ANATOMY

» FEMALE GENITAL ANATOMY
» Clitoris
» Corpus Spongiosum
» Peri-Urethral Glands
» Vagina
» Muscularis
» Adventitia
» Arterial Supply
» Innervation
» Urethra
» G-Spot
» Halban's Fascia
» Cervix
» Uterus

» MALE GENITAL ANATOMY
» Penis
» Arterial Supply
» Intracorporal Circulation
» Venous Drainage
» Lymphatic Drainage
» Nerves
» Cavernous Nerve Neurovascular Bundle


Female Genital Anatomy

There are multiple anatomical structures that comprise the internal and external female genital tract such as the clitoris, labia minora and corpus spongiosum (vestibular) erectile tissue, peri-urethral glans, urethra, G-spot, Halban's fascia, anterior fornix erogenous zone, pubococcygeus muscle and cervix. There are also multiple non-genital peripheral anatomic structures involved in female sexual responses such as salivary and sweat glands, cutaneous blood vessels and nipples.

The vagina consists of a tube of autonomically-innervated smooth muscle (longitudinal outer, inner circular layer) lined by stratified squamous epithelium and a sub-dermal layer rich in capillaries. The vaginal wall consists of an inner glandular mucous type stratified squamous cell epithelium supported by a thick lamina propia. This epithelium undergoes hormone-related cyclical changes including slight keratinization of the superficial cells during the menstrual cycle. The smooth muscles of the muscularis lie deep to the epithelium. There is a deeper surrounding fibrous layer above the muscularis which provides structural support to the vagina, and is rich is collagen and elastin, to allow for expansion of the vagina during sexual stimulation. Three sets of skeletal muscles surround the vagina including the ischiocavernosum, bulbocavernosus, transverse perinei and levator ani and pubococcygeus muscles.

The vulva includes the labia minora, labia majora, the clitoris, the urinary meatus, the vaginal opening, and the corpus spongiosum erectile tissue (vestibular bulbs) of the labia minora. The labia majora are fatty folds covered by hair-baring skin that fuses anteriorly with the mons verenis, or anterior prominence of the symphysis pubis, and posteriorly with the perennial body or posterior commissure. The labia minora are smaller folds covered by non-bearing skin laterally and by vaginal mucosa medially, that fuses anteriorly to forms the prepuce of the clitoris, and posteriorly in the fossa navicularis.

The corpora cavernosa of the clitoris measures up to 5 inches in length. The body of the clitoris consists of two paired erectile chambers composed of endothelial-lined lacunar spaces, trabecular smooth muscle and trabecular connective tissue (collagen and elastin) surrounded by a fibrous sheath, the tunica albuginea. The arteries include the dorsal and clitoral cavernosal arteries, which arise from the iliohypogastric pudendal bed. The autonomic efferent motor innervation occurs via the cavernosal nerve of the clitoris arising from the pelvic and hypogastric plexus.

CLITORIS
The clitoris is formed from the tubercle of the undifferentiated common tissue anlagen in the embryo. The clitoris consists of a midline shaft lying in the medial sagittal plane about 2-4 cm long and 1-2 cm wide which bifurcates internally into paired curved crura 5-9 cm long (attached to the under surface of the pubic symphisis). The clitoris is capped externally with a glans about 20-30mm long with a similar diameter. The glans is covered by a clitoral hood formed in part by the fusion of the upper part of the two labia minora. The erectile tissue of the clitoral shaft consists of two parallel corpora cavernosa surrounded by a fibrous sheath (tunica albuginea). The clitoral cavernosal erectile tissue consists of smooth muscle and connective tissue. The percentage of clitoral cavernosal smooth muscle in age group of 6 months to 15 years was 65 ± 1.5, in 44 to 54 years was 50 ± 1.2 and in 55 to 90 years was 37 ± 1.3 (ANOVA, p=0.0001). These studies, which revealed a strong link between increase in age and decreased clitoral cavernosal smooth muscle fibers, illustrate that aging women undergo histologic changes in clitoral cavernosal erectile tissue which may play an as yet undetermined pathophysiology in age-associated female sexual dysfunction. Because the shaft and the glans of the clitoris have no subalbugineal layer between the erectile tissue and the tunica albuginea the organ becomes tumescent or swollen with effective sexual stimulation but does not become erect or rigid. Nevertheless, human clitoral erectile tissue has the capacity to develop drug-induced priapism that responds by detumescing following administration of a-adrenergic agonists. The corpora cavernosa of the shaft do not extend into the glans.

Although the erogenous function of this organ has been known since antiquity, remarkably, the detail of its highly vascular anatomical structure is still in dispute. It is formed from the tubercle of the undifferentiated common tissue anlagen in the embryo. In the presence of androgens this develops into the penis while in their absence the clitoris is formed. Current dissections of adult female human cadavers have been interpreted to indicate that the organ is a triplanar complex of erectile tissue with a midline shaft lying in the medial sagittal plane about 2-4 cm long and 1-2 cm wide which bifurcates internally into paired curved crura 5-9 cm long (attached to the under surface of the pubic symphisis) and externally is capped with a glans about 20 -30mm long with a similar diameter.

The erectile tissue of the shaft consists of two parallel corpora cavernosa surrounded by a fibrous sheath (tunica albuginea) and the whole structure is covered by a clitoral hood formed in part by the fusion of the upper part of the two labia minora while the lower parts meet beneath the clitoris. The clitoral cavernosal erectile tissue consists of smooth muscle and connective tissue. Tufan et al utilized computer assisted histomorphometric image analysis to determine the age-associated changes in clitoral cavernosal content of smooth muscle and connective tissue. Human clitorises were obtained from fresh cadavers (age: 11 to 90 years) and from patients undergoing clitoral surgery (age: 6 months to 15 years). The percentage of clitoral cavernosal smooth muscle in age group of 6 months to 15 years was 65 ± 1.5, in 44 to 54 years was 50 ± 1.2 and in 55 to 90 years was 37 ± 1.3 (ANOVA, p=0.0001). These studies, which revealed a strong link between increase in age and decreased clitoral cavernosal smooth muscle fibers, illustrate that aging women undergo histologic changes in clitoral cavernosal erectile tissue which may play an as yet undetermined pathophysiology in age-associated female sexual dysfunction.

The paired, so-called vestibular (vaginal) bulbs of erectile tissue, which have normally been illustrated on either side of the vagina practically as if in the labia minora, are actually closely applied anteriorly on either side of the urethra. In the male the corpus spongiosum is a single tubular structure of erectile tissue that ensheaths the urethra ending internally as the penile bulb and externally as the penile glans pierced by the urinary meatus. The location and extent of the female corpus spongiosum is contentious. It has been described as being the vascular tissue surrounding the female urethra, as the bilateral vestibular bulbs and as the tissue between the bladder and anterior vaginal wall (Halban's fascia). Most authors claim that the clitoris has no spongiosus tissue. However, the extension of the corpus spongiosus tissue into the clitoris has been described by van Turnhout, Hage & van Diest from their dissections and histology of the adult female cadaver. They observed that the bilateral vestibular bulbs unite ventral to the urethral orifice to form a thin strand of spongiosus erectile tissue connection (pars intermedia) that ends into the clitoris as the glans. The corpora cavernosa of the shaft do not extend into the glans.

Because the shaft and the glans of the clitoris have no subalbugineal layer between the erectile tissue and the tunica albuginea the organ becomes tumescent or swollen with effective sexual stimulation but does not become erect or rigid. Nevertheless, human clitoral erectile tissue has the capacity to develop drug-induced priapism which responds by detumescing following administration of a-adrenergic agonists. The earliest attempt to characterise the possible mechanism(s) by which the crura and vestibular bulbs changed from the flaccid to the tumescent state was published first in diagrammatic form by Danesino & Martella in Italian. Their working hypothesis, based on the early mechanisms suggested for penile erection, was that during sexual excitement smooth muscle polsters ("cushions") in the arteries supplying the two vestibular bodies became relaxed. Those polsters in the draining veins became contracted as did those in the a-v anastomoses. This diverted blood into the lacunae, filling them and creating tumescence. For detumescence, the arterial polsters contracted while those in the veins and a-v anastomoses relaxed, reducing the flow to the lacunae and allowing the blood restricted in them to flow away. Despite this mechanism being published in English for over 23 years, no independent confirmation of either the mechanism or the polsters in the female arteries and veins have yet appeared. It must be regarded as a speculative working hypothesis.

The finding that human clitoral tissue has nitric oxide synthase (NOS) present in nerves and blood vessels suggests that nitric oxide (NO) may be involved in controlling clitoral blood flow as it does in the penis. Park et al have further examined the possible role for nitric oxide in the regulation of human clitoral corpus cavernosum smooth muscle contractility. In this study, cGMP and cAMP hydrolysis by phosphodiesterases were characterized in the high speed supernatant fraction (cytosol) and in partially purified preparations of human clitoral corpus cavernosum smooth muscle cells. Sildenafil was found to inhibit PDE type 5 cGMP-hydrolytic activity, in the crude extract (Ki=7 nM) and in partially purified preparations (Ki=5-7 nM) in a competitive fashion. Synthesis of cyclic nucleotides was also carried out in intact cells in culture in response to sodium nitroprusside (NO donor) and forskolin (direct adenylate cyclase activator). Intracellular cGMP was increased by 35% in presence of sildenafil (10nM) in intact cells in culture. The results of this study support a role for nitric oxide in regulation of human clitoral corpus cavernosum smooth muscle tone.
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CORPUS SPONGIOSUM
The paired corpus spongiosum, or vestibular bulbs of erectile tissue practically in the labia minora but are actually more closely applied anteriorly on either side of the urethra. The extension of the corpus spongiosus tissue into the clitoris has been described. The bilateral vestibular bulbs unite ventral to the urethral orifice to form a thin strand of spongiosus erectile tissue connection (pars intermedia) that ends into the clitoris as the glans.
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PERI-URETHRAL GLANDS
Unlike the clitoral glans the male glans is pierced by the urethra. It has been suggested that there are really two glans in the female, a clitoral glans) and a glans that surrounds the urethra (periurethral glans). The periurethral glans is defined as the triangular area of mucous membrane surrounding the urethral meatus from the clitoral glans to the vaginal upper rim or caruncle. The periurethral glans is mobile and has been shown to be pushed into and pulled out of the vagina by penile thrusting during coitus.
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VAGINA
The vagina is a fibromuscular tube that connects the uterus with the vestibule of the external genitalia. It acts in transport of sperm to the uterus and in expulsion of the newborn.

The vagina is a potential space with its anterior and posterior walls usually in apposition. The vaginal walls can be easily separated because their surfaces are normally "just moist", lubricated by a basal vaginal fluid (approximately 1ml). In the intermenstruum, basal vaginal fluid can consist of multiple secretions that collect in the vagina from peritoneal, follicular, tubal, uterine, cervical, vaginal, Bartholin's and Skene's gland sources. The vaginal wall consists of three layers: the mucosa, muscularis and adventitia.

The vagina has three layers: the internal mucosal layer, the intermediate muscularis layer and the external adventitial layer.

The internal mucosal layer: had traverse folds, or rugae. The epithelium is nonkeratinized stratified squamous epithelium. The epithelium has no glands so there is no mucus secretion.

The mucosa consists of a thick stratified squamous epithelium devoid of glands. The superficial cells of the epithelium undergo hormone-related cyclical changes such as slight keratinization, or increased glycogen production during the menstrual cycle. In the sexually unstimulated state, vaginal fluid has a higher K+ and lower Na+ concentration compared to plasma throughout the phases of the menstrual cycle. The actual basal vaginal transudate that percolates through the vaginal epithelium from the plasma circulating in the capillary tufts supplying the epithelium is modified by the limited Na+ lumen-to-blood reabsorptive transport capacity of the vaginal epithelial cells. The reabsorption of Na+ by the vaginal epithelium is presumably the ionic driving force for the reabsorption of the vaginal fluid and maintains its level under basal conditions to the "just moist"condition. Autologous plasma placed in a subject's vagina for up to 5 hours shows increased K+ and decreased Na+ concentrations indicating that the epithelium is capable of undertaking such ion transfer in vivo. The basal lubrication is usually not sufficient to allow painless penile penetration and thrusting so an enhancement of the lubrication is essential for coitus.

The lamina propria has many thin-walled blood vessels that contribute to diffusion of vaginal fluid across the epithelium.

The lamina propria of the mucosa contains many elastic fibers as well as a dense network of blood vessels, lymphatic and nerve supply. Transudate from these blood vessels, combined with cervical mucus, provide lubrication during sexual arousal and intercourse. Sexual arousal induces a neurogenic transudate that filters through the labyrinthine pathways of the epithelium and saturates its limited Na+ reabsorptive capacity. It appears within seconds of successful sexual arousal initially on the surface of the vagina as bead-like droplets that then coalesce to create a lubricative film that can partially decrease the acidity of the vaginal basal fluid. The smooth, slippery quality of the formed fluid is probably due to its pick up of sialoproteins coating the vaginal epithelium from the cervical secretion. On sexual arousal the blood supply to the vaginal epithelium is rapidly increased by neural innervation via the sacral anterior nerves S2-S4 and at the same time the venous drainage is probably reduced creating vasocongestion and engorgement with blood. Vaginal lubrication during sexual arousal does not occur from any increased secretion of vaginal glands (nonexistant), cervical fluid or from Bartholin's glands. The enhanced blood flow is activated by the VIPergic innervation of the large vessels supplying the epithelium and the transudation possibly aided by the CGRP (calcitonin gene regulating peptide) enhanced permeability of the capillary tufts. NPY, neuropeptide Y, a known vasoconstrictor, may be involved in constricting the venous drainage. There appears to be very little NOS in the blood vessels of the pre-menopausal vagina and none in the post-menopausal. After orgasm or the cessation of sexual stimuli, the continuous lumen-blood transfer of Na+ by the epithelium slowly reabsorbs the excess fluid of the neurogenic transudate by osmotic drag and resets the vagina back to its just moist basal state.

The vaginal epithelium responds to hormonal changes. Glycogen, stored in the epithelial cells, reaches maximal levels at ovulation after which time the glycogen-rich superficial layer of epithelium is shed. Breakdown of the glycogen by bacteria in the vagina produces lactic acid, causing the vaginal environment to have an acid pH of about 3. This inhibits growth of other bacteria, bacterial pathogens and fungus. It also limits the time in which sperm can survive in the vagina. The intermediate muscularis layer: inner circular and outer longitudinal which is continuous with the corresponding layer in the uterus.
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MUSCULARIS
The muscularis, consists of autonomically innervated smooth muscle fibers arranged into an outer longitudinal and inner circular layer. In the basal or sexually quiescent state the smooth muscle of the vagina is active especially perimenstrually when it contracts periodically to expel the uterine/vaginal contents. These vaginal smooth muscle contractions are normally not consciously recognized. They only become obvious if they reach painful, spasmotic levels (dysmenorrheic pain). During arousal to orgasm, there is an increasing vaginal luminal pressure. The smooth muscle layers contain a great variety of classical and peptidergic transmitters including 5HT, nor-epinephrine, acetyl choline, dopamine, VIP, NPY, GRP, TRH, CGRP, somatostatin, substance P, oxytocin, cholecystokinin (CCK) and relaxin, but the exact function of each neurotransmitter is unknown.
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ADVENTITIA
The adventitia is rich in collagen and elastic, provides structural support to the vagina, and allows for expansion of the vagina during intercourse and childbirth. Surrounding the adventitia are three sets of powerful pelvic striated muscles (1, superficial- ischiocavernosus and bulbocavernosus; 2, the transverse perineii and 3, deep- the levator ani forming the pelvic diaphragm across the anterior of the pelvis of which the largest medial portion is classified as the pubococyggeus). At orgasm a series of pelvic, clonic, striated muscle contractions occur at approximately 0.8 second intervals which gradually get longer and the contractions weaker. They can last for 5-60 seconds. These contractions are concommittant with the subjective feeling of orgasm. Voluntary contractions of the pelvic striated muscles do not give a feeling of intense pleasure but are often used to enhance arousal. During sexual arousal up to orgasm, individual uterine contractions may occur while at orgasm a series occurs mediated by the sympathetic system via the hypogastric nerve. It has been proposed that sexual satiation in the female occurs only when the orgasmic uterine contractions are intense but there has been no quantitative studies to back up this speculation.
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ARTERIAL SUPPLY
The main arterial supply to the vagina arises from the three sources. The upper vagina is supplied by vaginal branches of the uterine artery. A branch of the hypogastric artery, the vaginal artery (also know as the inferior vaginal artery), supplies the middle vagina. Finally the middle hemorrhoidal and the clitoral arteries send branches to the distal vagina.
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INNERVATION
Autonomic efferent innervation to the upper two thirds of the vagina is through the utervaginal plexus. Autonomic efferent innervation to the upper two thirds of the vagina is through the utervaginal plexus, which contains both sympathetic and parasympathetic fibers. Sympathetic efferent fibers from the lumbar splanchnic nerves travel first through the superior hypogastric plexus, and then through the bilateral hypogastric nerves to reach the inferior hypogastric plexuses, and finally the uterovaginal plexus. Parasympathetic effernt input to the uterovaginal pelxus in from the pelvic splanchnic nerves. Nerves from the uterovaginal plexus travel within the uterosacral and cardinal ligaments, to supply the proximal two-thirds of the vagina. Autonomic efferent innervation to the lower vagina is carried through the pudendal nerve (S2, 3, 4) which reached the perineum through Alcock’s canal. Autonomic afferent fibers from the upper vagina travel through the pelvic splanchnic nerves to sacral spinal cord segments. Autonomic afferent fibers from the lower vagina leave the sacral spinal cord through the pudendal nerve. Somatic sensation exists primarily in the distal one third of the vagina and is also carried by the pudendal nerve to the sacral spinal cord.
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URETHRA
The female urethra is a short conduit (approximately 3-5 cm long) running from the base of the bladder and exiting in the periurethral glans area to the outside. For nearly its entire length it is surrounded by numerous venous/sinus channels that constitute the corpus spongiosum of the urethra. This submucosal vascular tissue contributes approximately one third of the normal urethral closing pressure and becomes further vasocongested during sexual arousal converting the urinary urethra into the sexual urethra. Scattered in the lining lumenal epithelium are cells containing 5-HT (serotonin). Their function is unknown but they are thought to be chemosensing or mechanoreceptor paracrine cells that release the 5-HT on being stimulated by stretch or luminal chemicals. In the animal urethra, 5-HT sensitizes neural mechanisms. It may be that the stretching or massage of the human female urethra by the thrusting penis during coitus causes the release of 5-HT from the urethral paracrine cells enhancing neural afferent input from the organ.
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G-SPOT
The G-spot may be considered a general excitable area along the whole length of the urethra running along the anterior vaginal wall. Grafenberg reported that the digital stroking of the anterior vagina along the urethra, especially in the region of the base of the bladder, sexually aroused female subjects greatly. In a number of women this area swelled up to the size of a kidney bean and projected into the vaginal lumen. The G-spot may be considered a general excitable area along the whole length of the urethra running along the anterior vaginal wall. When this is stimulated manually, the sexual arousal induced is almost immediate. This erotic sensitive area is located in closer relation to the bladder base than the urethra. The G spot represents that part of the urethra that contains the periglandular or paraurethral tissue, corresponding to the female equivalent of the prostate. These glands are present to a greater or lesser degree in about 90% of women. In some women, when stimulated sexually, a fluid secretion claimed to be dissimilar to urine or vaginal fluid can be produced which is controversially "ejaculated" from the urethra.
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HALBAN'S FASCIA
Halban's fascia is the space between the trigone of the bladder and the anterior part of the vaginal wall. It is filled with mesenchymal lamina, a fibro-elastic sheet made up of collagen, elastic and muscular fibres with a rich blood supply and a nerve supply with Krause bodies or pseudo-corpuscular nerve endings. On stimulation this space becomes vasocongested and creates an erotic pleasurable response.
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CERVIX
The cervix is a relatively insensitive structure. The cervix is a relatively insensitive structure with no erotogenic capabilities per se but it has been implicated by some authors as being important when jostled or buffeted by deep penile thrusting so that the uterus is pushed or rubbed against the peritoneal lining. This is claimed to create sexually pleasurable feelings but in others it creates discomfort. In some women who have had their cervix/uterus removed, a significant loss in sexual arousal and orgasm by coitus occurs. Penile-cervix contact rarely occurs. Penile-cervix contact is not observed in the missionary or face-to-face position but it could occur in the rear-entry sideways and rear-("doggie") positions. An intriguing aspect of the cervix is that it has the second highest concentration of VIP of the female genitals yet no function has been ascribed to the Vipergic innervation. Its possible role in the secretion of mucus by the infolded crypts of the cervical epithelium has not been investigated.
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UTERUS
The uterus, composed of three layers of smooth muscle, is situated in the lower pelvic part of the abdomen. The motility patterns of these organs, especially during sexual arousal to orgasm, have been studied infrequently, rarely measured and are poorly characterised [40, 58, 59, 69, 70, 71]. Their activity is usually monitored either by small luminal balloons or pressure catheters or by electrodes (needle or surface) that pick up the electromyographic activity (EMG) that increases when the muscles contract [69]. Because of the setting of the vagina, smooth muscles amongst striated, contraction of either or both will influence the pressure motility pattern obtained and the interpretation of the records often relies on the fact that at orgasm the striated motility dominates. No studies have been published that record simultaneously, but independently, both the striated and the smooth muscle activity thus allowing their interaction to be better interpreted and characterized.

In the basal or sexually quiescent state the striated muscle plays little or no role but the smooth muscle of the uterus and vagina is active especially peri-menstrually when it contracts periodically to expel the uterine/vaginal contents. These uterine and vaginal contractions are normally not consciously recognized [40, 71, 72]. They only become obvious if they reach painful, spasmotic levels (dysmenorrhoeic pain). During arousal to orgasm, the few records obtained show an increasing vaginal lumenal pressure [40]. At orgasm a series of pelvic, clonic, striated muscle contractions occur at approximately 0.8 second intervals which gradually get longer and the contractions weaker [58, 69]. They can last for 5-60 seconds. These contractions are concomitant with the subjective feeling of orgasm. Voluntary contractions of the pelvic striated muscles do not give a feeling of intense pleasure but are often used to enhance arousal. Few records of the intrauterine pressure exist and those that do could well be influenced by the size of the devices used to measure the intrauterine pressure (see Levin [40] for discussion). During sexual arousal up to orgasm, individual uterine contractions may occur while at orgasm a series occurs mediated by the sympathetic system via the hypogastric nerve. These have been implicated by some to be important in rapid sperm uptake into the uterus/fallopian tubes but this ignores the effect of vaginal tenting on cervical elevation from the ejaculated pooled semen (see previous section on cervix and Levin [59] for discussion). It has been proposed that sexual satiation in the female occurs only when the orgasmic uterine contractions are intense but there has been no quantitative studies to back up this speculation.

Two studies have reported that vaginal distention induced by rapid increases in volume by inflation of luminal balloons cause i) contractions of the bulbocavernous and ischiocavernous muscles [73] and ii) an increase in the velocity of clitoral arterial blood interpreted as an increase in flow [74]. The volume increase used was between 100 to 300ml although the normal volume of the human penis is about 70 ml. Thus penile volume per se would have little effect, but penile thrusting would stretch the vaginal walls and cause the reflex actions. The enhanced clitoral flow and its engorgement and introital tightness around the penile shaft are all features suggested to enhance the pleasure of coitus for both male and female partners.
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Male Genital Anatomy

The penis is composed of 3 spongy cylinders. The three cylinders consist of paired corpora cavernosa and a single corpus spongiosum. The crural (roots) of the corpora cavernosa attach at the under surface of the ischiopubic rami as two separate structures. Such anatomy prevents the erect penis from sinking into the perineum when faced with an axially-oriented vaginal compressive load during intercourse. This unique anatomic arrangement, however, unfortunately places the penile crus at great danger from crush injuries during blunt perineal trauma.

The tunica albuginea consists of layers of collagen that can accommodate a considerable degree of intracavernosal pressure prior to rupture. To function effectively, these fascial layers must provide the penis with a wall container capable of withstanding a high degree of rigidity and axial strength when erect, yet be supple when flaccid. The tunica must be able to elongate symmetrically and increase in girth with tumescence, assuring a straight erection. The tensile strength of the tunica is approximately 1200 - 1500 mmHg making this fascia one of the strongest in the body. Approximately 5% of the tunica is elastin that enables the penis to develop elongation. The average volume increase of the erect penis from the flaccid volume is 3-fold with a range from 1.7 - 5 fold. The mechanical properties of the tunica that allow for maximum volume changes of the erect penis are called tunica dispensability. Regions of the tunica with focal poor dispensability cause the erect penis to bend. This focal tunical abnormality in dispensability is called Peyronie’s disease.

The substance of the corpora cavernosa (erectile tissue) consists of numerous sinusoids (lacunar spaces) among interwoven trabeculae of smooth muscles and supporting connective tissue. The corpora cavernosa sinusoids are widely communicative and larger in the center of the corpora, having a Swiss-cheese appearance. This fact enables the blood within the penis to transfer easily from the top to the bottom of the corpora. This also enables the penis to have a common intracavernosal pressure and a common penile rigidity. The sinusoids are smaller in the periphery and have a grape-like appearance. Peripheral sinusoids have a greater individual surface area than central sinusoids. These characteristics aid in the passive process of corporal veno-occlusion by sub-tunical venule compression against the tunica albuginea. All lacunar spaces are lined with endothelial cells, thought previously to have only a slippery surface preventing blood clotting. Recent research has revealed that endothelial cells have secretory function and synthesize factors involved in the regulation of corporal smooth muscle tone.

The paired internal pudendal artery, a branch of the hypogastric artery is the main source of arterial blood supply to the penis.

The internal pudendal artery terminates when the artery divides into the scrotal and common penile artery.

The common penile artery defines the condition whereby all red blood cells in the artery somehow end up in the penis. The common penile artery branches into 3 arteries, the bulbourethral, the dorsal and the cavernosal arteries. The common penile artery has direct apposition to the ischiopubic ramus. This artery is therefore commonly injured during blunt perineal traumatic events such as falling onto the top tube of a bicycle.

The penis is innervated by autonomic (parasympathetic and sympathetic) and somatic (sensory and motor) nerves.

The cavernosal nerves are branches of the pelvic plexus that innervate the corpora cavernosa of the penis. Injury to this branch may occur during radical prostatectomy, during urethral surgery, such as internal urethrotomy and from electrocautery injury during transurethral surgery.

PENIS
The penis is the common output tract for urine and sperm. It is a structure that is under the control of a complex series of reflexes, neuronal and humoral control. It contains several aggregations of "cavernous" tissue that under certain conditions can become engorged with blood, causing the penis to become rigid. In this state the penis is capable of delivering the genetic material contained in the sperm during coitus.

The penile erectile apparatus consists of paired vascular spongy organs (corpora cavernosa) that are closely attached to each other except in the proximal third. The corpus spongiosum with the urethra is related to the ventral aspect of the penile shaft and expands distally to from the glans penis. The pendulous part of the penis if 4-6 inches (10.2-15.2 cm) long. The penile skin is continuous with that of the lower abdominal wall and continues over the glans penis to form the prepuce; it then folds itself to reattach at the coronal sulcus. The penile skin envelopes the shaft and can be moved freely over the erect organ. The underlying fascial layer or dartos fascia (Colles’ fascia) is continuous with Scarpa’s fascia of the lower abdominal wall; inferiorly, it continues as the dartos fascia of the scrotum and Colles’ fascia of the perineum and attaches to the posterior border of the perineal membrane. The superficial dorsal vein is seen in this layer of the fascia. Buck’s fascia is the deep layer of the penile fascia that covers both the corpora cavernosa and the corpus spongiosum in separate fascial compartments. Proximally, Buck’s fascia is attached to the perineal membrane; distally, it is tightly attached to the base of the glans penis at the coronal sulcus, where it fuses with the ends of the corpora. The ischiocavernosus and the bulbospongiosus muscles lie beneath Colles’ fascia, but superficial to Buck’s fascia, to which their intrinsic fascia is loosely attached. Buck’s fascia has a dense structure and is composed of longitudinally running fibers; it is firmly attached to the underlying tunica albuginea and encloses the deep dorsal vein, dorsal arteries and dorsal nerves.

The fundiform ligament is a thickening of the superficial penile fascia, deep to which is the suspensory ligament that is a continuity with Buck’s fascia. The attachment of the ligament to the pubic symphysis maintains the penile position during erection. Severance of this ligament will lead to a lower angulation of the penile shaft during erection.

The tunica albuginea forms a thick fibrous coat to the spongy tissue of the corpora cavernosa and corpus spongiosum. It consists of two layers, the outer longitudinal and the inner circular. The tunica albuginea becomes thicker centrally where it forms a groove to accommodate the corpus spongiosum. As the crura diverge proximally, the circular layer provides the support. The corpora are separated in the center by an intercavernous septum. The septum is incomplete distally, perforated on its dorsal margin by vertically orientated openings in the pectiniform septum that provides communication between the corpora. Along the inner aspect of the tunica albuginea, numerous flattened columns or sinusoidal trabeculae composed of fibrous tissue, elastin fibers and smooth muscle surround the endothelium-lined sinusoids or cavernous spaces. In addition, a row of structural trabeculae arises near the junction of the three corporal bodies and inserts in the wall of the corpora about the mid-plane of the circumference. The tunical albuginea provides a tough uniform backing for the engorged sinusoidal spaces. The tunical albuginea of the corpus spongiosum is thinner and contains smooth muscles that aid ejaculation. The glans is devoid of tunica albuginea. The corpus spongiosum becomes bulbous where it is covered by the bulbospongiosus to form the urethral bulb.

The ischiocavernosus is a paired muscle that arises from the inner surface of the inschial tuberosity and inserts into the medial and inferior surface of the corpora. These muscles increase penile turgor during erection beyond that attainable by arterial pressure alone. They are supplied by the perineal branch of the perineal nerve (S3-4).

The bulbospongiosus muscle invests the bulb of the urethra and distal corpus spongiosum. It arises from the central tendon of the perineum. The fibres run obliquely upwards and laterally on each side of the bulb and insert in the midline dorsally. The muscle is supplied by a deep branch of the perineal nerve and helps to empty the last few drops of urine and to ejaculate semen.
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ARTERIAL SUPPLY
The arterial supply to the erectile apparatus originates from superficial and deep arterial systems. The superficial arterial system arises as two symmetrically arranged vessels arising from the inferior external pudendal artery (a branch of the femoral artery). Each of these vessels divides inito a dorsolateral and ventrolateral branch that supply the skin of the shaft and prepuce. At the coronal sulcus there is a communication with the deep arterial system. The deep arterial system arises from the internal pudendal artery, which is the final branch of the anterior trunk of the internal iliac artery. This passes dorsal to the sacrospinous ligament at the level of the ischial spine and passes through Alcock’s canal. As it emerges, it divides into the perineal and penile arteries, running deep to the superficial transverse perineal muscle and pubic symphysis. It pierces the urogenital diaphragm meddial to the inferior ramus fo the ischium close to the bulb of the urethra and then divides into three branches—the bulbourethral artery, the urethral artery and the cavernous artery or deep artery of the penis; it terminates as the deep dorsal artery of the penis. An accessory internal pudendal artery may arise from the obturator, inferior vesical or superior vesical and may be damaged during radical prostatectomy in as many as 50% of patients. The bulbo-urethral artery supplies the bulb of the urethra, the corpus spongiosum and the glans penis. It may arise from the cavernous, dorsal or accessory pudendal arteries. The urethral artery commonly arises as a separate branch form the penile artery, but may arise from the artery to the bulb, the cavernous or the dorsal artery. It runs on the ventral surface of the corpus spongiosum beneath the tunica albuginea.

The cavernous artery (deep artery of the penis) usually arises form the penile artery, but may originate from the accessory pudendal. It runs lateral to the cavernous vein along he dorsomedial surface of the crura to enter the erectile tissue where the two corpora fuse; it then continues in the center of the corpora cavernosa.

The dorsal artery of the penis is the termination of the penile artery; it runs over the respective crus and then along the dorsolateral surface of the penis as far as the glans between the dorsal vein medially and dorsal nerve of the penis laterally. This artery has tortuous configuration to accommodate for elongation during erection. It may arise from the accessory internal pudendal artery within the pelvis, and thus may be at risk during radical pelvic surgery. On its way to the glans, it gives off circumflex arteries to supply the corpus spongiosum. Distally, the dorsal artery runs in a ventrolateral position near the sulcus prior to entering the glans. The frenular branch of the dorsal artery curves around each side of the distal shaft to enter the frenulum and glans ventrally.
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INTRACORPORAL CIRCULATION
Arterial blood is conveyed to the erectile tissues in the deep arterial system by means of dorsal, cavernous and bulbo-urethral arteries. The cavernous artery (deep artery of the penis) gives off multiple helicine arteries among the cavernous spaces within the center of the erectile tissue. Most of these open directly into the sinusoids bounded by trabecular, but a few helicine arteries terminate in capillaries that supply the trabeculae. The petiniform septum distally provides communication between the two corpora. The emissory veins at the periphery collect the blood from the sinusoids through the subalbugineal venous plexuses and empty it into the circumflex veins that drain into the deep dorsal vein. With erection, the arteriolar and sinusoidal walls relax secondary to neurotransmitters and the cavernous spaces dilate, enlarging the corporal bodies and stretching the tunica albuginea. The venous tributaries between the sinusoids and the subabugineal venous plexus are compressed by the dilating sinusoids and the stretched tunica albuginea. The direction of blood flow could be summarized as follows: cavernous artery -> helicine arteries -> sinusoids -> post-cavernous venules -> subalbugineal venous plexuses -> emissary vein.
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VENOUS DRAINAGE
The venous drainage system consists of three distinct groups of veins—superficial, intermediate and deep. The superficial drainage system consists of venous drainage from the penile skin and prepuce which drain into the superficial dorsal vein that runs under the superficial penile fascia (Colles’) and joins the saphenous vein via the external pudendal vein. The intermediate system consists of the deep dorsal vein and circumflex veins that drain the glans, corpus spongiosum and distal two-thirds of the corpora cavernosa. The veins leave the glans via a retrocoronal plexus to join the deep dorsal vein that runs in the groove between the corpora. Emissary veins from the corpora join the circumflex veins; the latter communicate with each other at the side by lateral veins and corresponding veins from the opposite side, and run under Buck’s fascia before emptying obliquely into the deep dorsal vein. The latter passes through a space in the suspensory ligament and between the puboprostatic ligament and drains into the internal iliac veins. The deep drainage system consists of the cavernous vein, bulbar vein and crural veins. Blood from the sinusoids from the proximal third of the penis, carried by emissary veins, drains directly into the cavernous veins at the periphery of the corpora cavernosa. The two cavernous veins join to form the main cavernous vein that lies under the cavernous artery and nerves. The cavernous vein runs between the bulb and the crus to drain into the internal pudendal vein; it forms the main venous drainage of the corpora cavernosa. The crural veins arise from the dorsolateral surface of each crus and unite to drain into the internal pudendal vein. The bulb is drained by the bulbar vein, which drains into the prostatic plexus.
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LYMPHATIC DRAINAGE
The lymphatics from the penile skin and prepuce run proximally towards the presymphyseal plexus and then divide to right and left trunks to join the lymphatics from the scrotum and perineum. They run along superficial external pudendal vessels into the superficial inguinal nodes, especially the superomedial group. Some drainage occurs through the femoral canal into Cloquet’s node. The lymphatics from the glans and penile urethra drain into deep inguinal nodes, presymphyseal nodes and, occasionally, into external iliac nodes.
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NERVES
Somatic innervation arises from sacral spinal segments S2-4 via the pudendal nerve. The perineal branch of the pudendal nerve supplies the posterior part of the scrotum and the rectal nerve to the inferior rectal area. The pudendal nerve continues as the dorsal nerve of the penis, which runs over the surface of the obturator internus under the levator, runs deep to the urogenital diaphragm, and passes through the deep transverse perineal muscle to run along the dorsum of the penis accompanied by the dorsal vein and dorsal artery. In epispadia and exstrophy the dorsal nerves are displaced laterally in the middle and distal portion of the penile shaft. Cultaneous nerves to the penis and scrotum arise form the dorsal and posterior branch of the pudendal nerve. The anterior part of the scrotum and proximal penis is supplied by the ilioinguinal nerve after it leaves the superficial inguinal ring. The pudendal nerve supplies the ischiocavernous and bulbocavernous muscles. It branches into the inferior rectal nerve and the scrotal nerve and continues as the dorsal nerve of the penis.

Autonomic nerves consist of sympathetics that arise from lumbar segments L1 and L2 and parasympathetics from S2-4 (nervi erigentes or pelvic nerve). Lumbar splanchnic nerves join the superior hypogastric plexus over the aortic bifurcation, left common vein and sacral promontory. From this plexus, right and left hypogastric nerves travel medial to the internal iliac artery to the inferior hypogastric plexus. The pelvic plexus adjacent to the base of the bladder, prostate, seminal vesicles and rectum contain parasympathetic fibers as well. Nerves from the inferior pelvic plexus supply the prostate, seminal vesicles, epididymis, membranous and penile urethra and bulbo-urethral gland.
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CAVERNOUS NERVE NEUROVASCULAR BUNDLE
The cavernous nerves arise from the pelvic plexus from the lateral surface of the rectum. These nerves run posterolateral to the apex, mid-portion and base of the prostate anterior to Denonvilliers’ fascia between the posterolateral surface of the prostate and the rectum to lie between the lateral pelvic fascia and the prostatic fascia. The branches from the cavernous nerve accompany the branches of the prostatovesicular artery and provide a macroscopic landmark for nerve-sparing radical prostatectomy. The cavernous nerve leaves the pelvis between the transverse perineal muscles and membranous urethra before passing beneath the pubic arch to supply each corpus cavernosum; it also supplies the corpus cavernosum and penile urethra, and terminates in a delicate network around the erectile tissue.
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