Iinkqubo ze-neurobiological zokunxulumana nentlalo kunye nokudibanisa isibini (2015)

Yiya e:

Abstract

intshayelelo

Ubudlelwane esibumba kunye nosapho, izihlobo, kunye namaqabane othandana ayinto ebalulekileyo kwimbutho kunye nokusebenza koluntu. Nangona zintsonkothile, ezi zibophelelo zenza iinkqubo zecandelo ezinokuphandwa kuzo zonke iintlobo. Elinye ithuba elihle lokuphanda ngesiseko se-neurobiological zezinto eziqhotyoshelweyo yindalo ezimeleyo yokuziphatha kwesibopho esihlanganayo kuyo yonke i-taxa. Ngelixa uhenyuzo lwesini sisicwangciso esiliqili sokukhulisa izilwanyana (kuboniswe i-95-97% yezilwanyana ezincancisayo), iindlela zobuchule bokutshatisa ezithandanayo ziye zenzeka ngeendlela ezahlukeneyo, kubandakanya izilwanyana ezingenamqolo, iintlanzi, izilwanyana ezirhubuluzayo, iintaka kunye izilwanyana ezincancisayo. Ezi nkqubo zibonakaliswa ngokunyamezelana, rhoqo ubomi, ukukhetha ukunamathela phakathi kwamaqabane okutshatisa, nangona kungasoloko kungabekelwa ecaleni ngokwesini. Isiseko se-neurobiology yala mabhondi sisentloko sophononongo.

Ukuphanda ngebhondi kunye

Amathuba anamandla okuphanda ibhayoloji yendlela yokuziphatha ikhula: okokuqala, iintlobo zephylogenetically ezikude zibonakalisa indlela yokuziphatha eguqukayo, okwesibini, iintlobo ezinxulumene ngokusondeleyo zibonisa isimilo sokuphambuka ngendlela ephambeneyo, okwesithathu, iintlobo ezizodwa zibonisa amanqanaba aphezulu okuhlukahluka kwendlela yokuziphatha. Ngaphakathi kwezi meko, iindlela zothelekiso zinokukhawulezisa ukuphawulwa kwezinto ezichasene ne-neurobiological, genetic, development, development, kunye nezindululo eziphantsi kokuziphatha okubuzwayo [1]. Uninzi lokuqonda kwethu i-neurobiology yokubhangqa kokubini kuvela kwiindlela zokuthelekisa kuzo zontathu iimeko, ngakumbi ngophando lwendawo yokuncoma, Microtus ochrogaster. I-neurobiology yokubhangqa kokubini kwi-prairie voles iya kuba yinto ephambili, kodwa ingakhethi, kugxilwe kolu hlaziyo.

Uhlobo I-Microtus ziquka iindidi ezininzi zemibutho eyahlukeneyo yentlalo. Ii-Prairie voles zibonisa indlela yokuziphatha enoluntu ekuhlaleni kunye nokhathalelo lomzali kubantwana, ukukhetha ubundlongondlongo kubantu abangabaziyo besini esahlukileyo, kunye nokuziphatha okuxinzelelekileyo emva kokuphulukana neqabane [I-2,3]. Ii-Prairie voles zikwabonisa amanqanaba aphezulu okuguquguquka kwe-intraspecific kwezi ndlela zokuziphatha; Umzekelo, bobabini abesilisa nabasetyhini banokubonakalisa phenogypes 'wokubhadula' oziphethe kakubi.

Interrogation of couping bonding in the laberals yaqalwa ngokusebenzisa uthotho lovavanyo kusetyenziswa iqabane lokukhetha uvavanyo, apho isifundo sinokulisebenzisa ngokukhululekileyo ixesha laso kunye neqabane laso eliqhelekileyo lokwazana, inoveli eyamkelayo ngokwesini, okanye eyedwa kwindawo [4]. Olu vavanyo lubonise ukuba emva kweeyure nje ze-24 zokuhlalisana kunye neqabane, ii-voes zodonga-ngokungafaniyo nokuziphatha okubi okanye ii-voes ze-meching - zidibene ngokukhethekileyo neqabane labo. Ezi 'zinto zikhetha iqabane' ziye zaba yimetri yelebhu yokudala ukubumbana, kunye nokuxinana kwemithambo-luvo ngaphakathi kwesi paradigm zichonge imbonakalo eyodwa yeeseli zobuchopho be-prairie vole ubuchwephesha bokukhetha ukunamathela koluntu.

Ii-neuropeptides zentlalo ekubhanxeni kwezibini

I-neuropeptides oxytocin (OT) kunye ne-vasopressin (AVP) zilondoloze ngokufanelekileyo iindima kulawulo lokuziphatha kwabantu abathandana nabo kwi-invertebrate kunye ne-vertebrate taxa, kubandakanya nabantu [5]. Kwizilwanyana ezincancisayo, umbutho we-neuroanatomical ovelisa i-OT kunye ne-AVP evelisa ii-neurons kunye ne-axonal yazo eziqikelelweyo kuyo yonke ingqondo igcinwe ubukhulu becala, ngelixa ukuhanjiswa kwee-receptor zabo ekujoliswe kuzo - i-oxytocin receptor (OTR) kunye ne-arginine-vasopressin receptor 1a (AVPR1a) - ziyahluka ngaphakathi nangaphaya kwiintlobo [6,7]. Ubukho beplastiki kwezi nkqubo kucingelwa ukuba bube negalelo kubuntu obahlukeneyo obujongwe kwindalo [7-9].

Ukuhanjiswa kwe-OTR kunye ne-AVPR1a kwingqondo ye-prairie vole yahluke kakhulu kwizilwanyana ezinxulumene ngokusondeleyo, kunye nokwahlukana okujolise kwimimandla ethile yomvuzo we-mesolimbic kubandakanya i-preortal cortex (PFC), i-nucleus accumbens (NAcc), i-ventral pallidum (VP), kunye ne-seralum yecala emva kwexesha ( I-LS) [10]. Kwii-prairie voles, i-blockade ye-OTR okanye i-AVPR1a kwezi ndawo ngexesha lokuhlala-ngakumbi i-OTR blockade kwi-NAcc okanye i-PFC okanye i-AVPR1a blockade kwi-LS okanye i-VP - ithintela ukubophela isibini [10].

Ukuthelekiswa kohlalutyo lwesine I-Microtine Iindidi zifumanise ukuba i-monogamous prairie kunye ne-pine voles zibonisa i-AVPR1a ephezulu kwi-expression ye-VP ngokuthelekiswa ne -isciscuous montane kunye ne-meadow voles, iphakamisa ukuba ukwandiswa kwentetho ye-AVPR1a kwi-VP kunokuba yindlela yokwenza igalelo ekudalekeni kokubumbana kokubini IiMicrotines [10]. Ukuxhasa le hypothesis, overexpression ye-AVPR1a kwi-VP ye-voisc voous meadow voles induces pair bond, kunye ne-RNA yokunqunyulwa kwe-AVPR1a kwi-VP ye-prairie voles inhibits couping bonding [I-11]. Nangona kunjalo, zombini iimpuku ze-monogamous and non-monogamous deice zine-AVPR1a ephezulu yokubonisa kwi-VP [12], ecebisa ukuba iindlela ezininzi zinegalelo ekudalekeni kobuchule bokutshatisa obugwenxa kumashini; kodwa kunokwenzeka ukuba ukubonakaliswa okuphezulu kwe-AVPR1a kwi-VP yinto eyimfuneko yokuvela kwe-monogamy kufuna olunye uphando oluthelekisayo.

Njengee-prairie voles, ii-marogamous marmosets ziye zaphakamisa uxinano lwe-OTR kwi-NAcc, ngelixa iimonogamous zethusi ze-brigit titi ziye zaphakamisa i-AVPR1a kule ndawo [13,14 ••]. Uhlalutyo lwezicubu zomntu zisebenzisa i-ligands ezibonisa ubuqhetseba kuzo zombini i-OTR kunye ne-AVPR1a [15], kodwa kuxelwe uxinzelelo oluphezulu kwi-VP [16]. Kuya kuba kubalulekile ukuphinda nihlalutye izicubu zomntu ngokusebenzisa ubuchwepheshe obuthathaka ngakumbi kunye nokukhetha; Nangona kunjalo, kulwazi lwethu lonke, zonke iintlobo zezilwanyana ezithandekayo ezihlolwe kwangoku zibonise i-OTR ephezulu kunye / okanye i-AVPR1a intetho kwisekethe ye-NAcc-VP; Ezi datha zikhuthaza uphando olongezelelekileyo kummiselo we-neuropeptidergic wale sekethe kwindalo yokuzibophelela koluntu.

I-OT kunye ne-AVP ilungelelanise indlela yokuziphatha ngokudibeneyo kumacala ahlukeneyo [17]. I-blockade ye-OTR kunye ne-AVPR1a i-Homologs kwi-monogamous cichlid fish is kunciphisa ukuziphatha okudibeneyo ngexesha lokwakhiwa kwebhondi [18], kunye nonikezo lwangaphandle lwe-OT lukhuthaza indlela yokuziphatha ehambelana nolwalamano kunye namaqabane oluntu ezinjeni [19]. I-OTR ye-homolog iguqula indlela yokuziphatha yokuhlangana kwi-zeograous zebra Finches [20,I-21,22]. I-OT kunye ne-AVP ilawula indlela yokuziphatha yokuhlangana kwi-marmosets kunye ne-browny titi nyani, ngokulandelelana [23-25]. Abantu babonisa uluhlu lwendlela yokuziphatha yomtshato wesibini, kwaye I-OXTR kwaye I-AVPR1A ukuhlukahluka kohlobo kudityaniswa nemeko yobudlelwane [26,27]; Amanqanaba e-plasma OT axela kwangaphambili amazinga okuphumelela kubudlelwane bezothando [28]; kwaye kumadoda aqhotyoshelwe zothando, i-OT ye-Intranasal yandisa umsebenzi we-NAcc ngelixa ijonga ubuso beqabane kwaye yonyusa umgama ophakathi wokukhetha phakathi kwabasetyhini abangaqhelekanga [29 ••,30].

Inkqubo ye-dopamine ye-Mesolimbic ekwakhiweni kweqhina lesibini

Zonke iinkqubo ze-nerve zijongana nomceli mngeni wokucoca, ukuguqula, kunye nokuhlaziya ulwazi oluqhubekayo lweemvakalelo zemultimodal kwimibutho efundileyo ekhokelela ekuguqukeni kokuziphatha [8]. MnaI-vertebrates, inkqubo yembuyekezo ye-mesolimbic-equka kwinxalenye yoqhagamshelo phakathi kwendawo yokuhamba yangaphakathi (VTA), PFC, NAcc, VP, kunye ne-LS - yinkqubo ye-neural elondolozwe ngokwendalo eyenza ukuba le nkqubo ivelise inkuthazo yokufuna umvuzo kunye nokuthintela ukuphambuka, nangokwabela iindawo ezihambelana nezi ziphumo. Isenzo se-dopamine (DA) ngaphakathi kule nkqubo sibalulekile kule misebenzi [31].

Inkqubo ye-mesolimbic ye-DA ibalulekile ekuhlanganiseni izibini. Ukwenziwa kokukhethwa kwabalingane kuququzelelwa ngokuvuthwa, okunyusa ukukhutshwa kwe-DA kunye nokungena kwi-NAcc[10]. DIntshukumo kwenye yeenjongo zayo ekujoliswe kuzo, i-D2R, kwi-NAcc iyimfuneko ekwakheni ibhondi [10]. Emva kokudibana, i-receptor yesibini ye-DA, i-D1R, inyuswe kwi-NAcc kwaye ibalulekile kulondolozo lwebhondi ngokulamla ubundlongondlongo kubantu abangabaziyo besini esahlukileyo [32].

Ukunyusa i-D1R: D2R umqondiso wokubonisa kwi-ventral striatum nayo ibe nefuthe kwizidakamizwa kunye nokuxhaphazwa [33]. Kwiigundane, ukuphathwa okuphindaphindiweyo kwe-cocaine kwandisa umlinganiso we-D1R: D2R expression expression kwi-NAcc, kwaye oku kulungelelaniswa kwakhona kulungelelanisa inkqubo yokuziphatha elandelayo ye-plastiki. Njengoko kukhankanyiwe, kwii-voes praes, ii-couping coupon zonyusa umda we-D1R: D2R expression expression kwi-NAcc, kwaye oku kulungelelaniswa kwakhona kulungelelanisa phakathi kwenkqubo yokuziphatha ye-plastiki [32]. Ezi datha zinegalelo kwingcamango yokuba izibini zontengiso zimele ukubakho kweziyobisi phakathi kwezithandani, eziya kuthi zimanyaniswe yinxalenye yombutho kunye neplastic yeendlela ze-mesolimbic ze-DA. [33]. Izifundo kwi-zebra finch kunye ne-browny titi monkey ikwaze nengxelo yokudityaniswa kwakhona kwiindawo ezinomvuzo emva kokudalwa kweebhondi, iphakamise ukuba ukubambelela kwe-neural kwezi ndlela zinokuba yinto egcinwe ngokuzimela yokudibanisa izibini [34,35].

Iinkqubo zecandelo kunye nezinye iinkqubo ekwakheni ubumbano

Iinkqubo ezininzi zecandelo zinegalelo kwisibambiso. Ukusebenzisana kwasekuqaleni kuxhomekeke ekunyamezeleni ukusondelana, indlela yentlalo, kunye nokuthintelwa kwezenzo zokuthintela / zokwala. Njengoko amaqabane esebenzisana, indlela yokuziphatha ehambelana noluntu, ukwamkelwa kwentlalo, kunye neenkqubo zomvuzo zinegalelo ekwakheni ibhondi. Emva koqhawulo-mtshato, ukuxabana ngokwasentlalweni, ukukhusela umntu otshatileyo, iimpembelelo ezimbi ngexesha lokwahlukana, kunye nememori yentlalo esebenzayo egcina ugcino lwebhondi.

I-Microtine iintlobo ziyahluka kunyamezelo lwasentlalweni kunye nobudlelwane, mhlawumbi ngokuyinxenye ngenxa yantlukwano kwintlangano ye-mesolimbic D1R. Kwii-prairie voles, ezinamanqanaba asisiseko asezantsi e-NAcc D1R, inyanzeliso engaphezulu ye-NAcc D1R iguqula utshintsho olubi kwindlela yokuziphatha kubantu abangabaziyo ngokwesini (ukusuka kunxulumana ukuya ekuthinteleni / ekhatyweni), nasekunyuleni i-NAcc D1R ku sebenza Ngexesha lokuhlala unhibits bonding [10]. Kwiindawo zokudibanisa, ezinesiseko esiphezulu se-NAcc D1R uxinano kwaye ayihambelani, ikhethe i-NAcc D1R ibhloko yonyusa indlela yobudlelwane [10]. Kwimigqibelo ye-zebra, ukusebenza kwe-mesolimbic DA iinkqubo kulungelelana nendlela, ukuthintela, kunye nokuziphatha okumanyeneyo ngexesha lokunxibelelana kwamadoda nabasetyhini [36], ecebisa ukuba i-mesolimbic ye-DA yeendlela inokuguqula ukunyamezelana kwentlalo kunye nobudlelwane kuzo zonke iintlobo.

Ukubhanqa i-Pair nako kuxhomekeke ekubonakalweni kwentlalo, inkqubo ye-neural ethi, kwiigundane, isekwe ngokuzinzileyo kwaye ixhomekeke kwi-OT kunye ne-AVP isayine. Ii-OTR zisasazwa ngokusetyenziswa kwe-nuclei yokulungisa i-olixion kwiintonga, ngelixa i-primates - apho ukuphicothwa kunye nombono ziimpawu eziphambili zeemvakalelo -e-OTR zibonakaliswa kwimimandla ebaluleke kakhulu ekuqwalaselweni kokubonwa kunye nokuphicothwa, kuphakanyiswa ukuba ukutyhilwa kwe-OT kunokuguqula imiba yokulungisa uvakalelo iintlobo, nangona iindlela ezithile zingahluka [13,14 ••,37]. Iyahambelana nale ntuthuzelo, i polymorphisms zabantu ngaphakathi I-OXTR Ukuqikelela ukukwazi ukubona ubuso, ucebisa ukuba indima ye-OT yokwamkelwa kwentlalo inokulondolozwa phakathi kweentonga nabantu, kuyo yonke imo yeemvakalelo [38 ••].

Iinkqubo ezongezelelweyo ze-neural zivumelana ngezinto ezongezelelekileyo zokusekwa kwebhondi kunye nokugcinwa. Ukubonakaliswa kwe-Opioid kubalulekile ngexesha lokufunda komvuzo kwaye kulawule ukwakhiwa kunye nokugcinwa kweebhondi kwiindawo zokuncoma, mhlawumbi ngokulawula ii-hedonics ezilungileyo ngexesha lokuqulunqwa kunye ne-hedonics emibi ngexesha le -enenanc.e [I-39,40]. Inkqubo yokukhululwa kwe-corticotrophin (CRF) ikhokelela ekwakheni ubumbano kunye nokuziphatha okuxinzelelweyo kulandela ukuhlukana / ilahleko yeqabane [3,41]. Ukuzonwabisa kwezentlalo kuququzelela ukugcinwa kwebhondi; kwii-prairie voles, kulandela amava oxinzelelo, ukukhutshwa kwe-OT ebusweni beqabane kunciphisa amanqanaba oxinzelelo kunye nokuziphatha okunexhala [I-42]. Kwimo yendalo, ukugcinwa kwebhondi kuxhomekeke kwinkumbulo echanekileyo yomhlaba weqela. Izifundo zakutsha nje zibonisa ukuba i-OTR kunye ne-AVPR1a ebopha uxinano kwindawo ezihambahamba / indawo yememori iqikelela ukusetyenziswa kwendawo, iqhinga lokutshatisa ('umhlali' okanye 'umhambi'), kunye nempumelelo yokuzala kwi-voes adresi, ngokunokwenzeka ngokuququzelela ukudityaniswa kolwazi lwentlalo (umz. Iindawo yeqabane, abakhuphisana ngokuzala, kunye / okanye amathuba okuzala) kwiimephu zakwindawo [43,44,I-45].

Ezinye iindlela zokuncamathela ekukhetheni indawo yokuhlala (umz. Iibhondi zomama) zifuna uninzi lwezinto ezincinci ze-neural substrates kunye neenkqubo zecandelo - ngokuchanekileyo i-OT, i-DA, i-CRF, i-opioids kunye nokwamkelwa kwentlalo-njenge-bond ekhethiweyo phakathi komtshato[17]. Ngapha koko, kuyacingelwa ukuba ukuqhawulwa kwesibini esitshatileyo kunokuvela ngokuqokelela iinkqubo zokuqhawula umtshato zakudala ngexesha lokunxibelelana kwabantu besini esinye ukwenza ubudlelwane obukhethayo neqabane lokutshatisa [46,47].

Amava kunye nokubhangqa kokubini

Kwiindawo zokuncoma, njengabantu, ubomi bomi bokuqala banganefuthe kwibhondi yabantu abadala [48], into enokuthi inqamlezwe lutshintsho lwe-neuroplastic kwiinkqubo ezibalulekileyo zokuxinana. Iyahambelana nolu luvo, ukukhuthaza ubomi bokuqala kwenkqubo ye-melanocortin receptor (MCR), enxibelelana ne-neuropeptide kunye neenkqubo zomvuzo, kuququzelela ukwakheka kweqhina elidala ebudaleni [49]. Mnan ngokungafaniyo, ukukhetha kwe-D1R kusebenze kwi-neonates kuphazamisa ukuqina kwabantu abadala [50]. Into ebangela umdla kukuba, ulawulo olungapheliyo lwe-OT ye-intranasal kumachiza athile ifunyenwe luthintela ukubophelelwa kwabantu abadala abangamadoda [51]; nangona kunjalo, ukuphucula ukubonakaliswa kwe-NAcc OTR kumabhinqa angaphambi kokushicilela axhasa ukuqina kwabantu abadala [52], kunye Ukuncitshiswa kobudala bukatata (ubuqili obunokuthi kunciphise ukubonakaliswa kwe-OT) kuphazamisa ukubopheleleka kumakhaya abantu abadala kunye nee-mandarin voles [48,53]. Ezi datha zikhuthaza uphando olongezelelekileyo kwindima yamava obomi bokuqala kunye nokutyikitywa kwe-OT kumsebenzi wentlalontle yabantu abadala. Into enomdla kukuba ukuvimba i-histone deacetylation ngexesha lokuhlala ndawonye kwandisa i-NAcc OTR kunye ne-AVPR1a intetho kunye nokwenza lula ukudityaniswa kwi-voiri ye-prairie, iphakamisa ukuba ukuguqulwa kwe-epigenetic yindlela enye apho amava anokubumba iinkqubo ze-neural modulting bonding behaviour [54].

Njengabantu, iziyobisi zokuxhatshazwa zenza ukuba abantu bangakwazi ukuzibamba kwiindawo ezifanelekileyo, ngenxa yotshintsho lweplastikhi kwi-neural circuits. Kwii-prairie voles, i-amphetamine-indased upregation ye-D1R inhibits bonding; kwaye emva kokudibana, i-D1R iyakhula kwaye ihlulele ukuhla kwexabiso lomvuzo wama-amphetamines (ngokuthandabuzekayo, umbandela ophinde ubuyiselwe kunyango lwe-OT) [55-57]. Okubangela umdla kukuba, ukusela utywala ngokuzithandela kuthintela ukubopheleleka kumabhodlo endoda kodwa hayi ababhinqileyo, kwaye ukuququzelela intlalontle kunye nokuthintelwa kotywala obunxilisayo kwenzeka kwababesini-nye kodwa kungengombini oyindoda-nomntu obhinqileyo. [58,59].

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