I-Neurophysiology ye-Erectile Function: Iimpembelelo ze-Androgenic
UARTHUR L. BURNETT
Ijenali yeAndrology, Vol. 24, No. 90060, 2003
Ilungelo lokushicilela © American Society of Andrology
Ukusuka kwiSebe leUrology, iJames Buchanan Brady Urological Institute, iSibhedlele saseJohns Hopkins, eBaltimore, eMaryland.
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Ukwakhiwa kwePenile yinkqubo ekhethekileyo ye-vascular biologic efuna ukulawulwa kolawulo. Phakathi kweendlela zokulawula ezinegalelo kulo mmiselo, inkqubo ye-nervous ibonwa njengeyona ndlela iphambili yokulawula enika olu lawulo. Amanqanaba amaninzi e-neuroaxis, ukusuka kwingqondo kunye nentambo yomgogodla ukuya kwimithambo-luvo ephela ngaphakathi kwelungu lobudoda, iqala kwaye idlulisele iimpembelelo ze-neurochemical ezivelisa impendulo ye-erectile.
Kwangaxeshanye, kuyaziwa ukuba i-androgens idlala indima ebalulekileyo ekuphuhliseni nasekugcinweni kweesekethe ze-neuronal ezibandakanyekayo ekusebenzeni ngokwesondo kwamadoda. Ezi ndima zenziwa kumanqanaba enkqubo ye-nervous central kunye ne-peripheral. Uphononongo lweempembelelo ze-androgenic luye lwavavanya ngokuyintloko iindlela ezingundoqo, ngokugxila ekuhlukeni ngokwesondo kwingqondo ye-vertebrate. Iindlela zeeselluli phantsi kolawulo lwe-androgenic ziquka i-neurogenesis, ukuhlukana kweeseli, ukufuduka kweeseli, ukubunjwa kwe-synapse, ukupheliswa kwe-synapse, kunye nokufa kweeseli. Ugxininiso olongezelelweyo lunikezelwe kutshanje kwimiphumo ye-androgenic kwiindlela ze-peripheral neuronal. Ubume beziphumo ze-androgenic kwi-peripheral neurons zibhekiselele ikakhulu kulwakhiwo kunye notshintsho lwe-biochemical olwenzeka ngakumbi ekufikiseni nasemva kokuthenwa, nangona uphando lwakutsha nje luchonge utshintsho lwe-electrophysiologic phakathi kwezi zinto ziphazamisayo.
Lo mboniso ubonelela ngophononongo olufutshane lwangoku lokulawulwa kwe-neural ye-penile erection kumanqanaba aphakathi kunye ne-peripheral, ngokuqwalaselwa okukhethekileyo kwimpembelelo ye-androgenic kwi-neuroanatomy, i-neural pathways, kunye neendlela ze-neurological ezibandakanyekayo ekuphenduleni kwe-erectile. Ngokufanelekileyo, ubungakanani be-androgens sisebenzisa isiseko solawulo oluhlekisayo kwinkqubo yolawulo oluphambili lokwakhiwa luya kuhlolwa. Ukuqonda okuthathwe kolu hlaziyo kunokubonakalisa ukuba ukungenelela kwe-pharmacotherapeutic ebandakanya inkqubo ye-nervous usebenzisa i-testosterone supplementation inokwenzeka ukunyanga ukungasebenzi kwe-erectile yamadoda. Kuyaqatshelwa ukuba uninzi lwesiseko solwazi malunga nalo mbandela luvela kwizifundo zezilwanyana zebhubhoratri, kwaye ubungqina obuthe ngqo beziphumo ze-androgenic kwi-penile erection yabantu zihlala zilinganiselwe. Ngapha koko, iiyantlukwano zokwenyani kwiziphumo ze-androgenic ziyaziwa ukuba zikhona phakathi kwezilwanyana zaselabhoratri ezisetyenziswa ngokuqhelekileyo kunye neeprimates, ezinjengomahluko wokuba i-aromatized metabolites i-androgens ayinampembelelo kwi-dimorphism yesini kwiiprimates (Cooke et al, 1998). Nangona kunjalo, ulwazi oluchazwe apha lunika ithuba lokuqwalasela imiba ye-androgenic yokulawulwa kwe-neurophysiologically mediated penile erection. Iingqikelelo ezilinganiselweyo malunga nomsebenzi wobudoda bomntu unokuzotywa ngcono ngeli xesha.
IiNdlela eziPhakathi
Iinuclei ezininzi zobuchopho zicetywayo ukuba zithathe inxaxheba kwi-supraspinal control of penile erections (Burnett, 2000). Amaziko okulungelelanisa (kwindlela ehlayo ye-neuroaxis) abandakanya iindawo ze-cortical kunye ne-subcortical, i-diencephalon equkethe indawo ye-hypothalamic ye-medial (i-nucleus ye-paraventricular, indawo ye-preoptic ye-medial [MPOA], kunye ne-dorsal hypothalamic area), kunye nesiqu sobuchopho ( bona iThebhile). Kwinqanaba lomgogodla, i-nuclei ebalulekileyo ibandakanya ikholomu yeseli ye-intermediolateral (ebizwa ngokuba yi-sacral parasympathetic nucleus kumanqanaba e-lumbosacral), elawula igalelo lokuzimela kwi-pelvis; i-medial dorsal horn kunye ne-dorsal commissural nucleus, njenge-sensory input circuit ye-spinal reflexes; kunye ne-Onuf's nucleus (ekwaziwa ngokuba yi-spinal nucleus ye-bulbocavernosus) ekwi-ventral horn kumanqanaba e-lumbosacral, ekhusela izihlunu ezidibeneyo ezihambelana ne-penile. Engqondweni, i-nuclei ye-diencephalon inokuthi iqhagamshelane namanqanaba entambo yomgogodla edibanisa i-reflexive penile erections kunye nezinye iindawo zobuchopho ezibonisa umsebenzi wayo ekuhlanganiseni ulwazi olunyukayo kunye nokwehla kwe-neural. Intambo yomgogodla idibanisa i-neural reflexive loops phakathi kwe-penile afferents kunye ne-autonomic kunye ne-somatic efferents efunekayo kwiimpendulo ze-erectile. Umbindi we-neurochemistry wokulamla ngokuziphatha ngokwesondo kubandakanya ii-arhente ezahlukeneyo, ezisekwe ngezifundo ezahlukeneyo ze-morphological kunye ne-pharmacological kwizilwanyana. Iindima ezibekayo zikhona kwi-monoamines (i-dopamine, i-norepinephrine, kunye ne-5-hydroxytrypatmine), i-decarboxylated amino acids, i-neuropeptides (i-oxytocin, i-prolactin, i-adrenocorticotropin, i-opioids), kunye ne-molecule ye-gaseous.
IiNkqubo zePeripheral
I-neuroregulation ye-penile erection peripherally ifuna ukulungelelaniswa kwe-parasympathetic, uvelwano, kunye ne-somatosensory neural pathways (Burnett, 2000). Ezi ndlela zichaza uqikelelo olucacileyo kunye nolwahlukileyo. Iprojekti esebenzayo echaphazelekayo kwi-penile erection ibhekisela kwi-thoracolumbar sympathetic (T10-L2) kunye ne-sacral parasympathetic (S2-S4) yahlula inkqubo ye-nervous autonomic, kunye ne-sacral somatic (S2-S4) inkqubo ye-nervous system. Igalelo le-autonomic limelwe ngokuyinhloko yi-nerve ye-cavernous, evela kwi-plexus ye-hypogastric ephantsi, kwaye i-somatic input imelwe yi-pudendal nerves, leyo ikhosi evela kwi-sacral plexus. I-afferent projection echaphazelekayo kwi-penile erection ibandakanya i-sacral innervation (S2-S4) kwaye imelwe yi-dorsal nerves ye-penis, amasebe oluvo lwe-pudendal nerves.
Ngokudibeneyo, ezi ndlela ze-neural zilawula ulandelelwano lokungena kwegazi kunye nokungena kwe-penis, kwaye zilungelelanisa umsebenzi we-penile-associated striated muscles, ekhontrakthi yokwandisa ubulukhuni be-penile. Iindawo ezisebenzayo zesenzo ngaphakathi kwepeni ziyi-vascular smooth muscle components, i-vasculature enikezela i-penis, kunye ne-trabeculae equka izicubu ze-erectile ze-penis (Burnett, 2000). Ezi zixhobo ziphendula kwi-neuronal stimulus efanelekileyo ngokuvelisa idigri ye-vascular smooth muscle tone echaphazela imeko yokumisa i-penis. Ngexesha le-penile tumescence kunye nokulungiswa, i-penile vasculature vasodilates kunye nezicubu ze-trabecular ziyakhululeka. Ngexesha le-tonic state of flaccidity, i-penile vasculature i-vasoconstricted kwaye i-trabecular tissue ibanjwe. Iingcamango zangoku zixhasa indima ephambili ye-nitric oxide, i-molecule ye-gaseous isithunywa, njengomlamli oyintloko we-penile erection ekhonza zombini njenge-neurotransmitter kunye ne-endothelial effector (i-Burnett, i-2002), nangona ezinye ii-neurotransmitters ziyaziwa ukuba zinefuthe kwi-neuroregulation ye-penile erection. . Ukuququzelelwa kweendlela ze-proerectile neurotransmitter kunye nokuvinjelwa kweendlela ze-antierectile neurotransmitter zihlangene zibandakanyeka ekuveliseni ubuninzi be-penile erection.
Androgenic Regulation
Ingcamango ye-steroid-sensitive neurons kunye ne-neuronal regulatory pathways imiselwe kakuhle kwinkqubo ye-nervous central. Owona mmandla mkhulu wokufunda ubhekiselele kwi-dimorphism yezesondo, enxulumanisa iimpembelelo zehomoni kunye nemimandla yobuchopho elawula ukuziphatha kwamadoda okanye okwabasetyhini. I-Androgens kunye ne-estrogens zenza iziphumo ngokusebenzisa imimandla ebonisa ii-steroid receptors ezilawula iipropathi ezahlukeneyo ze-neuronal ngexesha lokuvuthwa kunye nakubudala bonke. Le mimandla imelwe kuyo yonke inkqubo ye-nervous ngaphandle kokugxila kwisakhiwo esisodwa okanye kwiziko le-neuronal.
Mhlawumbi eyona nto ibonakaliswe kakuhle kwi-sex dimorphic neuromuscular system kwizilwanyana ezanyisayo ibandakanya i-spinal nucleus ye-bulbocavernosus muscle (SNB). Le nucleus ichaza iqela le-motoneurons kwintambo esezantsi ye-lumbar spinal innervate striated ischiocavernosus, bulbocavernosus, kunye ne-levator ani izihlunu ezincanyathiselwe kwipipi. I-rat isebenze njengemodeli yezilwanyana eziphambili ukuphanda le nucleus, ehlala kwindawo ye-dorsomedial ye-ventral horn kumacandelo e-lumbar 5 kunye ne-6 kule ntlobo. Iigundane zamadoda ziphethe ubuncinci amaxesha e-3 uninzi lwe-SNB motoneurons njengabasetyhini, kwaye ezi motoneurons ziphindwe kabini ubukhulu kumadoda xa kuthelekiswa nabasetyhini (Breedlove kunye noArnold, 1980). Nangona zombini iigundane zamadoda nabasetyhini zineeseli ze-SNB ezihambelana nezihlunu ze-bulbocavernosus ngaphambi kokuzalwa, izihlunu kunye ne-motoneurons yazo zihlala ziwohloka kungekudala emva kokuzalwa kwiigundane zabasetyhini (i-Breedlove kunye no-Arnold, i-1983). Le nkqubo yophuhliso iseka umahluko wesini sabantu abadala kwi-SNB motoneurons kunye ne-fiber target muscle. Iindlela ezahlukeneyo zobungqina ezibandakanya ukuvezwa kwehomoni kunye nokuhoxiswa kwe-perinatally kwiigundane zesilisa kunye nesetyhini ezikhokelela kwiinguqu ezicacileyo ze-morphological ziye zaqinisa olu qwalaselo (uphononongo olongezelelweyo, bona uCooke et al, 1998).
Ubungqina obunjalo bokulinga bubonisa ngokucacileyo ukuba i-androgens ichaphazela ukuxolelaniswa kwe-penile reflexes ehambelana ne-SNB. Yintoni indlela yolawulo olunjalo? Izifundo eziphuhlisayo zithanda i-hypothesis yokuba i-SNB motoneurons igcinwe phantsi kweempembelelo ze-androgens zigcina izihlunu ekujoliswe kuzo, ngaloo ndlela zikhokelela ekugcinweni kwayo yonke into ehambelanayo yokuhlala (Breedlove kunye noArnold, 1983). Izinto ze-neurotrophic zicatshangelwa ukuba zilawulwe okanye zinyanzeliswe yi-androgens ukugcina imisipha kunye ne-motoneurons ehambelana nayo (Forger et al, 1995; Al-Shamma noArnold, 1997). Impembelelo ye-androgenic nayo ikholelwa ukuba yenza indima yokulondoloza ekukhuleni kwabantu abadala (iRand kunye neBreedlove, i-1995; i-Nanasaki kunye ne-Sakuma, i-2000). Isiseko seziphumo zetrophic xa umntu emdala asihambelani kuphela nenkxaso ye-androgenic ye-penile-associated striated muscles. Kule nqanaba lokuvuthwa, i-SNB i-motoneurons iye yaphuhliswa ukubonisa i-receptors ye-androgenic ukuba iphendule ngokuthe ngqo kwi-androgen exposure ukugcina ubukhulu be-somata (Freeman et al, 1996).
Mhlawumbi i-sexual dimorphic neural center ifundwe kakhulu yi-MPOA, ngolwazi oluthathwe kwiintlobo ezahlukeneyo, kuquka nabantu. Ngaphakathi kule nucleus, iqela le-sexual dimorphic ye-neurons liye lachongwa kuzo zonke iintlobo zeentlobo, kwaye kwiigundane zilingana namaxesha e-5 amakhulu ngobukhulu kumadoda kunabasetyhini (Gorski et al, 1980). I-cytoarchitecture yale neurons iphinda ihluke phakathi kwabasetyhini kwiintonga, ngokubhekiselele kwiinkalo ze-synaptic kunye nokudibanisa (i-Raisman kunye ne-Field, i-1973). Izifundo ezibophelela i-receptor ziye zahlala kwiindawo ze-steroid kwezi neurons zikhethekileyo, kunye noxinaniso olukhulu lweendawo zokubopha i-estrogen receptor-binding kwiimpuku zabasetyhini kuneempuku zamadoda kunye nomlinganiselo omkhulu weendawo ezibophelela i-androgen receptor kwiimpuku zamadoda kuneempuku zabasetyhini (Jacobson kunye nabanye, ngo-1987). Nangona kunjalo, umsebenzi wale ndawo ye-sexual dimorphic neural centre kwizilwanyana ezincelisayo uhlala ungaqinisekanga, kunye nedatha elinganayo nakwinqanaba le-rodent: emva kokutshatyalaliswa kwendawo, akukho lutshintsho kwiindlela zokuziphatha zamadoda kwiigundane, ngelixa ukuncipha kwezi ziphatha kukhokelela kwii-gerbils. Cooke et al, 1998).
Ukulawulwa kwe-androgenic ye-MPOA ye-sexual dimorphic neurons ibambelela kuyo yomibini intlangano (yophuhliso) kunye ne-aromatization hypotheses kwizifundo ze-rat. Impembelelo ye-humoral kwi-neurogenesis kunye nokukhula kwe-neuronal kubomi bomntwana kunye nomntwana wasemva kokubeleka kuthetha iziphumo zophuhliso (Gorski et al, 1980). Utshintsho kwi-gonadal steroid exposure ekukhuleni aluchaphazeli i-morphology yale nucleus (Gorski et al, 1980). Nangona kunjalo, ukutshintshwa kwe-testosterone yexesha elide kwiigundane ezikhulileyo kubonakaliswe ukugcina ukuziphatha okunyukayo kunye nomsebenzi we-dopaminergic kwi-MPOA (Sato et al, 1998), ebonisa ukuba ukuvezwa okuzinzileyo kwe-androgenic kunokuba nemiphumo ebudala ngaphaya kwenqanaba le-morphological. . Okubangel 'umdla kukuba, unyango lwe-estrogen lusebenza ngokufanelekileyo ukuvuthwa okuqhelekileyo kolu hlobo lwabantu abakhethiweyo be-neurons kunokuba unyango lwe-androgen kwiigundane, ebonisa ukuba i-metabolites enevumba elimnandi ye-androgen yanelisa ngokwaneleyo ezi neurons (Dohler et al, 1984).
Eminye i-nuclei embindini we-sexual dimorphic nayo ichazwe kakhulu ecaleni kwezizathu ze-morphological kwaye ngokuyintloko kwiintlobo zeempuku. Ezi ziquka inkqubo ye-olfactory limbic system (i-medial amygdala, i-nucleus yebhedi ye-stria terminalis, indawo ye-preoptic), i-hippocampus, kunye ne-assorted hypothalamic nuclei.
Umbutho we-androgens kunye nomsebenzi we-peripheral autonomic neurons usandul 'ukuza kuphononongwa. I-Androgen receptors kunye ne-androgenic sensitivity ibonakaliswe kwi-lumbosacral preganglionic neurons, i-sympathetic kunye ne-parasympathetic postganglionic neurons, kunye ne-sensory postganglionic neurons (Schirar et al, 1997b). Umdla omkhulu unikiwe kwiiseli ezizimelayo ze-pelvic ganglion, equka umxube we-neuron ezinovelwano kunye ne-parasympathetic. Kwiigundane, ii-neurons ezinovelwano zitshintsha ngobukhulu ngokuphawulekayo emva kokuchithwa okanye ulawulo lwe-testosterone (Keast kunye neSaunders, i-1998). Ngokuhambelana noku kuqwalaselwa, ukubonakaliswa kwe-testosterone kuhambelana ngokuthe ngqo kwi-membrane capacitance ye-neurons enovelwano kwi-pelvic ganglia yeegundane (Kanjhan et al, 2003). Nangona ubungqina bubonisa ukuba i-androgens ichaphazela ngokuyinhloko ukukhula komzimba we-neuronal kunye nobukhulu, ubungqina obongezelelweyo bubonisa umphumo wabo kwiindlela ze-ion kunye neendlela zezithunywa zesibini ezinxulumene ne-pelvic ganglion neurons. Ngokomzekelo, kwiigundane, ukuthenwa kukhokelela kumxholo we-neurochemical ophelile we-nitric oxide synthase, eyenza i-nitric oxide neuronally, ngelixa ukutshintshwa kwe-testosterone emva kokuchithwa kubuyisela eli nqanaba le-neurochemical (Schirar et al, 1997a). Iziphumo ezifanayo ziye zaboniswa kwi-postganglionic cavernous kunye ne-dorsal nerve terminations kwi-rat penis (Giuliano et al, 1993; Baba et al, 2000).
Ukufaneleka Kwabantu
Imigca emininzi yobungqina bovavanyo iseka indima enkulu yeehomoni ze-steroid kulwahlulo lwezesondo lweenkqubo ze-neural kwizilwanyana. Ngaphandle kobungqina be-morphological, ingakanani i-sexual dimorphic neural amaziko asebenzisa iziphumo zokusebenza ebantwini? Umba oqinisekileyo kulo mba kukuba kunzima ukwahlula kubuchopho bomntu omdala ukuba ngaba i-gonadal steroids ejikelezayo i-akhawunti ye-dimorphism yezesondo engahambelani nentshukumo yokuqala, ye-fetus steroid. Ukongezelela, ukungafani kwesakhiwo se-dimorphic ngokwesondo kunokuchazwa ngesiseko samava omntu, mhlawumbi ngexesha lophuhliso okanye ebudaleni, kwakhona ngaphandle kokuhambelana nesenzo se-fetal steroid. Isisombululo sobunzima obunjalo sinokufezekiswa ukuba i-dimorphisms yezesondo ibonakala ngokucacileyo ekuzalweni, nangona imizekelo embalwa ikhona ukuxhasa le ngxabano. Inkxalabo yokuba ubunzima bobuchopho bomntu bukhulu kumadoda, njengomzekelo, kunokudibaniswa nefuthe le-androgens ekukhululweni kunye nesenzo sezinye izinto ezifana ne-hormone yokukhula ngaphandle kwesenzo esithe ngqo se-steroids. Kuyacaca ukuba, uninzi lweeyantlukwano ze-dimorphic ngokwesondo ebantwini zinokubangelwa kukuvezwa kwe-androgen ngexesha elithile elibalulekileyo emva kokubeleka. Ubungqina ebantwini abugqibeki ngokupheleleyo ukunika ingxelo malunga nokuba ukuvezwa kwehomoni yomntwana kumisela indlela yokuziphatha ngokwesondo. Akukho bungqina bokuxhasa i-hypothesis ye-aromatization ebantwini, kwaye umzekelo womntu we-XY ongenalo i-androgen ongabonisi ukuziphatha kwamadoda ngaphandle komphandle wobufazi ngokuqinisekileyo uya kuphikisa ngokuchasene nale ngcamango. Ke, ubungqina obuqinisekileyo bokuba ii-steroids ezisebenza kwangethuba kuphuhliso zenza ubudoda ngokuthe ngqo kwingqondo yomntu zihlala zingekho.
Ngako oko, ngesiseko sobungqina bovavanyo kulo mbandela, ukuzimisela malunga nokulawulwa kwe-androgenic phezu kwe-neuroregulation yokumiswa komntu kufuneka kuthintelwe. Kwelinye icala, kuba kuyabonakala ngokwezonyango ukuba ii-androgens zinefuthe kwindlela yokuziphatha kwamadoda ngokwesondo, kusekho umdla ocacileyo wokuphonononga ukuba iihomoni ze-steroid zilawula njani ingqondo yomntu omdala. Ubungqina obuxhasa imiphumo ye-androgenic kwi-neuroregulation yokwakhiwa kwabantu inikezelwa yidatha yeklinikhi ebonisa ukubaluleka kwe-androgens kwi-erections elinganiswe ngexesha lovavanyo lwe-penile tumescence yasebusuku kwi-laboratory yokulala (Cunningham et al, 1990). Ukuqonda kwangoku kukuba ii-erections ezinxulumene nokulala zibandakanya ukusebenza kweempembelelo zokulawula i-supraspinal ezisebenza ngexesha lokulala lokuhamba ngokukhawuleza kwamehlo (Lue, 2002). Ukugcinwa kokulala okuhambelana nokulala ukuya kwinqanaba elikhulu kumadoda ane-eugonadism kunamadoda ane-hypogonadism iphakamisa ngamandla ukumodareyithwa kwe-androgenic yale ndlela. Nangona kunjalo, ezinye izibonakaliso ze-erection neurophysiology, ezifana neempendulo zokuvuselela ngokwesondo okanye ukuvuselela iqabane ngesondo, akunakusebenza phantsi kolawulo olucacileyo lwe-androgenic. Ewe, amadoda ane-hypogonadism anokufikelela kwi-erections ukuvumela umsebenzi wesondo, kwaye amadoda ane-eugonadism anokufumana ubunzima ekufezekiseni ii-erections ukwenza ngokwesondo. Uluhlu lwezinto ezilawulayo ngokuqinisekileyo zinokubandakanyeka phantsi kwezi meko zahlukeneyo kwaye zenze iindima eziphambili kungakhathaliseki izinga lenkxaso ye-androgenic.
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