Tlhahlobo. Tlhokomelo e kahare ea penile erection: Ho boela ho etela karolo ea oxytocin le ho sebelisana le dopamine le glutamic acid lipakeng tsa banna (2011)

Neurosci Biobehav Rev. 2011 Jan; 35 (3): 939-55. doi: 10.1016 / j.neubiorev.2010.10.014. Epub 2010 Nov 2.

Melis MR1, Argiolas A.

  • 1Bernard B. Brodie Lefapha la Neuroscience, Univesithi ea Cagliari, Cittadella Universitaria, 09042 Monserrato, CA, Italy. [imeile e sirelelitsoe]

NAHANA

Oxytocin ke inducer e matla ea penile erection ha e kenngoa mohareng oa methapo ea methapo. Likokoaneng tsa banna, boko bo botle ka ho fetisisa bakeng sa phello ea pro-erectile ea oxytocin ke mokokotlo oa phallo ea hypothalamus. Sebaka sena le libaka tse potolohileng li na le lihlopha tsa lik'hemik'hale tsohle tsa oxytocinergic tse hlalosang libaka tse ling tsa bokopi le tsa mokokotlo. Tlhahlobo ena e bontša hore oxytocin e kenyelletsa penile erection hape ha e kenngoa libakeng tse ling (mohlala, ventral tegmental sebakeng seo, ventral subiculum ea hippocampus, nucleus ea posteromedial cortical ea amygdala le thoraco-lumbar khoele ea mokokotlo). Lithuto tsa microinjection hammoho le microdialysis ea intra-cerebral le lithuto tse peli tsa immunofluorescence li bontša hore oxytocin libakeng tsena e sebetsa ka ho toba kapa ka tsela e sa tobang (haholo-holo ka glutamic acid) mesolimbic dopaminergic neurons. Dopamine e lokolloa nucleus accumbens e boetse e etsa hore ho be le litsela tsa neural tse lebisang ho ts'ebetso ea incerto-hypothalamic dopaminergic neurons mokokotlong oa pela. Hona ha e sebetse feela ka oxytocinergic neurons e hlahisang mokokotlo oa lipalesa le ho phelisana le penile erection, empa le eona e hlahisang libaka tse ka hodimo tsa hypothalamic, ho tsamaisa ka ho toba kapa ka tsela e sa tobang (ka ho sebelisa acid e nang le glutamic) mosebetsi oa mesolimbic dopaminergic neurons e laolang tšusumetso le moputso. Mesebetsi ena ea litsela tsa mahae e ka 'na ea e-ba potoloho e rarahaneng, e nang le karolo e seng karolo feela ea likarolo tsa ho kopanela liphate (mosebetsi oa erectile le phetoho), empa hape le likarolong tse susumetsang le tse khotsofatsang tsa karolo ea tebello ea boitšoaro ba thobalano.

1.Ka hlahisa

Penile erection ke thobalano e motona ea botona kapa botšehali e phethang karolo ea bohlokoa ho hlahisa liphoofolo tse jang liphoofolo tse akarelletsang motho, 'me seo se ka boela sa hlokomeloa ka maemo a fapaneng le a amanang le ho ikatisa. Ho itšetlehile ka moelelo oa moo penile erection e hlahang teng, mekhoa e fapaneng ea bohareng le ea meriana e mengata le / kapa ea boipheliso e kopanela molaong oa eona (bona Meisel le Sachs, 1994; Argiolas le Melis, 1995, 2004, 2005; Sachs, 2000, 2007; McKenna, 2000; Giuliano le Rampin, 2000, 2004; Andersson, 2001; Melis le Argiolas, 1995a, 2003; Hull et al., 2002). Har'a li-neurotransmitters le li-neuropeptides tse ka sehloohong tse laolang penile erection, tse tsebahalang haholo dopamine, serotonin, li-amino acid tse thahasellisang, nitric oxide, adrenocorticotropin, oxytocin le peptides ea opioid. Li ka tsamaisa kapa li thibela phoeni ea penile ka ho sebetsa libakeng tse 'maloa tsa boko, ke hore, sebaka sa lipakeng tsa pele ho moo, mokokotlo oa li-hypothalamus, sebaka sa ventral tegmental, hippocampus, amygdala, nucleus ea bethe ea stria terminalis, nucleus accumbens, medulla oblongata le mokokotlo oa mokokotlo (Tableau 1) (bona Meisel le Sachs, 1994; Witt le Insel, 1994; Stancampiano et al., 1994; Argiolas le Melis, 1995, 2005; Argiolas, 1999; Bancila et al., 2002; Giuliano le Rampin, 2000, McKenna, 2000; Andersson, 2001; Hull et al., 2002; Coolen et al., 2004).

Oxytocin, peptide ea neurohypophyseal e tummeng haholo bakeng sa karolo ea eona ea li-hormone ho lactation le parturition, e teng ho basali le ba batona, eseng feela ka li-neurons tse nang le lihlopha tse nang le nuclei e phahameng le e phahameng ea hypothalamus e hlahisang neurohypophysis, empa hape le li-neurone tse hlalosang ho tloha mokokotlong oa majoe le mehaho e potolohileng ho ea libakeng tse ling tsa boko (e leng, septum, sebaka sa ventral, hippocampus, amygdala, medulla oblongata le mokokotlo oa mokokotlo). Li-neurone tsena li nahanngoa hore li ameha mesebetsing e mengata e ka sehloohong, e kang ho hopola, ho ithuta, ho kopanela liphate le boitšoaro ba likamano tsa botona le botšehali, ho kenyelletsa le penile erection le boitšoaro bo hlephileng (sheba Buijs, 1978; Sofroniew, 1983; Argiolas le Gessa, 1991; Pedersen et al 1992; Carter, 1992; Wagner le Clemens, 1993; Ivell le Russel, 1995; Carter et al., 1997; Tang et al., 1998; Veronneau-Longueville et al., 1999). Ha e le hantle, oxytocin e thusa mosebetsi oa erectile le boitšoaro ba banna ba thobalano ka litoeba, likhoto, le mebutlanyana le litšoene (sheba Argiolas le Gessa, 1991, Carter, 1992; Sena se ka 'na sa etsahala hape ho batho, kaha plasma oxytocin e eketseha ka tšusumetso ea thobalano, haholo-holo ha ejaculation (Carmichael et al., 1992; Murphy et al., 1995) le ho sebelisoa ha lebese le liphatsa tsa lefutso, hangata li etsahala nakong ea thobalano ho kopanela liphate (Tindall, 2004).

Sepheo sa phetoho ea oxytocin ka boitšoaro bo bobe ba ho kopanela liphate e ile ea bontšoa ka lekhetlo la pele ka matla a ho phekola oxytocin ho fokotsa latency ho ea pele ho khaotsa le ho thibela ho felloa ke thobalano ea mebutlanyana e tona e kopanetsoeng le basali ba amohelehang (Melin le Kihlstrom, 1963). Leha ho le joalo, liphello tsa thobalano tsa oxytocin li ne li tsejoa ka ho hlaka feela lilemong tse mashome a robeli. Oxytocin e fanoeng ka nanogram e ngata e ile ea fumanoa e khone ho susumelletsa penile erection (Argiolas et al., 1985, 1986) le ho ntlafatsa boitšoaro ba ho kopanela liphate (Arletti et al., 1985) ka likhoto tsa banna, le ho eketsa lordosis lipakeng tsa basali (Arletti le Bertolini, 1985; Caldwell et al., 1986), kamoo ho bonahalang kateng ka ho sebetsana le li-receptor tsa mofuta oa uterine (bona Argiolas le Melis, 1995, 2004; Argiolas, 1999; Melis le Argiolas, 2003; Oxytocin e ntlafatsa boits'oaro ba thobalano eseng feela likhoto tsa banna tsa botona le botšehali (Arletti et al., 1985) empa hape le li-rats tse seng li tsofetse (Arletti et al., 1990), le tse ling tse ngata, empa eseng ka tlaase, banna ba li-squirrel (Winslow le Insel , 1991).

Phello ea pro-erectile ea oxytocin ke e itšetlehile ka testosterone, kaha e felisoa ke hypophysectomy le castration, 'me e tsosolosoa ka tlatsetso le testosterone kapa metabolite ea eona, estradiol le 5_-dihydro-testosterone e fanoeng hammoho (Melis et al., 1994a). Boko bo botle ka ho fetisisa bakeng sa ho kenngoa ha penile erection ke oktotocin ke mokokotlo oa motsoako oa hypothalamus (Melis et al., 1986), oo ho oona ho qalang li-oxytocineric tse ling tsohle tse ling tse hlahang holimo (see ka holimo). Mona, oxytocin e fumanoe e khone ho etsa hore penile erection (le ho khomarela) ha ejoe ka litekanyetso e le tlaase le 3 pmol (sheba Karolo ea 2.1 ka tlase). Oxytocin e kentse penile erection hape ha e kenngoa ka hare ho naha ea CA1 ea hippocampus, empa eseng ka tlas'a subiculum (sheba Karolo ea 2.3 ka tlaase), sebaka sa lateral, kakaretso e ka tlaase, sebaka sa lipakeng tsa pele, mokokotlo oa hypothalamus le mokokotlo oa supraoptic (Melis et al., 1986). Mabapi le mechine eo oxytocin e sebetsang ka eona mokokotlong oa ho etsa thobalano, lipatlisiso tse ngata li bontša hore oxytocin e sebelisa li-neurone tsa eona. Tumellanong le khopolo-taba ena, ho kopanela liphate ho eketsa FOS, sehlahisoa sa liphatsa tsa lefutso tsa pele tsa liphatsa tsa lefutso c-fos ka poneletso ea oxytocinergic neurons tse hlalosang mokokotlo oa sefate sa mokokotlo, tse amehang ho laola penile erection (sheba Witt le Insel, 1994 le litemana ho eona), le ho hloka matla ho kopanela liphate (mohlala, ho se khonehe ha motho e moholo e motona a ratanang le mosali ea amohelang) o sebelisitsoe ho rat e motona e nang le maemo a tlaase a oxytocin mRNA mokokotlong oa hypothalamus (Arletti et al., 1997).

Ho sa tsotellehe hore na oxytocin e susumetsa mohato ofe kapa ofe kapa mokhoa oa ho qetela oa boitšoaro ba ho kopanela liphate ha o hlahe hantle hona joale. Ha oxytocin e etsa hore penile erection e be le phello e ka sehloohong ea oxytocin ho itšoara ka mokhoa oa ho ikoetlisa ke ho fokotseha ha nako ea kamora ho khaola likokoanyana (Arletti et al., 1985), hoa utloahala ho nahana hore peptide e ntlafatsa ts'ebetso ea thobalano. Leha ho le joalo, kaha oxtocin e eketsa mofuthu oa ho kopanela liphate le batho ba bang (sheba Pedersen et al., 1992; Carter et al., 1997; Ivell le Russel, 1995), le bahanyetsi ba oktotocin ba thibela ho se lumellane ho itseng (Melis et al., 1999a), e leng li nkoa e le letšoao le tsosang takatso ea thobalano (bona Sachs, 1997, 2000, 2007; Melis et al., 1998, 1999b le litemana tsa eona), karolo e ka khonehang ea oxytocin ho tsosa takatso ea ho kopanela liphate le tšusumetso ea thobalano e ke ke ea felisoa.

Tlhaloso ena e akaretsa lipapatso le lipatlisiso tse sa hatisoang tsa lipatlisiso tsa moraorao, tse bontšang hore oxytocin e etsa hore penile erection ha e sebelisetsoe ho kenngoa mokokotlong oa hypothalamus, empa hape le libakeng tse ling tsa bokooa tse ling tse kang hyralthamic (Melis et al., 2007, 2009a; Succu et al., 2008), tke ventral subiculum ea hippocampase le mokokotlo oa kamore ea amygdala (Melis et al., 2009b, 2010), e leng likarolo tsa bohlokoa tsa tsamaiso ea limbicu 'me ho nahanoa hore e phetha karolo ea bohlokoa ho ts'oaroang le ts'ebetso ea moputso. Liphuputso tsena li senola hore oxytocin e kenya letsoho literekeng tsa neural, tse kenyelletsang li-neurotransmitters, tse kang dopamine le glutamic acid, le libaka tse ling tsa bokooa ntle le mokokotlo oa limela, mohlala, sebaka sa ventral tegmental, nucleus accumbens, hippocampus le libaka tse tla ba teng tse tsebahalang. Lipotoloho tsena li ka sebelisana hantle pakeng tsa mesolimbic le incerto-hypothalamic dopaminergic system, le ho bapala karolo e seng karolo feela ea boits'oaro ba boitšoaro bo bobe ba botona le botšehali (mohlala, ho koaloa ha palole le ho kopanela liphate), empa hape le tšusumetsong ea thobalano le thobalano ho tsosa, ka hona ho fana ka karolo ea neural substrate bakeng sa ho hlalosetsa litšobotsi tse susumetsang le tse khotsofatsang tsa thobalano.

2. Oxytocin e susumetsa penile erection ka ho sebetsa likarolong tse fapaneng tsa boko

2.1. Mokokotlo oa mokokotlo oa hypothalamus

Joalokaha ho hopoleloa ka holimo ho khanya ea hypothalamus e ile ea tsejoa e le sebaka sa boko bo botle haholo bakeng sa phello ea pro-erectile ea oxytocin. Ha o ne a entsoe ka unilaterally mokokotlong ona, oxytocin e fumanoe e sebetsa ka tekanyo e lekanang le 3 Ng (3 pmol) (Melis et al., 1986). Liphuputso tsa likamano tsa likamano-ts'ebetso li senola hore penile erection e entsoeng ka oxytocin e ne e sebelisana le li-receptor tsa uterinetype oxytocin, hammoho le tšusumetso ea Ca2 + ka har'a lisele tsa oxytocinergic neurons tse hlahisang libaka tse ling tsa boko bo feteletseng le ho sebelisoa ha nitric oxide-synthase. Nitric oxide e fetoha lenģosa la li-intracellular le mochine o sa tsejoeng (o sa amane le guanylate cyclase) o lebisa ho ts'ebetsong ea li-neurone tsa oxytocinergic tse hlalosang mokokotlo oa mokokotlo le libaka tse ling tsa boko, tse etsang hore penile erection (Fig. 1) (bona ka tlase le Argiolas le Melis, 1995, 2004, 2005 le litemana tsa eona). Bokhoni ba oxytocin bo kenyelletsa methapo ea eona

Setšoantšo sa 1. (MISSING) Tšoantšiso ea tlhahiso ea li-neurone tsa oxytocineric, tse hlahang mokokotlong oa hypothalamus le morero oa libaka tse ling tsa bokopi, tse kang molala oa mokokotlo, VTA, hippocampus, amygdala, joalo-joalo. dopamine, li-amino acids, excitatory amino acid, oxytocin ka boeona, li-peptide tsa hexarelin le li-peptide tse fumanoeng ka VGF li isa penile erection, e ka fokotsoang le / kapa ea felisoa ke ho tsosoa ha methapo ea GABAergic, opioid le cannabinoid CB1. Ts'ebetso ea oxytocinergic neurons ke ea bobeli ho ts'ebetso ea nitric oxide-synthase e teng hona joale ka neurons tsena. Ka sebele nitric oxide e tsitsitseng e entsoeng ka ho tsosolosa ha dopamine, e tsoetseng pele ea amino acid kapa oxytocin receptors kapa oksijene ea nitric oxide, kaha e fumanoe ho bafani ba nitric oxide e fanoeng ka ho toba mokokotlong oa phallo, e etsa oxytocinergic neurons ka mokhoa o sa tsejoeng, eo ho bonahalang eka ha o amana le khatello ea guanylate cyclase. Sena se etsa hore ho lokolloe oxytocin molaong oa mokokotlo le libakeng tse ling tsa boko. Lintlha tse ling tse mabapi le lisebelisoa tseo oxytocin e kenyang penile erection ha li lokolloa libakeng tsena, mohlala, VTA, ventral subiculum le amygdala li hlalosoa likarolong tse ling tsa bokooa. Mona, oxytocin e sebetsana le li-receptor ea eona mme e eketsa ha ho na tlhahiso, e lebisang ho penile erection e fumanoang PVN. Leha ho le joalo, ho fapana le PVN, ka Vala caudal ha e sebetse guanylate cyclase. Sena se baka kgolo ea cGMP e lebisang ho ts'ebetsong ea mesolimbic dopaminergic neurons le ho penile erection. Ho VS HA ha e sebetsane le li-neurone tsa glutamatergic tse hlahisang libaka tse ling tsa hippocampal, ho kenyelletsa le VTA. Letsoalo la Glutamic le VTA le hlahisa mesolimbic dopaminergic neurons e fumanoang ka oxytocin. Mekhoa e tšoanang le e hlalositsoeng ka holimo e ka 'na ea boela ea sebetsa ha penile erection e hlaha likarolong tsa' mele, e leng ha ha liphooe tsa banna li behoa ka pel'a botona ba basali ba sa amoheleng (mohlala, litloaelo tse sa lumellaneng) kapa nakong ea ho kopana.

mokokotlong ona o ne o tšehetsoa ke liphuputso tse bontšang hore: (1) oxytocin receptors li fumaneha mokokotlong ona oa hypothalamic (Freund-Mercier et al., 1987; Freund-Mercier le Stoeckel, 1995); (2) oxytocin e thusa ho lokolloa ha eona ka vitro le ka vivo (Freund-Mercier le Richard, 1981, 1984; Moos et al., 1984); le (3) oxytocin e thabisa li-neurone tsa eona ka ho etsa mokokotlo oa sepakapaka (Yamashita le al., 1987). Ho feta moo, li-synapses tsa oxytocineric tse hlahisang lihlopha tsa lisele tsa magnocellular oxytocineric neurons li boetse li fumaneha kahare e ka sehloohong le e phahameng ea hypothalamus (Theodosis, 1985). Qetellong, ho bolaoa ha li-neuron tse bohareng tsa oxytocinergic ke liso tsa electrolytic kapa lik'hemik'hale tse tsoetseng pele tsa mokokotlo, tse thibelang ka ho feletseng li-oxytocin tsamaisong ea methapo ea mokokotlo le mokokotlo, ha li felise feela phello e-erectile ea oxytocin, empa e boetse e senya lithethefatsi tse entsoeng penile erection le e nonactact erections (bona ka tlaase le Argiolas et al., 1987a, b; Liu et al., 1997 le litemana tsa eona). Liphello tse tsoanang le tse fumanoang ka lisele tsa mokokotlo oa libaka li fumanoa ka bahanyetsi ba matla le ba khethollang li-oxytocin receptor. Ka sebele, lik'hemik'hale tsena tse entsoeng ka har'a khalase e nkiloeng ka nano e ngata li thibela penile erection e kentsoeng ke oxytocin, athe ha e fuoa li-ventricles hamorao ha ea thibela feela penile erections ho susumetsoa ke oxytocin ka boeona, empa hape le penile erection e entsoeng ke lithethefatsi (bona karolo ea 3 e ka tlase le Argiolas le Melis, 1995, 2004, 2005 le litemana tsa eona) le ho se lumellane ho etsahalang (Melis et al., 1999a), hape ba atlehile haholo ho fokotsa boitšoaro bo hlephileng ba likhoto tsa banna tsa thobalano (Argiolas et al., 1988). Ho feta moo, ho kopanela liphate ho eketsa FOS, lihlahisoa tsa liphatsa tsa lefutso tsa liphatsa tsa lefutso tsa c-fos ka tsela e ikhethileng ea oxytocinergic neurons e hlalosang mokokotlo oa mokokotlo o amehang ho laola penile erection (sheba Witt le Insel, 1994 le litemana tsa eona). Qetellong, ho hloka matla a ho kopanela liphate (mohlala, ho se khonehe ha motlatsi e moholo e motona ho sebetsana le mosali ea bitsoang estrogen-progesteroneprimed h)e boetse e amahanngoa le rat e tona e nang le maemo a tlaase a oxytocin mRNA le ea nitric-oxide synthase mokokotlong oa phallo ea hypothalamus (Benelli et al., 1995; Arletti et al., 1997) (bakeng sa tlhahlobo e kholo ea lipatlisiso tsena bona Argiolas, 1999; Argiolas le Melis, 2004, 2005).

2.2. Sebaka sa ventral tegmental

Sebaka sa ventral tegmental se fumanoe morao tjena e le sebaka sa boko moo oxytocin e kenyang penile erection. Sebaka sena se na le likhaello tsa oxytocineric methapo e hlahang mokokotlong oa nucleus le oxytocin receptors (Freund-Mercier et al., 1987; Vaccari et al., 1998). Hantle haholo, oxytocin e fumanoe e khona ho iho fokotsa penile erection ha e entsoe ka unilaterally ho caudal, empa eseng sebakeng sa rostral ventral tegmental ka mokhoa o itšetlehileng ka tekanyo ea methapo (Melis et al., 2007). Meriana e sebetsang e ne e le e phahameng ho feta e hlokahalang ha e kenngoa mokokotlong oa sepakapaka le o tšoanang le o etsang hore penile erection e kenngoe ka ventral subiculum ea hippocampus kapa e kenngoe ka morao-rao ea coronate ea amygdala (bona ka tlase). Kamoo ho bonahalang kateng, phello ea pro-erectile e sebelisoa ke ho sebelisoa ha mesolimbic dopaminergic neurons e hlahisang khetla ea nucleus accumbens, e ntan'o hlahisa litsela tse sa tsejoeng tsa neural tse hlalosang incerto-hypothalamic dopaminergic neurons e phehelang ka paraventricular oxytocinergic neurons lipakeng tsa penile erection (Melis et al., 2007, 2009a).

Mabapi le mechine eo oxytocin e sebetsang ho eona ha e sebelisa dopaminergic neurotransmission sebakeng sa ventral tegmental, data e teng e bontša hore oxytocin e susumetsa li-oxytocinergic receptors tse ka har'a lihlopha tsa lisele tsa mesolimbic dopaminergic neurons. Sena se eketsa tšusumetso ea Ca2 + ka har'a litho tsa lisele tsa dopaminergic neurons, kahoo ho etsa hore nitric oxide-synthase (Succu et al., 2008) e sebelisoe. Ha ho fapana le mokokotlo oa sepakapaka (sheba Karolo ea 3 ka tlase), nitric oxide e etsa hore guanylate cyclase e khone ho eketsa mokokotlo oa GMP. Tumellanong le mokhoa ona, ebang ke (CH2) 5Tyr (Me) 2-Orn8-vasotocin, mohanyetsi ea matla oa oxytocin, kapa S-methyl-thio-l-citrulline, e leng inhibitor e matla ea neuronal nitric oxide-synthase, e kenngoeng ka sebaka sa caural tegmental sebakeng sa oxytocin, se felisitse penile erection le keketseho ea mahlaseli a dopamine a mangata-ngata ka har'a khetla ea nucleus accumbens e bakoang ke oxytocin. Ho feta moo, GMNUMX-bromo-cyclic GMP, e leng sebōpeho sa mofuta oa GMP e sa sebetsanang le phosphodiesterase, e etsa hore penile erection e kenngoe sebakeng sa ventral tegmental le ho eketsa mohopolo oa dopamine o eketsehileng molekong oa nucleus accumbens, e fumanoang ka oxytocin e kenngoa ka har'a sebaka sa caural tegmental (Succu et al., 8; Melis et al., 2008a) (bona le setšoantšo sa 2009).

Kamehla tumellanong le mokhoa ona, haloperidol, mohanyetsi o matla oa Dopamine D2, ea kenngoa ka har'a khetla ea nucleus accumbens e fokotsa penile erection e bakoang ke oxytocin e kenngoa ka ventral tegmental (Melis et al., 2007). Mokhoa ona o boletsoeng ka holimo o tšehetsoa ke lithuto tse peli tsa immuno-fluorescence, ho bontša hore oxytocin fibers impinge liphatleng tsa lisele tsa dopaminergic neurons sebakeng sa caural tegmental, seo pele ho neng ho ngotsoe ka retrogracer tracer Fluorogold e kenngoeng karolong ea nucleus accumbens ( Melis et al., 2007; Succu et al., 2008). Ts'ebetso ea li-neuron tsena tsa dopaminergic le tsa li-receptor tsa dopamine tse nucleus accumbens li etsa hore ho sebelisoe litsela tsa neural tse sa tsejoeng, e leng se susumelletsang incerto-hypothalamic dopaminergic neurons hore e lokolle dopamine mokokotlong oa pela, e leng se etsang hore li-oxytocinergic neurons li hlahise mokokotlo oa mokokotlo le ho lumellana le penile erection (bona ka holimo le Melis et al., 2007; Succu et al., 2007, 2008). Ha e le hantle, oxytocin e kenngoa sebakeng sa ventral tegmental ea caudal e neng e etsa hore penile erection e eketsehe, ho eketsa mahloriso a dopamine a eketsehileng ka har'a dialysate e fumanoeng feela ho tloha nucleus accumbens, empa hape ho tloha mokokotlong oa Succu et al., 2007).

2.3. Hippocampus

Tšimo ea CA1 ea hippocampus e ne e le sebaka se seng sa boko se nang le mafura a oxytocinergic le li-receptors tse khetholloang ke lithuto tsa pejana tseo ente ea oxytocin e bakileng penile erection (bona Bujis, 1978; Sofroniew, 1983). Leha ho le joalo, ha ho fapana le sekoti sa phallo, mona oxytocin e fumanoe e khone ho etsa hore penile erection e be e kenyelletsoeng habonolo feela ho feta e fumanoang e sebetsang mokokotlong oa meliski (Melis et al., 1986; Chen et al., 1992). Likokoana-hloko tsa oxytocin ka subiculum li ile tsa fumanoa li sa sebetse lithutong tsena tse fetileng. Leha ho le joalo, lithuto tsa morao-rao le tse hlokolosi haholo tsa microinjection li lebisa ho khetholla sebaka sa subralulum ea ventral eo ho eona ente ea oxytocin e neng e khona ho fokotsa penile erection ka mokhoa o itšetlehileng ka tekanyo ea methapo (Melis et al., 2009b). Phello ea pro-erectile ea oxytocin e kenngoa sebakeng sena sa boko e ile ea bonoa ka tekanyo e tšoanang le e fumanoang e sebetse sebakeng sa ventral tekaneng ka mor'a hore e se ke ea kenngoa mokokotlong (Melis et al., 2007), e fumanoang mokokotlong o ka pele. Kamoo ho bonahalang kateng, oxytocin e kenngoa ka ventral subiculum e etsa hore penile erection e sebelise li-receptor tsa oxytocinergic li-neurons tse nang le nitric oxide-synthase, e leng se etsang hore ho be le keketseho ea nitric oxide production. Nitric oxide ka nako e tšoanang e sebetsa e le lenģosa la li-intercellular le etsa hore motho a se ke a tšoaetsoa ke motsoako oa glutamic, o lebisang penile erection, mohlomong ka likhahla tsa neural (glutamatergic) tse fapaneng tse tsoang ho subralulum ea ventral ho libaka tse ling tsa bokopi ba hippocampal ho tsamaisa mosebetsi oa mesolimbic dopaminergic neurons (mohlala, sebaka sa ventral tegmental, prefrontal cortex, nucleus e ka pele) (bona ka tlase le Melis, 2007, 2009b; Succu et al., 2008).

Mokhoa ona oa ketso o tšehetsoa ke liteko tsa intra-cerebral microdialysis, tse bonts'ang hore oxytocin e kenngoa ka ventral subiculum ka tekanyo e etsang hore penile erection e eketsehe, e eketsa tlhahiso ea nitric oxide le maqhubu a extracellular glutamic acid ho dialysate ho tloha ventral subiculum (Melis et al. , 2010) le extracellular dopamine ka nucleus accumbens (Melis et al., 2007). Likarabo tsena li ne li sa sebelisetsoe feela ke mohanyetsi oa oxytocin (d) CH2) 5Tyr (Me) 2-Orn8-vasotocin, empa hape ke neuronal nitric oxide-synthase inhibitor S-methyl-thio-lcitrulline le nitric oxidous scavenger hemoglobin e fanoeng ho ventral subiculum metsotso e seng mekae pele oxytocin (Melis et al., 2010).

Ho feta moo, tumellanong le mokhoa ona oa ho nka khato, ho ts'oaroa ha glutamatergic neurotransmission ke NMDA e kenngoang ka subralulum ea ventral e kenyelletsa penile erection (Melis et al., 2010). The phenotype ea hantle e qalang ho tloha ventral subiculum, e leng se etsang hore ho sebelisoe ha mesolimbic dopaminergic neurons le keketseho ea dopamine e eketsehileng ea cellular nucleus accumbens, ha e tsejoe hona joale. Leha ho le joalo, kaha penile erection e susumelitsoeng ke oxytocin e kenngoa ka subralulum ea ventral e hlaha ka mokhoa o lumellanang le ho eketseha ha acide ea glutamic e ngata-ngata ho dialysate ho tloha sebakeng sa ventral tegmental, empa eseng ho tloha nucleus accumbens 'me e tšoaetsoa ke (+) MK-801, e leng mohanyetsi ea matla ea sa tsitsitseng oa li-receptor tse ntle tsa amino ea NMDA subtype (Woodruff et al., 1987), e kenngoa sebakeng sa ventral tegmental, empa eseng ka nucleus accumbens (bona setšoantšo sa 2 le Melis et al., 2009b) ,, ho ka etsahala hore lits'ebetso tsena li lebisa ho ts'ebetsong ea ho tlosoa ha glutamatergic neurotransmission sebakeng sa ventral tegmental, e leng se kenyang mesolimbic dopaminergic neurons se hlahisang nucleus accumbens. Ho sa tsotellehe hore na li-acute glutamic tse eketsehileng li fumanoa sebakeng sa ventral tectal ka mor'a ente ea oxytocin ka ventral subiculum e lokolloa li-neurons tse tsoang subiculum kapa libakeng tse ling tsa boko (mohlala, prefrontal cortex) ha e tsejoe hona joale. Leha ho le joalo, sena se baka ts'ebetsong ea mesolimbic dopaminergic neurons le ho ekolloa ho eketsehileng ha dopamine nucleus accumbens. Mona tšebeliso ea dopamine receptors e lebisa ho ts'ebetso ea incerto-hypothalamic dopaminergic neurons, ho lokolla dopamine mokokotlong oa sepakapaka, kahoo ho etsa hore li-neuron e sebetsane le oxytocinergic e hlalosang mokokotlo oa mokokotlo le ho fokotsa penile erection (bona ka holimo le Melis et al., 2007, 2009a; Succu et al., 2008).

2.4. The amygdala

Amygdala ke sebaka se seng sa boko bo nang le li-oxytocin fibers le receptors (bona Freund-Mercier le al., 1987; Vaccari et al., 1998; Uhl-Bronner et al., 2005). Oxytocin mona ho nahanoa hore e ameha mesebetsing e fapaneng, ho tloha maikutlong, mohopolong oa sechaba le ho tsebisoa, ho ithuta ka sechaba, ho utloela maikutlo ka maikutlo, khatello ea lifahleho le ho tšaba batho ho etsa mosebetsi oa erectile le boitšoaro ba thobalano (bona Kondo et al., 1998; Dominguez et al., 2001; Ebner et al., 2005; Huber et al., 2005; Domes et al., 2007; Petrovic et al., 2008; Lee et al., 2009; Donaldson le Young, 2009; Hurlemann et al. , 2010). Leha ho le joalo, bokhoni ba oxytocin bo etsa hore penile e kenngoe linthong tsa banna ha e kenyelletsoa kotsing ea posteromedial cortical ea amygdala e fumanoe morao tjena (Melis et al., 2009b). Karabo ena e ile ea etsoa ka mokhoa o ts'oanang le ho eketseha ha mohopolo oa dopamine o eketsehileng oa cellular ho dialysate e fumanoang sekhethong sa nucleus accumbens, e fumanoeng ka mor'a ente ea oxytocin ka ventral subiculum (Melis et al., 2009b). Mechine eo oxytocin e kenngoang ka eona karolong ea posteromedial cortical ea amygdala e etsa hore penile erection e se ke ea tsejoa hona joale. Ditaba tse fumanehang li bonts'a hore penile erection le ho eketseha ha mahlaseli a dopamine a eketsehileng-cellular ka dialysate e fumanoeng nucleus accumbens li kopanngoa ke ho sebelisoa ha li-oxytocinergic receptors, ha likarabo tseo ka bobeli li felisoa ke mohanyetsi oa oxytocin receptor d (CH2) 5Tyr ( 'Na) 2-Orn8-vasotocin e kenngoa mokokotlong oa amygdala metsotso e seng mekae pele oxytocin (Melis et al., 2009b).

Ho sa tsotellehe hore na oxytocin e sebetsa joang mokokotlong oa amygdala oa posteromedial ea amygdala, ho kopanela liphate ho susumelitsoeng ke peptide ho felisoa ke thibelo ea li-receptor tsohle tsa dopaminergic tse nang le cis-flupenthixol e kenngoang ka har'a khetla ea nucleus accumbens le ka thibelo ea li-receptors tsa NMDA tse nang le ( +) MK-801 e kenngoa sebakeng sa ventral tegmental, empa e se ka har'a nucleus accumbens, e fumanehang bakeng sa penile erection e bakoang ke oxytocin e kenngoa ka ventral subiculum (Melis et al., 2009b). Sena se fana ka maikutlo a hore oxytocin e kenngoa kahare ea posteromedial ea amygdala e etsa hore glutamic acid e amohelehe sebakeng sa ventral tegmental. Sena se etsa hore ts'ebetsong ea mesolimbic dopaminergic neurons e be teng, e lebisang penile erection. Ka lebaka la lipatlisiso tse bontšang litsela tsa neural li kopanya mokokotlo ona oa amygdala le ventral subiculum (Canteras et al., 1995; French le Totterdell, 2003), liphuputso tsena li hlahisa monyetla oa hore ho ka ba le puisano pakeng tsa libaka tsena tse peli tsa boko, le hoja e le e tobileng litsela tse tsoang ho amygdala ho ea fihla nucleus accumbens kapa sebakeng sa ventral tegmental se hlalositsoe (Kelley le Domesick, 1982; Witter, 2006).

2.5. Mokokotlo oa mokokotlo

Mokokotlo oa mokokotlo ke karolo e 'ngoe ea tsamaiso ea methapo e ka hare e nang le li-fibro-oxyretinic fibers le receptors (Freund-Mercier et al., 1987; Uhl-Bronner et al., 2005), moo oxytocin e kenyang penile erection (Tang et al., 1998 ; Veronneau-Longueville et al., 1999; Giuliano le Rampin, 2000; Giuliano et al., 2001). Joalokaha ho hopoleloe ka holimo, likhoele tsena tsa oxytocineric li simoloha mokokotlong oa hypothalamus 'me li tlatsetsa ho theoha litsela tse laolang li-neuron tsa mokokotlo tse ikemetseng tse phelisang penile erection. Ha e le hantle likhoele tsena li etsa hore li-synaptic li kopane le lenane la li-porsale le li-parasympathetic li-columns tsa thoraco-lumbar le lumbo-sacral tse nang le li-neuron tse sa tšoaneng tse nang le penile cavernous corpora (Marson le McKenna, 1996; Giuliano le Rampin, 2000; Giuliano et al. , 2001). Ts'ebetso ena ea synaptic e ile ea bontšoa ka ho ngolisa li-neurons tsa mokokotlo tse tsoang ka penis le ho fihla mokokotlo oa mokokotlo le li-tracers tse itseng tse kenngoa ka cavernous corpora, li kopantsoe le lithuto tse peli tsa immuno-fluorescence le li-confocal laser microscopy (Tang et al., 1998; Veronneau-Longueville le al., 1999). Tumellanong le lipatlisiso tsena, ho kenya manonyeletso ea intrathecal ea linonyana tse sa tsitsitseng a mangata a mangata a mangata a oxytocin a lumbo-sacral, empa eseng ka tekanyo ea thoraco-lumbar, e ile ea etsa hore khatello e ke keng ea lekanngoa e hlahe ka mokhoa o itšetlehileng ka tekanyo ea methapo. Liphello tsena li ile tsa felisoa ke thibelo ea oxytocinergic receptor le d (CH2) 5Tyr (Me) 2-Orn8-vasotocin le karolo ea methapo ea pelvic (Giuliano le Rampin, 2000; Giuliano et al., 2001). Liphello tsena li bontša hore oxytocin, e sebelisang molamu oa mokokotlo oa lumbo-sacral, e eketsa khatello ea 'mele,' me e fana ka maikutlo a hore oxytocin, e lokolloa nakong ea ts'ebetso ea 'mele ea mokokotlo oa mokokotlo ke motsoako o matla oa sepalesa sa spinal pro-erectile neurons se hlahisang likarolo tsa cavernous. Ho thahasellisang ke hore li-neuron tsena tsa pro-erectile tseo oxytocin li sebetsang ho tsona e le phello ea eona ea pro-erectile, li boetse li fumana li-synaptic tse tsoang ho serotoninergic neurons tse hlahang nucleus paragigantocellularis ea mantlha ea medulla oblongata (Marson le McKenna, 1992; Tang et al ., 1998). Ho timetsoa ha li-neruroninergic neurons tsena ho etsa hore ho be le li-rats le li-plele reflexes likokoanyana tse nyenyane (Marson le McKenna, 1992; Yells et al., 1992). Kaha lithethefatsi tse khothalletsang li-receptor tsa 5HT2C li etsa hore ho fanoe ka penile erection ha e fanoa ka sekhukhu, empa e se ka har'a mokokotlo oa limela, le lithethefatsi tse thibelang li-receptor tsa 5HT2C li boetse li fokotsa dopamine agonistand e entsoeng ke penile erection, ha bahanyetsi ba dopamine ba fokotsa penile erection ea 5HT2C e entsoeng ke agonist ( sheba Stancampiano et al., 1994 le litemana tsa eona), ho boetse ho bolelloa hore oxytocin e thusa mohato oa pro-erectile 5HT2C receptors ka tekanyo ea molamu oa mokokotlo oa lumbo-sacral (Stancampiano et al., 1994). Ntle le moo, oxytocin e ka 'na ea susumetsa tšebetso ea li-neuron tsa serotoninergic tsa mokokotlo ho theoha ka ho sebetsa ka ho toba ka nucleus paragigantocellularis, moo li-neurons tsena li simolohang teng (bona Stancampiano et al., 1994).

3. Likamano pakeng tsa oxytocin, dopamine le glutamic acid ka har'a tsamaiso ea methapo e meholo le ho emisa penile

Joalokaha ho boleloa ho Karolong ea 1, li-neuron tsohle tsa oxytocinergic tse teng tsamaisong ea methapo ea bohareng li tsoa mokokotlong oa limela le mehaho e potolohileng. Mosebetsi oa li-neurons tsena o tlas'a taolo ea li-neurotransmitters tse fapaneng le / kapa li-neuropeptides. E 'ngoe ea tse rutoang ka ho fetisisa li-dopamine, glutamic acid, gamma-aminobutyric acid (GABA), nitric oxide, endocannabinoids, peptides ea opioid, peptide ea ho hōla ha hormone, li-peptide tse amanang le VGF le oxytocin ka boeona. Dopamine, glutamic acid, li-pepide tse hlahisang li-pepide, li-peptide tse fumanoeng ke VGF le oktotocin li susumetsa, mohlala, metsoako ena le li-agonists li etsa hore penile erection e kenngoe ka har'a mokokotlo, ha GABA, peptides ea opioid le endocannabinoid e thibela, mohlala, metsoako ena kapa li-agonists tsa tsona li thibela penile erection (bona Meisel le Sachs, 1994; Witt le Insel, 1994; Argiolas le Melis, 1995, 2004, 2005; Giuliano le Rampin, 2000, 2004; McKenna, 2000; Andersson, 2001; Hull et al., 2002).

Litsela tse 'maloa tsa bopaki ba liteko li bontša hore li-neuron tsena tsa oxytocinergic le li-neurotransmitter tse ka holimo le li-neuropeptides li ameha ho laola mosebetsi oa erectile le boitšoaro ba ho kopanela liphate maemong a sa tšoaneng a mmele. Ho feta moo, oxytocin e lokolloa libakeng tse ling tsa boko, tse kang sebaka sa ventral tegmental, hippocampus le libaka tsa eona, amygdala le mokokotlo oa mokokotlo o ka ama tšebetso ea li-neurone tseo ho tsona li-syntose tsa oxytocineric synapses impinge. Hona joale, ke li-neurone feela tsa bohlokoa bakeng sa ho penile erection eo ho eona oxytocinergic synapses impinge, e tsejoang ka tieo, ke lihlopha tsa lisele tsa mesolimbic dopaminergic neurons tsa sebaka sa ventral tegmental sa caudal se hlahisang khetla ea nucleus accumbens (Melis et al., 2007 ; Succu et al., 2008), le methapo ea pro-erectile ea mongobo e hlahang ho tloha toropong ea lumbo-sacral ho ea ho corvernous corpora (bona Giuliano le Rampin, 2000; Giuliano et al., 2001) (bona le likarolo 2.2 le 2.5) . Ha e le hantle, le hoja li-synapses tsa oxytocineric le li-receptor li 'nile tsa fumanoa ka ventral subiculum, amygdala le mokokotlo oa mokokotlo, libaka tseo e leng tsa bohlokoa bakeng sa penile erection (bona ka holimo), libakeng tsena mofuta oa li-neurotransmitter / s e leng phekolo ea methapo ea oxytocineric impinge, e ntse e sa tsejoe.

Karolo ena ea tlhahlobo ena e akaretsa ka bokhutšoanyane lingoliloeng tsa morao-rao tse mabapi le mekhoa ea phello e-erectile ea oxytocin e kenngoa sebakeng sa ventral tegmental ea caudal, ventral subiculum ea hippocampase le mokokotlo oa mokokotlo. Tlhokomelo e khethehileng e fanoa ka ho sebelisana ha peptide e nang le dopamine le glutamic acid libakeng tsena le karolo eo karolo ena e ka bapalang ka eona ho laola karolo ea erectile. Tlhaloso e khuts'oanyane ea liphello tsa dopamine le glutamic acid ka oxytocinergic neurons mokokotlong oa seretse, o nang le karolo ea bohlokoa mosebetsing oa erectile o fanoa pele, e le ho etsa hore 'mali a hlokomele boemo ba pele ba lipatlisiso tšimong ena, joaloka lithuto tsena li se li ntse li hlahlojoa haholo (bona Argiolas le Melis, 1995, 2004, 2005; Melis le Argiolas, 2003). Hape tabeng ena, tlhokomelo e khethehileng e fanoa ka liphello tsa morao-rao, tse bontšang karolo ea bohlokoa bakeng sa linkano tsa dopamine-oxytocin le link ea glutamic acid-oxytocin eseng feela ho etsa thobalano (penile erection le copulation), empa le ka thobalano ho tsosoa le ho susumetsa thobalano.

3.1. Phetoho ea Dopamine-oxytocin mokokotlong oa sepakapaka

The bokhoni ba li-agonists tsa dopamine ho etsa hore penile erection e sebelise li-neuron tse bohareng o ne a etsa tlhahiso ea hang-hang ka mor'a ho sibolloa eo apomorphine bitsang penile tsogelwa ha tshelwa ka paraventricular khubung (Melis et al a., 1987) ha oxytocin receptor antagonist d '(CH2) 5Tyr (Me) 2-Orn8-vasotocin tshelwa intracerebroventricularly (icv) e ile ea fumanoa e khona ho fokotsa hoo e batlang e le ka ho feletseng penile erection e se ke ea susumetsoa feela ke oxytocin e fuoeng icv, empa e boetse ke apomorphine, e fanoeng ka tlase (Argiolas et al., 1987b). Liphello tsena li ile tsa lateloa ke tsa lithuto tse ling tse bontšang liphello tse tšoanang ha d (CH2) 5Tyr (Me) 2-Orn8-vasotocin e fanoe ka icv le apomorphine e fanoe ka ho toba mokokotlong o ka holimo (Melis et al., 1989b), lho etsa tlhahiso ea hore dopamine agonists e kenyelletsa penile erection ka ho etsa hore li-neurone tsa oxytocinergic neurone li hlahise likarolo tse ling tsa bokopi le ho khetheha mokokotlo oa mokokotlo (bona Argiolas le Melis, 1995, 2004, 2005). Tumellanong le khopolo ena, linthong tse sa tsitsitseng, ho thibela li-lumbo-sacral oxytocinergic receptors ke motho ea sa amoheleng li-peptide oxytocin receptor mohanyetsi o sa tsoa fumanoa a khonne ho felisa tsoalo ea apomorphine e bakoang ke khatello ea intracavernous e bakoang ke dopamine agonist apomorphine, ho fana ka bopaki ba paraventriculo-spinal oxytocinergic tsela e amehang penile erection (Baskerville et al., 2009).

Lithuto tse ikemiselitseng ho ho khetholla dopamine receptor e ikarabellang bakeng sa ho kenngoa ha penile erection, ho bontšitse hore hape le li-agonists tsa nucleus dopamine receptor tse kentsoeng ka tlhaho li etsa hore penile erection e sebetse ka li-receptor tsa dopamine tsa lelapa la D2, e fumanoang ka li-agonists tse amoheloang ke dopamine tse fanoeng ka mokhoa oa mokhoa (sheba Melis et al., 1987; Eaton et al., 1991; Melis le Argiolas, 1995a). Ka lebaka leo, apomorphine, e nang le matla a tsoakiloeng a D1 / D2 receptor agonist, le quinpirole, ea khethiloeng ka ho fetisisa oa li-agonist, empa e seng SKF 2, e khethiloeng ke D38393 receptor agonist, e kenngoeng ka nucleus ena e fumanoe e khona ho etsa hore penile erection ka mokhoa o itšetlehileng ka tekanyo , 'me karabo ea thobalano e susumelitsoeng ke li-agonists tsena tsa D1 e amoheloa ke bahanyetsi ba D2, joaloka haloperidol le l-sulpiride, empa eseng ka SCH 2, mohanyetsi ea khethileng oa D23390 (Melis et al., 1). Bokhoni ba apomorphine ho etsa hore penile erection e kenngoe ka har'a kakaretso e hlahelletseng e boetse e netefalitsoe ke lipatlisiso tsa telemetry tse bontšang hore dopamine agonist e fanoeng kahare e ka holimo e khona ho eketsa khatello ea intracavernous ho tsoha litoeba tsa banna ntle le ho fetola khatello ea mali ea systemic (Chen et al. , 1987; Giuliano le Allard, 1999), e fumanehang ka mor'a hore e hlajoe ke systemic (Bernabè et al., 2001). Lithuto tsena hape e netefalitse karolo e ka sehloohong ea li-receptor tsa D2, kahobane batho ba amohelang li-agonist ba D1 ba atisa ho fumanoa ba sitoa ho eketsa khatello ea intracavernous ha ba kenngoa mokokotlong oa pontšo (Chen et al., 1999).

Ho ne ho e-na le mela e 'maloa ea bopaki ba liteko e neng e fana ka maikutlo a hore li-receptor tse nang le D2, tse nang le tšusumetso e matlafatsang penile erection, li teng liphatleng tsa li-oxytocinergic neurons. Ntlha ea pele, mokokotlo ona o na le methapo ea dopaminergic ea methapo ea motsoako e bitsoang incertohypothalamic dopaminergic neurons. Lihlopha tsa li-neurons tsena li ka har'a sehlopha sa A13 le A14 ea Dahlstrom le Fuxe (1964), e qaqisa mekhoa e mengata ea li-hypothalamic, e kenyelletsoang lipono tsa oxytocinergic neurons tse hlahisang li-brain lepophysis le / kapa libakeng tse ling tsa bokopi (Buijs et al., 1984; Lindvall et al., 1984).

Ho ameha ha li-neuron tsena tsa dopaminergic ka tekanyo ea sekhahla ho laola penile erection le ho kopana ho tšehetsoa ke lithuto tsa microdialysis tse bontšang hore likhahla tsa extracellular dopamine le 3,4-dihydroxyphenylacetic acid (DOPAC), metabolite ea eona e ka sehloohong, li eketseha ka dialysate e fumanoeng mokokotlo oa litoeba tse motona ka ho kopanela liphate tse bontšang ho se lumellane ha ho kenngoa ka pel'a bothata ba esrogen + ea progesterone-primed e amohelang basali (Melis et al., 2003).

Ho eketseha ha likhahla tsa dopamine le tsa DOPAC li bile li phahame haholoanyane ha ho kopana le mosali ea amohelang ho lumelloa (Melis et al., 2003), e fumanoang sebakeng se ka pele ho moo (Hull et al., 1995) le nucleus accumbens (Pfaus le Everitt, 1995). Ntlha ea bobeli, lipatlisiso tse 'maloa li bontša hore penile erection e susumelitsoe ke ho tsosoa ha li-receptor tsa D2, e kopanngoa ke oxytocin e lokolloa libakeng tsena. Ka lebaka leo, apomorphine e fanoang ka tekanyo e etsang hore penile erection e fumanehe, e fumanoe e khone ho eketsa mohopolo oa oxytocin, eseng feela ka lero la mali la likhoto le litšoene (Melis et al., 1989a; Cameron et al., 1992), empa hape le boko bo bong bo feteletseng libaka tse kang hippocampus (Melis et al., 1990). Tumellanong le liphello tsena, apomorphine e kenngoa mokokotlong o ka holimo ho mohato o etsang hore penile erection e sa tsoa bontšoa e khone ho eketsa mohopolo oa dopamine o eketsehileng oa cellular nucleus accumbens, e leng phello e fokotsoang ke mohanyetsi oa oxytocin d (CH2) 5Tyr (Me) 2-Orn8-vasotocin e kenngoa sebakeng sa ventral tekano (Succu et al., 2007; Melis et al., 2009a) (bona le Karolo ea 4). Ntlha ea boraro, lisele tsa motlakase tsa motlakase oa motsoako oa bokaholimo, o batlang o felisa oxytocin ka ho feletseng libakeng tsa boko bo fetisisang (Hawthorn et al., 1985), o felisa penile erection (Argiolas et al., 1987a) ea apomorphine, le bahanyetsi ba oxytocin ho fanoa ka li-ventricles, empa eseng ka har'a mokokotlo oa ponele, ho fokotsa penile erection ka mokhoa o itšetlehileng ka tekanyo ea tekanyo ea potoloho e nang le potency e tšoanang le ea metsoako ena e thibelang oxytocin receptors (Melis et al., 1989b). Li-antagonist tsa Oxytocin li boetse li matla haholo ho fokotsa ts'ebeliso ea boitšoaro bo bobe ba ho kopanela liphate, eseng feela ka oxytocin, empa le ka apomorphine (Argiolas et al., 1988, 1989).

Mabapi le mechine eo li-receptor tsa D2 tse sebelisoang ke dopamine kapa li-agonists tsa dopamine, li eketsa mosebetsi oa li-neurone tsa oxytocineric, kahoo li lokolla oxytocin libakeng tsa bokooa tsa extrahypothalamic le mokokotlo oa lisebelisoa tsa sesebelisoaho tšehetsa khopolo-taba ea hore ho susumetsoa ke li-receptor tsa D2 ho eketsa mohopolo oa li-ionine tsa Ca2 + tse entracellular ka har'a lisele tsa oxytocinergic neurons, e leng se etsang hore ho sebelisoe nitric oxide-synthase, e leng CaxNUMX + -czydulin e itšetlehileng ka enzyme, e teng lihlopheng tsena (Vincent le Kimura, 2; Torres et al., 1992; Sanchez et al., 1993; Sato-Suzuki et al., 1994). Ho eketseha ha nitric oxide tlhahiso ho etsa hore ho sebelisoe li-neuron tsa oxytocineric. Ka lebaka leo, (1998) apomorphineinduced erection erection e thibetsoe ke li-calcium-channel blockers le ka -_-conotoxin GVIA, e leng thibelo e matla le e khethiloeng ea mabone a Ca1 + a N-type (McCleskey et al., 2), e fanoeng ka mokhoa o hlahelletseng nucleus (bona Argiolas et al., 1987, le litemana tsa eona); (1990) penile erection e entsoeng ke apomorphine e ne e thibeloa ke nitric oxide-synthase inhibitors tse fanoeng ka lekaneng le lekaneng (Melis et al., 2c); le (1994) apomorphine le li-agonists tse ling tsa D3 tse amoheloang ka litekanyetso tse etsang hore penile erection e eketsehe ka nitric oxide tlhahiso ka dialysate ea tlhaho e fumanoang ke microdialysis ea intra-cerebral, keketseho e fokotsoang ke li-inhibitor tsa paraventricular nitric oxide-synthase e fokolang meriana e fokolang D2 e amohelehang ka penile erection (Melis et al., 2). Mechine e sebelisoang ke nitric oxide e hlahisang li-oxytocinergic neurons, e ntse e sa tsejoe, le hoja boitsebiso bo fumanehang bo bontša hore nitric oxide e sebetsa e le lenģosa le se nang palo le hore guanylate cyclase ha e amehe. Ha e le hantle, sebōpeho se sebetsang sa phosphodiesteraseresistant ea cyclic GMP, 1996-bromo-cyclic GMP, se fumanoe se sitoa ho etsa hore penile erection e fanoe ka nucleus e sa bonahaleng (Fig. 8) (bona Melis le Argiolas, 2b le litemana tsa eona).

Tlhaloso e ka holimo e atisa ho nkoa e sa khotsofatse, haholo-holo hobane ts'ebetso ea dopamine D2 receptors hangata e kopanngoa le ho thibela ho e-na le ho tsosoa ha lihlopha tsa li-neurons tse nang le li-receptor tsena ka mefuta e fapaneng ea liprotheine tsa G (sheba Sokoloff le Schwartz, 1995). Leha ho le joalo, moelelo o ka khonehang oa khethollo ena, e lumellanang le ho tsosolosoa ka ho totobetseng ha li-oktootermic neurons ka dopamine, ho 'nile ha boleloa hore haufinyane ho fumanoa ha G protheine-e kopantseng dopamine D4 mofuthu, setho sa lelapa la amohelang D2 (D2, D3 le D4), ts'usumetso ea eona e eketsang tšusumetso ea Ca2 + ka litokisetso tsa seleng tse nang le phetolelo e hlakileng ea mokamo ona (Moreland et al., 2004). Habohlokoa le ho feta, D4 receptor agonist (mohlala, ABT 724) (N-methyl-4- (2-cyanophenyl) piperazynil-3methylbenzamide maleate) e fumanoe e khone ho etsa hore penile erection ka likhoto tsa banna ha e fuoa mokhoa (Brioni et al., 2004). Phello ena ha e fumanoe ka khetho ea D2 receptor subtype agonist PNU-95666E (R-5,6-dihydro-N, N-dimethyl-4Himidazo [4,5,1-i] quinolin-5-amine) (Hsieh et al., 2004), e leng o ne a boetse a sa khone ho eketsa tšusumetso ea Ca2 + seleng ea litokisetso tse nang le phetolelo e hlalositsoeng ea D4 receptor subtype (Brioni et al., 2004; Moreland et al., 2004). Tumellanong le khopolo-taba e ka holimo le lipatlisiso, PD 168,077 (N-methyl-4- (2-cyanophenyl) piperazynil-3methylbenzamide maleate), PIP-3EA (2- [4- (2-methoxyphenyl) piperazin-1-ylmethyl] imidazo [1,2-a] pyridine) le tse ling tse khethiloeng ke li-agonists tsa D4 (Heier et al., 1997; Melis et al., 2006b; Löber et al., 2009), li fumanoe li khone ho etsa hore penile erection e sebelisoe ka tsela e nepahetseng, mokokotlo oa bokaholimo, le hoja o sa sebetse hantle ho feta apomorphine. Phello lan-erectile tsena D4 receptor agonists ile thibeloa ke L-745,870 (3- (4- [chlorophenyl] piperazin-1-yl) -methyl-1H-pyrrolo [2,3-B] pyridine trihydrochloride), le khetha D4 receptor mohanyetsi (Patel et al., 1997; Melis et al., 2005, 2006b; Löber et al., 2009).

Qetellong, phello lan-erectile ho tse ka holimo D4 receptor agonists ne a boetse a fokotswa ke nitric inhibitors oxide e-trehalohydrolase, nehelanoa ka paraventricular khubu, 'me ke le d (CH2) 5Tyr (Me) 2-Orn8-vasotocin, ka khetha oxytocin receptor antagonist e fanoe ka icv empa eseng mokokotlong o ka holimo. Liphello tsena li lumellana le khopolo ea hore li-agonists tsa D4 li boetse li susumetsa li-neurone tsa oxytocinergic ka ho sebelisa nitric oxide-synthase, le ho lokolla oxytocin libakeng tsa bokooa tsa extrahypothalamic, e leng tsona tse thusang penile erection, joalokaha ho bontšitsoe bakeng sa apomorphine le li-agonists tsa khale tsa D2 (Melis et al ., 2005, 2006b; Löber et al., 2009).

Liphuputso tse boletsoeng ka holimo li tšehetsa khopolo ea hore dopamine e etsa hore penile erection e sebetsane le li-receptor tsa D4 tse ka lihlopha tsa lisele tsa oxytocinergic neurons, e leng se bakang ho eketseha ha Ca2 + ho lihlopha tsa lisele tsa oxytocinergic neurons, e leng se lebisang ho hlahisoang ke oxide e ngata. Nitric oxide le eona e sebelisa oxytocinergic neurons ho lokolla oxytocin libakeng tse ling tsa boko le tse ling tsa mokokotlo, joalokaha se se se tšohloa. Ntlheng ena, hoa hlokomeleha hore li-receptor tsa dopamine li 'nile tsa fumanoa liphatleng tsa lisele tsa oxytocineric neurons mokokotlong oa phallo haufinyane tjena ka lithuto tse peli tsa immuno-fluorescence tse nang le li-D2, D3 le D4 tse amohelang likokoana-hloko le li-antibodies. Liphuputso tsena li bontšitse polelo ea mefuta eohle e meraro ea D2receptor subtypes (D2, D3 le D4), e kopantsoeng ka thōko ho lihlopha tsa lisele tsa oxytocinergic neurons (ka hare ho nucleus) (hape le sebakeng sa supraoptic le sebaka se ka pele ho moo) (Baskerville le Douglas, 2008; Baskerville et al., 2009).

Sena se fana ka ts'ehetso e matla ea phekolo ea meriana e le hore monyetla oa hore li-agonists tsa dopamine le dopamine tse amohelang mofuthu oa mofuta oa D2 li hlahise penile erection ka ho etsa hore li-neurone tsa oxytocinergic li hlahelle ka holimo li hlahisitse libaka tsa bokopi tsa extrahypothalamic tse boletsoeng ka holimo, mohlala, mokokotlo oa mokokotlo, sebaka sa ventral tegmental, hippocampus le amygdala. Leha ho le joalo, liphuputso tsena ha li fane ka thuso leha e le efe bakeng sa ho khetholla litlhaloso tsa D2 receptor / s, tseo ho susumelletseha ho baka karabo ea erectile. Ka bomalimabe, ha ho na thuso e ka fumanoang esita le ho lithuto tse reretsoeng ho khetholla li-oxytocinergic neurons tse nkiloeng ke dopamine receptor agonists mokokotlong oa pontšo. Ha e le hantle, ho sa tsotellehe mosebetsi o fapaneng o fapaneng oa dopamine receptor subtypes, ebang ke li-agonists tse tsoakiloeng tsa dopamine (mohlala, apomorphine), kapa li-agonists tse khethollang D2 (mohlala, quinpirole, e sebetsang ho bohle ba amohelang D2 receptor subtypes) phetoho ea oxytocinergic neurons, kaha e lekantsoe ke keketseho ea protheine ea FOS ka li-neurone tsa parvocellular oxytocinergic tsa khalase ea libaka (Bitner et al., 4). Leha ho le joalo, sena se fumanoe morao tjena, kaha protheine ea FOS e eketseha ka poneletiki e-oxytocinergic neurons e fumanoe feela ha penile erection e bakoa ke quinerolane, e sebetsang haholo ka D2006 le D2 receptor subtypes, empa eseng ka PD 3, e leng D168077 receptor agonist, ho sa tsotellehe bokhoni ba lik'hemik'hale tse peli ho etsa hore motho a arabe ka thobalano (Baskerville et al., 4).

Liteko tse ling tse nang le li-agonists tse khethollang tse ling tsa D2 receptor subtypes (haholo-holo D2 le D3) joale li hlokeha ho tseba karolo e tobileng ea dopamine receptor subtype e laolang mosebetsi oa erectile moemong o hlahelletseng. Tabeng ena, joalokaha ho se ho hopoloa pele, apomorphine, e sebetsang ka matla holim'a dopamine receptor subtypes (sheba Brioni et al., 2004, le litemana tsa eona), e sebetsa ho feta D4 receptors agonists ho fokotsa penile erection ha e kenngoa ka ntle nucleus. Sena se ka 'na ho hlalosa ke amana se hodimo sa apomorphine ka D4 receptors ha ho bapisoa le hore ba e lekiloeng D4 receptor agonists, kapa ho seng jwalo, D4 receptor agonists e ka sebetsa jwalo ka D4 receptor agonists leeme, kapa Tshebediso ya pataganeng ea fapaneng subtypes dopamine receptor ke apomorphine ka ho hlahisa phetoho e phahameng ea li-neuron tsa oxytocinergic tse kopanelang ho emisa penile, ho feta ts'ebetso ea D2 ea ho amohela li-agonists ea D4 receptor subtype feela.

Likamano pakeng tsa dopamine D1 le li-receptor tsa D2 li se li hlalositsoe ho laola boitšoaro ba thobalano sebakeng sa sebaka se ka hare sa sebaka sa pele (sheba Hull et al., 1989). Tabeng ea hore ho se khone ho khetha batho ba nang le bothata ba D2 ba nang le mekhoa e metle ho etsa hore penile erection (Hsieh et al., 2004) e tiisetsoe (empa bona Depoortère et al., 2009), ka mohlala esita le ka mor'a hore e entsoe ka lik'hemik'hale tse ka hare ho mokokotlo , karolo e kholo bakeng sa li-receptors tsa D3 feela kapa hammoho le ea li-receptors tsa D4 ha ho sebelisoa li-neurone tsa oxytocinergic ho phehella likhahla tsa penile li lokela ho hlahlojoa ka ho qaqileng (sheba Baskerville et al., 2009). Ka bomalimabe, batho ba khethiloeng ke D2 le D3 receptor agonists (mohlala, e fapaneng ka kamano ea bona bakeng sa tsena tse peli tsa receptor subtypes bakeng sa bonyane litaelo tse 'nè kapa tse hlano tsa in vitro) ha li fumanehe hona joale. Ka lebaka lena, tlhahiso ea morao tjena ea hore batho ba amohelang D3 ba lumellane le penile erection e susumelitsoeng ke li-agonists tsa khale tsa D2, tse thehiloeng haholo-holo bokhoni ba bahanyetsi ba D3 ba amohelang litlhahlobo tse hlalositsoeng ka in vitro, ho fokotsa penile erection e bakoang ke li-agonists tsa khale tsa D2, tse kang apomorphine, quinpirole le pramipexole, e sebelisang ka matla dopamine D2 receptor subtypes (Collins et al., 2009), ka sebele e hloka ho netefatsoa ka liteko tse ling. Ho netefatsa sena hoa hlokahala hape hobane ha ho na ts'ebetso ea D4 ea ho amohela li-agonist ka penile erection e fumanoeng thutong ena, ka phapang e tsotehang le liphello tsa lithuto tse boletsoeng ka holimo, e leng se bontšang phello ea pro-erectile ea li-agonists tsa li-D4. Ha e le hantle, esita le bokhoni ba apomorphine bo ka etsa hore penile erection (le ho hlajoa) e be le litoeba tsa ho kokota ho D4 tse nang le potency e tšoanang le e bonngoeng ke litoeba tsa ho amohela likhomo tsa D4 le bokhoni ba bahanyetsi ba D3 ba ho felisa karabo ea apomorphine liphoofolong tsena ( Collins et al., 2009) ha e khone ho nkoa e le bopaki bo totobetseng ba karolo e khethollang ea D3 receptor subtype ho tswa ho penile erection ea D2 e amoheloang ke agonist. Mefuta e se tšoane ntle, lithuto tsa ka neurotransmitter / neuropeptide le / kapa neurotransmitter / neuropeptide receptor phatsa ea lefutso ablation (neurotransmitter / neuropeptide le / kapa neurotransmitter / neuropeptide receptor knockout liphoofolo tse) di hangata eketsa pherekano ka ho eketsehileng le mathata a ka netefatso ya putative karolo ea thobalano ea neurotransmitters le / kapa li-neuropeptides le li-receptors tsa tsona. Ka tsela e joalo, oktocin ea liphatsa tsa lefutso e hlahisa oxytocin ho tsoa litoeba tseo molekane le ho li phethang ka tloaelo, joalokaha eka oxytocin e ne e sa hlokahale bakeng sa ho nyalana le ho kopana. Hape basali ba homozygous oxytocin knockout mice show e tloaelehileng ea ho kopanya le ho arolelana likarolo, le hoja ho e-na le ho holofala ha lebese (Nishimori et al., 1996; Young et al., 1996). Ho tlisoa ha liphatsa tsa lefutso tse kenyang neuronal nitric oxide synthase, hape ho hlahisa nitric oxide synthase ho tsuba litoeba tseo molekane ebe li li phethang ka tloaelo (Huang et al., 1993). Leha ho le joalo, liphuputso tsena li ka 'na tsa bontša tšobotsi ea bohlokoa ea thuto ea ho ikatisa, e leng, ho hlophisoa ha litsamaiso tse amanang le taolo ea eona bohareng le boemong ba palo.

Tlhaloso e joalo e na le tšimoloho ea ho iphetola ha lintho, kaha e fana ka tiiso ea hore liphatsa tsa lefutso li tla hlaha ho moloko o latelang bakeng sa ho phela ha mofuta ona. Ka hona, taba ea hore phekolo ea liphatsa tsa lefutso ea D4 ha e fetole phello ea pro-erectile ea apomorphine, e fana ka maikutlo a hore li-receptor tsa D4, tse kang oxytocin le nitric oxide, ke baemeli ba 'maloa feela ba sebetsang tsamaisong e laolang erectile, ho e-na le ho fana ka maikutlo a hore ha ho na karolo bakeng sa li-receptors tsena tse laolang penile erection le boitšoaro ba thobalano. Ho hlōleha ha li-agonists tsa D4 ho etsa hore penile erection ha e fanoe ka mokhoa o tloaelehileng ho likhoto tsa banna tsa mefuta e sa tšoaneng e sa tsoa tlalehoa ke thuto e 'ngoe (Depoortère et al., 2009). Leha ho le joalo, ho fapana le mosebetsi oa Collins et al. (2009), 'me ho etsa setšoantšo setšoantšong sa mefuta e fapaneng ea li-D2 receptor subtypes tse laolang penile erection le ho feta pherekano, tlhatlhobo ena e boetse e bontša hore setative e khethollang bahanyetsi ba D3 ba fanoang ka mokhoa o hlophisitsoeng ba ne ba sitoa ho fokotsa ho emisoa ha penile ka apomorphine li-rats tsa bothata li utloisisa phello ea pro-erectile ea apomorphine, ha ho itšoara ka thobalano (le ho tsoma) ho ne ho hanyelitsoe ke mohanyetsi ea khethileng oa D2 L-741,626 (3 - [[4- (4-chlorophenyl) -4-hydroxypiperidin -1-yl] methyl-1H-indole), e leng ho etella bangoli ho fana ka maikutlo a hore li-receptors tsa D2, ho e-na le li-receptors tsa D3 le D4, ke tsona tse bapalang karolo e kholo penile erection ea D2 e entsoeng ke agonist (Depoortère et al., 2009). Qetellong, ho na le monyetla oa hore liphetoho tse entsoeng ke dopamine tse amohelang likokoana-hloko tsa oxytocinergic neurons tse lumellanang le penile erection, bonyane likarolong tse itseng, li sa tobahale ho e-na le ho toba, mohlala, ho buuoa kapa ho susumelitsoe ke liphetoho tse ling tsa li-neurotransmitters neuropeptides tse khonang ho fetola mosebetsi oa oxytocinergic neurons mokokotlong oa seretse, o ke ke oa felisoa ka ho feletseng.

3.2. Phetolelo ea Glutamic acid-oxytocin mokokotlong oa sepakapaka

Motsoako oa phallo ea hypothalamus o na le li-synapses tse nang le monate oa amino acid joaloka motsoako oa li-neurotransmitter (mohlala, glutamic acid le aspartic acid) (Van Den Pol, 1991). Li-amino acid tse thahasellisang ka nucleus ena li kenyeletsa mesebetsing e mengata, ho kopanyelletsa le penile erection le boitšoaro ba thobalano (Roeling et al., 1991; Melis et al., 1994b, 2000, 2004b). Ka lebaka leo, N-methyl-d-aspartic acid (NMDA), e leng mohloli oa li-agonist oa NMDA receptor subtype, empa eseng (±) -_- amino-3-hydroxy-5-methyl-isoxazole-4-propionic acid (AMPA) , motsoako oa li-agonist oa AMPA receptor subtype kapa (±) -trans (1) -amino-1,3-cyclopentane dicarboxilic acid (ACPD), e leng agonist e khethollang ea metabotropic receptor subtype, e fumanoe e khona ho etsa hore penile erection e kenngoe ka har'a mokokotlo oa likokoanyana tse lokolohileng (Melis et al., 1994b). Phello ea pro-erectile ea 948 MR Melis, A. Argiolas / Neuroscience le Litlhahlobo tsa Boipheliso 35 (2011) 939-955 NMDA e thibetsoe ke (+) MK-801, mohanyetsi oa NMDA ea sa amoheleng tlhōlisano (Woodruff et al., 1987 ), e kenngoa mokokotlong oa limela (Melis et al., 1994b). Tumellanong le liphetho tsena, lipuisong tsa telemetry tse lebisitseng ho hlahloba khatello ea intracavernous, NMDA e fumanoe e le mafolofolo haholo ho feta ba agonists ba e meng ea monate ea amino acid receptor subtypes ha ba kenngoa mokokotlong oa phallo ka ho eketseha khatello ea intracavernosal ho tsoha kapa ho haelloa ke likhoho tsa banna (Zahran et al., 2000; Chen le Chang, 2003).

Joalokaha ho hlalositsoe ka holimo bakeng sa oxytocin le dopamine, ho ka etsahala hore li-receptors tsa NMDA tse tsamaisang penile erection li ka har'a lihlopha tsa li-oxytocinergic neurons, kaha litlhaloso tsa meriana ea amino acid e qetella e e-na le lihlopha tsa oxytocinergic sekokotlong (Van Den Pol, 1991). Ka papiso seo fumanoa le agonists dopamine receptor, phello lan-erectile tsa NMDA e Kamoo ho bonahalang kateng go ranolelwang kwa ke Tshebediso ya ya neurotransmission oxytocinergic, ho felisa ka ho feletseng ke oxytocin antagonist d '(CH2) 5Tyr (Me) 2-Orn8-vasotocin icv fanoeng, empa eseng mokokotlong oa sepakapaka (bona Argiolas le Melis, 1995, 2004, 2005 le litemana tsa eona). Ka mokhoa o ts'oanang, ts'ebetso ea NMDA e susumelitsoeng ke neurotransmission e boetse e le ea bohlokoa ho sebelisoa ha nitric oxide-synthase, kaha NMDA-e bakoang ke penile erection e thibeloa ke nitric oxide-synthase inhibitors (N-Nitro-N-methyll-arginine methyl ester le N- methyl-thio-l-citrulline) e fanoeng ka kakaretso, 'me NMDA e kenngoa mokokotlong oa mahlaseli a etsang hore penile erection e eketsehe, e eketsa tlhahiso ea nitric oxide mokokoting oa hypothalamic (bona Argiolas le Melis, 1995, 2004, 2005 le litemana tsa eona ). Ha e le li-agonists tsa dopamine, li-NMDA li entse hore ho be le tšebeliso ea nitric oxidesynthase e ka boela ea e-ba bobeli bakeng sa ho eketseha ha Ca2 + meleng ea oxytocineric cell body ka li-receptor tsa NMDA tse nang le khanya ea Ca2 +, joalokaha ho bontšoa litokisetsong tse 'maloa tsa tlhaho (bakeng sa tlhahlobo bona Snyder, 1992; Southam le Garthwaite, 1993; Schuman le Madison, 1994 le litemana tsa eona). Nitric oxide le eona e sebelisa oxytocinergic neurotransmission (bona ka holimo). Ha ho tsejoe hore na ho na le bopaki ba mofuta ofe oa li-glutamatergic, tse etsang hore li-neuronic neurons tse hlahisang penile erection li se ke tsa tsejoa, le hoja bopaki bo itseng ba kankere ea motlakase le ea electrophysiological bo bontša hore e ka 'na ea e-ba karolo ea karolo ea hippocampus (Saphier le Feldman, 1987; Chen et al., 1992) . Le hoja ho hlokahala mosebetsi o eketsehileng ho bontša hore na ho na le tšimoloho efe ea glutamatergic to the core core, ho kenella ha acutamic e ka sehloohong e ka tlas'a taolo ea penile erection le boitšoaro ba thobalano ho tšehetsoa ka ho hlaka ke lithuto tsa microdialysis. Ka lebaka leo, libaka tse eketsehileng tsa lisele tsa glutamic acid le aspartic acid li ile tsa eketseha ka dialysate e fumanoang mokokotlong oa litoeba tsa banna tse bontšang ho se lumellane ha li kenngoa ka pel'a likokoanyana tse ling tse amohelang li-estrogen + tsa progesterone-primed (Melis et al., 2004b), li-penile erections tse kenyelletsanoang le ho sebelisoa ha phetisetso ea motsoako oa oxytocineric (Melis et al., 1999a, b). Keketseho e joalo e ile ea fumanoa e phahame ha ho kopana le mosali ea amohelang ho lumelloa (Melis et al., 2004a). Tumellanong le khopolo-taba ea hore mosebetsi o ntseng o eketseha oa amino acid e tsoetseng pele o hlaha mokokotlong oa penile nakong ea penile erection le copulation, bobeli ba ho se lumellane le boitšoaro ba boipheliso (nakong eo ka eona likhetho tsa penile li etsahalang) li fokotsoe ke ho thibeloa ha ba amohelang NMDA mokokotlong , 'me khaello ena e lateloa ke keketseho ea tlhahiso ea nitric oxide tlhahiso e hlahang mokokoting ona oa hypothalamic maemo ana a tlhaho (Melis et al., 2000). Ho eketseha ha extracellarlar glutamic acid mahloriso ho feta ho fokotseha GABA ho tsoa GABAergic methapo ea phekolo phehello ho susumelletsa ka excitatory synapses synapses e feteletseng ho oxytocinergic lihlopha tsa 'mele, o ile a boela a fumanoa ka kakaretso nucleus ka mor'a ho thibeloa ha cannabinoid CB1 receptors ka CB1 mohanyetsi SR 141761A, ho fanoa ka li-ventricles lateral kapa ka ho toba mokokotlong oa li-dose tse etsang hore penile erection e be teng (sheba Succu et al., 2006; Castelli et al., 2007). Khōlo e joalo e ile ea etsa hore ho sebelisoe nitric oxide-synthase liphatleng tsa li-oxytocinergic neurons, ho eketsa tlhahiso ea nitric oxide. Nitric oxide le eona e sebelisa oxytocinergic neurons ho phelisana le penile erection joalokaha ho hlalositsoe ka holimo. Tumellanong le mechine e joalo, SR XUMUMA e ile ea etsa hore penile erection e fokotsehe ka ho thibeloa ha li-receptors tsa NMDA le ka nitric oxide-synthase inhibitors, empa eseng ka thibelo ea dopamine kapa oxytocin receptors mokokotlong oa phallo, ha e thibeloa ke thibelo ea bohareng ba oxytocin receptors ke bahanyetsi ba oxytocin ba fumanoang icv

3.3. Oxytocin-dopamine puisano sebakeng sa ventral tegmental

Oxytocin e kenyelletsa penile erection ha e kenngoa karolong ea caudal ea sebaka sa ventral ho latela mokhoa o itšetlehileng ka tekanyo ea methapo (Melis et al., 2007). Leihlo le tlaase ka ho fetisisa le sebelisitsoeng ka lehlakoreng le le leng e ne e le 20 Ng, ha tekanyo e phahameng ka ho fetisisa e lekoa e ne e le 100 ng. Phello ea oxytocin e tsamaisana le ho sebelisoa ha li-receptor tsa oxytocin, kaha karabo ea thobalano e felisoa ke lejoe le pele la mohanyetsi oa oxytocin d (CH2) 5Tyr (Me) 2-Orn8-vasotocin sebakeng sa ventral tegmental. Li-receptor tsena li fumaneha libakeng tsa lisele tsa dopaminergic neurons, e leng morero o ka sehloohong ho shell ea nucleus accumbens. Ka hona, (1) lipatlisiso tse peli tsa immuno-fluorescence li bontša hore li-ventral tegmental sebakeng sa oxytocinergic fibers li amana haufi le lihlopha tsa dopaminergic neurons, tseo tse ngata tsa tsona li neng li ngotsoe ka tyrosine-hydroxylase 'me li na le retrograde tracer Fluorogold pele ho kenngoa ka har'a khetla ea nucleus accumbens (Melis et al., 2007), le (2) sebakeng se ka tlaase ho pentele erection e entsoeng ke oxytocin e hlaha ka mokhoa o khotsofatsang ka ho eketseha ha dopamine e eketsehileng-cellular ho dialysate e fumanoang sekhethong sa nucleus accumbens (Melis et al., 2007). Penile erection e entsoeng ke oxytocin e boetse e hlaha ka mokhoa o khotsofatsang ka ho eketseha ha nitric oxide tlhahiso sebakeng sa ventral tegmental, e leng likarabo tse peli tse sa sebetsanang le d (CH2) 5Tyr (Me) 2-Orn8-vasotocin le nitric oxide synthase inhibitor S- methyl-thio-l-citrulline, empa hape ke _-conotoxin, e leng CaxNUMX + channels blocker e itšetlehileng ka matla a motlakase, 'me ka ODQ (2H- [1] oxadiazole [1,2,4-a] quinoxalin-4,3-one), e leng inhibitor e matla ea guanylate cyclase, tsohle tse fanoeng sebakeng sa caural tegmental pele ho oxytocin (Succu et al., 1). Mokhoa o mongata oa Fluorogold o ngotsoeng ka dopaminergic lihlopha tse kopanetsoeng ke oxytocinergic fibers, e fumanoang e le ntle bakeng sa tyrosine hydroxylase sebakeng sa caural tegmental sa caudal, le tsona li ngotsoe ka katleho bakeng sa nitric oxide-synthase le guanylate cyclase (Succu et al., 2008), oxytocin -e nang le penile erection e ka sebelisoang ke mekhoa e latelang. Ts'ebetso ea oxytocinergic receptors meleng ea dopaminergic le peptide e eketsa khanya ea Ca2 + ka hare ho lihlopha tsa lisele tsa dopaminergic neurons. Sena se sebelisa nitric oxide-synthase, enzyme e itšetlehileng ka CaxNUMX + -calmodulin, kahoo e eketsa tlhahiso ea nitric oxide. Nitric oxide le eona e sebelisa guanylate cyclase, e leng se lebisang tlhokomelong e eketsehileng ea GMP ea maoto. Cyclic GMP e sebelisa li-neuron tse nang le dopaminergic tse hlahisang nucleus accumbens. Karolo ea phetoho ea GMP e hlahisoang ke penile e bakoang ke oxytocin e kenngoa sebakeng sa caural tegmental ea caudal e boetse e tšehetsoa ke bokhoni ba 8-bromo-cyclic GMP, e leng sebōpeho se sebetsang sa anti-phosphodiesterase ea cyclic GMP, ho etsa hore penile erection e kenngoe ka har'a sebaka sa caural tegmental, le ho eketsa dopamine e eketsehileng-cellular ho dialysate ho tloha nucleus accumbens (Succu et al., 2008; Melis et al., 2009a). Sena se fapane le mochine oo nitric oxide e etsang hore oxytocinergic neurons e sebetsane le nucleus, e leng 8-bromo-cyclic GMP e sitoang ho etsa hore penile erection e kenngoe ka nucleus (Melis le Argiolas, 1995b) (Fig. 2). Mabapi le litsela tsa neural tse sebelisoang ke dopamine nucleus accumbens e lebisang penile erection, tsena ha li tsejoe. Leha ho le joalo, boitsebiso bo fumanehang bo fana ka maikutlo a hore litsela tsena li kopa dopamine neurotransmission mokokotlong oa phallo ea hypothalamus. Ka lebaka leo, penile erection e susumelitsoeng ke oxytocin e hlaha ka mokhoa o khotsofatsang ho eketseha ha dopamine e eketsehileng ea cellular eseng feela ka nucleus accumbens, empa hape le mokokotlong oa holimo, 'me o hanyetsanoa ke mohanyetsi oa dopamine ea amohelang haloperidol a kenngoa ka lekaneng le nang le phallo (Melis et al., 2007). AHa e le hantle, liphello tsena li tšehetsa khopolo ea hore li-neurone tsa oxytocinergic tse hlahang mokokotlong oa li-ventral le ho hlahisa sebaka sa ventral tegmental, ha ho tlosoa oxytocin sebakeng sena, ka hona ho etsa hore ho be le mokhoa oa ho paka oa GMP oa NO-cyclic, e leng eona e kenyang mesolimbic dopaminergic neurons (Melis et al., 2007, 2009a; Succu et al., 2008). Dopamine e lokolloa nucleus accumbens e boetse e etsa hore ho be le litsela tsa neural tse etsang hore li-incerto-hypothalamic dopaminergic neurons li sebetse, e leng se etsang hore li-neurone tse nang le li-oktocinergic li se ke tsa hlahisa mokokotlo oa mokokotlo oa mokokotlo oa penile erection. Ka nako e tsoanang, dopamine e lokolloa mokokotlong o ka 'na oa etsa hore li-oxytocinergic neurons li hlahise likarolo tse ling tsa boko bo kang tsa ventral tepi, hippocampus, amygdala le mohlomong libaka tse ling tsa boko.

Joalokaha ho hopoleloe ka holimo, tumellanong le khopolo-taba ena, apomorphine e kenngoa mokokotlong o ka holimo ho lekaneng le etsang hore penile erection e kenye letsoho le ho eketsa mohopolo oa dopamine o eketsehileng nucleus accumbens, phello e fokotsoang ke mohanyetsi oa oxytocin d (CH2) 5Tyr (Me ) 2-Orn8-vasotocin e kenngoa sebakeng sa ventral tegmental (Succu et al., 2007; Melis et al., 2009a). Ka bobeli, litsela tse ka holimo tsa neural e ka 'na ea e-ba setsi sa potoloho se nang le mechine e nang le dopamine, oxytocin le li-neurotransmitter tse ling (mohlala, acid, glutamic acid, sheba ka tlaase). (sheba Karolo ea 4).

3.4. Ho sebelisana ha Oxytocin-glutamic acid ka subralulum ea ventral ea hippocampus

Oxytocin e kenyelletsa penile erection ha e kenngoa ka har'a ventral, empa eseng ka suborulum ea majoe, ka mokhoa o itšetlehileng ka tekano (Melis et al., 2009b). Tlhaloso ea thobalano e kopanngoa ke ho tsosoa ha li-oxytocin receptors, ho felisoa ke ente ea pele ea d (CH2) 5Tyr (Me) 2-Orn8-vasotocin e fanoeng sebakeng se tšoanang sa oxytocin, e fumanoang libakeng tse ling tsa boko (bona ka holimo) . Mabapi le ho fumana libaka tsena tse amohelehang, lintlha tse fumanehang li bontša hore li sebakeng sa lisele tsa methapo e nang le nitric oxide-synthase.

Ka lebaka leo, lipatlisiso tsa microdialysis li bontša hore penile erection e entsoeng ke oxytocin e hlaha ka mokhoa o khotsofatsang ka ho eketseha ha nitric oxide tlhahiso ka ventral subiculum, 'me keketseho ena e felisoa eseng feela ka ente ea pele ea nitric oxidesynthase inhibitor S-methyl-thio-l-citrulline le ka nitric oxide scavenger hemoglobin, empa hape le d (CH2) 5Tyr (Me) 2-Orn8-vasotocin, kaofela e fumanoang sebakeng se tšoanang sa oxytocin ka tekanyo e lekanang le penile erection (Melis et al., 2010). Habohlokoa le ho feta, penile erection e susumelitsoeng ke oxytocin e boetse e hlaha ka mokhoa o khotsofatsang ka ho eketseha ha mahlaseli a li-extra-cellular glutamic acid ka subralulum ea ventral, e hanyetsanoang feela ke mohanyetsi oa NMDA (+) MK-801 ea sa hlōliseng ventral subiculum (Melis et al., 2010). Ka kakaretso, liphello tsena li fana ka maikutlo a hore mokelikeli oa nitric o sa tsoa thehoa, ka ho sebetsa e le lenģosa le leng le le leng, o etsa hore motsoako oa glutamic neurotransmission o lebise penile erection, mohlomong ka sekhahla sa neural efferent ho tloha ventral subiculum ho libaka tsa boko bo eketsehileng ba hippocampal. Tumellanong le khopolo-taba ena, NMDA e kenngoa ka subiculum ea ventral e kenyelletsa penile erection ka mokhoa o itšetlehileng ka tekanyo ea methapo, 'me phello ena e hanyelitsoe ka ho feletseng ka lejoe le le leng pele ho (+) MK-801, empa eseng ka S-methyl-thio-l-citrulline , hemoglobin kapa d (CH2) 5Tyr (Me) 2-Orn8-vasotocin (Melis et al., 2010). Mabapi le mekhoa e metle ea li-neural e lebisang libakeng tse ling tsa bo-hippocampal tse entsoeng ke li-amino acid (e leng, glutamic acid) ka ventral subiculum, ho ka etsahala hore ebe tsena ke glutamatergic, joalo ka boholo ba li-hippocampal tse fapaneng. Ka motsotso ona, ho ka 'na ha fanoa ka maikutlo a hore lintlhoko tsena li fetola mosebetsi oa mesolimbic dopaminergic neurons, e leng tsona tse tsamaisang mosebetsi oa incerto-hypothalamic dopaminergic neurons mokokotlong oa paraventricular, e leng ho lebisang ho ts'ebetsong ea oxytocinergic neurons ho loantša ho koaloa ha penile joalokaha ho se ho ntse ho tšohloa (bona ka holimo).

Ka lebaka leo, penile erection e bakoang ke ventral subiculum oxytocin e hlaha ka mokhoa o khotsofatsang ka ho eketseha ha dopamine e eketsehileng-cellular sekheng sa nucleus accumbens, 'me keketseho ena, e kang penile erection, e felisoa ke d (CH2) 5Tyr (Me) 2- Orn8-vasotocin e fuoa subiculum ventral pele oxytocin (Melis et al., 2009b). Ho feta moo, kaha ventral subiculum oxytocin e kentse penile erection e boetse e fokotsoe ke (+) MK-801 e kenngoeng sebakeng sa ventral tegmental, empa e se ka nucleus accumbens (Melis et al., 2009b) 'me e hlaha ka mokhoa o khotsofatsang ho eketseha ha li-cellular limela tse peli tse felisoang ke d (CH2) 5Tyr (Me) -Orn8-vasotocin, e kenngoa ka ventral subiculum pele ho oxytocin (bona Fig. 3), ho ka etsahala hore ts'ebetsong ea mesolimbic dopaminergic neurons ke karolo ea bobeli ho ea ho eketseha ha pelo ho tsoa sebakeng sa ventral tegmental. Sena se fana ka maikutlo a hore ho sebelisana ha glutamic acid-dopamine ho laola penile erection ho sebakeng sa ventral tegmental. Liphuputso tse ling li hloka ho netefatsa hore na pro-erectile efferent glutametergic tsela e tsoang ho subiculum sebakeng sa ventral tegmental e tobileng kapa e sa tobang, ke hore, ka sekepe sa prefrontal kapa libakeng tse ling tsa boko (sheba Melis et al., 2009b le litemana tsa eona). Kaha mokokotlo oa seretse o boetse o fumana litekanyetso tsa glutamatergic ho tloha hippocampus (bona ka holimo le Saphier le Feldman, 1987), 'me glutamic acid e etsa hore ho be le li-neurone tsa oxytocinergic neurons tse kenyeletsang sebaka sa ventral tegmental (bona Argiolas le Melis, 2005 le litemana tsa eona), le oxytocin in the ventral tegmental sebakeng se etsa hore penile erection e eketsehe le ho eketsa mosebetsi oa mesolimbic dopaminergic neurons (bona ka hodimo), ho lekeha ho nahana hore paraventricular oxytocinergic neurons e ka 'na ea boela ea ameha, bonyane ka karolo e' ngoe, ha ho sebelisoa mesolimbic dopaminergic neurons ka oxytocin e kenngoa ka subralulum ea ventral (sheba Karolo ea 4).

4. Lipolelo tse qetellang

Lipatlisiso tse hlahlobetsoeng ka holimo li tiisa le ho atolosa liphuputso tsa pele tse bonts'ang hore linthong tsa banna tse oxytocin li phetha karolo ea bohlokoa ho laola bohareng ba penile erection ka lekaneng la mokokotlo oa phallo ea hypothalamus le ea mokokotlo oa mokokotlo. Ka ho khetheha, haholo ka ho fetisisaliphuputso tse lekholo li bontša hore oxytocin e susumetsa penile erection hape ka ho sebetsa likarolong tse ling tsa boko, e leng, sebaka sa ventral subiculum, le mokokotlo oa posteromedial of amygdala.

Sebakeng sa boemo bo phahameng, mohloli oa bohlokoa ka ho fetisisa ka ho fetisisa ke ho sibolloa ha polelo ea dopamine receptors ea lelapa la D2 (D2, D3 le D4) ka lihlopha tsa lisele tsa oxytocinergic neurons mokokotlong oa sepakapaka (le mokokotlong oa supraoptic le sebaka sa lipakeng tsa pele) (Baskerville le Douglas, 2008; Baskerville et al., 2009). Sena se fana ka methapo e matla ea kankere ho tšehetsa khopolo-taba ea hore li-agonists tse nang le dopamine le dopamine li ka etsa hore li-neurone tse sebetsang ka ho toba li sebetsane le erectile mosebetsing le hore li se ke tsa etsoa feela mokokotlong oa mokokotlo, empa hape le libakeng tse ling tsa boko bo botle ba hypothalamic. Tabeng ena, ke habohlokoa hape hore ho sibolloa hore ho eketseha ha li-intracavernous ho bakoang ke lefu la dopamine ho fokotsoe ke ho thibela li-receptor tsa oxytocinergic mokokotlong oa lesapo la sacrose (Baskerville et al., 2009). Ka sebele, le hoja bopaki bo joalo bo fumanoe linthong tsa banna tse sa tsitsitseng, ho fumanoa ho tiisa hore ho sebelisoa tsela e theohelang ea paraventriculo-spinal oxytocineric e kenyeletsang penile erection e entsoeng ke dopamine. HLeha ho le joalo, e ntse e lokela ho netefatsoa hore na penile erection e bakoang ke ho tsosoa ha methapo ea dopamine e fumanoang liphateng tsa oxytocinergic ke karolo ea bobeli ho ts'ebetso ea dopamine receptor subtype ea lelapa la D2 (D2, D3 kapa D4) kapa haeba li-receptor subtypes li sebelisana ho feto-fetola karabo ea erectile, mohlomong ka litsela tse fapaneng ho itšetlehile ka moelelo oo penile erection e leng teng (bona Moreland et al., 2004; Enguehard-Gueiffier et al., 2006; Melis et al., 2006a, b; Löber et al., 2009; Collins et al., 2009; Depoortère et al., 2009; Baskerville et al., 2009).

Ntho e 'ngoe ea bohlokoa e fumanoang ke hore oxytocin e etsa hore penile erection e kenngoe feela ka har'a mokokotlo oa mahlaseli kapa sebaka sa CA1 sa hippocampus, empa hape le sebakeng sa ventral tegmental, ventral subiculum le mokokotlo oa posteromedial cortical oa amygdala. Libaka tsena tsa boko ha lia ka tsa lekoa liphuputsong tse qalang tse bontšang hore oxytocin e eketsehile ka penile erection likarolo likokoanyana tsa banna, le hoja li amoheloa joaloka lisebelisoa tsa oxytocineric tsa lumbo-sacral mokokotlo ho tloha mokokotlong oa pontšo. Ha e le hantle oxytocin e fumanoe e khona ho eketsa likarolo tsa penile erection tse sa tloaelehang, tse etsahalang ho liphoofolo tse khōlō tsa banna ha ho se na ntho leha e le efe e ts'oanang ea thobalano, e kang e hlahang ho ba teng ho ratoang kapa ho sa khonehe ho fumana ratano ea basali kapa ho sebetsana le liphatsa tsa lefutso, ho kenngoa ka har'a mokokotlo oa mahlaseli le sebaka sa CA1 sa hippocampus, empa eseng ka suborulum ea mahlakoreng, sephahla sa lateral, sekoti se ka tlaase, sebaka se ka hare-pele, mokokotlo oa melumo le mokokotlo oa supraoptic (Melis et al. , 1986). Liphuputsong tsena tsohle penile erection e ne e atisa ho baloa ha penis e tsoa penile sheath ke mohlokomeli ea neng a sa tsebe mekhoa e fanoeng ka ho toba nakong ea teko kapa hamorao ka ho shebella teko e tlalehiloeng videoteng e nang le lisebelisoa tsa likhamera tsa video. E ngoe le e 'ngoe ea penile erection e nka 0.5-1 min mme hangata e tsamaisana le ho itlhoekisa ha penile le / kapa leqhoa la maqhubu. Ha ho liteko tse atisang ho etsoa lipotheng tsena ho netefatsa hore na boiphihlelo ba ho kopanela liphate ke bofe, lilemo kapa haeba litoeba li ka aroloa ka tlase kapa holimo ho arabela phello ea pro-erectile ea oxytocin e kenngoeng libakeng tse fapaneng tsa boko. Hona ho sebetsa hape le bongata ba lithuto tse mabapi le phello ea pro-erectile ea li-neuropeptides tse ling le lithethefatsi tse eketsang hangata penile erection lipapali, ho kenyeletsa dopamine agonists, li-amino acid tse thahasellisang, ACTH-MSH, hexarelin le li-peptide tsa VGF. Leha ho le joalo, phello ea pro-erectile ea lik'hemik'hale e 'nile ea tiisoa ka makhetlo ka mekhoa ea telemetry, e leng se khethollang ho hlaha ha penile erection ka keketseho ea khatello ea intracavernous e hlahelang ka boomo kapa ka mor'a ho tsamaisa metsoako ena ka litsela tse fapaneng, ke hore, tsamaiso ea tsamaiso, ho kena ka ho toba bokong ba nuclei, ka mor'a ho kenngoa ha khatello ea microtransducer ka ho toba ka har'a lik'hasepere (sheba Bernabè et al., 1999). Sebakeng sa ventral tegmental, subralulum ea ventral le mokokotlo oa posteromedial oa amygdala hape oxytocin e kenyelletsa penile erection ka ho etsa li-receptor oxytocinergic. Sena se lebisa ho ts'ebetsong ea mesolimbic dopaminergic neurons e hlahang sebakeng sa ventral tegmental le ho hlahella ho khetla ea nucleus accumbens, kaha li lekantsoe ke keketseho ea mahlaseli a dopamine a eketsehileng-cellular ho dialysate e fumanoang sekhethong sa nucleus accumbens le ka ho fokotsa karabo ea erectile e bakoang ke peptide e kenngoa libakeng tsena tse ling tse ngata, tse fumanoang ka mor'a ho thibela li-receptor tsa dopaminergic nucleus accumbens (sheba ka tlase). Mabapi le lits'ebetso tse susumelitsoeng ke ho susumetsoa ha li-receptor tsa oxytocineric libakeng tsena tsa boko, tse lebisang ho ts'ebetso ea mesolimbic dopaminergic neurons le ho penile erection, tse hlakileng ka ho fetisisa ke tse hlahang sebakeng sa ventral tegmental. Ha e le hantle, mona liphello tsa phekolo ea meriana le ea immuno-fluorescence li bontša hore oxytocin ea phekolo ea methapo e kenyelletsa lihlopha tsa lisele tsa dopaminergic neurons tse hlahisang khetla ea nucleus accumbens (Melis et al., 2007, 2009a; Succu et al., 2008). Bongata ba li-neurone tsena li na le nitric oxide synthase e ngata le guanylate cyclase. Ho tsosoa ha li-oxytocinergic receptors liphatleng tsa lisele tsa dopaminergic neurons tsena ho etsa hore ho sebelisoe nitric oxide synthase e lebisang tlhahiso e ngata ea nitric oxide. Nitric oxide le eona e sebelisa guanylate cyclase, kahoo e eketsa maqhubu a GMP, e leng ho lebisang ho ts'ebetso ea mesolimbic dopaminergic neurons le ho lokolloa ha dopamine nucleus accumbens, e lekantsoe ke keketseho ea dopamine e eketsehileng ea cellular ho dialysate ho tloha nucleus accumbens e fumanoang ke microdialysis ea intra-cerebral (Succu et al., 2008) ). Dopamine e lokolloa nucleus accumbens le eona e etsa hore ho be le litsela tsa neural tse lebisang penile erection. Sena se tšehetsoa ke bokhoni ba bahanyetsi ba dopamine ba amohelehang haloperidol le / kapa cis-flupentixol e kenngoang ka nucleus accumbens ho fokotsa phallo ea venteni e entsoeng ka oxytocin (Succu et al., 2008). E 'ngoe ea litsela tsa pro-erectile e bonahala e etsa hore li-incerto-hypothalamic dopaminergic neurons, haholo-holo tse hlahisoang ke lihlopha tsa lisele tsa oxytocinergic neurons. Ha e le hantle oxytocin e kenngoa sebakeng sa caural tegmental ea caudal e eketsa extracellular dopamine eseng feela ka nucleus accumbens empa hape le sekhutlong sa libaka, 'me ho thibeloa ha dopamine receptors mokokotlong o ka tlaase ho fokotsa palole erection (ventcu et al. , 2007, 2008; Melis et al., 2007, 2009a). Boteng ba linama tsena li bokella dopamine-paraventricular dopamine-paraventricular oxytocin-ventral tegmental sebakeng sa li-link tsa oxytocin-dopamine e boetse e boleloa ke bokhoni ba tekanyo ea pro-erectile ea apomorphine le ea D4 receptor aconist PD 168077 e kenngoang ka har'a kakaretso e ka holimo ho eketsa dopamine e eketsehileng ka seleng ea nucleus accumbens (Succu et al., 2007), karabo e felisoang ke d (CH2) 5Tyr (Me) -Orn8-vasotocin e fanoeng sebakeng sa ventral tegmental (Melis et al., 2009a, bona hape). Leha ho le joalo, mosebetsi o eketsehileng oa hlokahala ho khetholla litsela tsa neural tse hokahanyang nucleus accumbens ho incerto-hypothalamic dopaminergic system.

Mochine oo oxtocin o etsang hore penile erection e be teng le ho etsa hore li-neurone tsa mesolimbic dopaminergic li sebelisoe ha li kenngoa ka subiculum ea ventral kapa ho ea ka morao-raoana ea amygdala li utloisisoa ka nakoana feela. Libakeng tsena hape, oxytocin e sebelisa lits'ebeletso tsa eona tse etsang hore nitric oxide-synthase e sebelisoe, kahoo e eketsa tlhahiso ea nitric oxide. Nitric oxide e etsa hore ho se ke ha e-ba le li-projecti tse sa tsejoeng, tse bonahalang eka li eketsa tšollo ea mali holimo sebakeng sa ventral tegmental. Hona e le 'mele oa Glutamic e hlasimolla mesolimbic dopaminergic neurons e lebisang penile erection. Tlhahiso ena e tšehetsoa haholo ke bokhoni ba oxytocin e kenngoa ka subralulum ea ventral ho eketsa acid e mengata ea glutamic sebakeng sa ventral tektal (Fig. 3), le ea mohanyetsi oa NMDA (+) MK-801 ea sa amoheleng lihlahisoa (+) MK-2009 sebakeng sa ventral tegmental, empa eseng kucleus accumbens, ho fokotsa penile erection e bakoang ke oxytocin e kenngoa ka hare ho ventral subiculum kapa ho phallo ea posteromedial ea amygdala (Melis et al., XNUMXb). Ka motsotso ona, lintlha tse ling li fumaneha bakeng sa penile erection ea ventral subiculum e entsoeng ka oxytocin. Ha ho mona penile erection e entsoeng ka oxytocin e bonahala e le ea bobeli ba ho amohela li-receptor tsa oxytocinergic tse ka har'a litho tsa methapo ea nitric oxide-synthase-containing neurons. Sena se etsa hore tlhahiso ea nitric oxide e be e eketsehileng, e etsang hore motho a se ke a hlola a e-na le bothata ba ho ntša metsi ka ho etsa lintho ka nako e telele (sheba Snyder, 1992; Southam le Garthwaite, 1993; Schuman and Madison, 2004) . Tumellanong le khopolo ena ea phoeni e entsoeng ke oxytocin e etsahalang ka mokhoa o khotsofatsang ka ho eketseha ha acidide ea extracellular glutamic ho dialysate ho tswa ho ventral subiculum, le ho tsosolosa ha li-amino acid receptors ka subralulum ea ventral ka NMDA, ho kenyelletsa penile erection. Hape, dimutamic acid e etsa hore li-projection tsa neural efferent li sebetse hantle, e leng ho lebisang ho ts'ebetso ea mesolimbic dopaminergic neurons sebakeng sa ventral tegmental, joalokaha ho boletsoe ka holimo. Haeba lisebelisoa tsena li sebetsa hape le nucleus ea posteromedial ea amygdala ha e tsejoe hona joale. Ho feta moo, liphuputso tse ling li hlokahalang ho netefatsa hore (1) li-endrose tsa methapo ea mali le li-receptors ka subiculum ea ventral le mokokotlong oa posteromedial oa amygdala li sebakeng sa lisele tsa li-neurons tse nang le nitric oxide-synthase, (2) tlas'a taolo ea syninse ea amino acid (glutamatergic) le (3) ho khetholla mokhoa oa ho paka o sebelisitsoeng ke nitric oxide. Tabeng ena, hoa hlokomeleha hore bokhoni ba oxytocin bo eketsa ts'ebetso ea nitric oxide-synthase likarolong tsa lisele tsa dopaminergic neurons sebakeng sa caural tegmental le sa neurons tse sa tsejoeng ka subralulum ea ventral le mokokotlo oa posteromedial oa amygdala, oa hopola bokhoni ba oxytocin bo kenyelletsa nitric oxide-synthase liphatleng tsa lisele tsa oxytocinergic neurons mokokotlong oa pela (Melis et al., 1997). Leha ho le joalo, ha sebaka sa ventral tegmental tlhahiso e ntseng e eketseha ea nitric oxide ka liseleng tsa dopaminergic neurons e lebisa ho ts'ebetsong ha li-neurons tsena ka ho etsa hore guanylate cyclase e eketsehe le ho eketseha ha cyclic GMP, sena ha se hlahe mokokotlong. Ka lebaka leo, 8-bromo-cyclic GMP e kenngoa mokokotlong oa phallo ha e hlahise penile erection, ha e ntse e etsa ha e kenngoa sebakeng sa ventral tegmental. E 'ngoe ea tsela e fapaneng le ea nitric oxidecyclic GMP tsamaiso e kenngoa mofuteng oa phetoho ea li-oxytocinergic neurons e kopanelang ho emisa penile ka oxide e nang le mamello e kholo le / kapa e sa tloaelehang (Melis le Argiolas, 1995b; Melis et al., 1997). . 1). Ka lehlakoreng le leng, GMP ea cyclic sebakeng sa ventral tegmental e bonahala e phetha karolo ea bohlokoa ho ts'ebetsong ea mesolimbic dopaminergic neurons le ho eketseha ha dopamine e eketsehileng ea cellular e etsahalang ho dialysate e fumanoang sekhethong sa nucleus accumbens ea likokoanyana tse khethiloeng bakeng sa ho bontša kapa ho se bonts'a li-penile erections ha li kenngoa ka ho ba teng ha motho a sa amohelehe ka tsela e amohelehang (e tšoaroang ke estrogen + progesterone) rat ea basali. Maemong ana a liteko, lintlheng tsa banna tse bontšang li-penile erections tse sa sebetsane, keketseho ea mahloriso a dopamine a eketsehileng a fumanoa ho lekanngoe, 'me keketseho ena e ntse e eketseha, le hoja ka mokhoa o itekanetseng, ke phosphodiesterase inhibitors tse fanoeng ka ventral tegmental sebakeng sa Sanna et al., 2009).

Bokhoni ba oxytocin bo kenngoa sebakeng sa ventral tegmental, ka ventral subiculum le ka kamoreng ea posteromedial cortical ea amygdala, hammoho le ea dopamine agonists e kenngoa mokokotlong oa phallo, ho etsa hore penile erection le ho kopa mesolimbic dopaminergic neurons li tšoane . Ntlha ea pele, mekhoa e tšoanang le e boletsoeng ka holimo e ka 'na ea sebelisoa ha penile erection e hlaha likarolong tsa' mele, joalo ka nakong ea ho kopola (nakong ea li-copula penile erections) kapa nakong ea li-penile erections. Lisebelisoa tsena ke li-penile tse khetholloang ke pheromone tse sa tsejoeng ho tsoa ho ba susumelitsoeng ke lithethefatsi kapa oxytocin, e etsahalang ha linko tsa banna tse nang le likamano tsa botona le botšehali li kenngoa ka pel'a bothata ba ho se amohelehe (estrogen + progesterone primed) palo ea basali 'me e nkoa e le letšoao la thobalano ho tsosa (Sachs, 1997, 2007). Ha e le hantle, le hoja liphello tsena li sa bontše hore oxytocin libakeng tsena e phetha karolo ea ho hlahisa penile e etsahalang meleng ea boipheliso kapa ka morao ho tsamaiso ea lithethefatsi, li eketsa matla ho liphuputso tsa pele li fana ka maikutlo a hore libaka tsena tsa boko ke tsa bao oxytocin e fanang ka liketso tse eketsehileng eseng feela ka tlhaho ea penile erection liketsahalo tse bonngoeng ka mor'a lithethefatsi tsa pro-erectile, empa hape le ho ntlafatsa boitšoaro bo botona ba banna (le basali) (bona Argiolas le Melis, 2004 le litemana tsa eona). Ka hona, maqhubu a oxytocin a eketseha hippocampus ea likhoto tse tona tse tšoaroang ka tekanyo ea pro-erectile ea apomorphine, ea khale ea dopamine agonist (Melis et al., 1990) le d (CH2) 5Tyr (Me) -Orn8-vasotocin, e thibelang oxytocin li-receptors, li atleha ka ho fetisisa, eseng feela ho fokotsa boitšoaro bo hlephileng (Argiolas et al., 1987a) empa hape le phello ea boipheliso ea apomorphine boitšoarong ba banna ba ho kopanela liphate (Argiolas et al., 1987b) ka litoeba tse matla tsa ho kopanela liphate, nakong eo ka pulole erection ho etsahala. d (CH2) 5Tyr (Me) -Orn8-vasotocin e boetse e matla haholo ho fokotsa likarolo tse sa sebetsaneng tsa likhoto tsa banna tsa thobalano, ha li fuoa nanogram e le li-ventricles, empa e seng ka PVN (Melis et al., 1999a).

Ntlha ea bobeli, mahlaseli a dopaminergic neurons a phetha karolo ea bohlokoa litšusumetsong tsa tlhaho, tse kang lijo, metsi le thobalano (Fibiger le Phillips, 1988; Wise le Rompre, 1989; Everitt, 1990). Haholo-holo, dopamine e lokolloa li-neurons tsena e nahanoa hore e fetisa tšebetso ea likarolo tse susumetsang tsa tlhaho ho latela mekhoa ea boitšoaro, ka mohlala tabeng ea thobalano, ho batla molekane oa thobalano le ho kopanela liphate ho fihlela moputso le khotsofalo (Goto le Grace, 2005). Ka lebaka leo, maqhubu a extracellular dopamine a eketseha ka dialysate ho tloha nucleus accumbens ea litoeba tse nang le matla a ho kopanela liphate nakong ea ha motho a pepesehela lesela la mosali ea amohelehang la estrogen + ea progesterone-primed, 'me keketseho e joalo e bile e phahameng ha raleshanyana e lumelloa ho sebetsana le basali ba amohelehang (Pfaus le Everitt, 1995).

Ntlha ea boraro, liphello tsa hona joale li tšehetsa khopolo ea hore potoloho ea neural e kopanya mokokotlo oa sepakapaka le sebaka sa ventral ka ho toba kapa ka tsela e sa tobang (ho sebelisa subiculum ea ventral le / kapa kopo ea posteromedial ya amygdala) le nucleus accumbens, le ho tloha mona ho sa tsejoe mekhoa ea ho khutlela mokokotlong oa ho thibela tšebetso ea li-neuronicic neurons tse hlalosang mokokotlo oa mokokotlo o kopanyang penile erection le ea oxytocinergic neurons e lebisang sebakeng sa ventral tegmental, ventral subiculum le mokokotlo oa posteromedial cortical oa amygdala, o tsamaisang ka tsela ena ketsahalo ea mesolimbic dopaminergic neurons (setšoantšo sa 4). Ts'ebetso ena e rarahaneng ea neural e ka ba le karolo ho kenyelletsong ea mesebetsi ea li-neural e kenyelletsoeng ho laola mokhoa oa ho phekola (erectile-ejaculatory) le litebello (tse susumetsang le tse khotsofatsang) likarolong tsa thobalano ka thobalano. Ha e le hantle, dopamine e eketsehileng ea lisele e eketsa nucleus accumbens (Pfaus le Everitt, 1995) le mokokotlong oa litoeba tse motona ka ho kopanela liphate nakong ea ho pepeseha ho ratana ea basali e sa amoheleng, ha ho e-na le likarolo tse sa lumellaneng tse etsahalang, mohlala, ha ho hlaha li-penile erections (Melis et al., 2003). Ka hona, le hoja liphuputso tse eketsehileng li hlokahala ho hlakisa karolo ea phekolo ea oxytocin e sa feleng, sebakeng sa ventral subiculum le amygdala ka nako ea ho se lumellane le boitšoaro ba thobalano, e ka 'na ea e-ba ho utloahalang ho nahana hore potoloho ena ea mahlaseli, ha e ntse e tlatsetsa likarolo tse senyang tsa boitšoaro ba thobalano, ka nako e ts'oanang li ka boela tsa kenya tsamaiso ea mesolimbic dopaminergic e fanang ka substrate ea neural bakeng sa ho hlalosetsa litšobotsi tse thabisang tsa thobalano (Everitt, 1990; Pfaus le Everitt, 1995). Mabapi le taba ena, hoa hlokomeleha hore tsamaiso ea dopaminergic e sebelisoang ke mesotombic e sebelisoang ke oxytocin e kenngoa sebakeng sa ventral tekete e tšoana le e sebelisoang ke lithethefatsi tse sebelisoang hampe tse kang opiates, cannabinoids, amphetamine, cocaine le joala (Tanda et al., 1997), le hore oxtocin e fumanoe e khona ho fokotsa ho mamella le ho itšetleha ka cocaine, morphine, joala le li-cannabinoids (Kovacs et al., 1998; Cui et al., 2001). Qetellong, ho bonahala eka oxtocin e lokolloa sebakeng sa ventral tegmental feela, empa hape ho ventral subiculum le kakarese ea coronal posteromedial ea amygdala, e ka etsa hore ho be le methapo e kholo ea dopaminergic neurons, e ka kenang liphellong tse thabisang le tse putsang tsa thobalano . Ts'ebetso ea mesolimbic dopaminergic neurons e ka ba ka ho toba, ka li-receptor tsa oxytocinergic tse ka har'a lisele tsa mesolimbic dopaminergic neurons, kapa ka tsela e tobileng ka ho sebelisoa ha methapo ea mali ea glutamic sebakeng sa ventral tegmental.

Dopamine e lokolloa nucleus accumbens shell e etsa hore mosebetsi oa incerto-hypothalamic dopaminergic neurons o se ke oa e-ba teng mokokotlong o etsang hore e be penile erection (ka ho sebelisoa ha oxytocinergic neurons e hlalosang mokokotlo oa mokokotlo), kapa tšusumetso ea thobalano le moputso (ka tšebeliso ea oxytocinergic li-neurons tse lebisang sebakeng sa ventral tegmental, subralulum ea ventral kapa kopo ea posteromedial cortical ea amygdala). Ho tloha ha dopamine e boetse e lokolloa nucleus accumbens shell le mokokotlong oa phallo ha penile erection e hlaha maemong a mmele (mohlala, ho se lumellane le ho kopana) (Succu et al., 2007; Melis et al., 2003, 2007), ke ho ka etsahala hore karolo e bohareng ea oxytocinergic neurons e kenella lipotlelong tsa neural tse tsamaisang ho sebelisana pakeng tsa mesolimbic le incerto-hypothalamic dopaminergic system. Ts'ebetso ena ea mahlaseli a marang-rang e ka 'na ea se ke ea e-ba le karolo feela karolong e qetellang ea boitšoaro ba thobalano (mohlala, ho tlosoa ha penile le ho kopana), empa hape le tšusumetsong ea thobalano, tsoho ea thobalano le moputso oa thobalano.

 

Setšoantšo sa 4. Erectile physiology

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Tšoantšiso ea setemane ea potoloho ea methapo ea li-neural e amanang le oxytocin e susumetsang tšusumetso ea thobalano, e putsang le ts'ebetsong ea thobalano, joalokaha ho bontšitsoe ke liphello tsa khaolo ena le litlaleho tsa pele. Oxytocinergic neurons e hlahang mokokotlong oa sepakapaka le ho hlahisa mokokotlo oa mokokotlo ha o sebelisoa ka mohlala ke dopamine le glutamic acid (empa hape ke li-neurotransmitters le / kapa li-neuropeptides), ho etsa hore penile erection le ts'ebetso ea ho kopanela liphate ka ho etsa li-oxytocinergic neurons tse hlalosang mokokotlo. Dopamine le glutamic acid (empa hape le li-neurotransmitters le neuropeptides) mokokotlong oa sepakapaka hape li etsa hore oxytocinergic neurons e hlahise sebaka sa ventral tegmental, kahoo e etsa hore ho be le li-neuron tsa mesolimbic neurus, tse etsang hore ho be le tšusumetso ea thobalano le moputso. Dopamine e lokolloa nucleus accumbens (NAs) e ntse e etsa hore litsela tsa neural tse sa tsejoeng li ntse li sa tsejoe, tse eketsang ts'ebetso ea incerto-hypothalamic dopaminergic neurons (e hlahang lihlopheng tsa Ahlnumx-A13 tsa Dahlstrom le Fuxe) tse nang le li-neuron tsa oxytocinergic, mokokotlo oa mokokotlo, o lebisang penile erection. Ho potoloha ho ka boela ha nkoa ke oxytocin eseng feela sebakeng sa caural tegmental sebakeng sa caudal feela, empa le ka ventral subiculum le ho amygdala, e leng eona e fumanoang ho boloka boikutlo ba oxytocineric ho tloha mokokotlong oa phallo, mohlomong ka tsela e tobileng kapa e sa tobang ea ho hlahisa matla sebaka sa teptional, se lebisang ho feto-fetoha ha maikutlo a mabeli a ho kopanela liphate le ho fokotsa penile. Qetellong, liphutheho tse ka holimo li ka boela tsa ts'oaetsoa ke li-pheromone tsa ho kopanela liphate le lipheromone, kaha dopamine e eketsehileng ea cellular le glutamic acid e eketsa mokokotlo oa sepakapaka (le sebakeng se ka hare ho sebaka se ka pele) nakong ea li-pheromone-mediated non -act contractions (coping references) Lethathamo la litšupiso).

 

Tlhaloso Mosebetsi ona o ne o tšehetsoa karoloana ka thuso ea Lefapha la Kopano ea Italy le la Patlisiso ho AA le MRM

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