Indlela yokuziphatha ngokwesondo kwi-Rod Rods (2007)

UElaine M. Hull noJuan M. Dominguez

Abstract.

Imiba yehomoni kunye ne-neural circuitry elawula ukukopana iyafana kuzo zonke iintlobo zeempuku, nangona kukho umahluko kwiipatheni ezithile zokuziphatha. Bobabini i-estradiol (E) kunye ne-dihydrotestosterone (DHT) igalelo ekusebenziseni ukukhwelana, nangona i-E ibaluleke kakhulu kwi-copulation kunye ne-DHT, kwi-reflexes yesini. Ukusebenza kweHormonal yendawo ye-medial preoptic (MPOA) yeyona nto isebenzayo, nangona ukufakelwa kwi-amygdala ye-medial (MeA) nako kunokukhuthaza ukunyuka kwi-castrates. Igalelo le-Chemosensory ukusuka kwiinkqubo eziphambili kunye ne-accessory olfactory systems zezona zinto zibalulekileyo zokukhwela kwiigundane, ngakumbi kwii-hamsters, nangona igalelo le-genitosensory nalo linegalelo. I-Dopamine agonists iququzelela ukuziphatha ngokwesondo, kunye ne-serotonin (i-5-HT) ngokuqhelekileyo inhibitory, nangona i-5-HT ethile ye-receptor subtypes iququzelela ukulungiswa okanye ukukhupha. I-Norepinephrine agonists kunye ne-opiates zineempembelelo ezixhomekeke kwidosi, kunye needosi eziphantsi eziququzelelayo kunye needosi eziphezulu ezivimbela ukuziphatha.

Amagama angundoqo: iigundane, iigundane, i-hamsters, i-guinea pigs, i-estradiol, i-dihydrotestosterone, i-testosterone, indawo ye-medial preoptic, i-medial amygdala, i-reflexes yangasese

Intshayelelo.

Ukuziphatha kokuzala kunye nolawulo lwabo lwe-neural kunye nehomoni luyahluka ngokubanzi kuzo zonke iintlobo. Ukanti uphando oluninzi lujolise kwizilwanyana ezimbalwa ngokwentelekiso. Sichaza indlela yokuziphatha kweempuku zamadoda kunye ne-neural, hormone, kunye nolawulo lwamava. Siqala ngeempuku, izifundo eziqhelekileyo zophando lwebhubhoratri. Emva koko sichaza ukuziphatha kweempuku zamadoda, i-hamster, kunye neehagu ze-guinea, siphawula ukufana kunye nokwahlukana phakathi kweentlobo. Ukuziphatha ngokwesondo kusebenzisana kakhulu; apha sigxininisa kwindoda, sikhumbula ukuba igalelo lebhinqa libaluleke ngokulinganayo. Ngenxa yophando oluninzi malunga neempuku, kunye nemida yamaphepha alo mbhalo-ngqangi, sinokucaphula nje inxalenye encinci yayo. Ukufumana iinkcukacha ezongezelelweyo, nceda uqhagamshelane noHull et al. (2006) okanye uHull et al. (2002).

Inkcazo yeendlela zokuziphatha zokulinganisa iimpuku kunye ne-ex copula reflexes.

Iimpuku zamadoda zihlala ziqala ukudibana ngokwesondo ngokuphanda ubuso bemazi kunye nommandla we-anogenital. Omabini amaqabane anokukhupha amazwi avuselelayo angama-50 kHz. Inkunzi isondela ngasemva kwemazi, ikhwele, kwaye inike iityholo ezikhawulezayo ezikhawulezileyo (19–23 Hz) ngesinqe sayo; ukuba ubhaqa ubufazi bomntu obhinqileyo, utyhala ngokunzulu, afake incanca yakhe kwilungu lobufazi kangange-200–300 msec (Beyer et al., 1981). Emva koko ubuya umva ngokukhawuleza kwaye alungise amalungu akhe angasese. Emva kwe-7 ukuya kwi-10 intromissions, i-1 ukuya kwi-2 imizuzu ngaphandle, uya kukhupha. I-Ejaculation ibonakaliswe kukutyhala okude, okunzulu (750-2000 msec) kunye nokuhla kancinci (Beyer et al., 1981). Ihamba kunye ne-rhythmic contractions ye-bulbospongiosus kunye nemisipha ye-ischiocavernosus kwisiseko se-penis, kunye ne-anal sphincter kunye ne-skeletal muscles (Holmes et al., 1991). Emva kokuphuma kwe-ejaculation, uyazilungisa kwaye emva koko aphumle ngexesha le-postejaculatory interval (PEI), enokuhlala kwi-6 ukuya kwi-10 imizuzu ngaphambi kokuba aqalise ukukhwelana. Ngexesha lokuqala le-50 - 75% ye-PEI, indoda ayiyi kuphinda iphinde idibanise kwaye ikhuphe i-22 kHz vocalizations ultrasonic. Ngexesha le-25% yokugqibela, unokuphinda aqalise ukukopa ukuba unikwe inoveli yabasetyhini okanye isivuseleli esibuhlungu esincinci. Emva kwe-7-8 ejaculations amadoda afikelela kwi-satiety kwaye ngokuqhelekileyo akayi kuphinda abuyele kwi-1 ukuya kwiintsuku ze-3. Amava esondo angaphambili anikezela "ubuchule" obukhulu kunye nokwanda kokuchasana nemiphumo yezilonda ezahlukeneyo, ukuchithwa, kunye noxinzelelo (uhlaziywe kuHull et al., 2006).

Ikhono lokubambisana lifunyenwe phakathi kwe-45 kunye ne-75 iintsuku zeminyaka yobudala (kuhlaziywa kwi-Meisel kunye ne-Sachs, i-1994). I-Prepubertal castration ithintele ukuqala kokuziphatha kokukhwelana, kwaye i-testosterone yangaphandle (T) okanye i-estradiol (E2) ikhawulezise ukuphuhliswa kwayo. Iigundane zamadoda eziguga zilahlekelwa ukukwazi ukukhupha i-ejaculate, engabuyiswanga yi-T exogenous (I-Chambers et al., 1991). Ukuhla kwe-estrogen receptors (ER) (Roselli et al., 1993), kodwa kungekhona i-androgen receptors (AR) (I-Chambers et al., 1991), inokuthi ibe phantsi kwentsilelo kumadoda amadala.

I-Ex copula reflexes inokubonwa kwiimeko ezininzi. Ukumiswa okuzenzekelayo okanye okubangelwa ngamachiza kwenzeka kwikheji yasekhaya okanye kwibala elingathathi hlangothi. Ivumba eliguquguqukayo elivela kwi-estrous female libangela ukumiswa okungaxhunywanga, okunokuba ngumzekelo wokwakhiwa kwengqondo ebantwini. Kwiigundane "i-touch-based" erections inokufunwa ngokunqanda indoda emqolo kwaye ihoxise i-penile sheath. Ezi zilungiso zibangelwa yi-engorgement ye-corpus spongiosum, evelisa i-tumescence ye-glans penis (ihlaziywe kwi-Hull et al., 2006; i-Meisel kunye ne-Sachs, i-1994). I-Anteroflexions nayo iyenzeka; ezi zibangelwa ukunyanzeliswa kwemisipha ye-ischiocavernosus kunye nokumiswa kwe-corpus cavernosum, ebangela ukuba i-penis iphakame kwindawo yayo eqhelekileyo ye-posteroflexed. Ngamanye amaxesha, ukukhutshwa kwamadlozi kwenzeka kulo mongo. Uxinzelelo oluqhubekayo lwe-sheath ehoxisiweyo malunga nesiseko se-penis sinika i-stimulus kwezi zi-reflexes ezisekelwe kwi-touch. Ekugqibeleni, i-reflex ye-urethrogenital iye yafundwa kwiigundane ze-anesthetized zamadoda nabasetyhini njengomzekelo we-orgasm ebantwini (McKenna et al., 1991). Ikhutshwe yi-distension urethral, ​​ilandelwa kukukhululwa; iqulethe i-clonic contractions yemisipha ye-perineal.

Imiba yeHormonal ekusetyenzisweni kokuziphatha kwamadoda empuku.

Ukuziphatha kwendoda ngokwesondo kuzo zonke iintlobo ze-vertebrate kuxhomekeke kwi-T, efihliweyo ngamaseli e-Leydig ama-testes kunye ne-metabolized kwiiseli ezijoliswe kuyo nokuba yi-E2 (nge-aromatization) okanye i-dihydrotestosterone (DHT, nge-5α-reduction). I-Plasma T ayibonakali kwiiyure ezingama-24 zokuthenwa (uKrey noMcGinnis 1990); nangona kunjalo, amandla okukopa ayancipha ngokuthe ngcembe kwiintsuku okanye iiveki. Iintsuku ezintlanu ukuya kwezi-10 ze-T zihlala zifunwa ukubuyisela ukukhwelana (McGinnis et al., 1989). Nangona kunjalo, i-E2 yonyusa uphando lwe-chemo kunye nokunyuka ngama-castrates ngaphakathi kwe-35 min (i-Cross and Roselli 1999). Ke ngoko, ngokukhawuleza, mhlawumbi kusekwe kwi-membrane, iziphumo zehomoni zinokuba negalelo kwimpembelelo yezesondo, kodwa iziphumo zexesha elide ze-genomic ziyafuneka ukuze kubuyiselwe ngokupheleleyo ukukhwelana.

Ihomoni enkulu yokuvula indlela yokuziphatha ngokwesondo kwiimpuku zamadoda yi-E2, njengoko kucetywayo yi "aromatization hypothesis" (ihlaziywe kuHull et al., 2006). I-DHT, engaqholiyo kwaye inobudlelwane obukhulu kwii-ARs kune-T, ayisebenzi xa isetyenziswa iyodwa. Nangona kunjalo, i-E2 ayigcinanga ngokupheleleyo ukuziphatha kwendoda yesini (uMcGinnis kunye noDreifuss, i-1989; uPutnam et al., 2003) okanye ukhetho lweqabane (Vagell noMcGinnis, 1997). Ke, i-androgens inegalelo ekukhuthazeni nasekusebenzeni kwaye ziyimfuneko kwaye zanele ukugcina i-ex copula genital reflexes (Cooke et al., 2003; Manzo et al., 1999; Meisel et al., 1984). Nangona i-E2 yayingasebenzi kakuhle ekugcineni i-ex copula reflexes, iye yagcina ukungena kwe-vaginal intromissions kwi-copula (O'Hanlon, 1981). U-Sachs (1983) ucebise ukuba u-E enze "i-cascade yokuziphatha" enokuthi ibangele i-reflexes yangasese kwi-copula, kodwa ayikwazi ukubathintela ngaphandle kwe-copula.

Iziphumo zamachiza alawulwa ngokwenkqubo kwindlela yokuziphatha kwempuku eyindoda.

Ii-transmitter zihlala zisebenza ngokubambisana kwiindawo ezininzi, kwaye indawo yesenzo ihlala isaziwa njenge-priori. Ngoko ke, ulawulo lweziyobisi lwenkqubo lunokuba luncedo. Itheyibhile yoku-1 ishwankathela iziphumo zokuziphatha ngokwesondo kweempuku zamadoda kunye nonyango oluchaphazela ukusebenza kwe-neurotransmitter kwindawo yobuchopho engaphezu kwesinye.

Itheyibhile 1- Iziphumo zamachiza alawulwa ngokwenkqubo kwindlela yokuziphatha ngokwesondo kweempuku zamadoda.

Iindawo zobuchopho ezilawula ukuziphatha kweempuku zamadoda ngokwesondo.

Igalelo leChemosensory elivela kwiinkqubo eziphambili kunye ne-vomeronasal mhlawumbi yeyona nto ibalulekileyo ekuziphatheni kweempuku zamadoda. I-bulbectomy ye-olfactory yamazwe amabini, esusa zombini iindlela eziphambili kunye ne-vomeronasal, ivelise ukuphazamiseka okuguquguqukayo kokukopa kunye nokungaxhunywanga kwe-erections, kunye namadoda angama-sexual naïve asemngciphekweni wokukhubazeka (uhlaziywe ku-Hull et al., 2006). Ulwazi oluvela kwiinkqubo eziphambili kunye ne-accessory olfactory systems zicutshungulwa kwi-amygdala ye-medial (MeA), kunye negalelo le-somatosensory elivela kwizitho zangasese, ezihanjiswa ngenxalenye ye-parvocellular ye-nucleus ye-subparafascicular (SPFp), ekwayinxalenye yesekethe ye-ejaculation kwiintlobo ezininzi. (ihlaziywe kwi-Hull et al., 2006). Igalelo elivela kwi-MeA, zombini ngokuthe ngqo kunye ne-nucleus yebhedi ye-stria terminalis (BNST), ukuya kwindawo ye-preoptic ye-medial (MPOA) ibaluleke kakhulu ekuhlanganiseni kwiigundane zamadoda (i-Kondo kunye ne-Arai, i-1995).

I-MPOA ngokuqinisekileyo yeyona ndawo ibalulekileyo yokucwangcisa ukuziphatha ngokwesondo kwamadoda. Ifumana igalelo leemvakalelo ngokungathanga ngqo kuzo zonke iinkqubo zoluvo kwaye ithumela unxibelelwano olubuyelayo emva kuloo mithombo, ngaloo ndlela ivumela i-MPOA ukuba ibe nefuthe kwigalelo eliyifumanayo (uSimerly noSwanson, 1986). Ikwathumela imveliso kwi-hypothalamic, midbrain, kunye ne-brain stem nuclei elawula iipateni ze-autonomic kunye ne-somatomotor kunye namazwe akhuthazayo (u-Simerly no-Swanson, i-1988). Izifundo ezininzi ziye zanika ingxelo yokuphazamiseka okuqatha kunye nexesha elide lokukopa emva kwezilonda ze-MPOA (zihlaziywe kuHull et al., 2006). Nangona kunjalo, iigundane zamadoda ezinezilonda ze-MPOA zaqhubeka zibonisa ukumiswa okungaxhunywanga (Liu et al., 1997) kunye ne-bar-press yokukhanya okuye kwadityaniswa nokufikelela kwibhinqa (Everitt, 1990). U-Everitt (1990) ucebise ukuba i-MPOA ibalulekile kuphela ukudibanisa, kwaye kungekhona ukukhuthaza ngokwesondo. Nangona kunjalo, izilonda ze-MPOA zonakalise inkuthazo yezesondo kwezinye iimeko, kubandakanya ukukhetha iqabane labasetyhini (uEdward kunye no-Einhorn, i-1986; uParedes et al., 1998) kunye nokusukela umntu obhinqileyo (Paredes et al., 1993).

Ngakolunye uhlangothi, ukukhuthazwa kwe-MPOA kuququzelele ukukopishwa, kodwa akuzange kubangele ukukhwelana kwindoda egcweleyo (uRodriguez-Manzo et al., 2000). Ukuvuselela kwakhona kwandisa uxinzelelo lwe-intracavernosal kumadoda ane-anesthetized (Giuliano et al., 1996) kwaye yakhupha i-reflex ye-urethrogenital ngaphandle kokuvuselela urethral (uMarson noMcKenna, i-1994). I-MPOA ayiqhubeki ngokuthe ngqo kwintambo yomgogodla esezantsi, apho ukumiswa kunye nokukhutshwa kwe-seminal kulawulwa; ngoko ke, kufuneka isebenze ezinye iindawo ezithi, nazo, zivuse ezo reflexes.

I-MPOA yeyona ndawo isebenzayo yokuvuselela ihomoni yokukhwelana kwiimpuku ezithenwe; nangona kunjalo, i-T okanye i-E2 implants kwi-MPOA ayizange ibuyisele ngokupheleleyo ukukopishwa, kwaye i-DHT implants yayingasebenzi (ihlaziywe kwi-Hull et al., 2006). Ngoko ke, zombini i-ER kunye ne-AR kwi-MPOA inegalelo kwisakhono sokubambisana kweegundane zamadoda; nangona kunjalo, iziphumo zehomoni kwenye indawo ziyafuneka ukuze kusebenze ngokupheleleyo ukuziphatha.

I-MPOA microinjections ye-classic dopamine (DA) i-agonist apomorphine iququzelele ukukopishana kwi-gonadally intact kunye ne-castrated rats kunye nokwandisa i-reflexes esekwe kwi-touch (ihlaziywe kwi-Dominguez & Hull, i-2005; i-Hull et al., i-2006). I-MPOA i-apomorphine iphinde yabuyisela ukukopana kumadoda anezilonda ezinkulu ze-amygdala (Dominguez et al., 2001). Ngakolunye uhlangothi, umchasi we-DA uthintele ukukopa kunye nokuchukumisa-okusekelwe kwi-reflexes kunye nokunciphisa ukukhuthaza ngokwesondo ngaphandle kokuchaphazela umsebenzi we-motor (uhlaziywe kwi-Dominguez kunye ne-Hull, i-2005; i-Hull et al., i-2006). Ezi ziphumo bezingqamene ne-anatom kunye nokuziphatha.

I-DA ikhutshwa kwi-MPOA ngaphambi nangexesha lokudityaniswa (Hull et al., 1995; Sato et al., 1995). Kwakhona, kwakukho zombini iimpawu zokuziphatha kunye ne-anatomical. Kutshanje, kodwa hayi ngaxeshanye, i-T yayiyimfuneko ekunyukeni kwe-DA kunye nokudibanisa (Hull et al., 1995). Into ephambili ekhuthaza ukukhutshwa kwe-MPOA DA yi-nitric oxide (NO), kuzo zombini iimeko ze-basal kunye ne-female-stimulated (zihlaziywe kwi-Dominguez kunye ne-Hull, i-2005; i-Hull et al., 2006). AKUKHO i-synthase immunoreactivity (NOS-ir) ilawulwa kakuhle yi-T kunye ne-E2 (i-Du no-Hull, i-1999; i-Putnam et al., 2005). I-NO nayo ibalulekile ekusebenzeni kokubambisana, njengoko i-NOS inhibitor (L-NAME) kwi-MPOA ivalwe ukukopishana kwi-naïve yamadoda, ukukhwelana okungahambi kakuhle kumadoda anamava, kwaye kuthintele ukulungelelaniswa okuveliswe kumadoda aphathwe nge-saline nge-7 pre-exposures kwi-estrous. female (Lagoda et al., 2004). Igalelo elivela kwi-MeA liyafuneka kwimpendulo ye-DA kumfazi, kodwa hayi kumanqanaba e-DA esisiseko (Dominguez et al., 2001). Ukuvuselelwa kweekhemikhali ze-MeA kubangele ukwanda kwe-DA engaphandle kwe-MPOA efaniswa nezo ziveliswa ngumfazi (i-Dominguez kunye ne-Hull, i-2001). Akukho zi-neurons ezine-DA kwi-amygdala yeegundane zamadoda; nangona kunjalo, ezinye iziphumo ezivela kwi-MeA ukuya kwi-MPOA, kwaye nangaphezulu ukusuka kwi-BNST, zibonakala ziyi-glutamatergic (Dominguez et al., 2003). I-Reverse-dialysis ye-glutamate kwi-MPOA yokwandisa ukukhululwa kwe-DA, umphumo uvalwe yi-NOS inhibitor (Dominguez et al., 2004). Ukongezelela, i-glutamate ye-extracellular yanda ngexesha lokudibanisa kwaye yanyuka ukuya kwi-300% yamanqanaba e-basal kwisampula yemizuzu emibini eqokelelwe ngexesha lokukhupha; I-dialysis eguquguqukayo ye-glutamate reuptake inhibitors iququzelele imilinganiselo emininzi yokubambisana (uDominguez et al., 2006). Ngokufanayo, i-glutamate microinjected kwi-MPOA yandisa uxinzelelo lwe-intracavernous (Giuliano et al., 1996) kunye ne-reflex ye-urethrogenital (uMarson noMcKenna, i-1994) kwiigundane ezine-anesthetized. Ngoko ke, umfanekiso ohambelanayo uvela, apho i-glutamate, ubuncinci inxalenye ye-MeA kunye ne-BNST, iququzelela ukudibanisa kunye ne-reflexes ye-genital, zombini ngokuthe ngqo kunye nokunyuka kwe-NO-mediated kwi-DA, nayo igalelo ekuqalisweni kunye nenkqubela phambili yokubambisana. Ezinye i-neurotransmitters kwi-MPOA enokuthi iququzelele ukuziphatha ngokwesondo kwendoda yi-norepinephrine, i-acetylcholine, i-prostaglandin E2, kunye ne-hypocretin / orexin (hcrt / orx), kanti i-GABA kunye ne-5-HT ingaba inhibitory. Amanqanaba aphantsi e-opioids angaququzelela, kwaye iidosi eziphezulu zivimbela ukukopishwa (kuhlaziywe kwi-Hull et al., 2006).

Ukurekhodwa kwe-Electrophysiological kubonise ukuba ii-neuron ze-MPOA ezahlukeneyo zifaka isandla kwinkuthazo yesondo kunye nokusebenza kokubambisana (Shimura et al., 1994). Ukukhwelana kwandisa i-Fos-ir kwi-MPOA (ihlaziywe ku-Hull et al., 2006), ngokunyuka okukhulu kwamadoda anamava ngokwesondo, xa kuthelekiswa ne-naïve, nangona amadoda anamava ayenama-intromissions ambalwa ngaphambi kokuphuma kwe-ejaculation (uLumley noHull, i-1999). Ke ngoko, amava ngesondo anokuphucula ukusetyenzwa kwezivuseleli ezifanelekileyo ngokwesondo.

Iphecana le-DA ye-mesocorticolimbic, ukunyuka ukusuka kwindawo ye-ventral tegmental (VTA) ukuya kwi-nucleus accumbens (NAc) kunye ne-prefrontal cortex, ibalulekile ekuqiniseni kunye nokuziphatha okunomdla. Ifumana igalelo kwi-MPOA (uSimerly noSwanson, 1988) kunye neminye imithombo emininzi. Izilonda ze-VTA okanye ze-NAc zandise ii-PEI kunye nokunciphisa i-erections engaxhunywanga, kodwa ayizange ichaphazele ukukopishwa (kuhlaziywe kwi-Hull et al., 2006). Ngakolunye uhlangothi, ukukhuthazwa kombane we-VTA kuququzelele ukukopishwa (Markowski kunye neHull, i-1995). Ukusetyenziswa kweziyobisi kwi-VTA okanye kwi-NAc ngokuyintloko kuchaphazeleke ukusebenza ngokubanzi, kunokuziphatha ngokwesondo ngokuthe ngqo (kuhlaziywe kwi-Hull et al., 2006). Ukukhwelana kusebenze i-Fos-ir kwi-NAc kunye ne-VTA, kwaye ukonyuka kwe-estrous okukhuthazwa ngabasetyhini kuye kwaphuculwa ngamava esondo angaphambili (uLopez no-Ettenberg, 2002a). Ukudityaniswa kunye / okanye ukuvezwa kwivumba lomfazi okwi-estrous ukonyusa ukukhutshwa kwe-DA kwi-NAc (kuhlaziywe kuHull et al., 2006). Ukubuyisela umva i-dialysis ye-5-HT kwindawo yangaphambili ye-hypothalamic (LHA) yehla i-basal DA kwi-NAc kwaye ithintele ukunyuka okwenzeka ngenye indlela ngokwaziswa kowesifazane (Lorrain et al., 1999). Ngenxa yokuba i-5-HT inyuswe kwi-LHA ngexesha lokukhupha (uLorrain et al., 1997), ukuhla okubangelwa yi-NAc DA kunokufaka isandla kwi-PEI.

I-nucleus ye-paraventricular (PVN) ye-hypothalamus iquka ulwahlulo lwe-magnocellular, olukhupha i-oxytocin kunye ne-vasopressin kwi-circulation ukusuka kwi-pituitary yangasemva, kunye ne-parvocellular division, eyenza iprojekthi kwiindawo ezininzi zobuchopho kunye nentambo yomgogodla. Izilonda ze-Excitotoxic zenxalenye ye-parvocellular zinciphise ukumiswa okungaxhunywanga kodwa azizange ziphazamise ukukopa (Liu et al., 1997). Izilonda ezifanayo zanciphisa inani le-semen ejaculated kunye nenani le-oxytocin-containing fibers kwi-spinal cord, kodwa kwakhona ayizange ichaphazele ukukopa (u-Ackerman et al., 1997). Izilonda ezibandakanya ulwahlulo zombini ziye zaphazamisa ukukopishwa, kunye ne-touch-based and noncontact erections (Liu et al., 1997). I-Argiolas kunye noMelis baye banikezela ngomfanekiso omhle apho i-DA, i-oxytocin, kunye ne-glutamate (u-Melis et al., 2004) yandisa imveliso ye-NO kwiiseli ze-oxytocinergic kwi-PVN, ethi emva koko ikhulule i-oxytocin kwi-hippocampus (Melis et al., 1992) , intambo yomgogodla (u-Ackerman et al., 1997), nakwenye indawo, ngaloo ndlela ukwandisa ukulungiswa kunye nokukhutshwa kwe-seminal kunye nokuphucula ukukopishwa (kuhlaziywe kwi-Argiolas kunye ne-Melis, i-2004). I-GABA kunye nee-opioids zithintela ezi nkqubo. Le lebhu ibonise kwakhona ukuba i-DA (Melis et al., 2003), glutamate, (Melis et al., 2004), kunye NO (Melis et al., 1998) bakhululwa kwi-PVN ngexesha lokukopa.

Iindawo ezininzi zobuchopho ezongezelelweyo zinefuthe kwindlela yokuziphatha kwempuku yamadoda. I-5-HT ikhutshwe kwi-LHA ngexesha lokukhupha, njengoko kuphawuliwe ngasentla, kunye ne-microinjection ye-SSRI kwi-LHA enqatshelwe ukukopishwa (Lorrain et al., 1997). Ke ngoko, le inokuba yindawo enye apho i-SSRI antidepressants yenza ukuthintela ukusebenza ngokwesondo. Ukongeza, i-hypocretin/orexin (hcrt/orx) i-neurons ihlala kwi-LHA kwaye iyasebenza (Fos-ir) ilandela ukukopishwa, kwaye amanani e-hcrt/orx neurons ehlile emva kokuthenwa (Muschamp et al., ifakiwe). Ngaphezu koko, i-5-HT inqanda i-hcrt / orx neurons kwi-LHA (Li et al., 2002). Ngoko ke, indlela enokwenzeka ngayo i-LHA 5-HT evimbela ukuziphatha ngokwesondo ngokuthintela i-hcrt / orx neurons, eya kususa impembelelo yabo yokuncedisa kwi-VTA DA cell firing (Muschamp et al., ifakiwe).

I-nucleus paragigantocellularis (nPGi) ye-medulla ngowona mthombo omkhulu wothintelo lokuziphatha ngokwesondo kwempuku yamadoda. Izilonda ziququzelele ukukopa kunye nokulibaziseka kokuhlutha ngokwesondo (Yells et al., 1992). Izilonda ezifanayo ziququzelele i-reflexes esekelwe kwi-touch (Holmes et al., 2002; Marson et al., 1992) kwaye yavumela i-reflex ye-urethrogenital ukuba ikhutshwe ngaphandle kwe-spinal transection (uMarson noMcKenna, i-1990). Uninzi lwama-axons avela kwi-nPGi ukuya kwi-lumbosacral intambo yomgogodla iqukethe i-5-HT (uMarson noMcKenna, i-1992). I-5-HT ye-neurotoxin yanciphisa ukuhla kwe-inhibition ye-urethrogenital reflex, kunye nokusetyenziswa kwe-5-HT kwintambo yomgogodla igxininise i-reflex kwi-rats-transected rats (Marson noMcKenna, 1994). Ngaloo ndlela, i-5-HT esuka kwi-nPGi yi-inhibitor enkulu ye-reflexes yangasese.

I-generator ejaculation kwi-lumbar spinal cord iquka i-galanin- kunye ne-cholecystokinin (CCK) ene-neurons, ebonisa i-Fos-ir kuphela emva kokukhupha (i-Truitt ne-Coolen, i-2002; i-Truitt et al., i-2003). Izilonda zezi neurons ziphazamise kakhulu ukuphuma kwe-ejaculation; ngoko ke, abathwali kuphela i-ejaculation-specific sensory input to the brain, kodwa baphinde bakhuphe i-ejaculation (Truitt kunye neCoolen, 2003).

Inkcazo ye-mouse yendoda yokuziphatha kokukopa kunye ne-penile reflexes.

Impuku iye yathandwa kwizifundo zokuziphatha, ubukhulu becala ngenxa yokukwazi kwethu ukuvelisa i-transgenics, i-knockouts, kunye ne-knockdowns (bona uBurns-Cusato et al., 2004, uphononongo oluhle kakhulu). Impuku eyindoda iqala ukudibana ngokuphanda indawo yemazi, isoloko iyiphakamisa okanye imtyhale ngempumlo. Inkunzi emva koko icinezela imilenze yangaphambili kwiphango lemazi ize yenze ukutyhala okukhawulezileyo, okungenzulwanga. Xa ilungu lobudoda lingena kwilungu lobufazi, ukutyhala kwakhe ngokuphindaphindiweyo kuya kucotha kwaye kunzulu. Emva kokumitha izihlandlo ezininzi, inkunzi ikhupha amadlozi, apho inokuba ngumkhenkce imizuzwana engama-25 phambi kokuba yehle okanye iwe phantsi kwemazi. Kukho iiyantlukwano ezininzi zoxinzelelo ekukhwelaneni kweempuku. Ngokomzekelo, i-ejaculation latencies isukela kwi-594 ukuya kwi-6943 imizuzwana, kwaye amanani e-intromissions angaphambili aphuma kwi-ejaculation ukusuka kwi-5 ukuya kwi-142. Ukwaziswa komntwana osetyhini omtsha (uMosig noDewsbury, 17). Kwindawo iimvavanyo ezikhethwayo zombini ii-intromissions kunye ne-ejaculations zibonakaliswe njengomvuzo (Kudwa et al., 60).

I-reflexes esekelwe kwi-touch nayo iye yabonwa kwiimpuku. Ngokungafaniyo neempuku, iimpuku eziziinkunzi ezingagungqiyo azizange zibonise ukuphonononga okuzenzekelayo ngelixa zithintelwe kunye ne-penile sheath yazo ihoxisiwe; nangona kunjalo, uxinzelelo lwesisu luye lwabangela ukulungiswa, kodwa kungekhona i-anteroflexions (Sachs, 1980). Isihlunu se-bulbospongiosus sinegalelo ekumiseni ngexesha lokungeniswa kwaye ngakumbi kwiikomityi (i-erections eqinile ebambe isidoda kumlomo wesibeleko sowesifazane), ezibalulekileyo ekukhuleni kwebhinqa (u-Elmore no-Sachs, i-1988).

Izinto zeHormonal ekusebenziseni ukuziphatha kwenkunzi yempuku.

I-T iyasebenza ngakumbi kune-DHT okanye i-E2 ekubuyiseleni ukuziphatha kwangaphambili kunye nokuziphatha kwiigundane ezithenwe, kunye novakalelo kwi-DHT kunye ne-E2 eyahluka ngokubanzi phakathi kweengxaki (zihlaziywe kwiBurns-Cusato et al., 2004). I-T inokuba neziphumo ezikhawulezayo, njengoko iququzelele ukunyuka kwimizuzu engama-60 kwi-castrates (uJames noNyby, 2002). I-Synthetic androgens (i-5α-androstanediols) enokuthi i-aromatized kwi-E, kodwa ingabi yi-5α-ethotyiweyo kwi-DHT, yayisebenza ngakumbi kune-T ekubuyiseleni ukuziphatha ngokwesondo (Ogawa et al., 1996). Olunye uhlobo, i-B6D2F1 hybrid, yaphinda yakwazi ukuphinda isebenze malunga neeveki ezintathu emva kokuthenwa ngaphandle kwamahomoni angaphandle (McGill kunye noManning, 1976). La madoda "aqhubekayo" axhomekeke kwi-E2; nangona umthombo we-E2 ungacaci, unokuveliswa kwingqondo (Sinchak et al., 1996).

Iindima zeehomoni kwiindawo ezithile zobuchopho zeempuku zamadoda.

Ukufakelwa kwe-T kwi-MPOA ebuyiselwe ngokupheleleyo kwi-vocalization ye-ultrasonic, ukuphawula umchamo ngokuyinxenye, kwaye ibe nefuthe elincinci ekunyuseni okanye ukukhetha umchamo (iSipos kunye neNyby, i-1996). Nangona kunjalo, ukufakelwa okongeziweyo kwe-T kwi-VTA, engazange isebenze yodwa, ivelise iziphumo ze-synergistic ekunyukeni kunye nokukhethwa komchamo. Ukufakelwa kwe-E2 kwi-MPOA yayisebenza njenge-T (Nyby et al., 1992).

I-Steroid receptor mutants.

Ukuguqulwa kwe-testicular feminization (Tfm, okanye i-androgen insensitivity) kwiigundane, kunye nezinye izilwanyana, ziphumela ekucinyweni kwesiseko esisodwa kwi-AR gene (ehlaziywa kwiBurns-Cusato et al., 2004). Amadoda e-Tfm abonakala engowasetyhini, akananzala, kwaye azibandakanyeki ekuziphatheni ngokwesondo ukuba avavanyiwe ngaphandle kwamahomoni angaphandle. Amasende amancinci akhupha amanqanaba aphantsi e-T kunye ne-DHT. Nangona kunjalo, ukuba aba besilisa bathenwa kwaye baphathwa ngeenaliti zemihla ngemihla ze-DHT, T, E, okanye i-E + DHT, baqala ukubonisa amanani ahlukeneyo okuziphatha ngokwesondo, kubandakanywa ukukhutshwa ngezikhathi ezithile (Olsen, 1992). Iigundane ezingenayo i-ERα (ERαKO) zibonisa ukuziphatha okuncinci kwezesondo, nangona xa zithenwa kwaye zatshintshwa nge-T (Rissman et al., 1999; Wersinger and Rissman, 2000a). Oku akubangelwa kukunqongophala kwamahomoni, njengoko amadoda e-ERαKO akhupha ngaphezulu kwe-T kuneempuku zohlobo lwasendle, ngenxa yokuncipha kwengxelo engalunganga ye-ER (Wersinger et al., 1997). I-Castration ye-ERαKO yamadoda kunye nokutshintshwa kwamanqanaba aqhelekileyo e-T (Wersinger et al., 1997) okanye ngaphezulu kwamanqanaba aqhelekileyo e-DHT (i-Ogawa et al., 1998) yanda ukunyuka, kodwa ayizange ibuyisele i-ejaculation. Iinaliti ezicwangcisiweyo ze-apomorphine ye-apomorphine ye-DA ibuyisele ukukhwelana kunye nokukhethwa kweqabane lamadoda e-ERαKO ukuya kwesiqhelo (Wersinger and Rissman, 2000b). Nangona kunjalo, i-apomorphine icv ibuyiselwe kuphela ii-mounts kunye ne-intromissions (echazwe kwiBurns-Cusato et al., 2004). Amadoda asePubertal aswele i-ERβ (ERβKO) afumene amandla okukhupha emva kwexesha kunamadoda e-WT, kodwa ayeqhelekile (iTemple et al., 2003). Amadoda asweleyo zombini ii-ERs awazange ahlangane konke konke xa i-gonadally intact (Ogawa et al., 2000). Nangona kunjalo, i-apomorphine yakwazi ukuvuselela ukunyuka kwizilwanyana ezininzi kunye nokungena kwisiqingatha; akukho nanye i-ejaculated (echazwe kwiBurns-Cusato et al., 2004). Amadoda e-Genetic angenayo i-AR kunye ne-ERα ayizange idibanise, nasemva kokuthenwa kunye nokutshintshwa kwe-T; nangona kunjalo, indibaniselwano ye-E2 yokutshintshwa kunye ne-apomorphine ye-systemic iye yavuselela ukunyuka kwezinye izilwanyana (echazwe kwiBurns-Cusato et al., 2004). Amadoda aswele i-aromatase (ArKO) awakwazi ukwenza i-E kodwa ane-receptors eqhelekileyo. Abambalwa abangamadoda be-ArKO bakhwela, bangena, kwaye baphuma, kwaye babenexesha elide xa besenza; nangona kunjalo, malunga nesinye kwisithathu kubo babekwazi ukukhwela iilitha xa bebekwe kunye nebhinqa ixesha elide (Bakker et al., 2002; Matsumoto et al., 2003).

Iziphumo zamachiza alawulwa ngokwenkqubo kwindlela yokuziphatha ngokwesondo kwempuku.

Nceda ujonge iTheyibhile 2 ngesishwankathelo seziphumo zenkqubo yeziyobisi kwiimpuku zamadoda kunye neehamster.

Iindima zeendawo ezahlukeneyo zobuchopho ekuziphatheni ngokwesondo kwempuku yamadoda.

Iimpawu zeChemosensory zibaluleke kakhulu ekuziphatheni ngokwesondo kwiimpuku zamadoda (zihlaziywe kuHull et al., 2006). Nangona kunjalo, inkqubo ye-vomeronasal ingaba nendima ebalulekileyo, kodwa ingabalulekanga, ekuzaleni. Izilonda ze-MPOA zonakalise kakhulu ukudibanisa kwiimpuku zamadoda, njengakwezinye iintlobo (zihlaziywe kwi-Hull et al., 2006). U-ERAKO wayene-nNOS-ir encinci kwi-MPOA kune-WT okanye iigundane ze-Tfm; ngoko ke, u-E ulawula i-nNOS-ir kwiimpuku (i-Scordalakes et al., 2002) kunye nakwiimpuku.

Inkcazo ye-hamster yendoda yokuziphatha kokukopa.

Ukuziphatha kokukhwelana kwee-hamster kwahluka ngeendlela ezininzi kwezo zeempuku kunye neempuku (zihlaziywe eDewsbury, 1979). I-hamster yegolide yaseSiriya ibhinqa ihleli kwi-lordosis posture ngokuqhubekayo ngokulandelelana okulandelelanayo. Ukukhwelana kuqhubela phambili ngokukhawuleza kuneegundane, kunye ne-inter-intromission intervals yemizuzwana ye-10 kuphela kwaye i-PEIs inyuka ukusuka kwi-~ 35 sec emva kokuphuma kokuqala ukuya kwi- ~ 90 sec emva kwesithoba. Ii-intromissions kunye ne-ejaculations zinde, ~ 2.4 kunye ne-3.4 sec, ngokulandelanayo. IiHamsters nazo zineejaculate ezininzi kuneempuku, zihlala ziyi-9 okanye i-10, zilandelwa luchungechunge lwe "intromissions ende," kunye ne-intravaginal thrusting kwaye akukho kudluliselwa kwesidoda, ngaphambi kokuhlutha. Uhlalutyo olunzulu lwepateni ye-hamster yokukhwelana, kusetyenziswa i-accelerometric kunye ne-polygraphic technique, ibonise ukuba oololiwe be-pelvis batyhula balinganiselwa malunga ne-1 sec, nangona izitimela ezinxulumene nokunyuka zazide kunezo zine-intromissions kunye ne-ejaculations (Arteaga & Moralí, 1997). Iifrikhwekhwezi zokutyhala kwi-pelvic kumndilili wokutyhala nge-15 ngesekhondi nganye, nangona oololiwe ngexesha lokunyuswa bebecotha. Ngexesha le-intromissions bekukho ixesha ngaphandle kokutyhala, kanti ngexesha le-ejaculation, ukutyhala bekukwi-frequency ephezulu (16.4/sec) kunye namandla angaphantsi. Ukungeniswa ixesha elide kuphawulwe nge- ~ 6 ukuya kwi-25 sec ye-intravaginal thrusting ecothayo (1 ukuya ku-2 ngesekhondi nganye). Ubude bexesha lokufakwa kwelungu lobudoda belilide kwi-ejaculate kune-intromissions, kodwa belimfutshane kunoko lide.

IHormones.

Ukunqongophala kwe-T ngexesha lokufikisa okungahambi kakuhle emva kokutshintshwa kwe-T ebudaleni, xa kuthelekiswa ne-castrates kunye nokutshintshwa kwe-T ngexesha lokufikisa (Schultz et al., 2004). Amava esondo aphindaphindiweyo awazange ahlawule ezi ntsilelo. Ivumba lomfazi owamkelayo lisebenze i-Fos-ir kwi-MPOA nangaphambi kokufikisa (i-Romeo et al., 1998), kodwa ayizange inyuse i-DA metabolite DOPAC (umlinganiselo womsebenzi we-DA) de kube semva kokufikisa (Schultz et al., 2003) ). Ke ngoko, ukufikisa kunokuba lixesha lesibini lombutho apho iihomoni ze-gonadal ziguqula ngokusisigxina ukusetyenzwa kwe-neural kwiindawo ezilawula indlela yokuziphatha ngokwesondo (Romeo et al., 2002; Schultz et al., 2004).

Iziphumo zeziyobisi ezilawulwa ngokusesikweni kwiihamster zamadoda.

Nceda ujonge iThebhile 2 isishwankathelo seziphumo zeyeza zenkqubo kwiigundane kunye neehamsters.

Iindima zeendawo ezahlukeneyo zobuchopho kwindlela yokuziphatha ngokwesondo ye-hamster.

I-bulbectomy ye-olfactory ye-Bilateral okanye i-diafferentation edibeneyo yeendlela eziphambili kunye ne-accessory olfactory systems zipheliswe ngokusisigxina ukuziphatha ngokwesondo (kuhlaziywe ku-Hull et al., 2006). Ukungahambelani kwenkqubo yokuvumba yokufikelela kuneziphumo eziguquguqukayo, kunye namadoda anamava angachaphazeleki kangako (Meredith, 1986). Ukonyuka okubangelwa kukukhwelana kwi-Fos-ir kwiibhalbhu ze-olfactory ezingundoqo kunye ne-accessory olfactory bulbs zazithe ngqo kwi-chemosensory stimuli, endaweni yokukhwelana (kuhlaziywe ku-Hull et al., 2006).

Nokuba i-T okanye i-E, kodwa hayi i-DHT, ifakela kwi-MeA ebuyiselweyo ekuziphatheni okukopayo kwi-hamster yamadoda ethenwe (Wood, 1996). Ke, ukusebenza kwehomoni ye-MeA kwanele ukubonakaliswa kokuziphatha ngokwesondo kwiihamster zamadoda. Uqikelelo olusuka kwi-MeA yokuhamba ngendlela yestria terminalis kunye ne-ventral amygdalofugal indlela eya kwi-BNST, MPOA, kunye nezinye iindawo. Ukusika i-stria terminalis kulibazisekile kwaye kwacotha ukukopa, kunye nokusikwa okudityanisiweyo kwazo zombini iindlela kuphelisa ukukopa (uLehman et al., 1983).

Njengezinye iintlobo ezininzi, i-MPOA ibalulekile ekuziphatheni ngokwesondo kwiihamster zamadoda. Nangona kunjalo, ukufakelwa kwe-steroid kwi-castrates kunemiphumo eguquguqukayo kwaye ayanele ukubuyisela ukuziphatha ngokupheleleyo (uWood kunye noNewman, i-1995). Iimpawu zeChemosensory zisebenze i-Fos kwi-MPOA yeehamster zamadoda (uKollack-Walker noNewman, 1997). I-nNOS-ir i-co-localized kunye ne-gonadal steroid receptors kwi-MPOA, kwaye i-castration iyancipha i-nNOS-ir (i-Hadeishi kunye ne-Wood, i-1996). Njengakwiimpuku, amanqanaba e-DA e-extracellular enyuka kwi-MPOA yeehamster zamadoda ezivezwa nge-estrous female; olu lwando luvalwe ngamacala amabini okanye i-ipsilateral, kodwa kungekhona i-contralateral okanye i-sham, i-bulbectomy (i-Triemstra et al., 2005).

Inkcazo yehagu yehagu yendoda yokuziphatha.

Iihagu ezingamadoda zibandakanyeka kwiintlobo ezininzi zokuziphatha kwangaphambili, kubandakanya ukufunxa uboya bemazi entlokweni nasentanyeni, ukufunxa indawo yayo ye-anogenital, kunye nokwenza izandi zamathumbu ngelixa ijikeleza imazi okanye iguqula ubunzima bayo ezinyaweni zayo ezimbini ezingemva ngelixa igcina imilenze yangaphambili. umile (Thornton et al., 1991). Indoda emva koko isondela kwimazi ukusuka ngasemva, ibeke isifuba sayo phezu komqolo wemazi ngelixa ibambe amacala ayo, kwaye iqala ukutyhala i-pelvic, okuhlala kukhokelela ekungeneni kwilungu lobufazi (Valenstein et al., 1954). Amadoda angakwazi ukungenisa ngesantya esimalunga ne-1 ngomzuzu (Thornton et al., 1991), kwaye i-80% inokukhupha kwi-15-min test test (Butera & Czaja, 1985.) Nangona indoda ekhupha i-ejaculate kunye nebhinqa elilodwa ngokuqhelekileyo yenza angaqalisi ukukopa kwiyure elandelayo, unokuphinda adibanise nomntu obhinqileyo owahlukileyo (Grunt & Young, 1952).

IHormones.

Ngokungafaniyo neempuku zamadoda, i-DHT elawulwa ngokucwangcisiweyo inokubuyisela ngokupheleleyo ukukopishana kwiihagu zehagu zamadoda (Butera & Czaja, 1985). Ngaphaya koko, ukufakelwa kwe-DHT kwi-MPOA kwakwanele ukuba kusebenze ukukopa kwi-castrates (Butera kunye neCzaja, 1989).

Isishwankathelo kunye nemibuzo engaphendulwanga.

Nangona kukho iiyantlukwano kwizinto zokukopa phakathi kweempuku, izinto zehomoni kunye ne-neural circuitry ezilawula ezo zinto ziyafana. Bobabini i-E kunye ne-DHT igalelo ekusebenziseni ukukhwelana, nangona i-E ibaluleke kakhulu ekudibaniseni kunye ne-DHT, kwi-reflexes yangasese yeegundane, iigundane kunye ne-hamsters. Ukusebenza kweHormonal ye-MPOA yeyona nto isebenzayo, nangona ukufakelwa kwe-MeA nako kunokuvuselela ukunyuka kwama-castrates. Amagalelo e-Chemosensory asuka kwiinkqubo eziphambili kunye ne-accessory olfactory systems zezona zivuselelayo zibalulekileyo zokukhwelana, ngakumbi kwii-hamsters, nangona igalelo le-genitosensory nge-SPFp nalo linegalelo. I-agonists ye-DA iququzelela indlela yokuziphatha ngokwesondo xa itofwe ngokwenkqubo okanye kwi-MPOA okanye kwi-PVN. I-5-HT i-agonists, ngakumbi i-5-HT1B, ithambekele ekuvimbeleni ukuziphatha, nangona i-5-HT2C i-agonists iququzelela ukulungiswa kunye ne-5-HT1A i-agonists iququzelele i-ejaculation (ngaphandle kweegundane). I-norepinephrine agonists kunye ne-opiates zineempembelelo ezixhomekeke kwidosi, kunye needosi eziphantsi eziququzelelayo kunye needosi eziphezulu ezivimbela ukuziphatha.

Imibulelo.

Ukulungiswa kwalo mbhalo wesandla kwaxhaswa yi-NIMH izibonelelo ze-R01 MH 40826 kunye ne-K02 MH 001714 kwi-EMH.

Imibhalo esemazantsi.

Iphepha elichazayo ukuba awusenanto oyipapashileyo: Le yifayile yePDF yemibhalo ebhaliweyo engamiswanga eyamkelweyo ukuba ipapashwe. Njengenkonzo kubathengi bethu sinikezela ngolu hlobo lwengqangi. Umbhalo obhaliweyo uza kukopishwa, ukuhlelwa kweesetingi kunye nokuphononongwa kobungqina obunokubakho ngaphambi kokuba upapashwe kwifom yokugqibela. Nceda uqaphele ukuba ngexesha lokuveliswa kwempazamo kunokufunyanwa iimpazamo ezinokuchaphazela umxholo, kunye nazo zonke izinto ezisemthethweni ezichazayo ezisebenza kwijenali leyo.

Izalathiso.

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