Optogenetic le chemogenetic tsebo mabapi le ho lemalla ho lemalla lijo (2014)

Ho nona hangata ho fumanoa ka mokhoa o bonolo ho latela boima le bolelele ba motho (index ea boima ba 'mele), empa ho amana le matšoao a mang a boitšoaro ao ho ka etsahalang hore ke a tlhaho. Lilemong tsa morao tjena, bo-rasaense ba bangata ba ile ba botsa hore na liphetoho tse tšoanang tsa boits'oaro le kelello li etsahala ho lemeng ha lithethefatsi le botena, ho alima ba bangata ho bua ka monyetla oa "ho lemalla lijo" Tsoelo-pele ea kutloisiso ea potoloho e ka bang boits'oaro bo matla ba ho fepa le ho lemalla lithethefatsi e ka re lumella ho nahana ka potso ena ka pono ea lipotoloho tsa neural, ho tlatselletsa lipono tsa boits'oaro. Mona, re hlahloba tsoelo-pele ea kutloisiso ea lipotoloho tsena 'me re li sebelisa ho sheba hore na ho ho bapisa ho bapisoa le bokhoba ba lithethefatsi ho ka thusa ho utloisisa mefuta e meng ea botona.

Bokhoba ba lithethefatsi ke bokuli bo sa feleng bo khutlisetsang matšoao a 'mele joalo ka ho mamella le ho ikhula, hammoho le matšoao a maikutlo le boits'oaro bo kang maikutlo a ho lakatsa le ho batla moputso ka potlako. Mamello e hlalosa ketsahalo eo ho eona ho hlokahalang hore litekanyetso tse phahameng tsa mori li fihlelle litholoana, ha matšoao a ho ikhula a hlalosa mefuta e fapaneng ea litlamorao tsa mmele le maikutlo tse bang teng ha motho a lemaletse ho lemalla ho sebelisa moriana. Liphetoho tsa boitšoaro tse amanang le bokhoba ba lithethefatsi li ka aroloa ka bongata ka mekhahlelo e meraro e meholo (Koob le Volkow, 2010). Taba ea pele, lithethefatsi le litloaelo tse tsamaeang le tsona li ba le tšusumetso e matla lits'ebetsong tsa ho matlafatsa, li khanna boitšoaro bo laeloang ke lithethefatsi hore bo be boima. Taba ea bobeli, bokhoba ba lithethefatsi bo tsamaea le mekhoa e meng e sitisang taolo ea thibelo. Qetellong, bokhoba ba lithethefatsi bo tlatselletsoa ke lipelaelo tse mpe tsa maikutlo tse kang ho tšoenyeha le khatello ea maikutlo, tse ka sebeletsang e le lisosa tsa ho khanna ts'ebeliso e eketsehileng ea lithethefatsi. Ho joalo, batho le liphoofolo tse lemalloang ke lithethefatsi ba kotsing e kholo ea ho khutla hape nakong ea khatello ea maikutlo kapa mathata (Epstein et al., 2006; Koob, 2008; Erb, 2010; Sinha et al., 2011). Mefuta ena e meraro ea matšoao e ka bonts'a liphetoho lipakeng tsa potoloho e fapaneng, e sebetsang hammoho ho tsamaisa tšebeliso ea lithethefatsi ho batho ba lemaletseng. Re tla hlalosa lithuto tsa morao-rao tsa optogenetic le chemogenetic tse faneng ka limmapa tse inahaneloang tsa hore na potoloho ena e ka ba eng.

Mantsoe a reng, “bokhoba ba lijo” a ile a hlahisoa ka har'a lingoliloeng ka bo-1950. 1956), empa ho bile le lithuto tse fokolang tse phatlalalitsoeng ka sehlooho sena lilemong tse 60 tse ileng tsa latela. Sebakeng seo, palo e kholo ea bafuputsi e ile ea sebetsana le bokhoba ba lithethefatsi nakong ena (Setšoantšo (Figure1) .1). Sena se fetohile lilemong tsa morao tjena, moo palo e nyane empa e ntseng e eketseha ea bafuputsi ba seng ba qalile ho etsa lipatlisiso ka ho lemalla ha lijo. Bafuputsi ba sejoale-joale ba boemong bo botle ba ho fuputsa khokahano ena, kaha United States le linaha tse ling tse ngata li kentsoe seoa sa lefu la botenya se lokelang ho sebetsoa (Centers for Disease Control, 2013), le kamohelo ea sechaba sechabeng ea "ts'ebeliso ea lijo" e atile, joalo ka ha ho pakoa ke palo e kholo ea lihlopha tsa tšehetso bakeng sa ho ja haholo, bongata ba tsona bo ipapisitse le methati e 12 ea methapo e sebetsanang le ho sebetsana le ts'ebeliso ea lithethefatsi le joala (Weiner, 1998; Russell-Mayhew le al., 2010). Ho joalo, mehato e 'maloa ea ts'ebeliso ea lithethefatsi (haholo-holo ho tsuba koae) naheng ea Amerika e ntse e fokotseha lilemong tse mashome tsa morao tjena, ha bongata ba botenya bo ntse bo eketseha ka sekhahla (Litsi tsa Taolo ea Mafu, 2013).

 

Setšoantšo sa 1

Palo ea lipampiri tse phatlalalitsoeng ka selemo ho tloha ka 1912-2012 tse nang le "ho lemalla lithethefatsi" kapa "ho lemalla lijo" ka sehlooho kapa tse ling. Liphetho ho tsoa Patlisisong e phatlalalitsoeng ka la 11/08/13, re sebelisa lisebelisoa tse tsoang Leseding la Neuroscience ...

Joalo ka ho lemalla lithethefatsi, botenya ke bothata bo rarahaneng bo nang le lisosa le matšoao a mangata. Mohlala, palo e nyane ea batho ba batenya haholo e na le liphetoho tse ling tse hlahisang 'mele o mong (joalo ka leptin le melanocortin receptors) tse bakang boima bo fetisisang (Farooqi le O'Rahilly, 2008). Leha ho le joalo, boholo ba botena bo bileng teng lilemong tse 30 tse fetileng ha ho lumeloe hore ke phello ea liphetoho tse bang teng bophelong, empa ho e-na le hoo, phetoho ea phepelo ea lijo le mekhoa ea rona ea bophelo nakong ena (Farooqi le O'Rahilly, 2008). Matšoao le boits'oaro bo amanang le botenya bona bo ka shejoa habonolo mekhahlelong e ts'oanang le ts'ebeliso ea lithethefatsi: ts'ebeliso e fetelletseng ea mathata, bothata ba taolo ea ho ja lijo, le ho hlaha ha maemo a mabe a maikutlo a kang ho tšoenyeha le khatello ea maikutlo (Kenny, 2011a; Sharma le Fulton, 2013;; Sinha le Jastreboff, 2013; Volkow et al., 2013). Ka hona, ho a khonahala hore potoloho e fetolang lits'ebetso tsena ho botenya li ts'oana le tse etsahalang nakong ea bokhoba ba lithethefatsi. Ke habohlokoa ho hlokomela hore, joalo ka bokhoba ba lithethefatsi, batho ba khethehileng ba batenya haholo ba bonts'a likarolo tse tlase tsa mathata ana, hoo motho a ka bonts'a matšoao a fapaneng le liphetoho lipakeng. Ntle le moo, ho fepa ho itšetlehile ka ho fepa ka lapeng ho bohlokoa ho pholohang, phapang e fapaneng le bokhoba ba lithethefatsi.

Taba ke hore ho fepa hangata ho 'nile ha nkoa e le sehlahisoa sa marang-rang a mabeli a ikemetseng a kopanyang le ho laola takatso ea lijo, tlala le thabo ea hedonic (Kenny, 2011b). Ntle le ho putsa li-circry tse ka tlatsetsang ho lemeng lithethefatsi le bothateng, tsamaiso ea homeostatic e boetse e laola tšebeliso ea lijo e thehiloeng ho tlhoko ea caloric ka ho potoloha lintlha tse tsamaeang le mali tse kang tsoekere, mafura a mahala, leptin, ghrelin le insulin (Myers le Olson, 2012; Adane, 2013; Hellström, 2013). Tsena li kenella lipotolohong tsa hypothalamic le brainstem ho khothaletsa kapa ho arabela likarabo tse ferekanyang, ka hona li tlatsetsa tekong e tloaelehileng ea matla. Ena ke tsela e 'ngoe eo botenya bo fapaneng le bokhoba ba lithethefatsi, kaha botenya bo ka bonts'a liphetoho lipakeng tsa homeryatic, ho kenyelletsa liphetoho lipakeng tsa meputso. Habohlokoa, ho entsoe lisebelisoa tsa rialo tse lumellang litsebi tsa methapo ea kutlo ho tsamaisa li-circuits ka mokhoa o hlakileng le taolo e sa lekanyetsoang (Fenno et al., 2011; Rogan le Roth, 2011; Tye le Deisseroth, 2012). Thutong ena, re hlahisa lipatlisiso tsa morao-rao mabapi le potoloho e sebelisang ho fepa le ho lemalla lithethefatsi, 'me re tšohla hore na tekolo-ts'oants'o ena ea potoloho ena e ka hlahisa leseli le lecha mabapi le ho tšoana le phapang lipakeng tsa botona le bokhoba ba lithethefatsi.

Potoloho e sebelisang phepelo ea lapeng

Ho ithuta mekhoa ea lijo tsa homeostatic ho thata ka lebaka la litekanyetso tse liehang tsa nakoana tsa litekanyetso tse sebelisang phetoho lipakeng tsa tlala le satiety. Lihormone li hloka ho lokolloa liseleng tse potileng, ho ea bokong le ho supa li-neuron tse nang le phepo ho tsamaisa boits'oaro ba ho batla lijo le ho bo sebelisa. Liphetoho tsena tsa nako e telele tsa khaello ea matla li senya haholo tlhahlobo ea likamano tse tlatselletsang lipakeng tsa methapo ea kutlo ea methapo le methapo ea methapo ea kutlo eo li e etsang. Ho kenyelletsa bothata bona, manong a mangata a lisele tsa kutlo tse nang le phepo e ngata a ka sebelisoa ho paka taolo e bohareng ea phepo. Ha ho se ho fumanoe, litsela tse kopanetsoeng le tse sebetsang tsa ho hlopha tlala le ho hlonepha li ka hlahlobisisoa ka botlalo (Sternson, 2013).

"Arcuate nucleus (ARC) ea" hypothalamus "e na le mefuta ea mefuta e fapaneng ea sele e fumanehang ka mokhoa o nepahetseng ho kopanya lipontšo tse tsamaeang le mali tse lokollotsoeng linthong tse ka holimo, ha ARC e phomola motheong oa boko bo haufi le mochini oa boraro oa boemo bo phahameng le bo bohareng. . Ka ho khetheha, likaroloana tse peli tse arohaneng tsa ARC, protheine e amanang le orexigenic-agouti (AGRP) le li-neurone tsa projenorgenic proopiomelanocortin (POMC) li hokahane haholo le liphetoho phetohong ea lijo. Li-subtypes ka bobeli tse matlafatsang li susumetsoa ka mokhoa o fapaneng le tse ling 'me li thibetsoe ke leptin e nkiloeng ka mafura e ngata (Myers le Olson, 2012) le matla a lipontšo tsa glucose (Claret et al., 2007; Fioramonti et al., 2007) le insulin (Konner et al., 2007; Hill le al., 2010). Ho feta moo, li-neuron tsa AGRP li kenngoa ka kotloloho ke moea o khothalletsang tlala o tsoang ka mpeng (Cowley et al., 2003; van den Top et al., 2004). Ho eketsa litlatsetso tsa bona ho ja, liente tsa mantlha bokong ba li-neuromodulator bo lokollotsoeng ke li-neuronodulator tsa AGRP, li-peptides AGRP le neuropeptide Y (NPY) li nyolla ka ho fepa (Semjonous et al., 2009), ha α-melanocyte e susumetsang hormone (α-MSH) le adrenocorticotrophic hormone (ACTH), e lokollotsoeng ho li-neurons tsa POMC, e fana ka lijo ka bongata (Poggioli et al., 1986).

Optogenetic kapa chemogenetic (Aponte et al., 2011; Krash et al., 2011, 2013; Atasoy et al., 2012) ts'ebetso ea li-neuron tsa AGRP e lekane ho potlakela ho fumana lijo tse matlafatsang, le liphoofolong tse nang le "calging" tse ngata, tse kopanyang tšebetso ea methapo ena kutlo le pono ea tlala le ho fepa ka morao ho moo. Habohlokoa, tekanyo ea tšebeliso e itšetlehile ka palo ea li-neurons tse ntle le khafetsa (Aponte et al., 2011). Ts'ebetso e sa feleng ea li-neuron tsena le hyperphagia e lebisang le tšebeliso ea matla e fokotsang ho lebisa ho phaello e kholo ea 'mele, e tsamaeang le mabenkele a eketsehang a mafura (Krash et al., 2011). Ntle le moo, li-neuromediator tse lokollotsoeng ke li-neurons tsa AGRP li tsamaisa liketsahalo tsa phepo e phahameng ka GABA le / kapa NPY e khothaletsa ho ja lijo tse matla ha peptide AGRP e hlophisa ts'ebeliso ea lijo ka tekanyo e liehang, e sa foleng (Atasoy et al., 2012; Krash et al., 2013). Ho khahlisang, liphoofolo tse nang le "li-neurons" tsa AGRP tse hlasimolohileng ka nako ea phomolo e tloaelehileng, ha ho se na lijo, li bonts'a ts'ebetso e matla e sa tsitsang e sa fetoheng ka botlalo ea lijo, e fana ka maikutlo a matla a ho nka karolo bakeng sa li-neurons tsena (Krash et al., 2011). Ntle le moo, ho kenella ka hare ho AGRP ho eketsa haholo boikemisetso ba phoofolo ba ho sebeletsa lijo sebakeng sa nalepoke assay ea khale (Krash et al., 2011).

Ho etsa lipatlisiso ka liphallelo tse tlase tsa ts'ebetso ea li-neuron tsa AGRP mabapi le ho fepa, likhakanyo tsa axon tsa bolelele bo ile ba fotoa 'me sephetho sa lijo tsa tlhahloboa. Ts'ebetso e khethiloeng ea terminal-field activate ea paraventricular (PVN) hypothalamus e tsosetse ho fepa ka boholo bo ts'oanang ho tataisa ts'ebetso ea AGRP ea somatic, e kenya letsoho la bohlokoa ho li-neurons sebakeng sena sa boko ho tsamaisa takatso ea ho saena (Atasoy et al., 2012). Ho bontša sena ka mokhoa o hlakileng, mefuta e 'meli ea chemogenetic inhibition e ile ea sebelisoa ho khutsisa boholo ba li-neuron tsa PVN, e ileng ea fella ka ho eketseha lib lib takatso ea lijo le tšusumetso ea ho sebeletsa lijo. Ntle le moo, lithuto tse ntle tsa mahlale a ntlafalitsoeng moo li-AGRP li amanang le PVN le li-neurons tsa PVN tse tlase tse tšoailoeng ke "oxytocin (OXT)" sepheo sa papatso li ile tsa kopanngoa ka tšebeliso ea Channelrhodopsin-2 (ChR2) 'me ka nako e tšoanang li fetotsoe, li khutlisetsa keketseho ea AgRP → PVN. ho ja lijo. Kamora nako, ka ho sebelisa mananeo a kopantseng optogenetic le chemogenetic ka pharmacology, mefuta e meng e tlase ea noka ea li-neuron tsa AGRP e ile ea kenella mofuteng oa boitšoaro oa ho fepa. Haufinyane, ho ile ha senoloa hore likhakanyo tsa axonal tsa AGRP ho ea mokokotlong oa bethe ea stria terminalis (BNST), lateroth hypothalamus (LH) kapa paraventricular thalamus (PVT), ntle le PVN, li lekane ho khanna ho fepa (Betley et al., 2013; hloka ho kenyelletsa PM Ref ena: 24315102). Habohlokoa, merero e fapaneng ea AGRP e hlahisang likarolo tse fapaneng tsa boko bo tsoang liphoofolong tse itseng, moo "konopo e le 'ngoe" ea "one-to-one" e sebelisang li-neuron tsa AGRP e laolang likhokahanyo tse theohelang (Betley et al., 2013).

Ka mokhoa o ts'oanang le liteko tsa ho lekola ho lekana ha AGRP, lisebelisoa tse sebelisoang ho hatella li-neurons tsa AGRP li senotse bohlokoa ba tsona ho fepa (Krash et al., 2011), e tsamaisanang le karabelo ea hypophagic liphoofolong tse latelang ho tlosoa ka sekhahla hoa lisele tsena (Gropp et al., 2005; Luar et al., 2005). Mokhoa ona oa neano oa ho tlosa moea ka matsoho o lebisitse ho khethollotsoeng ea potoloho ea anorexia sebakeng sa parabrachial nucleus (PBN; Wu et al., 2009), e amohelang mohopolo o tsoang ho li-neurons tsa AGRP (Atasoy et al., 2012) le kenyelletso e matla ea nyakallo e tsoang mohaleng oa sesebelisoa se le seng (NTS), se hlahisoang ka ts'ebetso ea serotonergic ho tsoa ho raphe magnus le obscurus (Wu et al., 2012). Haholo-holo, ho tšoaetsana ka mokhoa o hlakileng oa glutamatergic ho tsoa ho PBN ho eketsa takatso ea lijo, ho hatisa bohlokoa ba molumo o monate oa ho tsoa sebakeng sena se nang le boiphihlelo ho tataiseng boits'oaro ba phepo (Wu et al., 2012). Ho bonts'a pele PBI e na le molaoli oa bohlokoa oa takatso ea lijo, potoloho ea lipale, e tšoailoeng ke li-neurons tse amanang le liphatsa tsa lefutso tsa calcitonin, e reretsoeng ho fihla karolong e bohareng ea amygdala e bonts'itsoe ho fana ka likarabo tsa phepelo (Carter et al., 2013).

Manipulations a Direct POMC a na le litlamorao tse fapaneng le takatso ea lijo joaloka optogenetic e sa feleng le chemogenetic (Aponte et al., 2011; Zhan et al., 2013) ts'ebetso ea sechaba sena sa ARC e fokotsa ho ja haholo. Phello ena e hloka ho supa matšoao a melanocortin a hlakileng, joalo ka litoeba tse amohelang melanocortin-4 li sa sebetse ho hlahisa karabelo ena ea hypophagic (Aponte et al., 2011). Ho feta moo, ho hlohlelletsa ho hoholo ha li-neuron tsa POMC ho NTS ho latsoa tšebeliso ea lijo ka li-kinetics tse sebetsang ka potlako (lihora tse) le ts'ebetso ea butle-butle ea ARC-e hlalosang POMC neurons (matsatsi) (Zhan et al., 2013). Leha ho le joalo, ke tsa morao-rao feela tse hlokahalang bakeng sa ho tsamaisana le ho satiety joalo ka mokokotlo o matla oa li-neurons tse hlahisang PCC tsa ArC tse bakang hyperphagia le botenya (Zhan et al., 2013). Boithuto bo tsoelang pele bo etsang lipatlisiso ka lipolao tse tlase le liphula tse nyolohang tse laolang li-neurons tsa AGRP le POMC li hlokahala ho bula taolo ea takatso ea takatso ea wiring.

Le ha mosebetsi ona o khahlisang o hlakisitse boholo ba li-circry tsa bohlokoa tse laolang ho fepa hoa lapeng tlasa maemo a tlhaho, ha ho hlake hore na polasetiki e potolohileng ena e kenya letsoho phetohong ea boits'oaro e amanang le botena, le hore na ho tsepamisa likhetla lena ho ka sebetsa bakeng sa tahlehelo ea boima ba nako e telele ( Halford le Harrold, 2012; Alvarez-Castro et al., 2013; Hellström, 2013). Le ha batho ba batenya ho feta tekano ba ja ho feta, ha ho hlake hore na batho ba batenya haholo ba na le maikutlo a matla a ho lapa kapa maikutlo a fokotsitsoeng a takatso, ntle le tlhoko ea mmele ea ho ja ho feta ho boloka boholo ba mmele o moholo (French et al., 2014). Boithuto ba nako e tlang bo ka etsa lipatlisiso tsa ho thunya ho hoholo hoa liphoofotsoana tsena tsa neural, le mekhoa ea "plasticity" har'a li-neuron tsena ho sebetsana le sena. Ka mokhoa o tsotehang, phuputso ea morao-rao e bonts'itse phetoho ea liphatsa tsa lefutso ea AgRP ho tsoa ho nts'etsopele kapa kamora ho phahama ha methapo ea methapo ena e ntlafalitsoeng ea boitšoaro le likarabelo tse matlafatsang tsa koae, ho bonts'a hore liphetoho tse teng methapong ena ea methapo li ka kenya letsoho boits'oarong bo amanang le likarolo tse ling tsa boko (Dietrich et al. , 2012). Ho qhekelloa ho sa feleng ha lipotoloho tsena ho ka sebetsana le boholo ba lipotoloho tsena tse fetotsoeng boteng, hammoho le menyetla ea bona ea kalafo ea ho fokotsa boima ba nako e telele.

Ntle le ho fepa ka lapeng

Bopaki ba bokhoni ba liphoofolo ba ho kenella lijong tse seng tsa lehae bo bontšitsoe lithutong tsa motlakase tsa khale le liteko tsa lesion tsa hypothalamus ea morao-rao (Delgado le Anand, 1953; Margules le maqheku, 1962; Bohlale, 1974; Markou le Frank, 1987), e ka bakang litoeba ho ja hole le tlhoko ea homeostatic. Mosebetsi oa morao-rao o hlakisitse hore sena se kanna sa itšetleha ka lipakanyo tsa thibelo ho tsoa BNST, e tšoailoeng ke Vesicluar GABA transporter (VGAT) ho LH (Jennings et al., 2013). Tšusumetso ea Optogenetic ea likhakanyo tsena tsa GABAergic e entse hore motho a fepe matla a litoeba le nako e sebelisoang sebakeng se khethiloeng sa lijo, ha a ntse a sitisa merero ena ho fokotsa ho fepa litoeba tse lapileng. Ho khahlisang ke hore litlatsetso tsena tse fuoang li-optogenetic tse senotsoeng li senotse hore GABA ena^BNST→ Glutamate^LH potoloho e bile le tšusumetso e kholo litlamong tse susumetsang. Ho theola 'mila ona ka tsela ea orexigenic ho ile ha tsosa takatso ea lijo, likarabo tse khotsofatsang joalo ka ha li lekiloe ho latela boikhethelo ba sebaka sa' nete le boitšisinyo ba boits'oaro, ha boqhekanyetsi ka tataiso ea anorexigenic bo tsosa likarabo tse nyarosang (Jennings et al., 2013). Ho makatsang ke hore tlhahlobo e le 'ngoe e bonts'a tlhokahalo le khotsofalo bakeng sa batho ba fokolang ba glutamatergic ba li-neuron ho LH tse tšoailoeng ke polelo ea Vglut2 (glutamate transporter 2; 2013). Leha mananeo a thetso ea LH a ka hlahisa litlamorao tse fapaneng boits'ebetsong bo khothalletsoang (ho kenyelletsa ho khaotsa ho fepa ka botlalo) (Hoebel, 1971; Bohlale, 1974), ts'usumetso ea optogenetic ea VGAT ena^BNST→ VGLUT^LH likhakanyo kapa thibelo ea optogenetic e tobileng ea VGLUT^LH li-neuron ka ho khetheha li hlahisitse tloaelo e matla ea ho iphepa, ho fana ka maikutlo a hore likhakanyo tse hlakileng tsa tšebetso ea hypothalamic kapa palo ea li-neurons tsa LH e kanna ea ts'ehetsa likarolo tse fapaneng tsa boits'oaro. Ntlha ena e hlokometsoe ka mashome a lilemo (Wise, 1974), leha ho le joalo ho hlaha ha lisebelisoa le mahlale a morao-rao ho lumelletse bafuputsi ho utloisisa ka ho hlaka hore na lipalo tsa maqhubu a neural le projeke li tšehetsa likarolo tse fapaneng tsa boitšoaro ba ho fepa.

Ho lakatsa le ho phehella tšebeliso ea meputso ea lijo

Ho lakatsa ke karolo ea mantlha ea bokhoba ba lithethefatsi, boo ho lumeloang hore bo baka tšebeliso e mpe ea lithethefatsi tsa tlhekefetso (Koob le Volkow, 2010). Hangata batho ba nkoang ba le litakatsong tsa lijo ba batla ho ba le lijo, 'me ho potoloha ha tsona ho tsamaellanang le ho lakatsa botenya ho bonahala ho tšoana le ho lemeng ha lithethefatsi (Avena et al., 2008; Jastreboff et al., 2013). Sena se kenyelletsa dopaminergic circry, le ho ikamahanya le maemo mehahong ena ho kanna ha ikarabella ho phahamiseng litakatso tsa lithethefatsi le bokhobeng ka botlalo (Volkow et al., 2002;; Wang et al., 2002). Palo e kholo ka ho fetisisa ea li-dopaminergic neurons e lula bohareng ba mpa, sebakeng sa substantia nigra pars compacta (SNc) le sebakeng sa ventral tegmental (VTA). Optogenetic activation ea midbrain dopaminergic neurons ka har'a litoeba e nolofalitse ho matlafatsoa ha nako ea ho batla lijo molemong oa motho ea sebetsang (Adamantidis et al., 2011) ho kenyelletsa tlhahlobo e akaretsang ea khetho ea libaka (Tsai et al., 2009). Thepa e tšoanang e matlafatsang, joalo ka ha e hlahlojoa ke ho itšunya-tšunya ha methapo ea kutlo, ea li-neuron tsena e ile ea bonoa ho litoeba (Witten et al., 2011). Li-neurons tsa GABAergic tsa VTA ka kotloloho li thibela lisele tsa dopaminergic tsa VTA le ts'ebetso ea optogenetic ea pele e lekane ho khanna boemo ba boiketlo ba sebaka hammoho le boits'oaro bo kopanetsoeng (Tan et al., 2012; van Zessen et al., 2012). Ho makatsang ke hore maemong a sebelisitsoeng thutong ea Adamantidis, ho ts'oaroa ha litefiso tsa dopaminergic feela ho ne ho sa matlafatsoe, leha ho ne ho thusa ho matlafatsa mokhoa o tiileng oa boits'ireletso ba lijo (Adamantidis et al., 2011). Sena se fana ka maikutlo a hore kamano e ikhethileng e ka ba teng lipakeng tsa ho matlafatsoa ho fepa maemo, hore liphoofolo li na le monyetla o tlase oa ho ithuta ka tlhahisoleseling e amanang le lijo ho feta tlhaiso-leseling e ngoe.

Ketso e matlafatsang ea dopamine e kanna ea itšetleha ka polasetiki e itšetlehileng ka dopamine ho kapa ka har'a li-neuron tsa striatal tse fumanang mohopolo ho tsoa ho libopeho tsa dbaminergic tsa midbrain. Tsena ke boholo ba li-neuron tse bohareng tse hlalosang e le dopamine D1 kapa D2 receptor, e tsejoang ka hore ke tsela e otlolohileng (dMSNs) kapa tsela e sa tsamaelleng e pakeng tsa li-spiny neurons (iMSNs), ka ho latellana (Gerfen et al., 1990). Mohlala oa mokhoa ona oa ho laola liketso tsa batho ba batšo o hlahisitsoe ho elella bofelong ba bo-1980, mme ka linako tse ling o bitsoa "mohlala oa mantlha" oa basal ganglia circry (Albin et al., 1989). E thehiloe haholo lithutong tsa anatomical, bangoli bana ba tiisa hore ts'ebetso ea li-dMSN e nolofalitse tlhahiso ea makoloi, athe ts'ebetso ea li-iMSN e thibelitse tlhahiso ea makoloi. Liteko tse hlakileng tsa mofuta ona li e tšehetse, e bontša hore tsela e otlolohileng e phahamisa motsamao, athe tsela e sa tobang e sitisa motsamao (Sano et al., 2003; Durieux et al., 2009; Kravitz et al., 2010).

Leha ho le joalo, joalo ka ha dopamine e ka khothaletsa ho matlafatsa le ho sisinyeha, li-DMSN le li-IMSN le tsona li bonts'a tšusumetso e hanyetsanang ea matlafatso, e ka bonts'ang khokahano ea 'mele lipakeng tsa motsamao le matlafatso (Kravitz le Kreitzer, 2012). Dopamine D1 receptor ke Gs e nyarosang e kopantsoeng, ka hona dopamine e ka thabisa li-DMSN ka receptor ena (Planert et al., 2013), e ka bang karolo ea bohlokoa mefuteng ea matlafatso ea dopamine. Kannete, tšusumetso ea optogenetic ea li-dMSN e lekane ho tsamaisa matlafatso e sebetsang ho litoeba (Kravitz et al., 2012), le ho feto-fetoha ha tšebetso ea li-dMSN ho ka fetolela thepa e matlafatsang ea koae le amphetamine (Lobo et al., 2010; Ferguson et al., 2011) le meputso ea tlhaho (Hikida et al., 2010) ka tsela e lumellanang le litlamorao tsa ts'usumetso e tobileng ea dMSN. Dopamine D2 receptor ke inhibitory Gi e kopantsoeng, ka hona, dopamine inhibits iMSNs ka receptor ena (Planert et al., 2013). Ts'ebetso ea Optogenetic ea D2 receptor e hlalosang li-IMSNs e khothalletsa lehloeo (Kravitz et al., 2012), hape e fokotsa ho rata (Lobo et al., 2010), le ho itlhokomela ka koae (Bock et al., 2013). Se lumellanang le sena, chemogenetic inhibition ea li-neuron tsena e ntlafatsa thepa e ruisang ea amphetamine le cocaine (Ferguson et al., 2011; Bock et al., 2013). Ka mokhoa o ts'oanang, ha likhoto tse haelloang ke lijo li fuoa khetho pakeng tsa lijo tse fumanehang habonolo (li-biscuits tsa chokolete) le chow ea tsona e tloaelehileng, ngaka ea D1 agonist SKF 38393 e ile ea eketsa khetho ea bona bakeng sa lijo tse monate, ha D2 agonist quinpirole ea e fokotsa (Cooper le Al-Naser, 2006). Ka tsela ena, ho lokolloa ha dopamine ho ka khothaletsa ho matlafatsoa ka lipotoloho tse peli tse ikemetseng tsa basal ganglia. Dopamine e ka khothaletsa matlafatso ka ho kenya tšebetsong li-dMSN le tšebetso ka tsela e otlolohileng, hape ka ho thibela iMSN le tšebetso tseleng e sa tsejoeng (Kravitz le Kreitzer, 2012).

Le ha ho lokolloa ha dopamine hangata ho fokotsoa ha liphoofolo li ithuta likamano tsa matlafatso, ho ikopanya ha li-scrose ho pheta-pheta ho tsosa litekanyetso tse phahameng tsa tokollo ea dopamine, khafetsa ho fana ka letšoao la ho tiisa le latelang boits'oaro bo lebisitsoeng ho lijo tsena (Rada et al., 2005; Hoebel et al., 2009). Hore na ho lokolloa ha dopamine hangata ho etsahala ka mafura a mangata kapa lijo tse ling tse monate ha ho tsejoe. Ho lokolloa khafetsa ha dopamine nakong ea ho lutla ha sucrose ho ka tšoana le se etsahalang ka lithethefatsi tse lemalloang, tse tsoelang pele ho khothaletsa ts'ebetso ea dopaminergic ka liketso tsa pharmacological, ho sa tsotelehe hore na phoofolo e ithutile hantle hakae kamano pakeng tsa boitšoaro le phepelo ea lithethefatsi (Di Chiara le Imperato, 1988). Ka hona, ha liphoofolo li ja lijo tse joalo, ts'ebetso ea matlafatso ea dopamine e ka hlaha maemong a pheta-phetoang le a 'mele. Kannete, botenya bo amahantsoe le ts'ebetso e ntlafalitsoeng libakeng tsa boko tse sebetsanang le ho sesa le ho putsa ho latela tšusumetso ea lijo tse bonoang (Rothemund et al., 2007; Stoeckel et al., 2008; Jastreboff et al., 2013), leha lithuto tse ling li tlalehile lipatlisiso tse hanyetsanang ntlheng ena (Stice et al., 2010). Habohlokoa, haholoholo ha ho nahanoa ho tšoana le ho se tšoane lipakeng tsa bokhoba ba lithethefatsi le bokhoba ba joala, ts'ebetso e fapaneng ea li-neuron tsa ts'ebetso e kenngoa ha liphoofolo li ikatisa koae kapa lijo, ho bontša hore "likarolo tse fapaneng" tsa "gangal" tsa basal li ka boloka boitšoaro bo lebisitsoeng ho litsi tsa litlokotsi khahlanong le lijo (Carelli et al., 2000). Leha ho na le mokhatlo o sebetsang, ho ka etsahala hore liphetoho tse tšoanang tsa ts'ebetso ea methapo ea methapo ea dopamine li ka kenya letsoho ts'ebelisong e tlatselletsang ea ts'ebeliso ea likarolo tsa mekhatlo e fanang ka lijo le lithethefatsi. Lithuto tse kaholimo li hlakisitse litsela tse ka fetolang maemo a matlafatsang a lithethefatsi tsa tlhekefetso, mme li fana ka maikutlo a hore litsela tsena li ka fetoloa tšebelisong ea lithethefatsi. Leha ho le joalo, ona ke karolo e le 'ngoe feela ea bokhoba ba tai, e leng lefu le rarahaneng le amang likarolo tse ngata tsa boko. Ntle le matlafatso e kopantsoeng le lithethefatsi ka lipotoloho tsa basal ganglia tse hlalositsoeng ka holimo, lisekhete tse ling li kenella mokhoeng oa taolo ea thibelo, le ho hlaha ha maemo a mabe a maikutlo. Le ha tse boletsoeng ka holimo li hlakisitse karolo ea sisteme ea dopaminergic ho matlafatsong ea matlafatso, ho bohlokoa ho hlokomela hore ha se matlafatso ohle ke bokhoba. Mohlala, boholo ba batho ba nang le lithethefatsi tsa tlhekefetso ha ba lemale, leha ba fumana lithethefatsi li matlafatsa. Ka hona, liphetoho tse ling tsa potoloho li kenelletse tšebelisong ea lithethefatsi, joalo ka tse tlasa taolo ea taolo ea taolo ea boits'oaro, le ho hlaha hoa maemo a mabe a maikutlo.

Ho sitisoa ha taolo ea thibelo

Ho lemalla lithethefatsi ho tsamaisana le ho senyeha hoa ts'ebetso ea methapo ea mantlha le orbitofrontal, le ho baka bofokoli taolong ea taolo ea boitšoaro (Koob le Volkow, 2010; Volkow et al., 2013). Ho liphoofolo, phuputso ea morao-rao e bonts'itse hore ho itaola nako e telele ha ho etsoa ka cocaine ho fokotsa thabo ea cellular ea "cortical neuron" ea pele ho nako, ho supang mochine oa ts'ebeliso ea koae ka makhetlo a mangata. 2013). Ho leka ka kotloloho karolo ea li-neuron tsa PFC mabapi le ho ts'oara koae, lingoli tsena li ile tsa hlasimolla le ho thibela li-neurons tsena, tse ileng tsa tiisa kapa tsa eketsa ts'ebeliso ea koae, ka ho latellana (Chen et al., 2013). Le ha e le karolong e fapaneng ea boits'oaro, litlamorao tse fapaneng li tlalehiloe ka ho khutlisetsoa hoa cueine ho kopantseng, moo thibelo ea sebopeho sena e sitisitseng ho khutlisetsoa hoa cueine ho bakiloeng ke ho batla cocaine (Stefanik et al., 2013). Phapang ena e bonts'a hore liphokotso tse tlang pele lithutong tsa batho li kanna tsa se ke tsa supa ho fokotseha habonolo hoa ts'ebetso ea pele, empa ho e-na le liphetoho tse tobileng makaleng a ikhethileng a pele ka litsela tse ntlafatsang bokhoni ba ho khutlisa. Ka sebele, lithuto tsa ts'usumetso ea optogenetic li bonts'a hore li-neuron tse ikhethang tsa PFC tse hlahang ho fetisisa serotonergic dorsal raphe li khothaletsa ho sesa ka mafolofolo tekong e qobelloang ea ho sesa, ha ts'ebetso ea li-neuron tsohle tsa PFC e sa etse (Warden et al., 2012). Ho ka etsahala hore lipotoloho tse fapaneng tsa pele ho pele li-cortical li tsamaisa likarolo tse hlalositsoeng tsa boits'oaro bo amanang le lithethefatsi, mme ka lebaka leo, li ka senoloa ke lipapatso tse fapaneng tsa boitšoaro.

Khaello e tšoanang ea cortical le eona e ka amahanngoa le botena. Indasteri ea lijo e matlafatsoa ke ho se khonehe ha batho ho laola lijo tsa bona ntle le ho kenella ka ntle. Ho na le bopaki bo ntseng bo eketseha ba hore botenya bo amahanngoa le ho senyeha hoa ts'ebetso ea kelello, ho kenyelletsa le bofokoli mosebetsing oa motsamaisi, mohopolo oa ho sebetsa, le tlhokomelo (Gunstad et al., 2007; Bruehl et al., 2009; Mirowsky, 2011). Ts'ebetso tsena li etsoa ke "cortical circry" e fanang ka taolo ea "holimo-tlase" lipotolohong tsa bongoaneng tse boletsoeng ka holimo. Boithuto ba monahano oa Brain bo senotse likotsi tse ngata tsa sebopeho tse amanang le botenya, joalo ka ho fokotseha ha molumo oa lintho tse bosootho le tšebetso ea metabolic libakeng tse ka pele tsa batho ba batenya, mohlomong li bakang khaello ea bokhoni ba ho thibela ho ja (Le et al., 2006; Pannacciulli et al., 2006; Volkow et al., 2009; Smucny et al., 2012; Van den Eynde et al., 2012).

Boemo bo bong boo ho bona hangata batho ba iphumanang ba leka ho fana ka taolo ea thibelo ke nakong ea lijo. Motho ea jang lijo o leka ho boloka boemo bo bobe ba 'mele, ha a ntse a hanana le mekhoa ea ho tiisa (e boletsoeng kaholimo) le khatello ea maikutlo (e boletsoeng ka tlase). Mohlala oa phoofolo ena ke ho khutlisetsoa maikutlong ha motho a batla lijo. Paradigm ena, liphoofolo li koetliselitsoe ho hatella lijo, ka mor'a moo li tima empa li ka khutlisoa ka khatello ea maikutlo, ho kenyeletsa le khatello ea maikutlo ea meriana e amanang le pharmacological mimic yohimbine (le α2-adrenergic antagonist). Tšitiso ea Optogenetic ea moriti oa PFC nakong ea kalafo ea yohimbine e sitisitse botseteli bona, bo ts'oanang le litlaleho tse ts'oarelitsoeng tsa cocaine, tse fanang ka maikutlo a hore lits'ebetso tse ts'oanang li kanna tsa hlahisa sephetho ka bobeli (Calu et al., 2013; Stefanic et al., 2013). Hape, sena se bontša hore mathata a amanang le botenya ha a na liphetoho tse bonolo mesebetsing e felletseng, empa ho e-na le hoo ketsahalo e ikhethileng ea litekanyetso tse ikhethileng pele. Ka sebele, patlisiso ea ts'ebetso ea Fos maemong a mabeli a ho khutlisa lijo le khatello ea maikutlo e bontšitse hore li-neurons tse etelletsoeng pele li bonts'a liphetoho tse ikhethileng tse amanang le li-neurons tse sa sebetseng (Cifani et al., 2012). Sebaka sa sepheo sa lipatlisiso tsa nako e tlang se tla lekola lits'oants'o tsa terminal tsa li-neuron tsa pele ho tsa pele, tse bonts'itsoeng ho romella li-axons ho fana ka libaka tsa moputso joaloka VTA le konokono ea li-accumbens. Lithuto tse joalo li tla re lumella ho sebetsana le li-dysfunctions tsa pele pele li ts'oana kapa li fapaneng lipakeng tsa botona le bokhoba ba lithethefatsi.

Boemo bo bobe ba maikutlo

Maemo a fosahetseng a maikutlo a joalo ka ho tšoenyeha le khatello ea maikutlo e ka ba lisosa tse matla tse tsamaisang tšebeliso ea lithethefatsi ho bokhoba ba tahi. Likhoba tsa tlhekefetso li kotsing e kholo ea ho khutla nakong ea khatello ea maikutlo kapa khatello ea maikutlo, mme ts'ebeliso ea lithethefatsi e ka khothaletsa maemo a sithabetsang le a sithabetsang maikutlo (Koob, 2008). Mefuta e ts'oanang e ka hlaha le ho ja ho tlatsana le botenya, ho etsa hore bafuputsi ba belaele hore na khatello e tšoanang le ea li-circry e bakile ho lemalla lithethefatsi le ho lemalla lijo (Parylak et al., 2011;; Sinha le Jastreboff, 2013). Mohlala, linako tsa khatello ea maikutlo hangata li tsamaisana le ts'ebeliso ea lijo tse matlafatsang haholo, li hlahisa mantsoe "lijo tse khothatsang" le "ho ja maikutlong". Ntle le moo, liphoofolo tse motenya li bonts'a maemo a phahameng a ho tšoenyeha le khatello ea maikutlo, li fana ka maikutlo a hore lijo tsena ka botsona li kenya letsoho potolohong eo ho eona maemo ana a sa thabiseng a maikutlo a kenyang letsoho ho eketseheng ha ho ja (Yamada et al., 2011; Sharma le Fulton, 2013).

Lisebelisoa tse ngata tsa boko li laola maemo a sithabetsang a maikutlo, ho kenyelletsa le sisteme ea dopamine. Ho bonts'a dopamine e fetotsoeng e bile le tšusumetso e kholo ho botona ha batho ka bobeli ba mafura mme litoeba li na le litheko tse tlase tsa striatal dopamine D2 receptor (D2R) ha li bapisoa le batho ba fokolang le liphoofolo (Wang et al., 2001; Johnson le Kenny, 2010). Ntle le moo, li-polymorphisms tse geneng ea D2 receptor (Drd2) li kopantsoe le botenya le mefuta e mengata ea ts'ebeliso ea lithethefatsi (Blum et al., 1990; Noble et al., 1993;; Stice et al., 2008; Chen et al., 2012). Ho khahlisang ke hore, le ha bofokoli ba ho fumaneha ha D2R bo boetse bo hokahantsoe le bokhoba ba koae, koae, opiate le nikotine, litlatsetso tsena ha li tsamaisane le ho nona. Sena se fana ka maikutlo a hore litlamorao tsa ho sitisoa ha li-receptor tsa D2 ha li amane le ho nona ka bobeli, empa ho liphetoho tse fapaneng tsa boits'oaro tse tsamaeang le botena le bokhoba ba lithethefatsi. Polelo e le 'ngoe ea kamoo ts'ebetso ea D2R e fokotsehileng e ka kenyang letsoho phetohong ea boits'oaro e amanang le botenya le lithethefatsi ke hore liphoofolo li sebelisa haholo ho lefella likarabo tse tlisoang ke dopaminergic ka lebaka la litekanyetso tse theohileng tsa receptor (Wang et al., 2002;; Stice et al., 2008). Ka mantsoe a mang, liphoofolo li hloka maemo a phahameng a tlatsetso ea dopaminergic ho fumana phello e ts'oanang le phoofolo e nang le tlatsetso e felletseng ea li-dopamine receptors. Sena se ka phethoa ka mekhoa ea meriana 1988). Ntle le moo, e ka phethoa ka tšebeliso ea lijo tse matlafatsang, joalo ka lijo tse nang le tsoekere e ngata le mafura.

Ts'ebetso ea D2R e fokotsehileng e kanna ea boleloa esale pele ho phahamisa tšebetso ho li-iMSN, ha D2R e le sebui se kopaneng sa Gi. Ka hona, ho ka etsahala hore batho ba batenya ho feta ba jang lijo tse matlafatsang ho lokolloa ha dopamine ho thibela li-iMSN tsena tse sebetsang mme ba balehe maemong a sithabetsang a maikutlo. E lumellana le khopolo ena, liphoofolo tse hlalosang ChR2 li-iMSN li bonts'a ho hlohlelletsa lisele tsena (Kravitz et al., 2012). Ha ho hlahlojoa molemong oa moputso oa koae, ho hlohlelletsoa ha optogenetic le eona hoa senyeha (Lobo et al., 2010; Bock et al., 2013), ha chemogenetic inhibition ea li-neuron tsena e ntlafalitse mekhoa e tataisitsoeng ea koae (Ferguson et al., 2011; Bock et al., 2013). Tumellanong le liphumano tsena, keketseho ea thepa e khahlisang ea amphetamine e fumanoe ha li-neuron tsena li chechisoa (Durieux et al., 2009). Ka kopanelo, liphumano tsena li bonts'a hore phokotso ho polelo ea D2 e ​​ka hlahisa boemo bo bobe ba maikutlo, le hore liphoofolo li tla batla tokollo ea dopamine ea super-physiological ho pholoha mmuso ona.

Ntle le li-receptor tsa dopamine, liphetoho tse teng ho dopamine tse hlahisang li-neuron ho VTA li ka kenya letsoho ho hlaha ha maemo a mabe a maikutlo. Ka litlatsetso tsa bona ho VTA, li-efferents tse tsoang mokokotlong oa bongoana le habenula li hlahisa lipehelo tse ntle le tse mpe ka litoeba, ka ho latellana (Lammel et al., 2012; Stamatakis le Stuber, 2012). Thibelo e ikhethileng ea li-neurons tsa VTA DA e hlohlelletsa phenotypes e kang khatello ea maikutlo, joalo ka ha e hlahlojoa ka ho emisoa ka mohatla le liteko tse qobelloang tsa ho sesa, ntle le anhedonia, e hlalositsoeng ka tumellano ea khetho ea sucrose (Tye et al., 2013). Ho bonts'a taolo ea "li-neurons tsena" le ho lekana ha tsona ho hokahaneng le mekhoa ena, bangoli ba bonts'itse hore nakoana ka nakoana phasic photoactivation ea li-neurons tsa VTA DA e pholosa khatello ea maikutlo e bakoang ke khatello ea maikutlo e kang phenotypes (Tye et al., 2013). Ho etsa lipatlisiso tsa ho hlaseloa habonolo khahlanong le khatello ea boitšoaro e bakang khatello ea maikutlo sechabeng, ho tlalehiloe hore ts'ebetso ea optogenetic ea phasic, empa eseng tonic, ho thunya ho VTA DA neurons ea litoeba tse tsamaeang le paradigm ea ts'ebetso ea sechaba e khothalelitseng phemelo ea sechaba le ho fokotsa khetho ea sucrose, li-readouts tse peli tse ikemetseng tsa khatello ea maikutlo (Chaudhury et al., 2013). Dopamine neurons e VTA ke khale e tsejoa ho kenyelletsa moputso o kopanyang le meputso e lebisang moputsong (Bayer le Glimcher, 2005; Pan et al., 2005; Roesch et al., 2007;; Schultz, 2007). Lithuto tsa electrophysiological li boetse li hokahanya li-neurons tsa VTA DA le khatello ea maikutlo le linaha tse mpe (Anstrom et al., 2009; Wang le Tsien, 2011; Cohen et al., 2012) e totobatsa ho rarahana ha lets'oao la dopaminergic.

Kamora nako, ho batho, amygdala e hokahane le mathata a mabeli a ho tšoenyeha (Etkin et al., 2009) le ho lakatsa (Ngaka ea bana et al., 1999; Hlakola et al., 2008), ho tlatselletsa le mekhoa e mengata ea maikutlo. Lithuto tse 'maloa tsa optogenetic li qapile li-circuits tsa amygdala mabapi le boits'oaro bo bongata bo tsoang ho ba amanang le matšoenyeho (Tye et al., 2011; Felix-Ortiz et al., 2013; Kim et al., 2013) kapa ho tšaba (Ciocchi et al., 2010; Haubensak et al., 2010; Johansen et al., 2010) hammoho le tse amanang le ho batla moputso (Stuber et al., 2010; Britt le al., 2012). Le ha lithuto tsa electrophysiological li bonts'a hore li-neuron tsa amygdala li kenyelletsa valence e ntle le e fosahetseng (Paton et al., 2006; Shabel le Janak, 2009), ha ho e-so be le liphuputso tse khethollang matla a ho kopitsa a neural a likarolo tse sa koahelang letho tsa li-neuron tse etsang joalo. Le ha li-neural correlates tsa maikutlo a mabe a amanang le botenya li sa utloisisoe ka botlalo, tlhahlobo ea liphetoho tse kopaneng le tsa selikalikoe makaleng ana e ka ba sebaka se ts'episang ho sheba.

fihlela qeto e

Lilemong tsa morao tjena, paradigm ea ts'ebeliso ea lithethefatsi e sebelisitsoe lits'ebetsong tsa methapo ea methapo e amanang le botenya. Pono ena e hlahisitse kutloisiso ea bohlokoa, ha e ntse e hlokomela hore botenya bo na le phapang ea bohlokoa phapusing ea lithethefatsi. Haholo-holo, lijo lia hlokahala bakeng sa ho phela, e leng se etsang hore ho fepa ka matla le ho se sebetse hantle ha phepo e nepahetseng ha ho nahanoa ka mefuta ea phekolo e ka bang teng, kaha batho ba batenya ho feta tekano ba ke ke ba loha leano la ho qoba lijo ka ho felletseng, joalo ka motho ea lemaletseng lithethefatsi ho lithethefatsi. Kaha o na le bokhoni ba ho fepa boits'oaro ho hlokahala hore o pholohe hape o be kotsi ka ho fetisisang, ho utloisisa mekotla ea neural e amanang le tšebeliso ea litheko tsa lijo bakeng sa lisebelisoa tse hlakileng haholo, joalo ka mananeo a tataisoang ke mekhoa ea optogenetic le chemogenetic.

Khohlano ea polelo ea thahasello

Bangoli ba bolela hore lipatlisiso li ne li etsoa ka ho se be le likamano leha e le life tsa khoebo kapa tsa lichelete tse ka nkoang e le khohlano e ka 'nang ea e-ba le thahasello.

References