ikemiseng ha dopaminergic neurons. Ka lehlakoreng le leng, bopaki bo bong bo fana ka maikutlo a hore ntle le litlamorao tsa eona tse nang le oksijene e ngata, gut peptide ghrelin e ka ba le karolo ho e sebetseng kamora moputso oa morao oa prandial (Müller et al., 2015; Reichelt et al., 2015).

Phc ea ts'ebeliso ea lijo le polasetiki ea Neural

Ts'ebetso ea moputso oa post-prandial ts'ebeliso ea lijo e kenyelletsa dopamine efflux ho dorsal striatum (de Araujo et al., 2012). Ka litoeba, sebaka sena se na le lipotoloho tse fapaneng tse amanang le boitšoaro bo lebisitsoeng ho sepheo sa dorsomedial striatum, empa boits'oaro bo thehiloeng ho tloaelo, maemong a dorsolateral striatum (Setšoantšo. (Figure2B) .2B). Bohlokoa lits'ebetsong tsena tsa taolo ea ketso bo nahanoa hore bo baka mathata a mangata a amanang le tsamaiso ea methapo (Balleine le O'Doherty, 2010). Ehlile, ho na le phapang e pharalletseng lipakeng tsa li-circuits tsa neural tse qaliloeng ke lijo tsa PHc le lithethefatsi tsa tlhekefetso (Kenny, 2011a). Lilemong tsa morao tjena, ho entsoe boiteko ba ho senola hore na botenya le tšebeliso ea lithethefatsi li arolelana mekhoa e meng e tloaelehileng, mohlala, phetolelong ea nako e telele ea boitšoaro bo batlang moputso (Benton le Young, 2016). Ntlheng ena, potso e 'ngoe ea bohlokoa ke ho botsa hore na ho pepesetsoa lijo tsa PHc ho ka hlahisa liphetoho tsa polasetiki tsa nako e telele mohlaleng o tataisoang ke sepheo le tloaelo e thehiloeng ho tloaelo? Haeba lijo tsa PHc li baka mofuta o mong oa bokhoba, ho lebeletsoe phetoho e lebisang ho ts'ebetsong e thehiloeng litloaelong. Taba ena e sa tsoa rarolloa ke sehlopha sa bafuputsi ba ileng ba pepesa likhoto ho thibela ho fihlella lebese le kolobisitsoeng ka mokhoa o monate (ke hore, lijo tsa PHc) nakong ea libeke tsa 5 mme ba ba ba lekanya kutloisiso ea bona ea sephetho-devaluation (Furlong et al., 2014). Maemong ana, mosebetsi oa ho theola boleng o sebelisa paradigm ea ho ithuta eo ho eona liphoofolo li ithutang ho hatella molemong oa moputso oa lijo; hang ha mosebetsi o se o ithutile hantle sephetho-phepo ea lijo -ka mor'a moo e ananeloa ka ho e lumella ka bolokolohi ho eona kapa ka ho e taka ka sephetho se nyarosang se kang mala mala; ka hona, ho hatisa matsoho ho lebelletsoe ho fokotseha ho liphoofolo tse sebelisang leano le reriloeng. Ha mosebetsi o phetheloa ka leano le thehiloeng ho litloaelo ho le joalo, sephetho sa sephetho se ke ke sa ama karabo ea ts'ebetso joalo ka ho tobetsa ea lever. Ho khahlisang, ba hlokometse hore liphoofolo tse nang le tšebeliso ea lijo tsa PHc nakong e fetileng, li bontšitse ho phehella ho hatisang ka matla ha li bapisoa le taolo, li fana ka maikutlo a hore liphoofolo tseo li fumane leano le thehiloeng ho tsona. Hape ba bonts'itse ts'ebetso e ntlafalitsoeng ea dorsolateral striatum, e leng sebaka se kenyelletsang boitšoaro bo tloaelehileng. Ka hona, AMPA kapa dopamine (D) 1-receptors antagonism ho dorsolateral striatum e ile ea boloka boitšoaro ho ea boemong ba taolo. Ka lebaka leo liphetho tsena li bonts'a hore nalane ea tšebeliso ea lijo tsa PHc e ka tsamaisa phetoho taolong ea tloaelo ea boitšoaro (Furlong et al., 2014). Habohlokoa, haufinyane ho bonts'itsoe hore maikutlo a boits'oaro ho ntlafatso ea sephetho a boetse a sekisetsoa ho bahlankana ba batenya (Horstmann et al., 2015). Phuputso ea mohlala ea lijo tsa PHc ka har'a likhoto e bonts'itse hore ho pepesetsoa mafura a mangata ho tsoa ho batho ba baholo ho fokotsa bokhoni ba ts'ebetso le ho fokotsa kutloelo-bohloko ea sephetho, ho fana ka maikutlo a ho hloka ts'ebetso, ho eketsa boitšoaro, kapa bobeli ba (Tantot et al., 2017). Habohlokoa, liphokotso tsena tsa boits'oaro li ka felisoa ka ho koetlisa batho ba baholo ka mosebetsi o tiisang boits'oaro bo tataisitsoeng (Tantot et al., 2017).

Tšebeliso e sa feleng ea lijo tsa PHc, joalo ka lithethefatsi tsa tlhekefetso, e ka lebisa phetolo ea nako e telele methapong ea boko e kenyelletsang boits'ebetsong ba ho batla moputso (Kenny, 2011b; Volkow et al., 2013b). Empa ho kenyelletsa lijo, joalo ka ha re boletse, ke boits'oaro bo rarahaneng bo kenyeletsang "likarolo" tse ngata tsa meputso. Joale tšobotsi efe ea lijo tsa PHc e atisang ho baka liphetoho mokhoeng oa bokong, 'me qetellong ka boits'oaro? Ho sebetsana le bothata bona, phuputso ea morao-rao e lekotse hore na liphetoho tsa neuronal li hlokometsoe kamora tšebeliso e tsitsitseng ea PHc e tsamaellanang le boleng ba hedonic, kapa ka litaba tsa eona tsa caloric (Guegan et al., 2013). Bakeng sa sena, ba koetlisitse litoeba ho hatisa mochine oa moputso bakeng sa meputso ea lijo eo e ka bang lijo tse tloaelehileng, hypercaloric kapa lijo tse fumanehang habonolo le ho hlahlobisisa dendritic spine morphology. Ntle le moo, ba bapisa ho phehella ha boitšoaro ba ho batla lijo lihlopheng tse tharo tsa litoeba hang ha thibelo ea lijo e khutšoe. Ho khahlisang, litoeba tse koetliselitsoeng ho fumana lijo tse fumanehang habonolo li bontšitse ho phehella ho hatelletseng ho feta lihlopha tse ling tse peli, ha li ntse li fumana lijo ad libitum. Ntle le moo, mochine oa khatiso o sa lefelloeng o ne o phahame haholo ho litoeba tse hlahisitsoeng ka lijo tse matlafatsang tsa isocaloric, ho fana ka maikutlo a hore mokhoa ona oa ho ja o khothalelletsa boitšoaro bo sa ts'oaneng. Habohlokoa, phetoho ena ea boits'oaro e ne e sa bonoe ho litoeba tsa KO bakeng sa mofuta oa cannabinoid receptor 1 (CB1^{- / -}), ho fana ka maikutlo a karolo bakeng sa li-receptors tsena tsa endocannabinoid ho batla lijo ntle ho tšusumetso. Ha ba hlahloba dendritic morphology ka lihlopha tsena tse tharo, bangoli ba hlokometse hore methapo ea mokokotlo oa dendritic e eketsehile ka har'a methapo ea methapo ea methapo (PFC) le lepolanka la NAcc, libaka tse amanang le boitšoaro bo bobe, sehlopheng sa lijo tsa isocaloric, ha se bapisoa le litoeba tse jeleng lijo tsa hypercaloric. kapa chow e tloaelehileng. Nako le nako, ketsahalo ena e ne e boetse e bonts'oa e itšetleha ka li-receptors tsa CB1 (Guegan et al., 2013). Leha ho le joalo, tekanyo eo methapo ea methapo ea kutlo e tsamaeang ka pele ho moputso oa morao-rao e sebelisanang le tse amanang le ho ithuta botle ba tatso ea kutlo e sa ntse e lokela ho theoa. Hoa thahasellisa ho hlokomela hona joale, hore kalafo ea kalafo e bonts'itsoeng ho phekola botenya ho batho (mohlala, ts'ebetso ea bongaka ea bypass) e ka fokotsa takatso e monate ka ho kena-kenana le tlhahiso ea dopamine ea post-prandial striatal dopamine, joalo ka ha ho netetsoa ke lipatlisiso tsa thuto ea methapo (Han et al., 2016). Holim'a moo, Roux-En-Y gastric bypass surge ho litoeba ho ile ha bontšoa ho fetola tšebetso ea neural libakeng tsa boko tse amanang le tatso le moputso (Thanos et al., 2015).

Joalokaha re hlokometse, lintlha tse ling tsa tikoloho le litloaelo tsa boits'oaro li ka lebisa ho "bokhoba ba lijo" 'me qetellong la botenya. Ho feta moo, ho na le bopaki bo bong bo supang hore mefuta e meng ea liphatsa tsa lefutso e ka etsa hore batho ka bobeli ba tsebe ho ja ka mokhoa o feteletseng (DIO) le ts'oaetso ea boko (Heber le Carpenter, 2011). Ka hona, batho ba bang ba kanna ba rera ho monya mafura hantle haholo. Ntle le moo, ho hlahisoa ke DIO ka HFD ho sa tsoa bontšoa ho itšetleha ka neurotensin, neuropeptide e nang le ts'ebelisano e kholo ea dopaminergic, le lithuto tse telele ho batho li bontšitse hore pro-neurotensin plasma level ke ponelo-pele e ka tšeptjoang ea ho hola ka nako ea botenya (Li et al. , 2016). Leha pono e joalo ea lefutso ka botona e ka nyenyefatsa karolo ea boits'oaro le taolo ea ho ja ho nona, e totobatsa taba ea hore bophelo ba ho lula le ho ja le ho ja ka bophirimela ha bo lumellane le bokhoni ba rona ba ho iphetola ha lintho ka mokhoa o phethahetseng oa ho amohela mafura (Bellisari, 2008). Ntle le moo, e supa leano le hlakileng la mahlale a meriana a ka sebelisoang ho kenyelletsa liphetoho litloaelong le mokhoeng oa ho ja.

Liphetho tse lemohuo tsa Phatlalatso ea Lijo tsa PHc le Bofetoheli bo Eketsehileng

Ho tlalehiloe hore lijo tsa PHc tse lebisang ho botenya li amana le bokhoni bo fokotsehileng ba ho hlahisa polasetiki ea synaptic libakeng tse ling tsa boko tse amanang le cognition (Dingess et al., 2016; Klein et al., 2016; Tran et al., 2016). Mohlala, tšebeliso e sa foleng ea HFD e senya li-cascade tse kenang kahare ho polasetiki ea synaptic le insulin signaling / glucose homeostasis (Dutheil et al., 2016) hape e ama maemo a liprotheine a amanang le "protein" (Cai et al., 2016). Ho se leka-lekane ha phepo e nepahetseng ho bakoang ke phepo ena ho qetella ho ama lits'oants'o tsa glutamate neural, ho theha batsamaisi ba glial glutamate (GLT-1 le GLAST), tlase ho laola li-enzyme tse nyenyefatsang tsa glutamate, ho fokotsa phetiso ea basal synaptic le ho sitisa NMDA-induction LTD (Valladolid-Acebes et al. 2012).

Kamehla, lintlha tsa phepo ea phepo e ntle, tse kang carbohydrate e bonolo le mafura a nang le mafura a mangata, li hokahantsoe le ho senyeha ha memori le ho hlobaela ha hippocampal (Canada), 2012; Sobesky et al., 2014) mme bopaki bo fana ka maikutlo a hore boko bo ka ba tlokotsing ea lijo tsa "obesogenic" nakong ea methapo e matla ea methapo e kang methapo ea bongoana, bokhachane le bohlankana. (Figure2A; 2A; Valladolid-Acebes et al., 2013; Noble le Kanoski, 2016; Reichelt, 2016). Litonong, bopaki bo supa hore ho pepesetsoa hoa HFD ho senya mohopolo oa liteko tse fapaneng tsa boits'oaro, joalo ka ts'ebetso ea metsi ea Morris, mazes ea Barnes, radial arm maze, Y- le T-maze, le tsebiso ea ntho e ncha (Cordner le Tamashiro, 2015). Ho khahlisang ke hore, ha bopaki bo bongata bo bonts'a hore HFD e tebisa mohopolo oa nako e telele le ho fetoha ha kelello mesebetsing ea ho ithuta (e its'etleha botšepehing ba hippocampus) e matlafatsoa ke lijo tse joalo (setšoantšo (Figure2B) .2B). Mohlala phuputso ea morao tjena e fumane ho eketseha ha mohopolo oa maikutlo le polasetiki ea amygdala ho likhoto tse pepeselitsoeng HFD ho tloha ho khoesoa le ho ba motho e moholo, ka mochine o itšetlehileng ka li-glucocorticoid receptors ho amygdala (Boitard et al., 2015).

Boithuto ho batho bo bontšitse hore tšebeliso ea HFD, botenya le metabolic syndrome li amahanngoa le ts'ebetso e mpe ea kelello ho bana (Bauer et al., 2015; Martin et al., 2016) le batho ba baholo (Singh-Manoux et al., 2012; Papachristou et al., 2015; Lehtisalo et al., 2016; Yao et al., 2016), hape e eketsa kotsi ea nts'etsopele ea 'dementia' (Francis le Stevenson, 2013; Freeman et al., 2014). Ho ja HFD e kenyelletsang omega-6 e ngata le mafura a nang le mafura a mangata ho amahanngoa le ts'ebetso e mpe ho mesebetsi ea kelello (Kalmijn et al., 1997) le ka kotsi e eketsehang ea lefu la Alzheimer's (Kalmijn et al., 1997; Luchsinger et al., 2002) khatello ea mali le lefu la tsoekere (Fowler, 2016). Mabapi le sena, thibelo ea caloric e bontšitsoe ho khutlisetsa likarolo tsa HFD hanyane (Murphy et al., 2014). Batho ka bomong ba khomarelang lijo tse khahlanong le khatello ea mali tse kopantsoeng le lithibelo tsa caloric le boikoetliso li bonts'a ntlafatso e kholo lithutong ka bobeli tsa ts'ebetso ea ho hopola le ho potlaka ha psychomotor ha ho hlahlojoa likhoeli tsa 4 kamora ho kenella (Smith et al., 2010). Ho khahlisang, ho na le bopaki bo matla bo fanang ka maikutlo a hore thibelo ea phepo ea lijo lithutong tsa batho ba baholo bao e seng batho e na le litlamorao tse bolokang ts'ebetso ea ts'ebetso ea kelello nakong ea botsofali (Colman et al., 2009; Mattison et al., 2012). Ntle le moo, tlhahlobo ea morao-rao ea meta e bontšitse hore ts'ebetso ea bongaka ea bariatric hangata e lateloa ke mesebetsi e ntlafalitsoeng ea kelello ho bakuli ba batho (Handley et al., 2016), leha ho boetse ho lokela ho hlokomelisoa hore maemong a itseng, mathata a neuropsychiatric, joalo ka kotsi ea ho ipolaea a ka hlaha le kamora kalafo ena ea kalafo (Peterhänsel et al., 2013; Yen et al., 2014).

Patlisiso e ncha ka mefuta ea liphoofolo e se e qalile ho hlakisa mekhoa le mekhoa e ka fokolisang mafu a kelello a hlokometsoeng ke batho ba batenya (Castanon et al., 2015). Mohlala, bopaki ba morao-rao ho likhoto bo bonts'itse hore ho fetisoa ha mafura ho hlahisitse ts'ebetso ea microglial ho hippocampus ha lipectomy e bile le litlamorao tse fapaneng. Bangoli ba ile ba tsoela pele ho bonts'a hore cytokine interleukin (IL) -1 e lumellana hantle le maemo a adiposity hammoho le ho sitisoa ha kelello, 'me mohanyetsi oa IL-1 o pholositse bofokoli ba kelello bo hlokometsoeng ho liphoofolo tsena (Erion et al., 2014; Sobesky et al., 2014). Ntle le moo, ho pepesetsoa ha HFD ho sa tsoa bontšoa ho baka ho fokotseha ha letsoalo la mokokotlo oa hippocampal dendritic spine le hape le liphokotso tsa polasetiki ea synaptic ka lebaka la ho ts'oaroa ha straptic ke microglia, e ka khutlisoang ke ho emisoa ha lijo (Hao et al., 2016).

Thibelo le linako tse utloisang bohloko ho litlolo tsa tikoloho tse nang le phepo

Joalo ka maloetse a mang a mangata, ho bonahala ho na le linako tse mahlonoko tsa ho holisa botona. Boithuto ba pele bo thehile hore ho emara, nako e pakeng tsa 5 le 7 ea lilemo, le lilemo tsa bocha li bohlokoa bakeng sa kotsi ea ho ba le botenya ba nako e telele (Dietz, 1994), leha ho le joalo liphuputso tsa morao-rao tsa morao-rao li bontšitse hore botenya ba bongoana ka boeona bo its'epahalla haholo lijong tsa 'm'a nakong ea kemolo (Glavin et al., 2014). Liphuputso ho likhoto li bonts'itse hore malinyane a matamo a fepetsoeng ke HFD a na le mamello e phahameng ea leptin le ho se mamellane ha tsoekere hammoho le ho phahamisa maemo ho eketsehileng (Tamashiro et al. 2009). Ka mokhoa o ts'oanang le litoeba, malinyane a matamo a fepa a HFD a bonts'a mokhoa o ikhethileng bakeng sa tharollo ea sucrose le tharollo e se nang khalori ha e lekoa. Ho khahlisang, litoeba tsena li boetse li bonts'a kutloisiso e eketsehileng ea cocaine le amphetamine, hammoho le litekanyetso tsa basal dopamine ka har'a striatum le VTA, e lumellanang le tšusumetso e phahameng ea ho fumana moputso oa lijo (Peleg-Raibstein et al., 2016).

Boemong ba li-neurodevelopmental, lilemo tsa bocha li tšoauoa ka pruning e fetelletseng ea tsebo ea maiketsetso (Petanjek et al., 2011), hammoho le liphetoho ho gliogeneis le myelination (Masimong, 2005; Barbarich-Marsteller et al., 2013; Estes le McAllister, 2016). Ho feta moo, haufinyane tjena ho fanoa ka maikutlo a hore boemo ba khatello ea mali le mali bo ka eketseha ka ho pepesetsoa hoa HFD (Kanoski et al., 2010; Hsu le Kanoski, 2014) mme e fetoloa ka mokhoa o fapaneng nakong ea bohlankana (Brenhouse le Schwarz, 2016). Likarolo tse ling, joalo ka PFC, e ntseng e hola ho fihlela e le motho e moholo, li lula li ntlafalitsoe hape li sebetsa hantle ka nako ena (Reichelt, 2016). Lilemong tsa morao tjena, lilemo tsa bocha li boetse li thehiloe e le nako ea bohlokoa bakeng sa nts'etsopele ea botenya le bofokoli bo amanang le botenya ha mekhoa e meng e ka tlase ea neural e qala ho hlakisoa (Labouesse et al., 2016; Reichelt, 2016). Ka har'a liteko, litoeba li ile tsa feptjoa HFD lilemong tsa bocha 'me hamorao tsa lekoa mohopolong oa tlhokomeliso ea sebaka, e leng mosebetsi o itšetlehileng haholo ka ts'ebetso e nepahetseng ea hippocampus mme e shebang haholo ho fetoloa ha dorsal CA1 (Assini et al., 2009; Vogel-Ciernia le Wood, 2014). Ha ba ne ba lekoa e le batho ba baholo, litoeba tsena li ne li sa sebetse hantle ho feta balekane ba bona ba taolo mosebetsing ona mme phapano ena e ne e bonahala le kamora ho fetoloa thibelong ea lijo nakong ea beke ea 5. Ka lehlakoreng le leng, kalafo e tšoanang ea HFD e ne e se na phello ha e fanoa nakong ea ho ba motho e moholo. Ka mokhoa o tsotehang, ho senyeha hona mohopolong oa sepakapaka ho ne ho tsamaisana le molek'hule ea neural cell adhesion (NCAM, eo hape e tsejoang e le CD56) ho eketseha le ho hokahana ha methapo ea kutlo ea dendritic sebakeng sa hippocampal CA1 (Valladolid-Acebes et al., 2013). Haufinyane tjena, ho pepesoa ha mocha oa HFD ea lilemong tsa bocha ho ile ha boela ha bontšoa ho fetola maemo a extracellular matrix glycoprotein reelin le ho senyeha LTD ho li-PFC synapses (Labouesse et al., 2016). Hape ho hlokometsoe ho fokotseha hoa neurogeneis le boitšoaro bo botle mesebetsing e itšetlehileng ka hippocampus ho litoeba tse pepesitsoeng ke HFD nakong ea bocha (Boitard et al., 2012). Ho tšehetsa khopolo ea hore lijo tsa PHc li lebisa ho ho senyeha hoa kelello haholo-holo linakong tse tlokotsing, ho tlalehiloe hore HFD e tlatselitsoeng ka 10% sucrose e boetse e bontšoa ho hlahisa ho senyeha ha ho ithuta le ho hopola litloholo tsa bana (Xu et al., 2015). Haufinyane tjena, patlisiso e bonts'itse hore likhoto li fepuoa ka lijo tse bitsoang Bophirimela (ke hore, lijo tsa PHc) nakong ea bohlankana li bile le karabelo ea khatello ea maikutlo kamora ho ba batho ba baholo. Phuputso e boetse e bontšitse ho fokotseha ho hoholo ha lipalo tsa hippocampal hammoho le likhakanyo tsa morao-rao tse matlafalitsoeng liphoofolong tsena (Kalyan-Masih et al., 2016). Habohlokoa le ho feta, phuputso e ts'episang e bonts'itse hore ka ho hatella ho pepesetsoa hoa HFD nakong ea ho ba motho e moholo, ho senyeha ha methapo ea kutlo ho bonahala ho khutlisoa ka litekanyetso le ha ba ne ba pepesehetse lijo tsena ka nako ea bohlankana (Boitard et al., 2016).

Chebahalo, Ho phela kahare le ho bokella tikoloho ea tumello ea lumello

Ka kopanelo, lintlha tsena li fana ka mabaka bakeng sa litlamorao tse itseng tsa melemo ea ho kenella lithutong tsa pele ho tsa thuto le kelello, hammoho le letšolo le matla la ho lemosa litlamorao tsa tšebeliso ea lijo tsa PHc tse mabapi le linako tsa methapo ea kutlo; ke hore, boimana, bongoana le bohlankana. Mohlala, ho sa tsoa bontšoa hore ha ho hlahisoa phepo e nepahetseng ea phepo e le khetho e tsamaellanang le litekanyetso tsa bongoana (joalo ka boipuso ho batsoali kapa lipalo tse ling tsa bolaoli le tokoloho ho tsoa lipapatsong tsa bongata ke lik'hamphani tse fanang ka lijo tse se nang phepo e ntle, USA graders ba ne ba na le monyetla oa ho khomarela khetho e nepahetseng ea lijo (Bryan et al., 2016). Ntle le moo, likhakanyo tse mpe tse tobileng tsa chelete li ile tsa bonts'oa ho fetola khetho ea bareki ka lekhetho. Mohlala, moetlong o utloisisang o lekang ho laola ts'oaetso e phahameng haholo ea botenya / botenya, le ho nka hore lino tse tahang e ne e le mehloli e meholo ea matla hara bana le batho ba baholo (Stern et al., 2014), mmuso oa Mexico o phatlalalitse kenya ts'ebetsong lekhetho la 10% ho lino tse tsoekere tse tsoekere hammoho le lijo tse seng tsa bohlokoa tse nang le letsoalo le phahameng la khalori, ho qala ka Pherekhong 2014. Ka sebele, tlhahlobo ea morao-rao e netefalitse hore ka December 2014, thekiso e ne e se e theohile ka 12% mme tlhaiso-leseling e khothalelitse hore bareki ba Mexico ba ne ba fetohela mekhoeng e theko e tlase le e phetseng hantle (Colchero et al., 2016).

Ho eketsa thekiso, lijo tse tsoetseng pele indastering li matlafatsa thepa e ruisang molemo ka ho sebelisa letsoai le tsoekere, mafura, litatso le lijo tse ling ho kenyelletsa lijo tse joalo ka thepa e tlatsetsang. 2009; Gearhardt et al., 2011; Carter et al., 2016). Ka lehlakoreng le leng, melao e fokolang e tsoang liofising tsa 'muso tsa bophelo bo botle e beha meeli indastering ea lijo' me ho fihlela joale ha ho na temoso ea sechaba mabapi le tšibollo e ka bang teng le mathata a bophelo bo botle ba ts'ebeliso ea lijo tsa PHc. Ntlheng ena, joalo ka lintho tse ling tse lemalloang joalo ka nicotine kapa joala, tšehetso e eketsehileng ea sechaba e ka khothaletsa mekhatlo e etsang melao ho: (a) ho qala ho lemosa ka tšibollo e ka bang teng tlhahisong ea lijo tsa PHc; (b) ho laola tšebeliso ea lijo tsa bana bakeng sa bana, e le mohato oa pele oa ho fetolela phihlello ea batho ba baholo lijong tse tlatsetsang (Carter et al., 2016); (c) ho khothaletsa lipatlisiso tse eketsehileng tse mabapi le ho hlalosa likarolo tsa tlatsetso ea lisebelisoa tse fapaneng tsa tlatsetso tsa tlatsetso / lijana hammoho le motsoako oa eona; le (d) ho matlafatsa bareki ka ho fana ka tlhaiso-leseling e hlakileng le e hlakileng ea bophelo bo botle ka mabokoseng a lijo le lipapatsong tsa papatso.

Ka kakaretso, liteko tsa morao-rao empa tse ke keng tsa qojoa le tlhahlobo ea bongaka li tlaleha litlamorao tsa bophelo bo botle ba tikoloho e lumellang tikoloho eo linaha tse ngata tsa bophirimela li tobaneng le eona, joalo ka ha re se re hlahlobile mona, 'me ho hlakile hore e fetela ho bophelo ba kelello ka lebaka la taolo ho polasetiki ea neuronal (Setšoantšo. (Figure2A) .2A). Ho totobetse hore fisiks ea rona ea 'mele ha e ea hlaha ho tobana le liphephetso tse sa khaotseng tse hlahisoang ke tikoloho ea tikoloho, e bakang lefu la seoa la ho nona / ho ba botenya haholo (WHO, 2016a) e phephetsang litsamaiso tsa bophelo bo botle ka ho beha meroalo e mengata ea moruo (OECD, 2014). Kahoo ho potlakile ebile ho hlokahala hore ho ntšetsapele merero ea setsoalle ea nako e telele, e tšoarellang nako e telele ho laola le ho khutlisa tikoloho ka: (a) ho fa baahi matla a ho nka qeto e thehiloeng tsebong le ho ba bareki ba ikarabellang; (b) ho sireletsa bareki ba maemong a tlokotsing (ke hore, basali baimana, bana le bacha) ekaba ka lekhetho, molao kapa likotloi (WHO, 2016b); mme ho qetela empa e se bonyane (c) ho phahamisa kholo ea moruo e thehiloeng mekhoeng e meng ea lijo tse matlafatsoang tse ncha.

Mosebetsi ona o tšehelitsoe ke Lekhotla la Mexico bakeng sa Saense le Theknoloji (CONACYT) lithuso No. 273553 le 255399 ho GP-L, Dirección General de Asuntos del Personal Académico- Universidad Nacional Autónoma de México (DGAPA-UNAM) IN-203616-2 ho SD -C le Mokhatlo oa Nationalale de la Recherche et de la Technologie (ANR-15-CE17-0013 OBETEEN) ho GF. Mosebetsi ona o ne o tšehelitsoe ka lichelete ke lipatlisiso tsa lichelete tsa univesithi ea Metropolitan Autonomous University (UAM) le GP-L, le boithuto ba morao-rao ho J-PM. Ts'ehetso e eketsehileng ea lichelete e fanoe ke CONACYT, DAAD-PROALMEX fund No. 267761 ho GP-L, le Lekhotla la Saense le Theknoloji ea Naha ea Mexico (COMECYT) letlole la ho etsa lipatlisiso tsa mahlale le mahlale ho GP-L.

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