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Mol Endocrinol. The glans penis of the marsh rice rat is long and robust,  averaging 7. Mammalian Biology 72 6 : The dartos fascia is only loosely connected with B jd penis underlying tissue so it provides the skin mobility and elasticity of the penile skin. A urinoma F lies below the Dartos fascia black open arrowheads. B jd penis foreskin is the least sensitive hairless tissue of the body.
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The foreskin is the double-layered fold of smooth muscle tissue , blood vessels , neurons , skin , and mucous membrane part of the penis that covers and protects the glans penis and the urinary meatus.
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Even within the Vertebrata there are morphological variants with specific terminology, such as hemipenes. Among bird species with a penis are paleognathes tinamous and ratites  and Anatidae ducks, geese and swans. The phallus that male waterfowl have evolved everts out of their bodies in a clockwise coil and aids in inseminating females without their cooperation.
These structures make it harder for males to achieve intromission. The clockwise coils are significant because the male phallus everts out of their body in a counter-clockwise spiral; therefore, a clockwise vaginal structure would impede forceful copulation. Male and female emus are similar in appearance,  although the male's penis can become visible when it defecates.
The male tinamou has a corkscrew shaped penis, similar to those of the ratites and to the hemipenis of some reptiles. Females have a small phallic organ in the cloaca which becomes larger during the breeding season. As with any other bodily attribute, the length and girth of the penis can be highly variable between mammals of different species. In mammals the penis is divided into three parts: .
The internal structures of the penis consist mainly of cavernous, erectile tissue , which is a collection of blood sinusoids separated by sheets of connective tissue trabeculae. Some mammals have a lot of erectile tissue relative to connective tissue, for example horses. The urethra is on the ventral side of the body of the penis. In comparison, the human penis is larger than that of any other primate , both in proportion to body size and in absolute terms.
The penises of even-toed ungulates are curved in an S-shape when not erect. When mating, the tip of a male pronghorn 's penis is often the first part to touch the female pronghorn. Copulation time ranges from 7 to 35 minutes, averaging 11—15 minutes. Bulls have a fibro-elastic penis. Given the small amount of erectile tissue, there is little enlargement after erection. The male genitalia of mouse deer are similar to those of pigs. The purpose of this is not fully understood.
Cetaceans' reproductive organs are located inside the body. Male cetaceans whales, dolphins, and porpoises have two slits, the genital groove concealing the penis and one further behind for the anus. The penis on a right whale can be up to 2. Stallions male horses have a vascular penis. When non-erect, it is quite flaccid and contained within the prepuce foreskin, or sheath. Tapirs have exceptionally long penises relative to their body size.
All members of Carnivora except hyenas have a baculum. During copulation, the spotted hyena inserts his penis through the female's pseudo-penis instead of directly through the vagina , which is blocked by the false scrotum and testes. Once the female retracts her clitoris, the male enters the female by sliding beneath her, an operation facilitated by the penis's upward angle. Domestic cats have barbed penises, with about — one millimeter long backwards-pointing spines.
Lions also have barbed penises. The beech marten 's penis is larger than the pine marten's, with the bacula of young beech martens often outsizing those of old pine martens.
Raccoons have penis bones which bend at a 90 degree angle at the tip. Male walruses possess the largest penis bones of any land mammal, both in absolute size and relative to body size. The adult male American mink 's penis is 2. The baculum is well-developed, being triangular in cross section and curved at the tip.
Males of Racey's pipistrelle bat have a long, straight penis with a notch between the shaft and the narrow, egg-shaped glans penis.
In the baculum penis bone , the shaft is long and narrow and slightly curved. Copulation by male greater short-nosed fruit bats is dorsoventral and the females lick the shaft or the base of the male's penis, but not the glans which has already penetrated the vagina. While the females do this, the penis is not withdrawn and research has shown a positive relationship between length of the time that the penis is licked and the duration of copulation. Post copulation genital grooming has also been observed.
The glans penis of the marsh rice rat is long and robust,  averaging 7. The papilla nipple-like projection on the dorsal upper side of the penis is covered with small spines, a character the marsh rice rat shares only with Oligoryzomys and Oryzomys couesi among oryzomyines examined.
Some features of the accessory glands in the male genital region vary among oryzomyines. In Transandinomys talamancae ,  a single pair of preputial glands is present at the penis. As is usual for sigmodontines, there are two pairs of ventral prostate glands and a single pair of anterior and dorsal prostate glands. In Pseudorhyzomys, the baculum penis bone displays large protuberances at the sides. In the cartilaginous part of the baculum, the central digit is smaller than those at the sides.
In Thomasomys ucucha the glans penis is rounded, short, and small and is superficially divided into left and right halves by a trough at the top and a ridge at the bottom. The glans penis of a male cape ground squirrel is large with a prominent baculum.
Unlike other squirrel species, red squirrels have long, thin, and narrow penises, without a prominent baculum. Winkelmann's mouse can easily be distinguished from its close relatives by the shape of its penis, which has a partially corrugated glans. The foreskin of a capybara is attached to the anus in an unusual way, forming an anogenital invagination.
It has been postulated that the shape of the human penis may have been selected by sperm competition. The shape could have favored displacement of seminal fluids implanted within the female reproductive tract by rival males: the thrusting action which occurs during sexual intercourse can mechanically remove seminal fluid out of the cervix area from a previous mating.
The penile morphology of some types of strepsirrhine primates has provided information about their taxonomy. They are less densely packed than in Otolemur crassicaudatus. The adult male of each vervet monkey species has a pale blue scrotum and a red penis,   and male proboscis monkeys have a red penis with a black scrotum. Male baboons and squirrel monkeys sometimes gesture with an erect penis as both a warning of impending danger and a threat to predators.
The human penis is an external sex organ of male humans. It is a reproductive , intromittent organ that additionally serves as the urinal duct. The main parts are the root of the penis radix : It is the attached part, consisting of the bulb of penis in the middle and the crus of penis , one on either side of the bulb; the body of the penis corpus ; and the epithelium of the penis consists of the shaft skin , the foreskin , and the preputial mucosa on the inside of the foreskin and covering the glans penis.
The human penis is made up of three columns of tissue : two corpora cavernosa lie next to each other on the dorsal side and one corpus spongiosum lies between them on the ventral side.
It is a passage both for urine and for the ejaculation of semen. In males, the expulsion of urine from the body is done through the penis. The urethra drains the bladder through the prostate gland where it is joined by the ejaculatory duct , and then onward to the penis. An erection is the stiffening and rising of the penis, which occurs during sexual arousal , though it can also happen in non-sexual situations. Ejaculation is the ejecting of semen from the penis, and is usually accompanied by orgasm.
A series of muscular contractions delivers semen, containing male gametes known as sperm cells or spermatozoa , from the penis. There is controversy surrounding circumcision. As of [update] , a systematic review of 15, men, and the best research to date on the topic, as the subjects were measured by health professionals, rather than self-measured, has concluded that the average length of an erect human penis is Monotremes and marsupial moles are the only mammals in which the penis is located inside the cloaca.
Male echidnas have a bilaterally symmetrical, rosette-like, four-headed penis. The heads used are swapped each time the mammal copulates. The male echidna's penis is 7 centimetres 2. The penis of the bush hyrax is complex and distinct from that of the other hyrax genera. It has a short, thin appendage within a cup-like glans penis and measures greater than 6 centimetres 2. Additionally, it has been observed that the bush hyrax also has a greater distance between the anus and preputial opening in comparison to other hyraxes.
An adult elephant has the largest penis of any land animal. In giant anteaters , the retracted penis and testes are located internally between the rectum and urinary bladder. When the male armadillo Chaetophractus villosus is sexually aroused , species determination is easier. Its penis can be as long as 35 millimetres 1. During its waking hours, it remains hidden beneath a skin receptacle, until it becomes erect and it projects outside in a rostral direction.
Male turtles and crocodiles have a penis, while male specimens of the reptile order Squamata have two paired organs called hemipenes. Tuataras must use their cloacae for reproduction.
In some fish, the gonopodium , andropodium , and claspers are intromittent organs to introduce sperm into the female developed from modified fins. The record for the largest penis size to body size ratio is held by the barnacle.
The barnacle's penis can grow to up to forty times its own body length. This enables them to reach the nearest female for fertilization. A number of invertebrate species have independently evolved the mating technique of traumatic insemination where the penis penetrates the female's abdomen, thereby creating a womb into which it deposits sperm.
Some millipedes have penises. In these species, the penis is simply one or two projections on underneath the third body segment that produce a spermatophore or sperm packet. The act of insemination however occurs through specialized legs called gonopods which collect the spermatophore and insert it into the female. In male insects , the structure analogous to a penis is known as aedeagus. The male copulatory organ of various lower invertebrate animals is often called the cirrus.
The lesser water boatman 's mating call , generated by rubbing the penis against the abdomen , is the loudest sound, relative to body size, in the animal kingdom. In , entomologist Charles Linehard described Neotrogla , a new genus of barkflies. Species of this genus have sex-reversed genitalia.
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Penis - Wikipedia
MRI of the penis is an expensive test that is not always superior to clinical examination or ultrasound. However, it shows many of the important structures, and in particular the combination of tumescence from intracavernosal alprostadil, and high-resolution T 2 sequences show the glans, corpora and the tunica albuginea well.
In this paper we summarise the radiological anatomy and discuss the indications for MRI. In some cases of penile fibrosis and Peyronie's disease, it may aid surgical planning, and in complex pelvic fracture may replace or augment conventional urethrography. It is an excellent investigation for the malfunctioning penile prosthesis. Before discussing the benefits of MRI for a number of pathologies, we will briefly summarise the radiological anatomy and scanning techniques.
The corpus spongiosum and corpora cavernosa are of high signal on T 2 weighted sequences, and intermediate to low on T 1 weighted sequences [ 1 ].
The septum divides the two corpora cavernosa and is generally porous, allowing the passage of blood and agents injected intracavernosally from one side to the other [ 2 ]. T 2 a and T 1 b weighted sequences through the tumescent penis. Black arrowheads mark the tunica albuginea, and white arrowheads Buck's fascia.
In a the thick white arrow shows the superficial dorsal vein and the thinner white arrows the deep dorsal vessels. The cavernosal arteries are marked by black arrows. The urethra, lying in the middle of the corpus spongiosum, is marked by an asterisk. T 2 weighted sagittal section close to the midline a after intracavernosal alprostadil and b without tumescence.
Black arrows mark the tunica albuginea, large white arrows the corpus spongiosum, small white arrows the urethra within it, and small black arrows the bulbocavernosus muscle. The white arrow head indicates the entry of the urethra into the roof of the bulb, and an asterisk marks the glans. Note the considerably thicker tunica albuginea in the detumescent state, and the lower signal in corpus cavernosum; the glans is not in the midline sagittal plane.
Close to the tunica albuginea is Buck's fascia, a slightly less dense fibrous sheath that is only sometimes distinguishable from the deeper tunica [ 3 ].
It fuses with the deep perineal fascia, so that haemorrhage from a penile fracture which does not compromise Buck's fascia is confined to the penis, rather than spreading to the perineum.
Superficial to Buck's fascia is the thin, incomplete Dartos fascia not clearly seen on MRI , then loose subepithelial connective tissue and skin.
The deep parts of the corpora cavernosa are closely related to the inferior pubic ramus on each side, and surrounded by the ischiocavernosus muscle. The corpus spongiosum is continuous with the glans, and the two generally have the same signal characteristics on MRI. Side branches the helicine arteries are easily seen on ultrasound but are not usually visible on MRI.
The venous anatomy is variable but the largest branches are the superficial and deep dorsal, in the midline, separated by Buck's fascia. This is our routine protocol, with further sequences added as required.
T 1 weighted images may be useful for the detection of haemorrhage and thrombosis within the corpora or draining veins. Small field-of-view short-tau inversion—recovery STIR sequences may show inflammation, and sometimes the deep veins of the pelvis well. For imaging the cavernosal vessels and corporal enhancement, we use dynamic contrast-enhanced sequences, with 3—4-mm slice thickness and a small field of view, usually in the coronal plane, as it shows the vessels in the base of the penis best.
For assessment of cavernosal viability in priapism, the dynamic scans show early perfusion, but there should be one scan at least 10 min after contrast—ideally a small field of view spin echo T 1 weighted sequence, with an identical sequence obtained before contrast for comparison.
There are two reasons for using intracavernosal agents to produce tumescence. First, the normal corpora become uniformly high in T 2 signal, and the tunica albuginea is seen as a thin, regular low signal layer around them. This makes it much easier to see both intracavernosal fibrosis and tunical plaque, as well as other intracavernosal pathologies such as fistula or tumour. Second, positioning of the tumescent though ideally not fully so penis in the midline makes it considerably easier to scan in true sagittal, coronal and axial planes.
The great majority of penile lesions are primary squamous cell tumours, although melanoma, basal cell carcinoma, sarcoma and lymphoma have been reported, and metastases are common enough to account for several cases series, with bladder the commonest site of origin—either haematogenous or from urethral spread of transitional cell carcinoma [ 10 , 11 ].
Anterior urethral tumours are rare, and usually of the squamous rather than the transitional type [ 12 ]. There is some controversy about the current classification, especially within the glans, which is the site of the majority of tumours [ 13 ] Figure 4. It was noted in an early ultrasound study that the tunica albuginea in the glans becomes difficult to see and blends with subepithelial connective tissue [ 4 ].
The T3 classification is also problematic: involvement of the urethra is commonly near the meatus and may have a better prognosis than T2 involvement of the corpus spongiosum in the shaft [ 17 ]. Ulcerating lesion on the glans white arrowheads, with a white arrow showing the ulcerated part , pT2 on histology and correctly called T2 on MRI. CC, corpus cavernosum; S, the spongiosal part of the glans. Tumour is seen as low signal compared with corpus spongiosum or cavernosum on T 2 sequences, but usually higher than the tunica Figure 4.
In particular, several studies have shown that invasion of the corpora cavernosa is rarely missed [ 7 , 19 , 20 ], and proximal tumours, which are hard to scan on ultrasound and difficult to palpate, are shown well. No cases of priapism were seen, although in a previous smaller study priapism occurred in 1 out of 10 patients [ 7 ]. The use of contrast has been anecdotally described as useful [ 21 ], but the larger studies do not support its routine use [ 7 ].
For superficial lesions clinical examination is usually sufficient. MRI of the primary tumour can be combined with an examination of the pelvic nodes. Tumour in virtually all cases involves superficial groin nodes first, and from there spreads to deep groin and pelvic nodes [ 22 ].
The accuracy of size criteria is little better for pelvic nodes [ 26 , 27 ]. The first important distinction in priapism prolonged, often painful erection is the distinction between low- and high-flow states. Low flow is the commonest type and a form of compartment syndrome; as elsewhere in the body if untreated it leads to infarction and fibrosis [ 28 ].
In contrast, in high-flow priapism there is often an arteriolacunar fistula, the corpora are at least initially well oxygenated and there is usually no pain. The distinction between the two states is important, because low-flow priapism requires emergency treatment, but the diagnosis can normally be achieved either clinically, by measurement of blood gases in an aspirate, or on Doppler ultrasound [ 30 ].
There are two main indications for MRI. It is important to emphasise that perfusion of the corpora can be slow, and in addition to dynamically enhanced sequences to assess the patency of the cavernosal arteries, delayed sequences at 5 and 10 min after contrast are necessary to assess tissue viability.
T 1 weighted scan 10 min after caverject in low-flow priapism after a right-sided Winter shunt. The left corpus cavernosum L is infarcted, and a little expanded, but the right R enhances normally. This is an unusual pattern but can be produced by surgical intervention. In high-flow priapism, a fistula can be suspected on the dynamic post-contrast images when there is asymmetrical, early enhancement in one corpus, and is often seen as a focus of heterogeneous flow void on T 2 weighted sequences Figure 6.
Axial T 2 a and early dynamic post-contrast gradient echo images b in high-flow priapism. The fistula confirmed on ultrasound and angiography is shown by the arrowheads in each. It is seen as an area of heterogeneous signal with elements of flow void on T 2 , and is associated with earlier enhancement in the right corpus cavernosum R than the left L. It usually requires urgent surgery to prevent subsequent deformity and erectile dysfunction [ 32 ].
The hallmark of a fracture is an interruption of the low-signal tunica albuginea Figure 7 , usually best seen on T 2 weighted sequences. However, a T 1 spin echo sequence may show the associated haematoma best, and in one small series was the only sequence that showed the fracture well in three patients; enhancement was not necessary [ 33 ]. Transverse T 2 weighted image in a patient with surgically confirmed penile fracture.
The white arrowheads show the tunica albuginea, and the white open arrowheads a little of Buck's fascia. A urinoma F lies below the Dartos fascia black open arrowheads. The fracture is seen as a defect in the ventral aspect of both corpora cavernosa black arrowheads , with, in addition, disruption of the normally high-signal corpus spongiosum black asterisk. Identification of the fracture may be useful to the surgeon because a localised exploration may then be performed, rather than an extensive subcoronal degloving procedure [ 34 ], which probably has a higher post-operative morbidity [ 35 ].
Associated urethral injuries may be found in around a quarter of patients, and can be suspected on MRI [ 33 ], although urethrography remains the gold standard for their detection. MRI may also detect several pathologies that mimic penile fracture—in particular intracavernosal haematoma [ 34 ] and rupture of a superficial vein.
Although we have been able to diagnose suspensory ligament rupture in one case [ 3 ], we do not know the accuracy of MRI in this condition. Tumescence with intracavernosal agents both improves the sensitivity for plaque and intracavernosal fibrosis both seen as focal areas of low T 2 signal and demonstrates associated angulation or waisting [ 40 ] Figure 8.
Except at the base of the penis, MRI is still probably less sensitive for tunical plaque than clinical examination [ 41 ], and is probably not indicated routinely, although it can be useful for surgical planning. Peyronie's disease. T 2 coronal image showing tunical plaque in the distal right corpus cavernosum white arrowheads , with moderate distal waisting.
Intracavernosal fibrosis is seen on the left white arrows. Intracavernosal fibrosis both in Peyronie's disease and post traumatic may be seen as areas of low-signal stranding, but should not be overdiagnosed: some low-signal stranding in the distal corpora cavernosa is seen in normal males [ 3 ]. Inflatable implants are all safe in 1. Inflatable implants are well seen on T 2 and STIR sequences [ 44 ], and we scan with the device inflated when possible.
Although ultrasound is sufficient in many cases to check for adequate fluid in the device, several abnormalities are best seen on MRI. In particular, kinking from overlong cavernosal components can be hard to detect clinically, and in one study an anatomical abnormality was seen on MRI in all 14 patients with pain, apparent clinically in only 5 [ 45 ]. Crossover of cavernosal components is relatively rare, but well seen [ 3 ], as is aneurysmal dilation of the cavernosal components, which may lead to bulge and underinflation Figure 9.
Aneurysmal dilation of the penile prosthesis white arrow , causing a palpable lump and poor inflation. Note the tubing extending to the pump in the scrotum white arrowheads. No fluid is seen around it to indicate infection. We have found that infections that are convincingly shown on MRI are usually clinically apparent as well. Peyronie's disease and penile fibrosis can both cause erectile dysfunction and can be imaged on MRI.
However, clinical tests of nocturnal penile tumescence, response to phosphodiesterase inhibitors and intracavernsoal agents, together with penile Doppler ultrasound remain the mainstay of imaging investigation [ 48 , 49 ]. MR angiography may demonstrate the branches of the internal iliac vessels, and can be used to plan pelvic revascularisation, but is not of adequate resolution to show the penile vessels well, and conventional angiography is superior [ 50 ].
MR urethrography is technically feasible, but not necessarily straightforward. MR urethrography can be performed with T 2 sequences and saline or jelly [ 51 ] or T 1 gradient echo sequences around an hour after injection of intravenous gadolinium [ 53 ].
Periurethral enhancement correlates with inflammation but is as yet of uncertain clinical significance [ 56 ]. MRI can be used to define the location of injected silicone [ 57 ] in particular using silicone-suppressed sequences or other substances, and T 2 and STIR sequences will show dilated Cowper's ducts. Finally, MRI is an excellent tool for defining the anatomy in cases of ambiguous or abnormal genitalia [ 58 ].
MRI can contribute useful information for many different pathologies in the penis, but is in many cases not convincingly superior to clinical examination or ultrasound to justify its routine use. National Center for Biotechnology Information , U.
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