Tse ling tsa methapo ea kutlo ea ho ikopanya ka bobeli: lintlha tse tsoang ho motho ea nang le molekane ea nang le mekhoa e mengata (2011)

Front Neuroendocrinol. Sengoloa se ngotsoeng; e fumaneha ho PMC Jan 1, 2012.

E hatisitsoe ka mokhoa o qetelle o hlophisitsoeng e le:

PMCID: PMC3012750

NIHMSID: NIHMS227401

Moetso oa khatiso oa ho qetela oa mohatisi o fumaneha ho Front Neuroendocrinol

Sheba lihlooho tse ling ho PMC hore Qotsa sehlooho se hatisitsoeng.

Eya ho:

inahaneloang

Sebopeho sa likamano tse tšoarellang pakeng tsa balekane ba baholo (ke hore, maqhama a mabeli) ke karolo ea bohlokoa ea boitšoaro ba batho 'me e bile le tšusumetso bophelong ba' mele le ba kelello. Leha ho le joalo, ka lebaka la ho rarahana ha tlhaho ea litlamo tsena le ho raroloha habonolo ha tsona li bopiloeng ka mefuta e meng ea liphoofolo tse anyesang, re tseba ho fokolang ka tšibollo ea tsona ea tlhaho. Lilemong tse mashome a seng makae tse fetileng, sebaka sa thoriso (Microtus ochrogaster) e hlahile e le mohlala oa phoofolo ea maqhama a mabeli. Patlisiso e mabapi le mokhoa ona oa bophelo bo botle ba 'mele e fane ka leseli la bohlokoa mechineng ea methapo e laolang boitšoaro bo kopanang. Mona, re lekola lithuto tsena 'me re tšohla melaoana ea melaoana e meraro ea tlhaho e kopantsoeng: ho thehwa ha litakatso tsa balekane, nts'etsopele ea ho khetha mabifi ho kholiseho e sa tloaelehang, le tlhokomelo ea batsoali ba bacha. Re shebana le karolo ea vasopressin, oxytocin, le dopamine taolong ea boits'oaro bona, empa re boetse re bua ka ho kenya letsoho ha li-neuropeptides tse ling, li-neurotransmitters le lihormone. Lithuto tsena ha li na ho kenya letsoho kutloisisong ea tlamahano ea likamano tsa mefuta e 'meli feela, empa li ka fana ka leseli mabapi le lisosa tsa bofokoli ba boiketlo ba sechaba tse boletsoeng ho mafu a' mele a kelello.

Keywords: Khokahano ea kahisano, mabifi, boitšoaro ba bo-ntate, vasopressin, oxytocin, dopamine, prairie vole, monogamy

1. Selelekela

Khahleho e matla lipakeng tsa balekane, eo hangata ho thoeng ke maikutlo a lerato kapa a lerato, ke e 'ngoe ea matla a matla ka ho fetisisa a khannang boitšoaro ba batho, mme hangata e etella pele ho theha maqhama a ho mamella, maqhama a ikhethileng pakeng tsa balekane ba thobalano. Leha maqhama a joalo a setsoalle a atile haholo litsong tse tsoetseng pele tsa khoebo tse nang le mokhatlo o ikemetseng oa batho ba bangata, li etsahala hoo e batlang e le mekhatlong eohle ea batho, ho sa tsotelehe mokhoa oa ho iphelisa (mohlala, molisa, morui, joalo-joalo) kapa leano la ho hola (mohlala, sethepu le monogamy), le ke karolo ea bohlokoa ea boits'oaro ba motho. Leha tlhaloso ea maqhama a mabeli e fapana ho pholletsa le lingoliloeng, e hlalosoa ka mokhoa o fapaneng, mefuta eohle, e le setsoalle se tšoarellang se khethiloeng pakeng tsa batho ba baholo ba babeli ba nang le thobalano, 'me se khetholloa ke ho ikopanya, ho ikamahanya le ho qapa le molekane ka motho eo a sa mo tsebeng ( thatohatsi ea molekane) [105]. Ntle le khetho ea molekane, mekhoa e meng e fapaneng e kenelletse ka mokhoa o tebileng tlamong e rarahaneng ea bophelo. Mohlala, maqhama a mabeli ho batho, hammoho le mefuta e meng ea liphoofolo tse anyesang, a amahanngoa khafetsa le ho lebela balekane (mohlala, boitšoaro bo mabifi ho bahlabani ba thobalano) le tlhokomelo e ntle ea batsoali ka bana ba banyenyane [32, 86, 136]. Ho ba teng hoa mekhoa ena ho batho ba babeli ba kopaneng hoa utloahala ha ho shebiloe ka lenane la khopolo ea ho iphetola ha lintho, e fanang ka maikutlo a hore karolo e 'ngoe ea tlamahanyo e ile ea fetoha tlasa maemo moo ho neng ho hlokahala tsetelo e eketsehileng ea batsoali ho netefatsa kholiso e atlehileng ea bacha. [45, 85, 89, 105, 208]. Ho joalo, likhatello tse tšoanang tsa khetho e neng e hloka ho ba teng hoa batsoali ka bobeli bakeng sa pholoho ea bana li ka thusa ho theha ts'ebelisano pakeng tsa balekane [86] le mekhoa ea ho boloka tšebelisano-mmoho ena (mohlala, ho itšireletsa ha balekane).

Bohlokoa ba tšebetso ba tlamahano e 'meli ho batho bo tlalehiloe ka mokhoa oa setso. Batho ba ratoang haholo, haholo ba nang le likamano tse tsitsitseng tsa lenyalo, ba phela halelele ho feta balekane ba bona ba sa lefelloang, se fumanoeng se tsebahala ka lihlopha tsohle tsa baahi [116, 144]. Ho kenyelletsa moo, likamano tse phahameng tsa kamano pakeng tsa bobeli li hokahane ka mokhoa o hlakileng le maemo a mabe a kelello, joalo ka khatello ea maikutlo, mme e amana hantle le ts'ebetso ea boits'ireletso ba mmele le bophelo bo botle ba pelo [131, 212]. Monyetla o mong o amoheloang hohle ka tlamahano ea batho ka bobeli, joalo ka mefuta e meng, ke boiketlo ba bana le bophelo ba kelello, phello e ka bang teng ka lebaka la tšebelisano ea bonngoe le tlhokomelo ea batsoali le ba batjha. Kannete, ho nka karolo ha ntate ho tsa tlhokomelo ea bana ho hlokomelehile ka ho lekana joalo ka litšusumetso tsa bo-mme ho ntlafatso e atlehileng ea ngoana. Mekhatlong ea pele ho lenyalo le linaheng tse tsoelang pele, mohlala, moo lijo le tlhokomelo ea bophelo li sa fumaneheng habonolo, bana ba basali ba nyetsoeng ka mokhoa o nyane ba na le litekanyetso tse tlase tsa batho ba shoang ho feta bana ba basali ba sa nyalanang kapa ba leng mokhatlong oa polygynous [206]. Mekhatlong e tsoetseng pele ea indasteri, boteng ba bo-ntate ba tsotellang bo ntlafatsa bophelo bo botle ba maikutlo le ba kelello ba bana, joalo ka ha ho bontšoa ke maemo a holimo a katleho ea ngoana linthong tse fapaneng, ho kenyelletsa katleho ea thuto [41, 71, 83, 88, 181, 191] le thibelo le kalafo ea mathata a ho tšoenyeha [28], tlhokomelo-deficit / hyperactivity disorder (ADHD) [75], tšebeliso ea lithethefatsi, le boitšoaro ba botlokotsebe [200].

Leha ho mamella maqhama pakeng tsa balekane ba baholo ho bohlokoa bakeng sa bophelo bo botle ba 'mele le ba kelello ba batho le bana ba bona, hape ho ka ba le ts'usumetso ea botsitso sechabeng, ha re tsebe ho fokolang ka neurobiology ea pairing. Sena se bakoa ke taba ea hore litoeba tsa setso tsa laboratori tse sebelisitsoeng boithuto ba boitšoaro ba neuroendocrinology ka kakaretso ha li supe litšobotsi tsa sebopeho sa bonto, 'me ka hona li ka se sebelisoe joalo ka lits'ebetso tsa mohlala bakeng sa ho ithuta ka maqhama a mabeli. Ha mefuta e fapaneng ea liphoofolo e sa tloaelehang e hlahile ho ithuta boitšoaro bona bo sa tloaelehang, ho kenyeletsa le litšoene tsa marmoset le titi [15, 197] le litoeba tsa California [24-26, 59, 189], re tla tsepamisa maikutlo ho se seng se atileng haholo; The prairie vole (Microtus ochrogaster). Re tla qala ka ho hlalosa lithuto tsa lebaleng le tsa laboratori tse hatisang boitšoaro ba tlamahano ea tumellano. Ebe re tla tšohla mosebetsi oa pele ka laboratoring e hlalositseng mekhoa e metle ea boitlamo ba ho tlamahane ka nako ea lithatohatsi. Ka mor'a moo, re tla tšohla mekhoa ea methapo e amanang le mekhoa e meraro e arohaneng e amanang le ho tlamahana ka bobeli; theho ea likhetho tsa balekane, nts'etsopele ea mabifi a ikhethileng ho bonts'a maikutlo a sa tloaelehang, le tlhokomelo ea batsoali ka bana ba banyenyane-tse tsepamisang maikutlo haholo-holo tlhokomelong ea bo-mme kaha tlhokomelo ea basali e atile ho mefuta eohle ea liphoofolo tsa matsoalloa mme e hlahlobiloe haholo libakeng tse ling [31, 170, 171, 199]. Re tla shebisisa ho kenya letsoho ha li-neuropeptides arginine vasopressin (AVP) le oxytocin (OT) le neurotransmitter dopamine (DA) ka boits'oaro bona, empa re tla shebisisa le li-neurochemicals tse ling tse kentsoeng ts'ebetsong ea maqhama a mabeli. Qetellong, re tla hlahloba hore na li-neurochemicals tsena li ka sebetsa 'moho joang ho laola ho theha le ho boloka litlamo tsa maqhama.

2. Mohlala oa mofuta o motle oa tlotlo

2.1. Lithuto tsa masimong tsa boitšoaro

The prairie vole ke mofuta o mong oa bophelo bo botle ba litonanahali tse lulang merung ea United States bohareng [106]. Ho fanoe ka maikutlo a hore ho ikamahanya le tikoloho ena e thata, e nang le mehloli e fokolang ea lijo le phepelo e fokolang ea metsi [27, 92, 159], mohlomong e kentse letsoho phetohong ea leano le lecha la bophelo ba motho ho mofuta ona [38, 218].1 Lithuto tsa masimong tsa khale tse sebelisang maraba a ho hapa batho ba bangata li fane ka bopaki ba hore li-voes de prairie li theha litlamo tsa nako e telele 'me li tsamaea hammoho naheng, kaha lipere tsa banna le basali li ne li kopantsoe hammoho khafetsa [94]. Ntle le moo, ts'ebeliso ea radiotelemetry e kopantsoeng le ho ts'oasa libere li lumelletse ho bona hore lipakeng tsa banna le basali ba lula lihlaha le ho arolelana libaka tsa lapeng nakong ea nako ea bobeli ea ho tsoala le ho se tsoale.69, 94, 95]. Liphuputso tse ling li bonts'itse hore mafahla a joalo a ho tsoala a lula a le 'moho ho fihlela setho se le seng se e-shoa,' me maemong a mangata, molekane ea setseng ha a tsamaee le molekane e mocha [38, 96, 97]. Ntle le moo, li-voales tsa banna tsa tlholeho li kenya letsoho ho lebeleng sehlaha, ka ho khetholla tse tona le tse tšehali sebakeng se haufi le sehlaha le ntlo, hape li kenya letsoho moahong oa sehlaha [97, 205]. Leha boits'oaro ba botona ba botona ba banna bo ne bo le thata ho bona maemong a tlholeho, ka lebaka la liphumano tse hlalositsoeng kaholimo le tekanyo e phahameng ea letsete la bo-ntate e fumanoang mefuteng e meng ea monogamous, ho ne ho boletsoe esale pele hore li-voes tsa praherie tsa banna e ne e le tsa ntate haholo.205, 230], mme polelelopele ena e tiisitsoe lithutong tse latelang tsa boitshwaro tlasa maemo a laboratori.

2.2. Lithuto tsa laboratori tsa boits'oaro

Boitšoaro bo kopanyang ba li-Prairie vole bo tsebahala haholo ka laboratoring. Likamano tsa botona le botšehali tsa thobalano li na le katleho sechabeng haholo 'me li bontša boitšoaro bo kopanetsoeng bo kopanetsoeng ho li-Conspecifics [194]. Kamora ho lula hamonate le / kapa ho hloa mekoalaba, libaka tsa tumello li ntšetsa pele litabatabelo tsa setjhaba le tsa thobalano ho balekane ba li tloaetseng [68, 69, 102, 229]. Mokhatlo ona o ikhethileng (Setšoantšo sa 1A) e tsamaisana le mabifi a khethiloeng ho li-confpecifics tse sa tloaelehang [8, 99, 100, 124, 223, 224, 231]. Ntle le moo, balekane ba babeli ba arolelana sehlaha, ba lula ba le hammoho nakong ea kemaro, 'me ba bonts'a tlhokomelo e ntle ea motsoali le motsoali o mong nakong eohle ea kotlo [158, 174]. Ka tlase, re hlalosa ka botlalo boits'oaro bona le mekhoa ea boitšoaro e sebelisitsoeng ho li lekanya.

Setšoantšo sa 1 

Ts'oaetso ea laboratori ea boits'oaro bo amanang le ho tlamahana ka bobeli. (A) Litšoantšo li bontša monna ea motona le e motšehali ea bonts'a mahlakore a mabeli. (B) Lisebelisoa tsa likamore tse tharo tse sebelisetsoang tlhahlobo bakeng sa likhetho tsa molekane. Ho na le mekotla e meraro e tšoanang ...

Sebopeho sa likhetho tsa molekane ke lenane le tšepahalang la tlamahano ea maqhama, mme le tšoauoa ka kopano e ikhethileng, kamano, le ho qapanyetsana le molekane mabapi le osele [105]. Sebakeng se laoloang, boitšoaro bona bo ithutoa ho sebelisoa tlhahlobo ea likamore tse tharo ea balekane ba ratang pele e hlahisitsoeng ka laboratoring ea Dr. Sue Carter [229] mme ka mora moo e amohetswe ke laboratori tse ling tse ngata. Lisebelisoa tsa ho etsa liteko li na le k'harare e bohareng e hokahantsoeng ke methopo e meholo ho tse peli tse tšoanang, e 'ngoe e na le phoofolo e tloaetsoeng (molekane) ha e' ngoe e le phoofolo e sa tsejoeng (osele) (Setšoantšo sa 1B). Liphoofolo tsena tse peli tse susumetsang li kenngoa ka sekhukhu ka mekotleng ea tsona 'me ha li lumelloe ho kopana. Nakong ea tlhahlobo ea khetho ea molekane oa 3-hr, taba e kenngoa ka kamoreng e bohareng mme e lumelloa ho tsamaea ka bolokolohi ho lisebelisoa tsohle tsa liteko. Liranteng tse ling, ho sebelisoa lenaneo la "likhomphutha" le kopantsoeng le khanya ea khanya ea lifoto tse kentsoeng liphaepheng tse khubung e hokahanyang mabokose a marang-rang. Boitšoaro ba sechaba, ho kenyelletsoa ho tlolelana le thobalano ka lehlakoreng le leng, li khethoa ka vide nakong ea tlhahlobo ena 'me hamorao ea hlakoloa. Ho theha khetho ea molekane ho kenella ha taba e qeta nako e ngata ho hokahana le molekane ntle le moeti. Likopanong tsa botumo tsa banna le basali, lihora tsa 24 tsa ho lula hammoho ka ho tsofala li etsa hore ho khethoe balekane, athe lihora tsa 6 tsa ho phelisana ka bohanyetsi ka nako ea bocha ha li khothalletse boitšoaro bona [124, 125, 229] (Setšoantšo sa 1C). Paradigm ena ea boitšoaro e sebelisitsoe ka katleho lithutong tsa neuroanatomical, neurochemical, le pharmacological ho hlahloba neurobiology ea sebopeho sa khetho ea molekane [216, 237, 245].

Boitšoaro bo bong bo hlahang kamora ho hloa mekoallo ea lipina tsa boithabiso ke ho futuhela batho bao re sa ba tsebeng. Tlhekefetso ena e lebisoa ho banna le basali ba sa tsejoeng, empa eseng molekane eo ba mo tloaelang, ka hona e reiloe 'mabifi le khethiloeng'. Khethollo e khethiloeng libakeng tsa prairie e hlahlojoa ka laboratoring e sebelisa paradigm ea moahi ea tšoanang le e sebelisitsoeng litoeba [162, 231]. Paparong ena, phoofolo e sa tsejoeng e kopanetsoeng (intruder) e kenngoa kahara lehae la sehlooho sa taba eo (moahi). Litšebelisano tsa boitšoaro bo pakeng tsa moahi le intruder li bonoa nakong ea tlhahlobo ea metsotso ea 6-10, 'me khafetsa le nako e telele ea boits'oaro bo fapaneng bo khaotsoa kamora nako. Boithuto bo sebelisang paradigm ena bo bonts'itse hore li-voes tsa banna ba botona ba botona le botšehali li bonts'a maemo a tlase haholo a mabifi ho bahlaseli [124, 224, 231]. Leha ho le joalo, kamora lihora tsa 24 tsa ho se kopane ka tlhalohanyo, boits'oaro bo mabifi ho ba kenang bo ntse bo eketseha haholo [124, 224, 231]. Ha tlhekefetso ena e lebisoa ho ba batona le ba batšehali, boits'oaro bo matla bo hlaselang bo lemohuoa ho ba batona feela, empa eseng basali ba basele, ka nako ena [224]. Khethollo e mpe e ntse e tšoarella; e nka bonyane libeke tse peli kamora ho khethoa ha molekane [8, 99, 100]. Ho feta moo, ba batona ba roetse nako e telele ea nako (joalo ka banna ba babeli ba kopantsoeng), ho fapana le ba roalitsoeng ka lihora tsa 24 feela, ba bontša boits'oaro bo matla ho basali ba sa tsebeng, esita le ba amohelang bohlola ka ho hanana, mme ka ho etsa joalo ba nyala balekane ba bona ba bacha (Setšoantšo sa 1D) [8, 99, 100]. Ka hona ho fanoe ka maikutlo a hore mabifi a khethang ha a bapala karolo ea bohlokoa ho lebela molekane le tšimo [37, 38], empa e ka sebetsa ho boloka bonto e teng [8, 10] le ho fokotsa likopi tsa tse ling tse eketsehileng. Le ha mabifi a khethiloeng a lekoa feela ka mokhoa o hlophisehileng libakeng tsa basali ba rorisang banna, bopaki bo teng ho fana ka maikutlo a hore basali le bona ba ka bontša mokhoa ona oa boits'oaro [94]. Polelo e ts'epahalang ea khetho ea molekane le mabifi a khethiloeng ke li-prairie voles maemong a laoloang ka hloko a laboratori e totobatsa ts'ebeliso ea mohlala ona oa phoofolo lithutong tsa neuroendocrine tsa boits'oaro.

Li-voirie tsa maqhubu, tse tšoanang le mefuta e mengata e etsang maqhama a pakeng tsa balekane ba baholo [86], bonts'a tlhokomelo e ntle ea motsoali ho bana (ke hore, ntate le ntate ba thusa ho holisa bana) (1E ea setšoantšo). Ha tlhokomelo ea bo-mme e atile ho mefuta eohle ea liphoofolo tse anyesang, re tla tsepamisa moqoqo oa rona tlhokomelong ea batsoali le motsoali e le karolo ea ntate. Boitšoaro ba bo-ntate ho li-voes tsa prairie bo bonoe ka laboratoring ba sebelisa libaka tse ling tsa tlhaho ea tlhaho [104, 158, 174]. Kamora ho ts'oaroa ka matlakala, bo-ntate ba bonts'a mekhoa eohle ea boits'oaro ba batsoali bo bonahatsoang ke basali, ntle le booki [174, 205]. Tsena li kenyeletsa boits'oaro bo tobileng ba botsoali, joalo ka ho tsikinyeha (ho re, ho pikitla), boitlhopho, ho ikopanya le ho fumana malinyane le boits'oaro bo sa tobang, bo kang moaho oa sehlaha le ho ts'oara lijo [104, 174, 205, 230]. Bo-ntate ba bile ba tsoela pele ho bontša tlhokomelo e lerato ea ntate ho bana ba bona ba banyenyane ka mor'a hore ho be le litamati tse latelang [218, 220]. Leha ho le joalo, boteng ba bana, bo-ntate ba tlotloang ba qeta nako e fokolang ba le sehlaheng sa tlhaho ba bonahatsa boits'oaro ba ntate le nako e eketsehileng ea ho hlasela [93, 218]. Ho ba teng ha bana ho ka fokotsa tlhokeho ea tlhokomelo ea ntate ka kotloloho ke ntate, hobane bana ba setseng setsing sa tlhaho ka nq'ane ho weaning hangata ba kenya letsoho tlhokomelong ea matlakala a latelang —e leng boitšoaro bo bitsoang 'alloparenting' [104, 198, 218, 220, 222]. Boitšoaro bo sa tsotelleng bana le basali ba baholo ba hlomphuoang ea botona le botšehali bo tšoana le tlhokomelo ea ntate ho bo-ntate [198, 218, 220], mme boits'oaro bona ba bo-ntate bo matlafatsoa ke boiphihlelo ba kahare / tsa thobalano le mosali ea sa kopaneng [18]. Habohlokoa, boteng ba ntate le ponahatso ea bo-ntate ba bontšitsoe ho nolofatsa nts'etsopele ea 'mele le boits'oaro ba bana [4, 218, 220], ho fumanwa ho ts'oanang le litlamorao tse boletsoeng pejana tsa tlhokomelo ea ntate ho bana ba rona. Kahoo, ho utloisisa mekhoa e tataisang boits'oaro ba bo-ntate ho ka fana ka leseli la bohlokoa mabapi le tlhokomelo e nepahetseng ea batsoali ho liphoofolo tsa liphoofolo tse anyesang tse kenyelletsang bonngoe ba rona.

3. Lits'ebetso tse kopanetsoeng tsa tumellano ea tlholeho ea tlholeho

Boithuto ba pele bo neng bo etsa lipatlisiso tsa li-neural amanang le li-bonding bo bapisa li-neuropeptide le li-neurotransmitter pakeng tsa mefuta ea vole e bonts'itseng merero ea bophelo e fapaneng. Mefuta e mene e sebelisitsoeng e ne e le prairie, phaine (M. pinetorum)M. pennsylvanicus) le montane (M. montanus) maqhubu. Monogamous prairie le li-pine voles li theha maqhama a pakeng tsa balekane ba baholo mme li bonts'a tlhokomelo e ntle ea motsoali le bana ka nako e le 'ngoe ha li thehe maqhama a mabeli ebile li bontša tlhokomelo ea bo-mme feela [37, 82, 91, 95, 104, 124, 126, 127, 154, 155, 158, 174, 230]. Kamano e haufi ea taxonomic e arolelanoang ke mefuta ena, hammoho le phapang ea bona molemong oa bophelo e etsa hore litoeba tsena li be ntle bakeng sa lithuto tse bapisoang tse etsang lipatlisiso tsa boitšoaro ba sechaba (bakeng sa tlhahlobo, bona [237]).

Joalokaha AVP le OT li ne li tsejoa ho laola boitšoaro bo ikhethileng ba sechaba, ho kenyelletsa boitšoaro ba thobalano (bakeng sa tlhahlobo, bona [11]), mabifi [79], le tlhokomelo ea bo-mme [129, 176, 177], ho ne ho boletsoe esale pele hore lits'ebetso tsena tsa neuropeptide li tla fapana pakeng tsa mefuta e nyonyehang le e litšila [17, 122]. Ho leka tšebetso ena ea moea, mekhoa ea ho jala ea lisele tsa AVP le OT, likhoele le li-receptor li ile tsa kenngoa bokong ba vole. Ho mefuta eohle e hlahlojoang e hlokometsoe, ho sa tsotelehe leano la bophelo, li-neuron tsa AVP-immunoreactive (AVP-ir) li ile tsa fumanoa libakeng tse 'maloa tsa boko, ho kenyelletsa le "patventricular (PVN)" le "supioptic" (SON) nuclei ea hypothalamus. (BNST), medial amygdala (MeA), anterior hypothalamus (AH), le sebaka sa preoptic (POA) [17, 221, 223]. Li-fibers tsa AVP-ir li fumanoe ho lateral septum (LS), lateral habenular nucleus (LHN), diagonal band (DB), BNST, medial preoptic area (MPOA), le MeA [17, 223]. Lisele le likhoele tsa OT-immunoreactive (OT-ir) li ne li le libakeng tse 'maloa tsa boko mefuta e' ngoe le e 'ngoe, ho kenyeletsoa PVN, SON, MPOA, le BNST [223], 'me likhoele tsa OT-ir li fumanoe hape ka har'a nucleus accumbens (NAcc) [187]. Leha mefuta-futa ea tlhaho e fumanoe, ka kakaretso, mekhoa ea phepelo ea li-neuron tsa AVP-ir le OT-ir le li-polokelo li bolokiloe hantle lipakeng tsa mefuta e nyarosang le e litšila ea mofuta oa eona [187, 221, 223].

Phapang e sa tloaelehang ea mefuta e hlokomeloe, leha ho le joalo, ka mekhoa ea phepelo le methapo ea tikoloho ea li-AVP le li-OTR (li-OTR) Li-voirie voles, mohlala, li na le methapo e meholo ea li-AVP-V1a receptors (V1aRs) ho BNST, ventral pallidum (VP), bohareng (CeA) le basolateral (BLA) nuclei ea amygdala, le accessory olfactory bulb (AOB), hara libaka tse ling, ho feta li-vooms tsa montane, athe li-V1aR tse phahameng li bonoa ho LS le medial prefrontal cortex (mPFC) ea li-voes tsa montane ho feta li-voes tsa prairie [123, 145, 196, 225, 241] (Setšoantšo sa 2A). Ho khahlisang, ha mefuta e mengata ea li-vole e ne e bapisoa, libaka tse ikhethang tsa thorisoe le li-peine li bontšitse sebopeho se tšoanang sa V1aR, mme mokhoa ona o ne o fapane le oa mehendi e hlephileng le li-voes tsa montane [123, 145], e bonts'ang ho nka karolo ha V1aR e ikhethang ea tikoloho mesebetsing ea kelello le boits'oaro e amanang le maano a bophelo a fapaneng ka li-voles [123, 145, 196, 241]. Ka mokhoa o ts'oanang, mekhoa ea peo le boholo ba libaka tsa li-OTR le tsona li fapana pakeng tsa mefuta e nyonyehang le e litšila ea tlholeho. Li-monogamous prairie le li-voine tsa phaene, ka mohlala, li na le mathata a phahameng a OTR ho BNST, mPFC, le NAcc ho feta li-pearl le li-montane voice.Setšoantšo sa 2D), athe mokhoa o fapaneng o fumaneha maemong a OTR a tlamisang mofets'e oa moea oa "hypothalamus" (VMH), LS, le anterior cortical amygdala (AcA) [122, 196, 239]. Phapang ea mefuta ea phallo ea V1aR le OTR e tsitsitse ho pholletsa le nako ea bophelo [215, 225'me li batla li le receptor, hobane ha ho khethollo e joalo lits'ebetsong tsa benzodiazepine kapa opiate receptor [122]. Ka hona, ha re fuoa karolo ea AVP le OT ka boits'oaro ba sechaba, liphapang tsa mefuta ea li-V1aR le li-OTR ho nahanoa hore li amana ka ho khetheha le phapang ea mefuta ea boits'oaro bo amanang le mekhoa e fapaneng ea bophelo li-voles [107].

Setšoantšo sa 2 

Vasopressin (AVP) le taolo ea oxytocin (OT) ea sebopeho sa khetho ea molekane. (A) Mefuta ea phapang ea vasopressin receptor (V1aR) e tlamang kahare ea venral pallidum (VP) ea prairie le montane voles. Methapo e phahameng ea li-receptor e bontšoa ke tse ling ...

Phapang e atileng ea mefuta ea phallo ea li-neuropeptide receptor e hlalositsoeng ka holimo e ka ba ka lebaka la liphapang tse poteletseng tse boletsoeng libakeng tse khothalletsang tsa V1aR le OTR [239, 240, 242, 243]. Leha sebopeho sa liphatsa tsa lefutso tsa V1aR le OTR li-coding li tšoana hantle ka mefuta eohle ea li-vole [239, 240, 242, 243], li-prairie le li-pine voles li na le tatellano ea li-microsatellite tse ngata tse phetoang tikolohong ea papatso ea mofuta oa V1aR tse sa fumaneheng mefuteng ea mepopotlo kapa ea montane, 'me liphetoho tsena tse latellanang li ka baka phapang ea mefuta ea polelo ea li-receptor [107, 108, 242, 243]. Ho ts'ehetsa mohopolo ona, litoeba tse nkileng kh'outu bakeng sa V1aR, li bonts'itse lipaterone tsa V1aR tse ts'oanang le li-voles tsa prairie [243]. Ho khahlisang, ha e kenella ka AVP, litoeba tsena tsa transgenic li bonts'itse ts'ebelisano e ntlafalitsoeng ea sechaba, e bonts'a hore mekhoa ea phepelo ea li-receptor e ka susumetsa karabelo ea boko ho li-neuropeptides tsa endo native, mme ka tsela ena, e ka fetola boits'oaro ba sechaba [243].

Haufinyane tjena, liphuputso tse bapisang li fuputse lits'ebetso tse bohareng tsa DA ka mefuta e metle haholo, hobane DA, joalo ka AVP le OT, e bapala karolo e tsebahalang lits'ebetsong le boits'oarong bo amanang le ho tlamahana ha lenyalo, ho kenyelletsa ho ithuta le ho hopola [1, 23, 141, 234], olfaction [164], boitšoaro ba thobalano [22, 117], le boitšoaro ba motsoali [170, 171]. Lithuto tsena li hlokometse liphapang mokhoeng oa ho tsamaisa lisele tsa DA le li-receptor hammoho le phapang litekanyetsong tsa libaka, lipakeng tsa maeto a boitšoaro bo bobe le a boitšoaro bo bobe bo ka amanang le boitšoaro ba sechaba.

E lumellana le se fumanoeng mefuteng e meng ea litoeba [44, 114, 167, 210], Lisele tsa DA-tse ngotsoeng bakeng sa tyrosine hydroxylase (TH; tekanyetso e fokotsang enzyme ho catecholamine synthesis) ha ho e-na le DA beta hydroxylase (enzyme e fetolang DA ho ba norepinephrine) - e fumanoe libakeng tse ngata sebakeng se matla sa monogamous prairie vole bokong, ho kenyelletsa khubu ea mantlha ea BNST (pBNST), posterodorsal MeA (MeApd), le sebaka sa ventral tegmental (VTA) [99, 168]. Ntle le moo, letsoalo le phahameng la polokeho ea terminal ea DA le teng ho NAcc le caudate putamen (CP) [7], le liteko tsa morao-rao tse lekotsang liteko tsa banna li bontšitse hore liteishene tsena li tsoa li-neuron tsa proion ho VTA [101], joalo ka ha ho bontšitsoe ho mefuta e meng hape [34, 128, 193]. Leha ho le joalo, li-voales tsa boitšoaro bo bobe li na le lisele tse 'maloa haholo, haeba li teng, lisele tsa DAergic tse ho PBNST le MeApd [168], e bonts'ang phapang e teng pakeng tsa mefuta e nyarosang le e mpe ea tlholeho.

Ho ajoa ha Dopamine receptor (DAR) bokong ba vole le hona ho tsebahetse. Li-DAR li ka aroloa ka malapa a mabeli a mantlha, D1-like (D1R) le li-receptor tsa D2-like (D2R), tse arohantsoeng le meaho ea tsona ea limolek'hule, li-affinities tsa pharmacological, le litlamorao ho litseleng tse sa tsitsang tsa ho saena [163, 166]. Li-voles tsa prairie, li-D1R li fumanoa NAcc, CP, le mPFC, hammoho le libakeng tse ling tsa boko [8, 196] (BJ Aragona, Y. Liu, le ZX Wang, data e sa phatlalatsoang). Li-D2R, ha li le teng libakeng tsena tse tšoanang, li ka fumanoa le VTA le substantia nigra (SN) [8, 196] (BJ Aragona, Y. Liu, le ZX Wang, data e sa phatlalatsoang). Le ha karolelano ea li-receptor e ts'oana le e fumanoang mefuteng e meng ea litoeba, mesaletsa ea tsona e ts'oanehile 'me e kanna ea lumellana le phapang ea mefuta ea boitšoaro sechabeng [8, 196]. Mohlala, li-voogamous prairie voles li na le methapo e phahameng ea li-D2R le maemo a tlase a D1Rs mPFC ho feta li-voales tsa boitšoaro bo bobebe [196]. Ntle le moo, li-voles tsa mehendi li na le letsoalo le phahameng haholo la li-D1R ho NAcc ho feta li-voles tsa prairie, mohopolo o fumanoang o amana le boemo bo tlase ba boikamano ba sechaba bo boletsoeng li-voales voles [8]. Ka sebele, blockade ea pharmacological ea D1Rs ho NAcc e ile ea eketsa boits'oaro bo kopanetsoeng boitsebisong tsa mehendi [8].

Ha li nkuoa hammoho, liphuputso tsena li bonts'itse phapang lits'ebetsong tsa AVP, OT, le DA lipakeng tsa mefuta e mebe ka mekhoa e fapaneng ea bophelo. Ka lebaka leo, bafuputsi ba tsepamisitse maikutlo lits'ebetsong tsena bokong ba prairie vole ho hlahloba ka nepo ts'ebetso ea boits'oaro e amanang le tlamahano e pakeng tsa balekane, ho kenyelletsa sebopeho sa khetho ea molekane, tlhaselo e ikhethileng le boitšoaro ba ntate. Re tla tšohla taolo ea methapo ea kutlo e 'ngoe le e' ngoe ea boits'oaro bona, likarolong tse latelang.

4. Neurobiology ea sebopeho sa khetho ea molekane

Ts'ebetso ea Brain ea 4.1 e amanang le sebopeho sa khetho ea molekane

Mokhoa o mong o sebelisoang haholo ha ho etsoa lipatlisiso tsa tšebelisano lipakeng tsa boko le boits'oaro ke ho etsa polelo ea pele ea mofuta oa bokong kamora tlhahlobo ea boits'oaro. Mohlala, Fos ke sehlahisoa sa proteni ea mofuta oa pele, fos, e hlahisoang ka potlako ho li-neuron tse latelang ts'ebetso mme e ka bonoa habonolo ke immunocytochemistry. Ka hona, lits'oants'o tsa Fos immunoreactive (Fos-ir) li sebelisitsoe litekong tsa boitšoaro tsa neuroendocrine ho khetholla ts'ebetso ea "neuronal activation" bokong e amanang le pontšo ea boits'oaro bo itseng.

Likopanong tsa prairie, ho kopanya batho ba bong bo fapaneng, ho kopanya batho ba bong bo fapaneng, le / kapa ho kopanya ho tlatsitseng Fos-ir maemong a mangata a boko ho kenyeletsoa MeA, BNST, le MPOA ho ba batona le ba batšehali [56, 169]. Mating, haholo, e ne e amana le litekanyetso tse eketsehang tsa Fos-ir ho MeA, BNST, MPOA, le ntlha e hlakileng ea medulla oblongata, e fana ka maikutlo a libaka tsena tsa boko e le likarolo tsa ts'ebetso tsa potoloho ea mating tse ka kenyang letsoho ho rateng ha molekane [50, 51, 169]. Karolo ea MeA ho khetholliseng molekane oa mabotho e bontšitsoe hape ke lithuto tsa lesion, joalo ka ha mabala a tširelletsang maikutlo a MeA libakeng tsa praemia tsa banna a fokotsa boitšoaro ba bona bo kopanetsoeng ho mosali ea tloaelaneng empa a se na phello ho boits'oaro ba ho hlaka, ho hlonepha kapa ho etsa hampe. tlhotlhomiso [133].

4.2. Neuropeptide molaoana oa sebopeho sa khetho ea molekane

Bopaki ba pele bo bonts'ang hore AVP le OT li ka bapala karolo ea bohlokoa ho thehoeng ha balekane ho tsoa lipatlisisong tse lekolang litlamorao tsa boiphihlelo ba sechaba le tsa thobalano — litabatabelo tsa tlhaho ea tšusumetso ea molekane ka mokhoa oa tlhaho — lits'ebetsong tsena tsa neuropeptide bokong ba praeree vole. Liketelong tsa basali tsa basali, ho lula hammoho ka ho hola ho ile ha eketsa palo ea lisele tse ngotsoeng ke AVP mRNA ho BNST [214] le ho fokotsa letsoalo la likhoele tsa AVP-ir ho LS [18]. As BNST-AVP neurons projeke ho LS [60], datha tsena li fana ka maikutlo a hore ho hola ho thusa ho hlophisoa ha AVP ho BNST le tokollo ea AVP ho LS ea li-voes tsa banna tsa praemia [216]. Kaha ho nyalana ho bohlokoa molemong oa khetho ea molekane ho ba batona [124], datha tsena li fana ka bopaki bo lumellanang ba ho kenya letsoho ha AVP molemong oa khetho ea molekane. Ho basali, ho e-na le hoo, ho pepesetsoa litekanyetso tsa chemosensory ea banna tse fetotsoeng letsoalo la OTR ho AOB, ho bonts'a hore OT e ka bapala karolo ea ho khethoa ha balekane ho li-voes tsa basali tsa basali [233].

Bopaki bo tobileng ba karolo ea AVP le OT ho thehoeng ha lithahasello tsa balekane bo fanoe ke ho qhekella hoa litekanyetso tsa meriana. Tsamaiso ea Intracerebroventricular (icv) ea mohanyetsi oa V1aR o thibetsoe ho khetha khetho ea molekane litulong tsa banna ba rorisang, ha tsamaiso ea AVP e bohareng e susumetsa likhetho tsa molekane ha ho se letho la bona [43, 231]. Ka mokhoa o ts'oanang, tsamaiso ea li-icv ea AVP e kenyellelitseng thato ea molekane maikutlong a basali nakong ea 1 feela ea ho ikoetlisa ka monna, mme phello ena e ne e thibetsoe ke taolo e tšoanang ea mohanyetsi oa V1aR, e bonts'a hore AVP e laola mokhoa o ratoang oa molekane ho batšehali ka bobeli [43]. Kalafo ea OT e bile le tšusumetso ea khetho ea balekane ho ba batona le ba batšehali. Ka ho khetheha, tsamaiso ea icv OT e ile ea susumetsa khetho ea molekane ho ba batona le ba batšehali mme litlamorao tsena li ne li thibetsoe ke tsamaiso e kopanetsoeng le mohanyetsi oa OTR [43]. Le ha lintlha tsena li bonts'a hore AVP le OT li hlophisa sebopeho sa khetho ea balekane ho batona le ba batšehali, ho bohlokoa ho hlokomela hore litekanyetso tse sebetsang tsa li-neuropeptide li fapana pakeng tsa ba batona le ba batšehali [43].

Mananeo a ikhethileng a setsing esale a bonts'a libaka tse 'maloa tsa boko tse bohlokoa bakeng sa taolo ea AVP le OT ea sebopeho sa khetho ea molekane. Ho banna, tsamaiso ea mohanyetsi oa V1aR ka kotloloho ho LS kapa VP, empa eseng likarolo tse ling tsa boko, e thibetse ho thehwa ha litakatso tsa molekane oa mating, athe Tsamaiso ea AVP ka kotlolloho ho lithahasello tsa molekane tsa LS ka ho hloka mathoiso (Setšoantšo sa 2B) [146, 149]. Ho feta moo, tsamaiso ea mohanyetsi oa OTR ho kenella LS ea li-voes praeses tsa banna le eona e thibetse ho theha khetho ea molekane ea ratoang [149]. Ho basali, ho fapana le hoo, prelimbic cortex (PLC; karolo ea mPFC) le NAcc li kentsoe ts'ebetsong ea neuropeptidergic ea sebopeho sa khetho ea molekane [150, 244]. Maemo a OT a eketsehile ho NAcc nakong ea bolekane le monna [187]. Ntle le moo, ente ea OT ka kotloloho ho likhethong tsa molekane tsa NAcc ha e sa kopane, ha li-block tsa li-OTR sebakeng sena kapa PLC e thibetse ho theoa ha likhetho tsa molekane oa mating2E ea setšoantšo) [150, 244].

Boithuto bo 'maloa bo sebelisang phetiso ea vaerase ea maqhubu a viral ho fetisetsa le ho laola polelo ea liphatsa tsa lefutso libakeng tse itseng tsa boko bo tšehelitse liphumano tsa hore methapo ea methapo ea methapo ea methapo ea methapo ea methapo ea methapo ea methapo ea methapo ea methapo (VP le OT) ho NAcc ho laola likhetho tsa molekane ho li-voes tsa banna le tsa basali. . Ho banna, mohlala, vector e amanang le adeno e sebelisitsoe ho isa mofuta oa V1aR ho VP [183]. Joalokaha ho ne ho lebelletsoe, ho qhekella hona ho felletse ka bongata ba V1aRs tikolohong ena. Ho khahlisang, banna bana ba thehile lithahasello tsa balekane ho ba sieo ha ho nyalana, ho ts'ehetsa liphumano tse ntlafalitseng methapo ea kutlo ea VP ho VP li ka nolofalletsa khetho ea balekane molemong oa li-voes tsa banna prairie [183] (Setšoantšo sa 2C). Ho feta moo, V1aR overexpression ho VP ea li-voales tsa mehendi e tona, e kentse tlhoko ea khetho ea molekane ho mofuta ona o boitšoaro bo bobe ba sechaba [145]. Ka mokhoa o ts'oanang, OTR overexpression ho NAcc ea basali ba thobalano le basali ba thobalano le basali ba potlakisang ho hlekefetsoa ha ba bapisoa le taolo (Setšoantšo sa 2F), empa kalafo ena ha ea ka ea fetola khetho ea balekane ho li-voales tsa mehendi ea basali [188]. Ha li kopantsoe hammoho, liphuputso tsena li totobatsa bohlokoa le litlamorao tse ikhethileng tsa AVP le OT mabapi le theho ea khetho ea molekane lihlopheng tsa basali tsa basali le basali.

4.3. Taolo ea DA ea moralo oa khetho ea molekane

Mosebetsi oa morao-rao o bonts'itse hore khetho ea balekane ho li-voirie voles e boetse e laoloa ke DA bohareng, haholo ts'ebetso ea mesolimbic DA-sehlopha sa DA se hlahisang lisele tse tsoang VTA le projeke ea NAcc, mPFC, le likarolo tse ling tsa bokapele. Potoloho ena ea neural e nahanoa hore e nka karolo e kholo kabelong ea boleng bo susumetsang molemong oa tikoloho, e lebisang molomong oa boits'oaro bo tataisoang ke sepheo [120, 232]. Mohlala, mesolimbic DA e sa le nako e telele e qobelloa ho abela batho ba sa tsebeng letho ho khothaletsa lintho tse kang lijo le balekane ba amohelang litaba, ka tsela eo, mekhoa e kang ho fepa le ho ikatisa e bohlokoa bakeng sa ho pholoha [120, 232]. Ka mokhoa o ts'oanang, Mesolimbic DA e hlahisitsoe ho thusa ho khetha khetho ea molekane, ho etsa hore boiteko ba ho ts'oaroa bo tsepamise maikutlo ho li-confpecifics tse ratoang [80], khopolo-taba e tšehetsoang ke data e hlalositsoeng ka tlase. Ho kenya letsoho ha ts'ebetso ena molemong oa khetho ea balekane ho a utloahala boemong ba ho iphetola ha lintho, joalo ka ha khatello ea khetho e hlokang sebopeho sa tšebelisano pakeng tsa balekane e ka lebisa ho boleng bo eketsehang ba tšebelisano e fuoeng molekane, le kamano e ikhethileng eo e leng tšobotsi ea maqhama a mabeli.

Bopaki ba liteko ba pejana bo bonts'ang ho nka karolo ha DA molemong oa khetho ea balekane bo tsoa manollong a bongaka a meriana. Hopola hore lihora tsa 24 tsa ho tloaelana ho sebelisana le batho ba bang li etsa hore ba rate litlatse tsa banna le basali. Ha moralo oa khetho ea molekane o ne o sa angoe ke ente ea saline pele ho ho betla, kalafo ka mohanyetsi oa DAR, haloperidol, e thibetse likhetho tsa molekane tse khahlisang batho ba bong bo bong [7, 217]. Ho feta moo, kalafo e nang le tekanyetso e tlase ea apomorphine, agonist e sa sebetseng ea DAR, e nolofalitse ho thehwa ha lithahasello tsa molekane ka mor'a lihora tsa 6 feela tsa ho se sebetse ka nako ea ho ba sieo [7, 217]. Ha li kopantsoe hammoho, liphumano tsena li fana ka maikutlo a hore ts'ebetso ea DAR e bohlokoa bakeng sa kaho ea khetho ea balekane ho libaka tsa tumello ea tumellano.

Bopaki ba pele bo sebetsang ba ho kenya tšebetsong mesolimbic DA kamanong ea khetho ea balekane ke ho fumana hore ho tlolelana ha maraka ho eketsa ts'ebetso ea DA ho NAcc ea maqhubu a banna le ba basali ba rorisang [7, 98]. Ho basali, mohlala, maemo a DA a kantle ho naha a eketsehile hoo e batlang e le 51% kaholimo ho nako ea pele ea nako ea botona [98]. Ka mokhoa o ts'oanang, ba batona ba nang le balekane ba ne ba na le 33% ho tsoa ho DA tikolohong ena ha ho bapisoa le ba batona ba sa nyalanang [7]. Bopaki bo tobileng bakeng sa karolo ea NAcc DA popehong ea likhetho tsa balekane bo tsoa mananeong a itseng a bongaka a meriana le methapo ea kutlo ea DA. Microinjection ea haloperidol ho NAcc e thibetse ho hlophisoa ha litakatso tsa molekane oa mating, athe microinjection ea apomorphine e sebakeng sena e nolofalletsa ho ratoa ha balekane ha ho sa sebetse [7]. Litlamorao tsena li ne li ikhethile sebakeng sa marang-rang, joalo ka ho qhekelloa ha DAR ho CP, sebaka se bapileng le NAcc e amohelang bolulo ba DAergic ho tsoa libakeng tsa midbrain, ha e a ka ea fetola sebopeho sa balekane ba khethiloeng [7].

Liteko tse ling li sebelisitse li-agonists / antagonists tse ling ho bontša hore li-D1R le li-D2Rs ho NAcc li laola ka mokhoa o ratoang khetho ea balekane (Setšoantšo sa 3A & B). Ka ho khetheha, ts'ebetso ea NAcc D2R e nolofalitsoeng, le thibelo ea D2R e thibetsoe, sebopeho sa khetho ea molekane ho li-voes tsa basali le tsa basali tsa basali, se bonts'a hore ts'ebetso ea NAcc D2R e hlokahala ebile e lekane bakeng sa kaho ea molekane [8, 98]. Ka lehlakoreng le leng, ts'ebetso ea NAcc D1R e thibetse ho ratoa ha molekane le lipuisano tsa basali tsa D2R, ho bonts'a karolo ea thibelo ea NAcc D1Rs ka boitšoaro bona [8]. Taba ea bohlokoa ke hore manong ana a ne a sebetsa hantle feela ha a ne a tlisoa ka har'a khetla ea NAcc, empa e seng ea mantlha, e neng e supa melaoana ea likhetho tsa molekane ka har'a NAcc [8].

Setšoantšo sa 3 

Dopamine (DA) ho li-nucleus accumbens (NAcc) e tsamaisa sebopeho sa khetho ea molekane ho li-voes tsa prairie. (A) Cartoon e bontša potoloho ea mesolimbic DA. Lisele tse utloisang bohloko tsa DA sebakeng sa proven tegmental area (VTA) projeke ea NAcc le preortal cortex (PFC), ...

Taolo e ikhethileng ea DAR ea sebopeho sa likhetho tsa balekane ho NAcc haufinyane e se e hlahlobiloe maemong a makatsang. Li-D2R le D1R ka bobeli ke li-receptor tsa 7-transmembrane tseo litlamorao tsa eona tse kahare li kopantsoeng ke li-protein tsa Gter tse tlisang heterotrimeric (G-protein) (bakeng sa litlhahlobo, bona [163, 166]). Le ha li-D2R le D1R li e-na le litlamorao tse ts'oanang litseleng tse ling tsa ho saena, li laola ka mokhoa o fapaneng cyclic adenosine 3 ′, 5'-monophosphate (cAMP) e bonts'a Cascade ka alpha subunit ea liprotheine tsa G-tseo li sebelisanang le tsona [163, 166](Setšoantšo sa 3C). Li-D2R li tlama liprotheine tsa inhibitory G (Gcyi le Gcyo). Ha D2R e sebetsoa, ​​alpha subunit ea Gcyi / o inhibits adenylate cyclase (AC) shughuli, e lebisang ho thibelleng ha tlhahiso ea cAMP le ho fokotseha hoa tšebetso ea protheine kinase A (PKA) [163, 166]. Li-D1R, ho fapana le moo, li tlama ho protheine ea G-protein (Gcy)s le Gcyolf). Ts'ebetso ea D1R e lebisa keketseho ea ts'ebetso ea AC, tlhahiso ea cAMP le ts'ebetso ea PKA [163, 166]. Ha ts'ebetso ea D1R le D2R e ama phapang ea CAMP ka mokhoa o fapaneng, ho fanoe ka maikutlo a hore tsela ena e tšoaeang e ka tlasa taolo ea DAR e ikhethang ea sebopeho sa khetho ea molekane [9]. Ho ts'ehetsa ts'ebetso ena, phokotso ea ts'ebetso ea PKA ka har'a khetla ea NAcc, empa eseng ea mantlha, e nolofalletsang ts'ebetso ea khetho ea molekane ho li-voes tsa praemia tsa banna, sephetho se lumellanang le litlamorao tsa ts'ebetso ea D2R [8, 9] (Setšoantšo sa 3D). Ho feta moo, litekong tse peli tse arohaneng, ts'ebetso ea liprotheine tsa G-protheine le ts'ebetso ea PKA ts'ebetsong ea NAcc e 'ngoe le e' ngoe e thibetse ho thehoa ha likhetho tsa molekane oa mating, tse lumellanang le litlamorao tsa ts'ebetso ea D1R [8, 9] (Setšoantšo sa 3D). Habohlokoa, ho qhekelloa hona ha hoa ka ha fetola ho ruruha kapa nako ea ho ikopanya nakong ea lihora tsa 24, ho fana ka maikutlo a hore ho eketseha ha cAMP ho kena-kenana ka kotloloho le tlhahiso ea khetho ea molekane. Ha li etsoa hammoho, liteko tsena li bonts'a hore ho tšoaetsoa ha cAMP ho kenella ka hara khetla ea NAcc ho laola ho ratwa ha balekane, mme ho ka baka litlamorao tse tobileng tsa DAR ka boitšoaro bona.

5. Neurobiology ea mabifi a ikhethileng

Joalokaha ho se ho boletsoe pejana, kamora lihora tsa 24 tsa ho hola le ho theha khetho ea balekane, li-voes tsa praemia tsa banna li bonts'a maemo a phahameng a mabifi ho basele ba ikhethang, haholo banna bao ba sa ba tsebeng, empa eseng ho balekane ba bona [124, 224, 231]. Ntle le moo, kamora beke e le 'ngoe ho isa ho tse peli tsa ho lula hamonate le ho nyalana le balekane ba bona, mabatooa a mabili a kopantsoeng a bonts'a mofere-fere haholo ho ba batona le ba batšehali ho kenyelletsa basali ba amohelang thobalano, ka ho etsa joalo ba hana bao e ka bang balekane ba bona ba bacha [8, 99, 100, 231]. Tlhekefetso ena e khethiloeng e nahanoa hore e bohlokoa bakeng sa ho lebela molekane, tšireletso ea sehla, le ho boloka tlamahano e teng lipakeng tsa monna le molekane oa hae [8, 37, 38, 99, 100, 231]. Joalokaha ho hlalositsoe ka tlase, lithuto li bonts'itse libaka tse ngata tsa boko, le ho nka karolo ha AVP le DA molemong oa boitšoaro bona.

5.1. Ts'ebetso ea boko e amanang le mabifi a ikhethileng

Likarolo tse fapaneng tsa boko li bile le tšusumetso ho khethollo e mpe. Mohlala, ponts'o ea boitšoaro bona e kopantsoe le Fos-ir e phahameng ho MeA, BNST, MPOA, LS, le AH (Setšoantšo sa 4A) [99, 224]. Ho e 'ngoe ea libaka tsena, AH, ts'ebetso e khethollang e ile ea bonoa pakeng tsa ho pepesehela molekane eo u mo tloaetseng le motho eo a sa mo tsebeng [99]. Ka ho khetheha, li-voile tsa basali tse neng li pentiloe le mosali bakeng sa libeke tse peli (ke hore, tse kopantsoeng) ebe li pepesetsoa monna ea sa tsejoeng kapa oa mosali ea neng a e-na le letsoalo le holimo haholo la lisele tsa Fos-ir ho AH ho feta banna ba batona. senoloa hape ho molekane oa bona. Ho khahlisang, ba batona ba pepesitsoeng ke banna kapa basali ba kenang lipakeng le bona ba ne ba e-na le letsoele le phahameng haholo la lisele tse bitsitsoeng AVP-ir le Fos-ir tikolohong ena ea kelello ho feta banna ba pepesitsoeng ho balekane ba bona, ba fana ka maikutlo a hore AH AVP e ka laola mabifi a khethiloeng [99] (Setšoantšo sa 4B).

Setšoantšo sa 4 

Ho nka karolo ea Vasopressin (AVP) le dopamine (DA) ho mabifi a ikhethileng libareng tsa banna ba rorisang basali. (A) Photomicrograph e bonts'a 'mele le lisele tsa AVP-immunoreactive (AVP-ir) le li-fiber (brown cytoplasmic st lits), Fos-immunoreactive (Fos-ir) stain (lefifi la nyutlelie) ...

5.2. Neuropeptide molaoana oa mabifi a khethiloeng

Ka lebaka la karolo e tsebahalang ea AVP lipontšong tsa naha [79], le phapang ea kabo ea li-receptor ea AVP libakeng tse bonts'ang lipakeng tsa li-voogamous le polygamous voles [123, 225], AVP e ne e kopitsoe hore e kenelle taolong ea mabifi a khethiloeng. Boitekong ba pele ba ho leka mohopolo ona, Winslow et al. (1993) e fumane hore ente ea mohanyetsi oa V1aR, empa eseng mokelikeli oa "cerebrospinal fluid" (CSF), e kenang kahare ea moea nakong ea lihora tsa 24 ea ho hola e ile ea thibela ponts'o e ileng ea latela ea bofetoheli bo neng bo hlohlelletsa ho hlekefetsa ho hlekefetsoang ha banna. Ho kenyelletsa moo, ho kenella ha AVP ka har'a li-ventricles tsa morao-rao ho ile ha etsa hore motho a kenelle ka matla ho motho ea kenang lipakeng tsa botona le botšehali ba batona bao e seng basali. Melao e tšoanang ea sistimi ea OT ha ea ka ea fetola mekhoa e mabifi, e bonts'a hore AVP e bohareng, empa eseng OT, neurotransication e laola mabifi a khethiloeng ho li-voes tsa banna prairie [231].

Liphetoho tse ikhethileng tse mabapi le sebaka ho AH li tšehelitse maikutlo ana [100]. Ba batona ba botona le botšehali ba amohetseng ho kenella ka kotloloho ho AH ba bonts'a maemo a hlekefetsang haholo ho mosali e motona ho feta banna ba tšoaroang ka koloi kapa ka bobeli ba AVP le mohanyetsi oa V1aR, ho bonts'a hore UVP neurotransuction ho AH e ka baka moferefere libakeng tsa maqhubu a thorisoe (4E ea setšoantšo). Ho feta moo, ka li-voes tsa tlotlo tse tona tse kopantsoeng tsa banna, tokollo ea AVP ho AH e ne e phahame haholo lithutong tse pepesitsoeng phoofolo e sa tsejoeng ho feta tse pepesitsoeng ke balekane ba tsona (Setšoantšo sa 4C). Ho khahlisang, boholo ba phutuho ea AVP ho liphoofolo tsena bo ile ba lumellana hantle le khafetsa ba bona ba mabifi le hampe nakong ea ho hokahana. Ntle le moo, blockade ea V1aRs ho AH, empa eseng likarolo tse ling tsa boko, e thibetse pontšo ea mabifi a ikhethileng ho banna ba kopaneng, ka ho totobatsa AH AVP ka mokhoa ona (4E ea setšoantšo). Phuputsong e ts'oanang, ho fumanoe hore banna ba kopaneng ba na le li-density tse phahameng haholo tsa li-V1aR, empa eseng li-OTR, ho AH ho feta banna ba thobalano ba naïve (Setšoantšo sa 4D), ho fana ka tlhahiso ea hore bolekane bo kopane bo ka baka liphetoho tsa neuroplastic tsamaisong ea AH AVP e bakang ho hlaha ha mabifi a khethiloeng [100]. Khopolo-taba ena e ne e tšehelitsoe ke ho fumana hore maiketsetso a maiketsetso a V1aR ke phetisetso ea bong ea veter-mediated gene, ka tsa thobalano naïve prairie voles tse ntlafalitseng mabifi ho basali ba maoatle (Setšoantšo sa 4F)[100]. Ha li kopantsoe hammoho, lintlha tsena li bonts'a hore AVP ho AH e bapala karolo ea bohlokoa ho taolo ea mabifi a ikhethileng libakeng tsa basali tsa praemia.

5.3. Taolo ea DA ea mabifi a ikhethileng

Mesolimbic DA le eona e bile le tšusumetso ho bosholu bo ikhethileng, haholoholo, mabifi a bontšitsoeng ke ba batona ba tlamahaneng ka bobeli ho basali ba basele [8]. Litekong tse peli tse arohaneng, methapo ea DAR ka har'a masapo a li-voes tsa praemia tsa banna tseo ho neng ho etsoa thobalano le tsona e ne e bapisoa le ea ba batona ba hiriloeng le ba basali lihora tse 24 kapa libeke tse peli (ke hore, ba tlamahantsoeng) [8]. Le ha ho na phapang lipakeng tsa meno ea DAR e bonts'itsoeng lipakeng tsa banna ba nang le thobalano le banna ba neng ba ratana le basali ka lihora tsa 24, banna ba kopaneng ba ne ba na le maemo a holimo haholo a li-D1R, empa eseng D2R, ho NAcc, empa eseng CP, ho feta tsa bona tsa thobalano. balekane ba (naïve)Setšoantšo sa 4G le H). Joalo ka libeke tse peli, empa eseng lihora tsa 24, tsa cohabitation le mating li eketsehile NAcc D1R, liphetho tsena li bonts'a hore phetoho ena ea neuroplastic ha e hlokehe molemong oa ho theoa ha likhetho tsa molekane-sephetho se lumellanang le D2R, empa eseng D1R, molao oa sebopeho sa khethollo ea molekane se boletsoeng kaholimo-empa se supa ts'oaetso ea nako e atamelaneng ea moratuoa le molekane (ke hore8]. Ho khahlisang, keketseho ena ea maemo a D1R ho ba batona ba kopaneng e kopane le boits'oaro bo hlaha ba tlhaselo e nyarosang ho mosali osele (banna ba kopantsoeng le mosali libeke tse peli ba bonts'a mabifi a matla ho basali ba basele [8, 99, 100], athe banna ba etsang thobalano naïve kapa ba lumelletsoeng ho nyalana le mosali ka lihora tsa 24 ha ba [224]). Ka hona, e ne e le hypothesized hore maemo a eketsehileng a D1R ho NAcc ea ba batona ba kopaneng a ka laola pefo e khethang ho basali ba basele. Sethala se ikhethileng sa pharmacological ea NAcc D1Rs se sebelisitsoe ho leka tlhahiso ena ea maikutlo. Ha ba batona ba tlamelletsoeng ka bobeli ba tšoaroang ke CSF ba bonts'itse moferefere o matla ho mosali ea kenelletseng, ho kenella kahare-NAcc ka mohanyetsi oa D1R ho felisitse khalefo ena (Mofuta oa 4I). Ha re e kopanya, lintlha tsena li fana ka maikutlo a hore ho eketseha ha NAcc D1R ho ka baka phetoho e kholo ea boits'oaro e etsahalang mahaheng a banna ba rorisang ha ba ntse ba tsoela pele ho tloha boemong ba ho ba setho sa botona kapa botšehali ho ba maqhama a felletseng, e leng se lebisang ho moferefere ho basali ba basele le ho hlokomela maqhama a tiisitsoeng a [8]. Ka mokhoa o ts'oanang o ts'oanang le ts'ebetso ena, ho pepesetsoa hangata lithethefatsing tse tloaelehileng tsa tlhekefetso, amphetamine, ho eketsa mabifi ho khokahano le ho thibela sebopeho sa likhetho tsa molekane [100, 151]. Habohlokoa, liphetoho tsena tsa boits'oaro li tsamaisana le ho nyolisoa ha li-D1R ho li-NAcc le V1aR ho AH, ho bonts'a hore lithethefatsi tsa tlhekefetso li ka utsoa mefuta ea tlhaho ea neuroplasticity e iphetotseng ho boloka li-bond [100, 151].

6. Neurobiology ea Paternal Behavior

Boitšoaro ba bo-ntate-moholo bo tlalehiloe mefuteng e 'maloa ea liphoofolo tse anyesang tseo e seng batho, ho kenyelletsa le tamarins [246], marmosets [5], titis [160, 161], hamsters [118], likokoana-hloko [182], litoeba [24] le li-voles [174, 205]. Boithuto lithutong tse seng tsa botho bo shebile tšobotsi ea boits'oaro ba bo-ntate le litlamorao tse fetotsoeng ke tikoloho ea sechaba e bonts'ang mekhoa ena, 'me e fane ka leseli la bohlokoa la phetolelo bakeng sa bophelo ba motho. Boithuto lithutong, ho e-na le hoo, bo shebile haholo taolong ea boits'oaro ba bo-ntate 'me ba fane ka leseli la bohlokoa mabapi le mekhoa ea bo-ntate e ka tlas'a boits'oaro ba ntate. Leha panya ea California e bonahetse mofuta oa litoeba tse sebetsang bakeng sa morero ona [24-26, 59], mohlomong mofuta oa vole o sebelisitsoe haholo lithutong tsa neurobiology tsa boits'oaro ba ntate, mme dintlha tse tsoang liphuputsong tsena li akaretsoa ka tlase.

6.1. Ts'ebetso ea boko e amanang le boitšoaro ba bo-ntate

Joalo lipatlisisong tsa sebopeho sa khetho ea molekane le mabifi a khethiloeng, lithuto tsa pele tse neng li hlahloba boitšoaro bo botle ba bo-ntate li sebelisitse Fos-ir ho hlahisa libaka tsa boko tse hlahisoang ke ho pepeseha lipopae le ponts'o ea boitšoaro ba ntate. Kamora ho pepesehela popi e ts'oanang, li-voirs tsa banna tsa tumello li bontšitse ho eketseha ha Fos-ir libakeng tse ling tsa bokapele ho kenyeletsoa AOB, MeA, BNST, MPOA, le LS, ho fana ka maikutlo a ho ameha ha libaka tsena ho sebetsanang le likamano tse amanang le bana le / kapa ho taolo ea boits'oaro ba ntate [134, 222]. Karolo ea tsamaiso ea bokhelohi le MeA ho boits'oaro ba ntate e boetse e netefalitsoe ke lithuto tsa lesion libakeng tsa prairie. Tse tona tse fumaneng bulbectomy ea naha e 'ngoe li bontšitse phokotso e kholo boits'oaro ba bo-ntate hammoho le boits'oaro bo bong ba sechaba, ha bo bapisoa le ba batona ba fumaneng ts'ebetso ea sham [135]. Ho ekelletsa moo, mabala a pepesang a molumo oa ts'ebetso oa MeA a fokotsehile boits'oaro ba bo-ntate lithutong tsa banna ba rorisang ntle le ho ama boits'oaro bo bong bo kang ho phenyekolla, ho etsa lipatlisiso le ho etsa lipatlisiso ka ho hloka toka [133]. Qetellong, ho fapana le li-voes tsa prairie, ho pepesetsoa ha bana ha hoa ka hoa phahamisa haholo ho ngola Fos-ir ho MeA, BNST, MPOA, kapa LS ea li-voes tsa mehend tsa banna, ho fana ka tlhahiso ea bohlokoa ea libaka tsena tsa boko ho taolo ea tlhokomelo ea botona ea banna [222].

6.2. Taolo ea Neuropeptide ea bo ntate

Ntle le boits'oaro bo boletsoeng pejana, AVP e bohareng le OT le tsona li kentsoe tšebetsong ea boitšoaro ba batsoali, haholoholo basali. Ho kenngoa ha AVP ka har'a likhoto tsa basali tsa morao-rao ho etsa hore batsoali ba tsoelepele [177]. Ho feta moo, likhoto tse telele-Evans li bonts'a boits'oaro bo holimo ho botsoali ha bo bapisoa le mefuta e fapaneng ea AVP e haelloang, litoeba tsa Brattleboro [227]. OT, ka botlaaseng le bokong, e boetse e bapala karolo ea bohlokoa boits'oarong bo amanang le tlhokomelo ea basali, ho kenyelletsa le phello ea popelo ka popelong, ho tebela lebese nakong ea lebollo [87, 211] le taolo ea boitšoaro ba bo-mme ho basali:142, 177]. Joalo ka ha OT le AVP li bile le tšusumetso ho boits'oaro ba batsoali ho basali, le boits'oaro bo bong ba boiketlo ba basali le ba batona, bafuputsi ba ile ba qala ho lekola karolo eo li-neuropeptide tsena li e phethileng taolong ea boitšoaro ba motsoali oa monna.

Bopaki ba pele ba ho bonts'a karolo ea li-neuropeptides tsena boits'oarong ba bo-ntate li fanoe ke lithuto tse etsang lipatlisiso mabapi le kamano lipakeng tsa boiphihlelo ba bo-ntate le boteng ba AVP-ir fiber kapa polelo ea AVP / OT mRNA bokong. Li-voiri tsa Prairie tse neng li entsoe le mosali ka libeke tse peli kapa e ne e le bo-ntate ba nako ea pele ba bontšitse boits'oaro bo matla ba bo-ntate ebile ba na le methapo e fokolang ea methapo ea AVP-ir ka LS, empa eseng MPOA, ho feta balekane ba bona ba thobalano ba naïve [17, 18]. Ho khahlisang, phetoho ena ea letsoalo la LS AVP-ir fiber ha ea ka ea fumaneha ho bo-ntate ba se nang meeli-e bonts'ang litloaelo tse fokolang tsa bo-ntate ho bana - ho fana ka maikutlo a hore liphetoho LS AVP ka sebele li ka bapala karolo ea boitšoaro bo botle ba ntate oa ntate.17]. AVP ho PVN e boetse e kentse letsoho boits'oarong bo botle ba bo-ntate ba botona, kaha ho ngoloa ha AVP mRNA sebakeng sena ho ile ha eketseha ka li-voes tsa banna ba rorisang bao e sa tsoa ba bo-ntate, empa ba sa fetoheng ka tlhaho ka bo-ntate ba se nang bo-ntate ba montane vole [226]. Le ha ho tsejoe hanyane ka karolo ea OT ka boits'oaro ba bo-ntate lithutong tsa basali, ho na le bopaki bo bong ba hore neuropeptide ena e ka ameha. Mohlala, malinyane a boreleli a holisoang ke bo-'mè feela a amohelang likhalase tse nyane / ho koetlisoa le ho hola hanyane ka hanyane, ha a bapisoa le malinyane a holisitsoeng ke batsoali ka bobeli. Ha a le moholo, ea fetileng o bonts'itse boitšoaro bo botsoali bo sa tataisoang ke popi le ho eketsa polelo ea OT mRNA ho hypothalamus ho feta ea morao-rao, empa litlamorao tse joalo li ne li bonoa haholo ho basali.4]. Ha li nkuoa hammoho, lintlha tsena li bonts'a hore AVP le OT libakeng tse fapaneng tsa boko li ka laola boits'oaro ba ntate.

Ke lithuto tse fokolang tse ileng tsa hlahloba ka ho hlaka bohlokoa ba tšebetso ba AVP bohareng le OT ka boits'oaro ba ntate. Ho e 'ngoe ea lithuto tsena, liphetoho tse potileng li ile tsa bonts'oa ho boits'oaro ba botona ba botona le botšehali ba botona ba botšehali kamora ho tsamaisoa ha icv ho AVP kapa OT, athe ho kopanya phekolo ea icv ea OTR le mohanyetsi oa V1aR ho amme boitšoaro ba bo-ntate ka tsela e itšetlehileng ka mokhoa oa ho latela tekanyetso [13]. Ka litekanyetso tse tlase (1ng e 'ngoe le e' ngoe), bahanyetsi ba OTR / V1aR ba ne ba tloaetse ho eketsa bokhoni ba ho sutumetsa le ho ts'oaroa, ha litekanyetso tse phahameng (10ng e 'ngoe le e' ngoe) boits'oaro ba ntate bo fokotsoe haholo mme ho hlaha ha litlhaselo tsa malinyane ho ile ha eketseha haholo [13]. Le ha thuto ena e bonts'a hore AVP ea mantlha le OT li hlile li na le litlamorao tse sebetsang boits'oarong ba bo-ntate, liteko tse eketsehileng li hlokahala ho utloisisa karolo ea neuropeptide ka ngoe ho boits'oaro bo khethehileng ba bo-ntate le libaka tsa bona tsa ts'ebetso kahare ho bokong. Phuputsong e le 'ngoe feela ea ho etsa joalo ho fihlela joale, Wang et al (1994) o ile a hlahloba litlamorao tsa ho qhekella ha AVP ho LS ka mekhoa e mene e atileng haholo ea bo-ntate, ho kenyelletsa ho nyeka / boitlhopho, ho pikitla / ho koata, ho hokahana le ho khutlisa malinyane. Litaba tsa puisano ea botona le botšehali tse kenang ka thobalano tse kenang ka kotloloho kahare ho LS e sebelisitse nako e ngata haholo ho bontša liketso tsa bo-ntate, ho ikopanya ka ho khetheha le ho patela malinyane ho feta maqhubu a kentsoeng ka letsoai. Litlamorao tsena li ne li koetsoe ke ho entoa pele ke mohanyetsi oa V1aR ho LS, a fana ka maikutlo a hore LS AVP e hlokahala ebile e lekane ka taolo ea boits'oaro ba ntate [213].

Leha litlamorao tse ikhethileng tsa OT mabapi le boits'oaro ba bo-ntate li e-so ka li hlahlojoa, ho na le bopaki bo supang hore NAcc OT e ka ameha. Bopaki bona bo tsoa liphuputsong tse bapisang ho bonts'a phapang ea mefuta mefuteng ea NAcc OTR e lumellanang le phapang ea mefuta ea boits'oaro ba ntate [122], bohlokoa ba ts'ebetso ea OTR mekhoeng e meng e amanang le ho tlamahana ho ba batona (mohlala, sebopeho sa khetho ea molekane) [150, 244], le lithuto tse fapaneng tse tlalehang karolo ea NAcc OTR ka boitšoaro ba botsoali ba basali. Mohlala, mathata a NAcc OTR a amana le boits'oaro ba bo-'m'e nakong ea thobalano le basali ba baholo ba basali ba rorisang basali. Ka ho khetheha, basali ba neng ba bonahatsa boits'oaro ba basali ba basali ba ne ba e-na le mathata a phahameng a li-OTR ho NAcc ho feta basali ba sa kang ba bontša boits'oaro ba bo-mme kapa ho hlasela malinyane [173]. Kamano e tšoanang e tšoanang e ile ea bonoa pakeng tsa letsoalo la NAcc OTR le tlhokomelo ea bohle maemong a basali a manyane a basali ba rorisang [172]. Ho feta moo, letsoalo la OTR ho NAcc le kopantsoe hantle le boits'oaro bo bong bo amanang, ho kenyelletsa sebopeho sa khetho ea balekane [188]. Le ha ho so ka ho hlahlojoa ka kotloloho ho banna, datha tsena li bonts'a monyetla oa NAcc OT oa ho nka karolo boits'oarong ba bo-ntate.

6.3. Taolo ea DA ea bo ntate

Ha lipatlisiso tse ngata li hatisitse bohlokoa ba DA bohareng boits'oarong ba boimana (bona [171] bakeng sa ho lekola), lithuto tse fokolang li fuputse karolo ho DA e bohareng boits'oaro ba bo-ntate. Le ha e fokola ka palo, lithuto tsena li fane ka bopaki bo matla ba hore DA e nka karolo tlhokomelong ea batsoali ba banna.

Tekong e le 'ngoe feela ea tlhahlobo ea meriana ho hlahloba molao oa DAergic oa bo-ntate lithutong tsa bakhachane, Lonstein (2002) o bonts'itse hore DAR blockade e na le litlamorao tse fapaneng likarolong tse fapaneng tsa boits'oaro ba bo ntate (mohlala, ho ikopanya, ho nyeka le ho thothomela holim'a malinyane). Haholo-holo, ho thibeloa ha li-DAR le mohanyetsi ea morao-rao oa DAR, haloperidol, ho sitisitse boitšoaro bo bong ba ntate-ho kenyelletsa ho hokahana le malinyane — empa ho ntse ho matlafatsa ba bang, joalo ka ho tsikinyetsa bana. Le ha haloperidol e senya mosebetsi o akaretsang oa koloi ka tekanyetso e itseng [195], litlamorao tsa haloperidol boits'oarong bo bong ba bo-ntate, haholo-holo ho nyeka ha bana, li ile tsa bonoa ho litekanyetso tse sa fetoleng lintlha tse felletseng tsa mosebetsi, ho bonts'a hore ts'ebetso ea DAR e na le litlamorao tsa mantlha boits'oarong ba ntate. Ka hona, lintlha tsena ha li bontsane karolo ho DA boits'oarong ba bo ntate empa li supa hape hore taolo ea DAergic ea boits'oaro ba bo-ntate e ikhethile ka boits'oaro [153]. Ha ho na mananeo a tobileng a sebaka sa marang-rang a ntseng a sebelisoa ho senola likarolo tsa boko tse amehang taolong ea DAergic ea bo ntate. Leha ho le joalo, tlhahlobo ea 'mele e hlahisang ts'ebetso ea ts'ebetso ea methapo ho arabela ho malinyane e fane ka leseli le itseng mabapi le taba ena. Hopola hore bokong ba "prairie vole" bo na le sehlopha sa lisele tsa DAergic ka har'a pBNST le MeApd e nang le semorphic - banna ba na le lisele tse ngata tsa DAergic libakeng tsena ho feta tsa basali.168] - 'me lisele tsena li na le tšusumetso ho li-androgens le estrogens [40]. Ho khahlisang, lipalo tsena tsa lisele li hlahisoa (li bonts'itsoe ke Fos / TH-habeling habeli) bokong ba banna ba prairie vole kamora ho sebelisana le malinyane a Conspecific [168], mme ka hona a ka ameha boits'oarong ba bo-ntate.

Leha lithuto li se li entsoe ho fihlela mona, ho khothaletsoa hore NAcc DA le eona e ka bapala karolo ea boits'oaro ba bo-ntate. Joalo ka ha ho hlalositsoe pejana, Vole NAcc e na le liteishene tsa DA le litheolelo tse ngata ebile NAcc DA e bapala karolo ea bohlokoa ho taolo ea boits'oaro bo bong ba bophelo bo amanang le tlamahano, ho kenyelletsa ho thehwa ha balekane le tlhekefetso e khethiloeng maetong a banna ba rorisang [7, 8]. Ntle le moo, NAcc DA e bapala karolo e tsebahalang boits'oarong ba bo-mme ho mefuta e meng ea litoeba. Ho litoeba, mohlala, DA e lokolloa ho NAcc ho arabela sepheo sa bana [109] le liphetoho lipakeng tsa NAcc DA tšebetsong linako tsa kamora ho pepa li hokahanngoa le liphetoho mekhoeng e fapaneng ea boits'oaro ea motsoali ho tloha ho ho nyolla ngoana, mooki, ho nyeka / boitlhopho, le mohopolo oa bo-mme [3]. Ka hona ho ka ba molemo ho lipatlisiso tsa nako e tlang tsa boits'oaro ba motsoali oa monna ho hlahloba monyetla oa hore NAcc DA e bapala karolo ea bohlokoa.

7. Li-neurochemicals / lihormone tse ling li bohlokoa ho tlamahanong ea bobeli

Ntle le AVP, OT le DA, li-neurotransmitter le lihormone tse ling tse 'maloa li kentsoe tšebetsong ea boits'oaro bo amanang le ho tlamahana ha li-voes tsa prairie. Mohlala o mong o khahlisang o kenyelletsa li-neurochemicals tse amanang le hypothalamic-pituitary-adrenal (HPA) axis, sistimi e sebetsanang le likarabo tsa khatello ea maikutlo. Ka bokhutšoanyane, nakong ea khatello ea maikutlo, corticotrophin-releasing factor (CRF) e lokollotsoeng ho hypothalamus e tlama ho li-receptor tsa CRF ka pituitary ea anterior e lebisang ho synthesis ea adrenocorticotrophic hormone (ACTH) [143]. ACTH joale e ntšoa ka har'a mali ebe e sebetsa ho adrenal cortex ho hlahisa li-glucocorticoids, joalo ka corticosterone (CORT), e ka bang ea sebetsa ho li-glucocorticoid receptors (GR) bokong ho arabella likarabo tsa khatello ea maikutlo [143]. Li-voiri tsa Prairie li nkuoa e le litoeba tse loantšanang le glucocorticoid kaha li na le 5- ho ea 10-fold e kholo ea basal plasma CORT le 3-fold fold basal e kholo ea ACTH, hammoho le li-GRN tsa 10-fold tse tlase tsa kamano, haholo-holo mofuta oa I GR, hore li hlahisoa ka letsoalo le tlaase bokong, ha li bapisoa le likhoto le maqhubu a boitšoaro bo bobe [110, 204].

Tlhahiso-leseling e tsoang litekong tsa boits'oaro e bonts'a hore phello ea CORT ho tlamahanong ea balekane ke dimorphic ea thobalano. Liketelong tsa basali tsa basali, ho tloaelana ho kopana le e tona, e lebisitseng ho khethoeng ha balekane, ho fokotsehile haholo litekanyetso tsa serum CORT [63]. Ho feta moo, phokotso mosebetsing oa GR, ekaba ka ho fokotsa ho potoloha CORT ka adrenalectomy [63] kapa ka ho phekola liphoofolo ka mohanyetsi oa GR [52], e tataisa sebopeho sa khetho ea molekane. Ho fapana le seo, liente tsa CORT kapa teko ea ho sesa e hatellang, e ileng ea eketseha ho potoloha CORT [66], e thibetse nts'etsopele ea sebopeho sa khetho ea balekane [63]. Ka bobeli, tlhaiso-leseling ena e fana ka maikutlo a hore ho fokotseha hoa ts'ebetso ea li-axis tsa HPA ho thusa ho theha khetho ea molekane maemong a basali ba rorisang basali. Ho ba batona, ka lehlakoreng le leng, adrenalectomy inhibhibition preference ya molekane mme sephetho sena se ile sa khutlisoa ka phetoho ea CORT [64], ho bonts'a hore CORT ea hlokahala bakeng sa kaho ea khetho ea balekane ho banna. Ho ekelletsa moo, phuputsong e sa tsoa feta, tahlehelo ea molekane ea tlamahantsoeng e eketsehile haholo litekanyetso tsa CORT le boima ba adrenal gland ka li-voes tsa banna ba rorisang, a fana ka tlhahiso ea hore ts'ebetso ea axel ea HPA e ka akaretsa litlamorao tse tlatselletsang tsa karohano ea molekane ka hona, e bapala karolo ho bolokeheng le ho tokiso ea li-bond tse teng [29]. Mokokotlo oa HPA o bile le tšusumetso ho boits'oaro ba ntate. Ba batona ba pepesetsoeng khatello ea ho sesa ba qeta nako e ngata ba thellelana malulong le tloaelo ea ho qeta nako e ngata le ho koetlisa malinyane ho feta taolo e sa hatelloang [14]. Litlamorao tsena tsa boitšoaro ha lia ka tsa fumanoa libakeng tsa basali tse rorisang basali, ho bonts'a hore litlamorao tsa khatello ea maikutlo boits'oarong ba motsoali-joalo ka sebopeho sa khetho ea molekane-e kanna ea ba serorphic [14].

CRF e boetse ea kenella mekhoeng ea bolekane bo kopanetsoeng. Li-voes tsa prairie tsa banna tse fumaneng liente tsa CRF li bonts'itse khetho ea molekane ha ho se letho le emeng, mme boits'oaro bona bo kentsoeng bo thibetsoe ke tsamaiso ea kopanetsano ea mohanyetsi oa CRF receptor antagonist [67]. Libaka tsa bori tse kenyelletsang khokahanong ea CRF ea likhetho tsa molekane le tsona li fumanoe. Liente tsa CRF tsa lehae ka hara NAcc li thusitsoe, athe CRF receptor antagonists inhibhibitileng, sebopeho sa khethollo ea molekane ho li-voes tsa banna tsa prairie [148]. Ho feta moo, ho thatafalloa le mosali ho felletse ka keketseho ea CRF mRNA ho BNST ea li-voes tsa banna [29]. Kamora nako, tsamaiso ea li-icv ea urocortin-II, setho sa lelapa la CRF peptide, e ile ea eketsa boitšoaro ba motsoali ho basali le basali, empa kalafo ena e ne e se na litla-morao tsa ho tšoenyeha kapa boits'oaro ba locomotor [190].

Li-neurochemicals tse ling tse 'maloa le tsona li kenella tlamong ea sechaba ka libaka tsa prairie. Mohlala, ho li-voes tsa praemia tsa banna, tsamaiso ea intra-VTA ea NBQX, mohanyetsi oa AMPA receptor, kapa bicuculline, mohanyetsi oa GABA receptor, a susumetsa khetho ea molekane, o kenya li-amino acid tsena tumellanong e ikhethileng [53]. Tsamaiso ea serotonin reuptake inhibitor e khethiloeng, fluoxetine, e ekelitse morao-rao ho etsa boits'oaro ba botsoali maemong a mabeli a banna le basali, e fokotse mabifi ho banna, mme ha e na litlamorao ho boits'oaro bo bobe ba [209], e bonts'a hore serotonin e kanna ea kopanya boitšoaro bo amanang le botsoalle bo kopaneng. Li-steroid tsa Gonadal le tsona li ka eketsoa lenaneng lena. Phetoho ea testosterone kapa estrogen bekeng ea pele kapa ea bobeli ea bophelo e fetole haholo litloaelo tse amanang tsa morao-rao le / kapa boits'oaro bo botle ho li-volesile prairie voles [138, 184]. Estrogen receptor alpha (ERα) e kanna ea akaretsa tse ling tsa litlamorao tsa li-steroid tsa gonadal ho boits'oaro bo kopanyang maqhubu a prairie. Boithuto bo bontšitse hore basali ba na le lisele tse ngata tsa Ercy-ir libakeng tse ngata tsa boko ho kenyeletsoa MeA, BNST, MPOA, le VMH ho feta banna, le ho fokotseha ha maemo a ERcy-ir ho BNST, MPOA, le VMH ea basali e ne e amana le ho kenngoa ea thobalano [113]. Ho banna, polelo e ntlafalitsoeng ea ERcy ho MeA, ka phetisetso ea vevishene ea vaerase e kopaneng, e sitisitse pontšo ea boits'oaro bo kopanetsoeng le tšebeliso ea khetho ea molekane ea holofetseng [58]. Ka mokhoa o ts'oanang, banna ba nang le polelo e ntlafalitsoeng ea ERSI ho BNST e bontšitsoeng e fokotsehile kamano ea sechaba [140]. Lintlha tsena li bonts'a kamano e fapaneng lipakeng tsa polelo ea tikoloho ea ERcy le boits'oaro ba sechaba maemong a rorisang. Ha ho kopanngoa hammoho, lithuto tse hlalositsoeng kaholimo li bonts'a ho kenya letsoho ha li-neurochemicals tse ngata le li-hormone taolong ea boits'oaro bo kopanyang maqhubu a mabeli a banna le a basali.

8. Litloaelano tsa Neurochemical / hormone

Joalokaha ho hlokometsoe ka holimo, lits'ebetso tse fapaneng tsa methapo ea kutlo, li-neurotransmitter le li-hormone li kentsoe tšebetsong ea maqhama a mabeli. Leha ho le joalo, ha ho na monyetla oa hore litsamaiso tsena li iketsetse ka botsona ho laola boitšoaro bona bo rarahaneng ba sechaba. Karolong e latelang, re tla hlahloba lithuto tse sebetsanang le litšebelisano tse tsejoang lipakeng tsa tse ling tsa lits'ebetso tsena, ho kenyelletsa CRF, OT, AVP, DA, glutamate (GLU), gamma-aminobutyric acid (GABA), le lihormone tsa gonadal steroid, molatong oa li-pair. boits'oaro ba tlamahano, haholo-holo sebopeho sa khetho ea molekane.

Li-neurochemicals tse peli tsa pele tse khothalelitsoeng ho hokahana le motho e mong ka taolo ea moralo oa khetho ea balekane libakeng tsa prairie e ne e le CORT le OT. Hopola hore ho basali ba basali ba nang le thobalano, ho pepesehela monna ea sa tloaeloeng ho ile ha eketsa haholo tokollo ea OT [187] le litekanyetso tsa serum CORT tse fokotsehileng [63], litlamorao tse nahanoang ho tsamaisa tlhophiso ea molekane. Ho khahlisang, li-injection tsa icv tsa OT li hlahisitse ho fokotseha ho lekanang le maemo a serum CORT, a fana ka tlhahiso ea hore OT e ka sebelisana le axis ea HPA ho laola sebopeho sa likhetho tsa molekane [65] - mohopolo o ka lumellang lipatlisiso tsa nako e tlang ha ho fanoa ka tšebelisano pakeng tsa OT le CORT ka boits'oaro bo bong ba sechaba [6, 39, 42, 132].

OT e boetse e bontšitsoe hore e sebelisana le AVP ho ea ka tatellano ea khetho ea balekane, sephetho se sa makatseng ha ho thoe li-neuropeptide tsena li amana haufi ebile ha li arolelane feela likarolo tse tšoanang tsa lik'hemik'hale tse fapaneng ka li-amino acid tse peli feela. hape o sebelisane le li-receptors tse ling le tse ling [19]. Joalokaha ho hlalositsoe pejana, ente ea icv ea AVP kapa OT e ka susumetsa khetho ea balekane lithutong tsa banna le tsa basali kamora nako e ka bang hora ea 1 ea ho tloaelana le phoofolo e fapaneng ea bong bo fapaneng. Ho khahlisang, litlamorao tsa AVP mabapi le sebopeho sa khetho ea molekane li felisoa boteng ba mohanyetsi oa OTR, mme litlamorao tsa OT li felisoa boteng ba mohanyetsi oa V1aR, ho bonts'a hore AVP le OT ba ka sebelisana ho ikhetholla ho ratoa ke molekane [43]. Ho feta moo, liphetho tsena li bonts'a hore ho tsamaisoa ha moralo oa khetho ea balekane ho ka hloka ts'ebetso e ts'oanang ea li-V1aR le li-OTR [43]. Ho qhekella ka mokhoa o ikhethileng ka LS ea li-voes tsa banna tsa praemia ho tloha ka nako eo ho tšehetsa khopolo ena. Sebopeho sa litheko tsa Partner se susumetsoang ke AVP microinjection ho LS se ile sa koaloa ke tsamaiso e le 'ngoe ea mohanyetsi oa li-OT receptor [149]. Ha li kopantsoe hammoho, liphuputso tsena li fana ka maikutlo a hore li-system tsa OT le AVP tse bohareng li ka sebetsa ka bonngoe ho tsamaisana le sebopeho sa khetho ea molekane.

OT le AVP e boetse e bontšitsoe hore e sebelisana le lits'ebetso tse ling tsa neurotransmitter, tse kang DA, ho buella likhetho tsa molekane. Lipuong tsa basali tsa basali, mohlala, likhetho tsa molekane tse hlahisitsoeng ke NAcc D2R ts'ebetso li thibetsoe ke tsamaiso e tšoanang ea mohanyetsi oa OTR [150]. Ka lehlakoreng le leng, likhetho tsa molekane tse hlahisoang ke tsamaiso e bohareng ea OT li ne li thibetsoe ke tsamaiso e tšoanang ea mohanyetsi oa D2R ho NAcc [150]. Liphetho tsena li bontša hore ts'ebetso e tšoanang ea li-D2R le li-OTR ho NAcc lia hlokahala bakeng sa ho tsamaisa thato ea molekane maemong a basali ba rorisang basali. Litšebelisano tsa AVP-DA le tsona li bile le tšusumetso molemong oa khetho ea balekane. Phuputsong ea morao-rao, li-voales tsa tlhaho tsa basali tse sa sebetseng hantle-tse neng li ke ke tsa etsa khetho ea molekane le molekane-li amohetse mofuta oa veter-veter-mediated ka mofuta oa "prairie vole V1aR" ho tsoa VPNUMXaR tikolohong ena le sebopeho. ea likhetho tsa molekane ka mor'a lihora tsa 1 tsa ho ruruha [145]. Boitekong ba bobeli, likhetho tsena tse hlahisitsoeng ke vaerase tse hlahisitsoeng ke vaerase li ile tsa thibeloa ke tsamaiso ea mohanyetsi oa D2R pele ho nako ea ho nyalana, a fana ka tlhahiso ea hore AVP le DA li ka sebelisana ho etsa tšebelisano ea maqhama a mabeli [145]. Hypothesis ena e tšehetsoa ke khokahano e tsebahalang ea neuroanatomical lipakeng tsa libaka tsena tse peli, ha D2R e bua ka li-neuron tse bohareng ba projeke ea projeke ea NAcc ka kotloloho ho VP [90].

GLU, GABA, le CRF le bona ba khothalelitsoe hore ba sebelisane le DA ho tsamaisong ea likhetho tsa balekane [52, 53]. Blockade ea AMPA GLU kapa li-receptor tsa GABA, ka ente ea NBQX kapa bicuculline, ka ho latellana, likhethong tsa molekane tse etsoang ke VTA ha ho se na botsitso. Joalokaha VTA e fana ka mohloli o moholo oa lipuisano tsa DAergic ho libaka tsa boko ba mesolimbic, ho kenyelletsa le NAcc, ho fanoe ka maikutlo a hore litlamorao tsa bahanyetsi bana mabapi le sebopeho sa khetho ea molekane li kanna tsa ba pakeng tsa litlamorao tsa bona ho NAcc DAergic neurotransmission [53]. Phuputsong e arohaneng, tsamaiso ea peripheral ea RU-486, mohanyetsi oa GR, o ile a susumetsa likhetho tsa molekane ho li-voes tsa prairie tsa basali nakong ea ho ba sieo [52]. Litlamorao tsena li ne li koetsoe ke coadminication ea mohanyetsi oa D1R kapa D2R ho kena kamor'a morao-rao, a fana ka maikutlo a hore litlamorao tsa GR antagonism mabapi le sebopeho sa khetho ea molekane li ka kopanngoa ka tšebelisano le lits'ebetso tse bohareng tsa DA [52]. Teko e tsoelang pele e hlokahala ho hlakisa mofuta oa tšebelisano lipakeng tsa GLU, GABA, CRF, le DA ka ho theha khetho ea balekane.

Qetellong, li-steroid tsa gonadal li bapala karolo ea bohlokoa ho tlamahaneng ea bobeli mme ho nahanoa hore e sebelisana le lits'ebetso tse fapaneng tsa neuropeptide le neurochemical tse kenyellelitsoeng ketsong ena. Mohlala, ho pepesetsoa nako ea monna kapa lipontšo tsa chemosensory tse tona li eketsa maemo a potoloho ea estradiol mme kamora nako, boits'oaro bo botle, kapa boiphetetso ba thobalano ho li-voes tsa basali tsa basali [35, 36, 47, 203]. Thobalano ea thobalano le eona e ka kenngoa ho basali ba ovariectomised ka tsamaiso ea estrogen feela [35]. Ho khahlisang, litekanyetso tsa serum ea estrogen e eketsehang, e susumetsoang ke ho pepesetsoa hoa matšoao a chemocensory ea banna kapa tsamaiso ea estrogen, li ile tsa eketsa tlamo ea OTR haholo bokong ba prairie vole [233], e bonts'a hore estrogen le OT li ka sebelisana ho laola ho tlolelana — e leng e tataisang sebopeho sa khetho ea balekane ho basali [228]. Ho banna, testosterone e fumanoe e susumetsa litlamorao tsa AVP ho theho ea khetho ea molekane. Hopola, tsamaiso ea li-icv ea AVP e thusa ho theha lithahasello tsa balekane ho li-voes tsa praemia tsa banna tse latelang feela hora ea 1 ea ho phomola [43]. Ho khahlisang, tsamaiso ea AVP ha e khothalletse khetho ea balekane ho li-voils tsa banna ba baholo ba ileng ba beoa ka letsatsi la bona la tsoalo, ba fana ka maikutlo a hore litlamorao tsa testosterone lia hlokahala bakeng sa litlamorao tsa AVP ho thehoeng ha balekane ba ratoang [57]. Testosterone le AVP li ka sebelisana le taolong ea boits'oaro ba bo-ntate, le hobane li-voes tsa banna ba rorisoang ha li se li le batho ba baholo li ne li fokolitse letsoele la AVP-ir ka LS 'me le bontšitse boitšoaro bo fokolang ba ntate ho feta taolo e nang le li-gonads tse hlakileng [219] (bapisa le152, 153]).

9. Kakaretso le lintlha tse ling tsa tlatsetso

Ho tlamahana kahare ho li-voes tsa prairie ke boitšoaro bo rarahaneng ba sechaba bo kenyelletsang khokahano ea boitšoaro bo fapaneng bo fapaneng, ho kenyelletsa kamano e ikhethileng, ntoa e khethiloeng le tlhokomelo ea botsoali. Polelo e loketseng ea boits'oaro bona e hloka mesebetsi e mengata e hlokolosi, ho kenyelletsa ts'ebetso ea maikutlo, ho hlophisoa ha mohopolo, le tlhokomeliso ea motho ka mong, hammoho le tlhahiso ea koloi. Likarolo tse fapaneng tsa boko, joalo ka MeA, BNST, LS, NAcc, PFC, le AH, tse tsebahalang ka ho tsamaisana le lits'ebetso tsena, li bile le tšusumetso ho boits'oaro bo kopanyang maqhubu a praeree, joalo ka ha ho hlalositsoe kaholimo. Ho feta moo, li-neurochemicals tse ngata tse kentsoeng tšebetsong ea maqhama a kopanyang batho ba babeli, ho kenyeletsoa AVP, OT, DA, CRF, le GLU li tsejoa ho nka karolo libakeng tsena tsa boko ho hokahanya lits'ebetso tsa bohlokoa bakeng sa ho momahana. Mohlala, AVP ho LS [78] le OT ho MeA [77] buisana ka ho amoheloa sechabeng ka mefuta e meng, ketso e ipapisitseng ho thehoeng le ho bonts'a khetho ea balekane ba libaka tsa thorisoe. Mohlala o mong, DA ho NAcc e bapala karolo ea bohlokoa thutong ea moputso [23, 119], ketso e ka 'nang ea kenya letsoho ho theheng khetho ea molekane ka ho hokahanya setsoalle se ithutileng pakeng tsa thepa e matlafatsang ea ho tlolelana le lets'oao le ikhethileng la molekane oa [8, 10, 245].

Le ha tlhahlobo e hlokolosi e senotse bohlokoa ba o mong le o mong oa likarolo tsena tse arohaneng tsa boko le li-neurochemicals bakeng sa boits'oaro bo kopanyang, ha ho pelaelo hore boitšoaro bo bong le bo bong bo lekiloeng bo laoloa ke potoloho e kholo e kenyeletsang libaka tse ngata tsa boko le li-neurochemicals. Ha e e-so hlahlojoe ka botlalo li-voles, datha e tsoang ho mefuta e meng ea litoeba e bontšitse khokahano e hlakileng ea anatomical lipakeng tsa likarolo tsa boko tse boletsoeng ka holimo, 'me lihokela tsena li theha li-circuits tsa neural tse nang le mesebetsi ea discrete e amanang le ho tlama maqhama [245]. Mohlala, khokahano lipakeng tsa setho sa ho hlapa (VNO), AOB, le MeA li bapala karolo ea bohlokoa ho ts'ebetsong likhoele tsa chemosensory [30, 72, 121, 130, 192], mme ha ho makatse hore ts'ebetso ea bobeli ba VNO le MeA nakong ea boithuto ba sechaba le thobalano (ha litekanyetso tse hlakileng tsa bonono li le teng) li bohlokoa molemong oa ho theha khetho ea molekane [49, 133]. Khokahano lipakeng tsa MeA le AH li laola mabifi [33, 202], le ts'ebetso ea tsela ena e kanna ea laola mabifi a ikhethileng libakeng tsa praemia [99, 224]. Ntle le moo, sistimi ea "mesolimbic DA" e hlophisitsoeng hantle e nang le lisele tsa DAergic tse VTA morero oo o eang ho NAcc le mPFC, o laoloa ke lipono tsa GLUergic tse tsoang ho mPFC ho ea ho Nacc le VTA [34, 193]. Potoloho ena e tsamaeang ka neural e fana ka maikutlo a ho thusa ho susumetsa tikoloho e loketseng tikoloho [120, 232], ho tataisa boits'oaro bo tataisang bo phehellang, bo kang ho tsamaisana le molekane ea amohelang [22, 74, 179, 180] le ho khutlisetsoa ha malinyane [2, 112, 171]. Ho bohlokoa ho hlokomela hore joalo ka ha sistimi e ts'oanang e ka kopanya boitšoaro bo fetang bo le bong, joalo ka ha ho bonahatsoa ke mohlala oa morao, neurochemical e tšoanang e ka laola mekhoa e fetang e le 'ngoe. Mohlala, ts'ebetso ea methapo ea kutlo ea AVP, ho LS, VP, le / kapa AH e bapala karolo ea bohlokoa ho thehoeng ha balekane, ho khetha mabifi le boits'oaro ba ntate. Ho feta moo, ketsahalo e le 'ngoe ea boits'oaro e ka tsamaisoa ke potoloho e fetang e le' ngoe ea neural. Mohlala, boiphihlelo ba batho ba ratanang le ba bong bo fapaneng - litabatabelo tsa kaho ea khetho ea balekane - li etsa hore ho lokolloe tšebetso ea DA le OT ho NAcc [98, 187], Ho lokolloa ha AVP ka bobeli ho LS [216] le VP [245], hape ho khothalelitsoe ho lokolla OT ho PFC [245]. Ka hona, li-circuits tse ngata tsa neural le li-neurochemicals li sebetsa 'moho ho khothatsana ho laola boitšoaro bo amanang le ho tlamahana.

Kamora nako, lipuisanong tsa tlamahano ea bonono phuleng ea sebaka se rorisang, liphapang tsa thobalano ha lia lokela ho tlotsoa. Joalokaha ho tšohliloe kaholimo, melaoana ea boitšoaro bo kopanyang ba babeli e atamelaneng ka tsela e itseng maemong a mang ka thobalano (mohlala, Molao oa CORT oa mofuta oa khetho ea molekane [63, 64]). Ho feta moo, li-neuropeptide tse ling, tse kang AVP le OT, li kanna tsa ba le karolo e ikhethileng ea bong ka boits'oaro bo itseng bo amanang le ho tlamahana ka maqhama: AVP e laola tlhekefetso e khethiloeng [99, 100] le boits'oaro ba ntate149, 213] ho banna, ha OT e tsamaisa tlhokomelo ea basali ho basali.16, 76, 165, 173, 178]. Ho feta moo, ba batona le ba batšehali ba bonahala ba fapane maikutlong a bona ho AVP le OT. Leha li-neuropeptide ka bobeli li bile le tšusumetso molemong oa khetho ea molekane ho ba bong bohle [43], litekanyetso tse tlase tsa AVP li ne li lekane ho susumetsa khetho ea balekane ho banna ho feta basali mme tsamaiso ea peripheral ea OT e ne e sebetsa hantle ho susumetsa khetho ea balekane feela ho basali, empa e se banna, prairie voles [54]. Litšoantšo tsa botona le botšehali ho physiology le li-subctates tsa neural li ka baka liphapang tsena tsa boitšoaro. Mohlala, joalo ka ha o fumanoa mefuteng e meng ea litoeba [48, 61, 207], li-voles tsa banna li na le lisele tse ngolisitsoeng ka ho fetisisa tsa AVP mRNA ho BNST le MeA le bongata bo phahameng ba li-fiber tsa AVP-ir ho LS (boholo ba merero e ka etsahalang ho tsoa ho AVP e hlahisang lisele ho BNST le MeA) ho feta li-voles tsa basali [17, 221]. Ho khahlisang ke hore matsatsi a 3 a ho ikopanya le motho oa bong bo fapaneng a etsa hore ho be le keketseho ea palo ea lisele tse ngolisitsoeng tsa AVP mRNA ho BNST le ho fokotseha ha letsoalo la AVP-ir le ho koala LS ho banna, empa eseng basali, li-voes tsa prairie, tse bontšang litla-morao tsa botona le botšehali tsa boiphihlelo ba batho ba bong bo fapaneng liketsong tsa AVP, tseo, li ka nkang karolo taolong ea boitšoaro bo kopanang ba kopaneng lihlopheng tsa banna ba batšo. Phapang ea thobalano e boetse e fumanoe ho palo ea lisele tsa TH tse hlophisang tikoloho ea anteroventral periventricular preoptic [139] le amygdala e atolositsoeng [168] bokong ba prairie vole, leha ho le joalo karolo e sebetsang ea lisele tsena ka bonngoeng e ntse e hloka ho hlahlojoa. Ho tsebahala hore litekanyetso tsa botona le botšehali ho physiology le neural substrates li ka tlasa karohano ea botona le botšehali ea boitšoaro. Ntle le moo, liphapang tsa botona le botšehali bokong li ka lumella pontšo ea boits'oaro bo ts'oanang lipakeng tsa ba batona le ba basali ntle le 'mele ea bona e fapaneng [62]. Ka mantsoe a mang, lits'ebetso tsa methapo ea kutlo tsa thobalano li ka lumella banna le basali hore ba be le mekhoa e lumellanang e sebetsang le 'mele ea bona ho hlahisa litholoana tse tšoanang. Tlhahiso ena e lumellana le lithuto tse ithutoang khafetsa tse bonts'ang lits'ebetso tsa thobalano, joalo ka tsela ea AVP ho tloha MeA le BNST ho ea LS [17], nolofalletsa pontšo ea boits'oaro ba botsoali ho ba batona [216] ha lits'ebetso tsa OT li etsa boits'oaro bo ts'oanang ho basali [16, 173].

10. Liphello le litaelo tsa nakong e tlang

Leha ho ithuta ka maqhama a thehiloeng lipakeng tsa lipale tsa prairie ho ke ke ha re lumella ho utloisisa ka botlalo ho rarahana ha likamano tsa batho, ka nepo ba ka fana ka leseli ka mekhoa ea mantlha e ka hohelang ho hohela le ho hokahana le batho ba baholo. Lingoliloeng tse hlahlojoang ka holimo li amme mefuta e mengata ea li-neuropeptide, li-neurotransmitter, le lihormone maemong a taolo ea tlamahano ea maqhama a kopaneng a li-voes tsa prairie. Ka hona, ts'ebetso ea pele ho batho e ammetse lits'ebetso tsena tse tšoanang litloaelong tsa batho tsa bophelo. Mohlala, lipatlisiso tsa morao-rao tse sebelisang monahano o sebetsang oa matla a tlhaho ho lekanya ts'ebetso ea ts'ebetso ea boko ho batho li bontšitse hore ts'ebetso ea methapo ea DA e ka etsa hore motho a ikhethe le ho ikamahanya le eena [81]. Lithuto tsena li fumane hore libaka tse ruileng haholo tsa DA, joalo ka VTA, li ile tsa qaptjoa ha barupeluoa ba neng ba le mekhahlelong ea likamano tsa lerato le tebileng kapa kamanong e telele e lerato ba shebile setšoantšo sa baratuoa ba bona empa eseng ha ba bona litšoantšo tsa batho ba bang ba tloaetseng. batho ka bomong [12, 20, 21]. Boithuto ba morao-rao bo boetse bo hatisitse tsamaiso ea OT litšebelisanong tsa batho ba babeli. Litekong tse laoloang ke placebo, ho etsa mohlala, tsamaiso ea methapo ea OT - mokhoa oa ho fana ka tumello o lumellang neuropeptide ho fihlela bokong — o matlafalitse puisano e ntle pakeng tsa banyalani, joalo ka ha e bonts'oa ka leihlo, bohelehele / tlhokomelo, le lintlha tsa tumellano [70]. Ho kenyelletsa moo, OT e fumanoe e eketsa ts'epo ho batho - ntho ea pele e amanang le botsoalle [137]. Ho feta moo, AVP e kentsoe ts'ebetsong ea batho ba mabifi, joalo ka ha maemo a AVP ka har'a mokelikeli oa banna le basali a ne a amana hantle le nalane ea bophelo ba boitšoaro bo mabifi [46]. Ha li kopantsoe hammoho, lithuto tsena li bonts'a monyetla oa hore mekhoa e ts'oanang ea tlhaho e ka kopanyang boitšoaro ba batho le liphoofolo tse anyesang tseo e seng batho.

Tekanyo e phahameng haholo ea polokeho lipakeng tsa boits'oaro bo botle ba 'mele le boits'oaro bo botle ba batho ba sechaba e fana ka maikutlo a hore mofuta oa tlholo ea prairie e ka ba hantle bakeng sa lipatlisiso tsa mantlha le tsa phetolelo tse etsang lipatlisiso tsa methapo ea boitšoaro ea sechaba. Ka hona, lipatlisiso tsa li-voes tsa prairie li kanna tsa se re lumelle feela ho ithuta haholoanyane ka mabaka a etsang boitšoaro bo tloaelehileng sechabeng, empa hape li ka re nolofalletsa ho hlahloba lisosa tsa bofokoli ba boiketlo ba sechaba bo boneng likhatisong tse 'maloa tsa bophelo ba kelello. Mohlala o hlokomelehang o kenyelletsa tšebeliso ea mofuta oa mofuta oa viri bakeng sa ho ithuta ka litšoaetso tsa lefu la autism [147, 157], eo AVP, OT, le DA li seng li entsoe tšusumetso [111, 115, 156, 236]. Ntle le moo, viri ea prairie e sa tsoa theoa e le mohlala oa phoofolo bakeng sa khatello ea maikutlo, haholo khatello ea maikutlo e bakiloeng ke tahlehelo ea sechaba bathong ba baholo [29, 103]. Kamora nako, li-voes de prairie li sa tsoa sebelisoa haholo ho ithuta litlamorao tsa lithethefatsi tsa tlhekefetso ho bonngoe, 'me liphuputso tsena li bonts'itse hore ts'ebeliso ea ts'ebetso ea mesolimbic DA e kanna ea kenya letsoho phosong e bakiloeng ke lithethefatsi ea boitšoaro ba sechaba [151, 238]. Lithuto tsena le tse ling [84] bonts'a ts'ebeliso ea sebopeho sena sa phoofolo bakeng sa lipatlisiso tsa mekhoa ea neural e ka tlasa mekhoa e tloaelehileng le e sa tloaelehang ea sechaba, le lits'ebetso tsa tsona tse amanang.

lumela hore baa fokola

Re ka rata ho leboha C. Lieberwirth, K. Lei, MM Martin le AS Smith ka ho bala ha bona ho matla ha buka e ngotsoeng ka letsoho. Re boetse re leboha AS Smith ka lipuisano tsa hae tse molemo nakong ea ho ngola mongolo ona le ka menehelo ea hae ea lifoto. Re leboha ka kananelo C. Badland le J. Chalsters ka thuso ea bona lipalo. Lichelete bakeng sa mosebetsi ona li fanoe ke National Institutes of Health Grants DAF31-25570 ho KAY, MHF31-79600 ho KLG, le MHR01-58616, MHR21-83128, DAR01-19627, le DAK02-23048.

Mongolo o botlaaseng ba leqephe

1Le ha maqhubu a tumello a simolohile Illinois a bontša boits'oaro ba leano le letona la bophelo tšimong, 'me ka mokhoa o ts'epahalang a bonts'a boits'oaro bo kopanyang ba botsofaling maemong a laboratori, ho bohlokoa ho hlokomela hore mangolo a tumello a tsoang Kansas [55, 186] le Tennessee [175, 235] bonts'a phapang e poteletseng likarolong tse ling tsa boitšoaro ba bona le physiology [185] Liphapang tsena li tšehetsa khopolo ea hore maemo a fapaneng a tikoloho ea tikoloho a ka susumetsa boits'oaro ba liphoofolo le maano a ho hanana pakeng tsa batho ba mefuta e le 'ngoe [73]. Ka lebaka la phapang ena, li-voes tsa prairie tse tsoang Illinois li sebelisoa haholo lithutong tsa laboratori tsa neurobiology ea pairing. Lintlha tse tsoang lithutong tseo ke sepheo sa maikutlo a morao-rao.

Boitlhotlhollo ba Mohoeletsi: Ena ke faele ea PDF ea mongolo o ngotsoeng ka letsoho o sa amoheloang o amoheletsoeng bakeng sa ho hatisoa. Joaloka tšebeletso ho bareki ba rona re fana ka phetolelo ena ea pele ea mongolo o ngotsoeng ka letsoho. Mongolo o ngotsoeng ka letsoho o tla ba le ho kopitsa, ho kenya lihlopha le ho hlahloba bopaki bo hlahang pele o hatisoa ka mokhoa oa oona oa ho qetela. Ka kopo hlokomela hore nakong ea mekhoa ea tlhahiso ea lihlahisoa e ka fumanoa e ka amang litaba, le litsebiso tsohle tsa molao tse sebetsang koranteng e amanang le eona.

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