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In the present article, we summarize the current literature on the physiology of ejaculation.

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We describe the anatomy of the organs involved and the erection physiology. We discuss the physiology of orgasm and ejaculation as two separate physiological processes.

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In addition, we describe the neurochemical and hormonal regulation of the ejaculation process. The male genital system consists of external and internal reproductive and sexual organs such as the penis, prostate, epididymis, and testes. Figure 1 shows the gross anatomy of the ejaculatory structures. Table 1 provides a summary of the functional anatomy of these organs 2 - 5. Gross anatomy of the ejaculation structures.

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Reprinted with permission from Sheu G, Revenig LM, Hsiao W. Physiology of ejaculation. In: Mulhall JP, Hsiao W, eds. Men's sexual health and fertility: a clinician's guide.

New York: Springer; The penile erection results from complex neurovascular mechanisms. Several central and peripheral neurological factors in addition to molecular, vascular, psychological and endocrino-logical factors are involved, and the balance between these factors is what eventually determines the functionality of the penis. In this section, we summarize some of those mechanisms. Cerebrally controlled penile erections are induced through erotic visual stimuli or thoughts.

The main cerebral structures involved in erection are contained within the medial preoptic area MPOA and paraventricular nucleus PVN in the hypothalamus 6. Dopamine is the most important brain neurotransmitter for erection, likely through its stimulation of oxytocin release 7.

Another important neurotransmitter is norepinephrine, which is demonstrated through the erectogenic effect of the ? -2 agonist Yohimbine 8. Several other brain neurotransmitters are involved in the erection process to varying degrees such as nitric oxide NO? -melanocyte stimulating hormone ? -MSHand opioid peptides 9.

Parasympathetic stimulation is the main mediator for penile tumescence, although central suppression of the sympathetic nervous system also plays a role. Parasympathetic supply to the penis is derived from the sacral segments S2-S4 However, patients with sacral spinal cord injury still maintain erections through psychogenic stimulation, although of less rigidity than normal.

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These psychogenic erections do not occur in patients with lesions above T9 11suggesting that the main mechanism for these erections is central suppression of sympathetic stimulation Patients with lesions above T9 still may maintain reflexogenic erections.

This implies that the main mechanism for reflexogenic tumescence is the preservation of the sacral reflex arc, which mediates erection through tactile penile stimulation 13 The penis at baseline is in a flaccid state maintained by the contraction of corporal smooth muscles and constriction of cavernous and helicine arteries leading to moderate state of hypoxia with partial pressure of oxygen of mm Hg During sexual arousal, NO is released from cavernous nerve terminals through the action of neuronal NO synthase The NO activates guanylate cyclase, which in turn converts guanosine triphosphate to cyclic guanosine monophosphate 1517leading eventually to smooth muscle relaxation and vasodilation Although the initiation of tumescence is through neuronal NO synthase, the maintenance of erection is through endothelial NO synthase There is no standard definition of orgasm.

Orgasm is generally associated with ejaculation, although the two processes are physiologically different Orgasm is also associated with powerful and highly pleasurable pelvic muscle contractions especially ischiocavernosus and bulbocavernosus 23along with rectal sphincter contractions and facial grimacing There is also an associated release and elevation in PRL and oxytocin levels after orgasm; however, the significance of this elevation is not entirely clear Studies using positron emission tomography, which measures changes in regional cerebral blood flow, have identified areas of activation in the brain during orgasm.

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Primary intense activation areas are noted to be in the mesodiencephalic transition zones, which includes the midline, the zona incerta, ventroposterior and intralaminar thalamic nuclei, the lateral segmental central field, the suprafascicular nucleus, and the ventral tegmental area.

Strong increases were seen in the cerebellum. Decreases were noted at the entorhinal cortex and the amygdale Quality and intensity of orgasms are variable.

For instance, short fast buildup of sexual stimulation toward orgasm is associated with less intense orgasms than slow buildup. Early orgasms are less satisfying than later orgasms in life as the person learns to accept the pleasure associated with orgasms. Lower levels of androgen are associated with weaker orgasms, such as in hypogonadism or in older age It has been suggested that pelvic muscle exercises, particularly the bulbocavernosus and ischiocavernosus muscles, through contracting those muscles 60 times, 3 times daily for 6 weeks will enhance the pleasure associated with orgasm However, the effort and time associated with such exercises prevent their utilization.

The orgasm induced through deep prostatic massage is thought to be different from the orgasm associated direct penile stimulation. Although penile stimulation orgasms are associated with pelvic muscle contractions, prostatic massage orgasms are associated with 12 contractions.

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Prostatic massage orgasms are thought to be more intense and diffuse than penile stimulation orgasms, but they require time and practice and are not liked by many men 2026 Following orgasm in men is a temporary period of inhibition of erection or ejaculation called the refractory period. This is a poorly understood phenomenon, with some investigators suggesting a central rather than spinal mechanism causing it Elevated levels of PRL and serotonin after orgasm have been suggested as a potential cause; however, there is much debate about their exact role More research is still needed in the area of male orgasm Ejaculation is a physiological process heavily controlled by the autonomic nervous system.

It consists of two main phases: emission and expulsion. The main organs involved in ejaculation are the distal epididymis, the vas deferens, the seminal vesicle, the prostate, the prostatic urethra, and the bladder neck The first step in the emission phase is the closure of bladder neck to prevent retrograde spillage of the seminal fluid into the bladder.

Subsequently, the fructose-containing seminal vesicle fluid alkalinizes the final ejaculatory fluid. The organs involved in the ejaculation process receive dense autonomic nerve supply, both sympathetic and parasympathetic, from the pelvic plexus.

The pelvic plexus is located retroperitoneally on either side of the rectum, lateral and posterior to the seminal vesicle It receives neuronal input from the hypogastric and pelvic nerves in addition to the caudal paravertebral sympathetic chain The sympathetic neurons play the predominant role in the ejaculation process. The role of the hypogastric plexus in emission is best demonstrated clinically by the loss of emission after non-nerve sparing para-aortic lymph node dissection for testicular cancer 35and induction of emission in paraplegic men through electrical stimulation of superior hypogastric plexus Input from genital stimulation is integrated at the neural sacral spinal level to produce emission The emission phase of ejaculation is also under a considerable cerebral control, and can be induced through physical or visual erotic stimulation Expulsion follows emission as the process of ejaculation climaxes, and refers to the ejection of semen through the urethral meatus.

The semen is propelled through the rhythmic contractions of the pelvic striated muscles in addition to the bulbospongiosus and ischiocavernosus muscles To achieve antegrade semen expulsion, the bladder neck remains closed, whereas the external urethral sphincter is open. The external sphincter and the pelvic musculature are under somatic control; however, there is no evidence that voluntary control plays a role in the expulsion process The exact trigger for expulsion is unknown.

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It has been suggested that a spinal center is triggered during emission of seminal fluid into the prostatic urethra However, there is mounting evidence through clinical and experimental studies to suggest that this is not the case. This, in addition to the identification of spinal generator for ejaculation SGE in rats, led to the postulation that the process of expulsion is a continuum of the process initiated through emission, after reaching a certain spinal activation threshold 30 Ejaculation is heavily controlled by the nervous system.

Figure 2 summarizes the reflex circuit necessary to elicit ejaculation. Reflex circuit needed to establish ejaculation. The main sensory input from the penis comes from the dorsal nerve of the penis, which transmits sensation from the glans, prepuce, and penile shaft. It transmits signals to the upper and lower segments of the sacral spinal cord The glans contains encapsulated nerve endings, termed Krause-Finger corpuscles, whereas the remaining penile shaft contains free nerve endings.

Stimulation of these corpuscles potentiated by stimulation from other genital areas, such the perineum, testes, and penile shaft, play an important role in the ejaculation process A secondary afferent route is through the hypogastric nerve, which runs through the paravertebral sympathetic chain to enter the spinal cord through the thoracolumbar dorsal roots The sensory afferents terminate in the medial dorsal horn and the dorsal gray commissure of the spinal cord The efferent peripheral nervous system constitutes of sympathetic, parasympathetic, and motor nervous components The soma of the preganglionic sympathetic cell bodies involved in ejaculation are located in the intermedio-lateral cell column and in the central autonomic region of the thoracolumbar segments TL1 The preganglionic sympathetic fibers emerge from the ventral roots of the spinal cord and travel through the paravertebral sympathetic chain to relay either directly through the splanchnic nerve, or through relaying first in the celiac superior mesenteric ganglia and then through the intermesenteric nerve, to the inferior mesenteric ganglia The hypogastric nerve then emanates from the inferior mesenteric ganglia to join the parasympathetic pelvic nerve to form the pelvic plexus, which then sends fibers to the ejaculation structures The preganglionic parasympathetic cell bodies are located in the sacral parasympathetic nucleus.

The sacral parasympathetic nucleus neurons travel then in the pelvic nerve to the post-ganglionic parasympathetic cells located in the pelvic plexus. An additional spinal center is the SGE located in laminae X and VII of L3-L4 spinal segments The SGE contains spinal interneurons called lumbar spinothalamic cells, which project fibers to the parvocellular subparafascicular nucleus of the thalamus in addition to preganglionic sympathetic and parasympathetic neurons innervating the pelvis The SGE stimulation elicits a complete ejaculatory response resulting in collection of motile spermatozoa in anesthetized rats Further research on the SGE spinal center is still needed, and it is unclear whether it contains other cells than lumbar spinothalamic cells.

Sensory and motor areas in the brain play an important role in the ejaculation, which requires a highly coordinated and integrated central process. The study by Holstege et al. Furthermore, specific areas in the brain have been involved in the ejaculation process, as demonstrated in animal immunohistochemical studies examining Fos protein pattern of expression 53 - 56and confirmed using a serotonin 1A subtype receptor agonist proejaculatory pharmacologic agent in rats These are discrete areas within the posteromedial bed nucleus of stria terminalis, the parvicellular part of the subparafascicular thalamus, the posterodorsal preoptic nucleus, and the posterodorsal medial amygdaloid nucleus.

There are reciprocal connections that link those areas to the MPOA of the hypothalamus, a brain area with a well-established role in controlling sexual behavior as demonstrated by anatomical and functional studies 5455 Electrical or chemical stimulation of the MPOA elicited ejaculation 59 - 62whereas an MPOA lesion was shown to abolish both phases of ejaculation No direct connections of MPOA to the spinal centers for ejaculation were found on neuroanatomical studies; however, there are projections of MPOA to other regions in the brain involved in ejaculation, such as PVN, the periaqueductal gray, and the paragigantocellular nucleus nPGi 64 - The PVN projects to pudendal motor neurons located in the L5-L6 spinal segment in addition to autonomic preganglionic neurons in the lumbosacral spinal cord in rats 4567 It also projects to nPGI in the brainstem Bilateral lesions of the PVN with N-methyl-D-aspartate NMDA results in a one-third reduction of the seminal ejaculate material weight The parvicellular part of the subparafascicular thalamus was found to send projections to bed nucleus of stria terminalis, medial amygdala MeAand MPOA 7172 and receives input from lumbar spinothalamic cells The precise role of these regions is still unclear but they are likely involved in relaying genital signals to MPOA 53 The brainstem regions nPGI and periaqueductal gray have recently received increasing attention.

The nPGI nucleus likely plays an inhibitory role in ejaculation as evidenced through the urethrogenital reflex experimental model, a rat model for the expulsion phase of ejaculation 73 Using the same model, the periaqueductal gray was found to be important for the ejaculation process, likely by acting as a relay between MPOA and nPGI Midbrain structures have a significant role in ejaculation; however, much is still unknown about their exact role and further research is needed.

Figure 3 summarizes the putative brain structures involved in ejaculation. Putative brain structures involved in ejaculation. Reprinted with permission from Clement P, Giuliano F. In: Mulhall JP, Incrocci L, Goldstein I, Rosen RC, eds.

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Cancer and sexual health. Many neurotransmitters are involved in the ejaculation process. Defining the exact role of these neurotransmitters is difficult given the variety of sexual parameters affected, the different sites of action within the spinal and the supraspinal pathways, and the presence of multiple receptor types. Some of the molecules that received special attention for their role in ejaculation are mentioned later. Dopamine is known to be important for normal male sexual response 76 Two families of dopamine receptors exist, D1-like D1 and D5 receptors and D2-like D2, D3, and D4 receptors In rats, D2-like receptors are known to stimulate ejaculation 7879and trigger ejaculation even in anesthetized rats 80 Systemic injection of the D3 receptor agonist 7-OH-DPAT has been shown to trigger ejaculation in rats without affecting arousal 82 It also triggers ejaculation in anesthetized rats when injected directly into the cerebral ventricles or MPOA with the effect being specifically reversed by the D3, not the D2 antagonist 84 The D3 receptor blockage has been shown to inhibit the expulsion phase of ejaculation and lengthen ejaculation latency in rats Evidence suggests that serotonin 5HT inhibits ejaculation Selective serotonin reuptake inhibitors increase 5HT tone resulting in impairment of ejaculation, which led to their clinical use in premature ejaculation.

This inhibitory effect is likely to occur in the brain 88as 5HT effect on ejaculation in the spine is likely stimulatory The amphetamine derivative p-chloroamphetamine leads to a sudden release of 5HT in synaptic clefts triggering ejaculation in anesthetized rats with complete spinal cord lesion Intrathecal serotonin or selective serotonin reuptake inhibitor injection leads to enhancement of the expulsion phase of ejaculation There are 14 receptor subtypes for 5HT, with 1A, 1B, and 2C being the ones involved in ejaculation It is difficult to designate one influence for each receptor subtype, as each receptor could either activate or inhibit ejaculation depending on its location within the central nervous system The role of NO in ejaculation has received special attention after the introduction of type-5 phosphodiesterase PDE5 inhibitors and using them for premature ejaculation.

Nitric oxide has an inhibitory role on the ejaculation process 1.

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Centrally, intrathecal sildenafil results in elevation of NO and cyclic guanosine monophosphate levels in MPOA causing a decreased peripheral sympathetic tone and inhibition of ejaculation N -Nitro- l -arginine methyl-ester injection, an NO synthase inhibitor, was shown to increase the number of seminal emissions and reduce latency to first seminal emission in rats Peripherally, nitronergic innervation and NO synthase were found in the seminal vesicle, vas deferens, prostate, and urethra 93 - Therefore, drugs such as PDE5 inhibitors or NO donors are associated with reduced seminal vesicle contraction and inhibit seminal emission The administration of NO inhibitors, such as l -nitroarginine-methylester, diminishes human seminal vesicle contraction 98inhibits vasal contraction in guinea pigs 99and decreases latency to ejaculation in rats Furthermore, reduced latency to emission was found in knockout mice for the gene encoding endothelial NO synthase compared with their wild-type counterparts Although male sexual function is heavily regulated by the hormonal system, there are few clinical studies performed to evaluate hormonal regulation of ejaculation, and the knowledge about hormonal effect on ejaculation is still lacking.

We discuss some of the studies examining the effect of different hormones on ejaculation. Oxytocin is an oligopeptide synthesized in the supraoptic and PVN of the hypothalamus and released from the posterior pituitary gland.

Pharmacologic oxytocin administration in humans and animals results in increased ejaculated spermatozoaconfirming that oxytocin has a role in male genital tract motility. It was specifically found to augment powerful epididymal contractions and sperm motilityan important effect blunted by pretreatment with the oxytocin antagonist des Gly-NH2d CH2 5-[d-Tyr2,Thr4] ornithine vasotocin Peripheral oxytocin receptors were found to be highly expressed in the epididymis and tunica albuginea in smooth muscles more than epithelial cellsand to a lesser extent in the vas deferens and seminal vesicle Oxytocin has a synergistic action on the epididymis with endothelin-1, where they augment epididymal contraction and propel spermatozoa forwar Injection of oxytocin into the cerebral ventricles in male rats facilitated ejaculation by shortening the ejaculation latency and postejaculatory refractory periodswhereas these effects were curbed using the oxytocin receptor antagonist d CH2 5-Tyr Me -[Orn8]vasotocin injected into the cerebral ventricles Despite these encouraging findings and some anecdotal evidence suggesting that intranasal oxytocin can facilitate orgasm in an anorgasmic malea double-blind placebo-controlled clinical study failed to demonstrate an effect of intranasal oxytocin on sexual behavior.

Hyperprolactinemia has a marked inhibitory effect on male sexual desire Some investigators have hypothesized that a low PRL level is a cause of premature ejaculation, where PRL levels were similarly low in those men with lifelong or acquired premature ejaculation Further research is needed on this issue.

The relationship between thyroid hormonal abnormalities and ejaculatory dysfunction has been well documented - In rats, l -thyroxin administration has been shown to increase bulbospongiosus contractile activity and seminal vesicle contraction frequency Clinically, the prevalence of suppressed TSH, which is a marker of hyperthyroidism, was found to be twofold higher in patients with premature ejaculation than in patients who reported normal ejaculatory timing Another single-center prospective study by Cihan et al.

It also identified a positive correlation of TSH with intravaginal ejaculation latency time. Ozturk et al. However, Waldinger et al.

A meta-analysis by Corona et al. They also showed an increase in intravaginal ejaculation latency time by These findings suggest that thyroid hormones do not only affect the ankle reflex, but also the ejaculatory reflex, and screening patients with ejaculatory dysfunction for thyroid hormone abnormalities is warranted Cortisol F levels in several animal studies were found to be elevated during arousal and ejaculation - In horses and donkeys, F was elevated 30 minutes after ejaculation, with unknown significance of this finding In addition, F levels were sharply elevated after electroejaculation in several anesthetized animal studies In humans, however, there was no change in F levels whether during sexual stimulation or orgasm - Although hypercortisolism in men was associated with reduced libido, no effect was identified on orgasm or ejaculation Replacement of F in Addison disease was associated with improvement in overall sexual function including orgasm Data in humans are still too preliminary to draw final conclusions, and further research is needed.

Estradiol plays an important role in the regulation of the emission phase of ejaculation through the regulation of epididymal contractility, luminal fluid reabsorption, and sperm concentration This role in the epididymis is the reason for recommending Tamoxifen as a first-line treatment for idiopathic oligospermia by the World Health Organization Finkelstein et al.

Testosterone, through its central and peripheral androgen receptors, has a well-known role on male sexual function, particularly on libido Low T levels are associated with delayed ejaculation, whereas high levels were associated with premature ejaculation This is likely because the emission phase of the ejaculation relies on the NO-PDE5 system, which is influenced by T Testosterone facilitates the control of the ejaculatory reflex through its androgen receptors in the MPOA and other areas in the central nervous system Furthermore, pelvic floor muscles involved in ejaculation are androgen dependent There are likely multiple mechanisms involved in T action and further research is needed to identify specific targets for treatment in the ejaculatory reflex.

Table 2 summarizes the neurochemical and hormonal regulation of ejaculation. In conclusion, ejaculation is a complex process involving several anatomical structures and under extensive neurochemical and hormonal regulation. Orgasm, although associated with ejaculation, is a distinct physiological process, different from ejaculation. Many cts of these physiological processes are still unknown and further research is needed to identify treatments for ejaculatory dysfunction.

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Journal List HHS Author Manuscripts PMC Fertil Steril. Author manuscript; available in PMC Jun 7. PMCID: PMC NIHMSID: NIHMS Amjad AlwaalM. BreyerM. LueM. Amjad Alwaal a Department of Urology, King Abdulaziz University, Jeddah, Saudi Arabia b Department of Urology, University of California, San Francisco, California Find articles by Amjad Alwaal. Benjamin N. Breyer b Department of Urology, University of California, San Francisco, California Find articles by Benjamin N. Tom F. Lue b Department of Urology, University of California, San Francisco, California Find articles by Tom F.

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Abstract Orgasm and ejaculation are two separate physiological processes that are sometimes difficult to distinguish. Keywords: Erectile function, male sexual function, ejaculation, orgasm. Open in a separate window. FIGURE 1.

TABLE 1 Summary of the functional anatomy of the male genital organs. Forms the rete testis inside the testis mediastinum Rete testis: gives rise to efferent ductules Epididymis Posterior and superior to the testicle Composed of head, body, and tail Efferent ductules unite to form the convoluted duct of the epididymis Becomes the vas deferens at the end of the tail Vas deferens Muscular tube; typically 45 cm long and has a 2.

Cerebral Control Cerebrally controlled penile erections are induced through erotic visual stimuli or thoughts. Autonomic Control Parasympathetic stimulation is the main mediator for penile tumescence, although central suppression of the sympathetic nervous system also plays a role. Molecular Mechanisms The penis at baseline is in a flaccid state maintained by the contraction of corporal smooth muscles and constriction of cavernous and helicine arteries leading to moderate state of hypoxia with partial pressure of oxygen of mm Hg Emission The first step in the emission phase is the closure of bladder neck to prevent retrograde spillage of the seminal fluid into the bladder.

Expulsion Expulsion follows emission as the process of ejaculation climaxes, and refers to the ejection of semen through the urethral meatus. FIGURE 2.

Peripheral Nervous System Afferents The main sensory input from the penis comes from the dorsal nerve of the penis, which transmits sensation from the glans, prepuce, and penile shaft. Efferents The efferent peripheral nervous system constitutes of sympathetic, parasympathetic, and motor nervous components Brain network Sensory and motor areas in the brain play an important role in the ejaculation, which requires a highly coordinated and integrated central process.

FIGURE 3. Dopaminergic Control Dopamine is known to be important for normal male sexual response 76 Serotonergic Control Evidence suggests that serotonin 5HT inhibits ejaculation Nitric Oxide The role of NO in ejaculation has received special attention after the introduction of type-5 phosphodiesterase PDE5 inhibitors and using them for premature ejaculation.

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Oxytocin Oxytocin is an oligopeptide synthesized in the supraoptic and PVN of the hypothalamus and released from the posterior pituitary gland. Prolactin Hyperprolactinemia has a marked inhibitory effect on male sexual desire Thyroid Hormones The relationship between thyroid hormonal abnormalities and ejaculatory dysfunction has been well documented - Glucocorticoids Cortisol F levels in several animal studies were found to be elevated during arousal and ejaculation - Estrogens Estradiol plays an important role in the regulation of the emission phase of ejaculation through the regulation of epididymal contractility, luminal fluid reabsorption, and sperm concentration Androgens Testosterone, through its central and peripheral androgen receptors, has a well-known role on male sexual function, particularly on libido TABLE 2 Neurochemical and hormonal regulation of ejaculation.

Footnotes A. Sheu G, Revenig LM, Hsiao W. In: Mulhall JP, Hsiao W, editors. Men's sexual health and fertility. Springer Science; New York: Bella AJ, Shamloul R. Functional anatomy of the male sex organs.

In: Mulhall JP, Incocci L, Goldstein I, Rosen R, editors. Meacham R, Lipshultz L, Howards S. Male infertility. In: Gillenwater JY, Grayhack JT, Howards S, Duckett JW, editors.

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Adult and pediatric urology. Mosby; St. Louis: Hinman F. Normal surgical anatomy. In: Thomas Thomas AJ, Nagler HN, editors. Atlas of surgical management of male infertility. Romanes G. The pelvis and perineum.

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In: Romanes G, Cunningham D, editors. Cunningham's manual of practical anatomy. Tang Y, Rampin O, Calas A, Facchinetti P, Giuliano F. Oxytocinergic and serotonergic innervation of identified lumbosacral nuclei controlling penile erection in the male rat. Danjou P, Lacomblez L, Warot D, Puech AJ. Assessment of erectogenic drugs by numeric plethysmography.

J Pharmacol Methods.

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Clark JT, Smith ER, Davidson JM. Testosterone is not required for the enhancement of sexual motivation by yohimbine. Physiol Behav. Andersson KE. Mechanisms of penile erection and basis for pharmacological treatment of erectile dysfunction.

Pharmacol Rev. Lue TF, Zeineh SJ, Schmidt RA, Tanagho EA. Neuroanatomy of penile erection: its relevance to iatrogenic impotence. J Urol. Paick JS, Lee SW. The neural mechanism of apomorphine-induced erection: an experimental study by comparison with electrostimulation-induced erection in the rat model.

Chapelle PA, Durand J, Lacert P. Penile erection following complete spinal cord injury in man. Br J Urol. Dean RC, Lue TF. Physiology of penile erection and pathophysiology of erectile dysfunction. Urol Clin North Am. Courtois FJ, Charvier KF, Leriche A, Raymond DP. Sexual function in spinal cord injury men. Assessing sexual capability. Sattar AA, Salpigidis G, Schulman CC, Wespes E. Relationship between intrapenile O2 lever and quantity of intracavernous smooth muscle fibers: current physiopathological concept.

Acta Urol Belg. Prieto D. Physiological regulation of penile arteries and veins. Int J Impot Res. Pharmacology of penile erection. Walsh MP. The Ayerst Award Lecture Calcium-dependent mechanisms of regulation of smooth muscle contraction.

Biochem Cell Biol. Hurt KJ, Musicki B, Palese MA, Crone JK, Becker RE, Moriarity JL, et al. Akt-dependent phosphorylation of endothelial nitric-oxide synthase mediates penile erection. Proc Natl Acad Sci. Levin R. Physiology of orgasm. Masters W, Johnson V. Human sexual response. Little Brown; Boston: Levin R, editor.

Heart rate responses can be used to differentiate simulated from real orgasms in the human male: a pilot study. Proceedings of the first conference on orgasm. VRP Publishers; Bombay: Gerstenberg TC, Levin RJ, Wagner G. Erection and ejaculation in man-assessment of the electromyographic activity of the bulbocavernosus and ischiocavernosus muscles.

Sex Relat Ther. Holstege G, Georgiadis JR, Paans AM, Meiners LC, van der Graaf FH, Reinders AS. Brain activation during human ejaculation. J Neurosci. Hite S. In one instance she is alleged to have performed a sex act on him in her car - and again in a classroom.

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On another occasion the boy claims she contacted him at midnight and drove to his grandparents' house so he could sneak out to be with her.

Zamora allegedly responded: "I know baby! I want you every day with no time limit If I could quit my job and have sex with you all day long, I would. The boy's distraught parents are suing Liberty Elementary School District claiming that school authorities did not do enough to protect their son. They are seeking ?1. We pay for your stories! Do you have a story for The Sun Online news team? Email us at tips the-sun. uk or call You can WhatsApp us on We pay for videos too.

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