Testicle
Testicle | |
---|---|
Lumbar lymph nodes | |
Identifiers | |
Latin | testis |
MeSH | D013737 |
TA98 | A09.3.01.001 |
TA2 | 3576 |
FMA | 7210 |
Anatomical terminology] |
A testicle or testis (pl.: testes) is the male gonad in all bilaterians, including humans. It is homologous to the female ovary. The functions of the testicles are to produce both sperm and androgens, primarily testosterone. Testosterone release is controlled by the anterior pituitary luteinizing hormone, whereas sperm production is controlled both by the anterior pituitary follicle-stimulating hormone and gonadal testosterone.
Structure
Appearance
Males have two testicles of similar size contained within the scrotum, which is an extension of the abdominal wall.[1] Scrotal asymmetry, in which one testicle extends farther down into the scrotum than the other, is common. This is because of the differences in the vasculature's anatomy.[1] For 85% of men, the right testis hangs lower than the left one.[1]
Measurement and volume
The volume of the testicle can be estimated by palpating it and comparing it to ellipsoids (an orchidometer) of known sizes. Another method is to use calipers, a ruler, or an ultrasound image to obtain the three measurements of the x, y, and z axes (length, depth and width). These measurements can then be used to calculate the volume, using the formula for the volume of an ellipsoid:
However, the most accurate calculation of actual testicular volume is gained from the formula:[2]
An average adult testicle measures up to 5 cm × 2 cm × 3 cm (2 in × 3⁄4 in × 1+1⁄4 in). The Tanner scale, which is used to assess the maturity of the male genitalia, assigns a maturity stage to the calculated volume ranging from stage I, a volume of less than 1.5 cm3; to stage V, a volume greater than 20 cm3. Normal volume is 15 to 25 cm3; the average is 18 cm3 per testis (range 12–30 cm3).[1]
The number of spermatozoa an adult human male produces is directly proportional to testicular volume, as larger testicles contain more seminiferous tubules and
Internal structure
Duct system
The testes are covered by a tough fibrous shell called the
Primary cell types
Within the seminiferous tubules, the germ cells develop into
Between tubules (interstitial cells) exist
Immature Leydig cells and interstitial
Blood supply and lymphatic drainage
The testis has three sources of arterial blood supply: the
- The paired testicular arteries arise directly from the abdominal aorta and descend through the inguinal canal, while the scrotum and the rest of the external genitalia is supplied by the internal pudendal artery (a branch of the internal iliac artery).[13][14]
- The testis has collateral blood supply from the cremasteric artery (a branch of the inferior epigastric artery, which is a branch of the external iliac artery), and the artery to the ductus deferens (a branch of the inferior vesical artery, which is a branch of the internal iliac artery).[15][16] Therefore, if the testicular artery is ligated, e.g., during a Fowler-Stevens orchiopexy for a high undescended testis, the testis will usually survive on these other blood supplies.[17]
- Lymphatic drainage of the testes follows the testicular arteries back to the
Layers
Many anatomical features of the adult testis reflect its developmental origin in the
The blood–testis barrier
Large molecules cannot pass from the blood into the lumen of a seminiferous tubule due to the presence of tight junctions between adjacent Sertoli cells.[13] The spermatogonia occupy the basal compartment (deep to the level of the tight junctions) and the more mature forms, such as primary and secondary spermatocytes and spermatids, occupy the adluminal compartment.[13]
The function of the blood–testis barrier may be to prevent an
Injection of sperm antigens causes inflammation of the testis (auto-immune orchitis) and reduced fertility.[13] The blood–testis barrier may reduce the likelihood that sperm proteins will induce an immune response.[19]
Temperature regulation and responses
Carl Richard Moore in 1926 [20] proposed that testicles were external due to spermatogenesis being enhanced at temperatures slightly less than core body temperature outside the body. The spermatogenesis is less efficient at lower and higher temperatures than 33 °C. Because the testes are located outside the body, the smooth tissue of the scrotum can move them closer or further away from the body.[5] The temperature of the testes is maintained at 34.4 °C, a little below body temperature, as temperatures above 36.7 °C impede spermatogenesis.[1][5] There are a number of mechanisms to maintain the testes at the optimum temperature.[21]
The cremasteric muscle covers the testicles and the spermatic cord.[22] When this muscle contracts, the cord shortens and the testicles move closer up toward the body, which provides slightly more warmth to maintain optimal testicular temperature.[22] When cooling is required, the cremasteric muscle relaxes and the testicles lower away from the warm body and are able to cool.[22] Contraction also occurs in response to physical stress, such as blunt trauma; the testicles withdraw and the scrotum shrinks very close to the body in an effort to protect them.[23]
The
Gene and protein expression
The
Development
There are two phases in which the testes grow substantially. These are the embryonic and pubertal phases. During mammalian development, the gonads are at first capable of becoming either
Testes follow the path of descent, from high in the posterior fetal abdomen to the inguinal ring and beyond to the
The testes grow in response to the start of spermatogenesis. Size depends on lytic function, sperm production (amount of spermatogenesis present in testis),
Clinical significance
Protection and injury
- The testicles are very sensitive to impact and injury. The pain involved travels up from each testicle into the abdominal cavity, via the spermatic plexus, which is the primary nerve of each testicle.[31] This will cause pain in the hip and the back. The pain usually fades within a few minutes.
- Testicular torsion is a medical emergency. This is because the longer it takes to access medical intervention with respect to extending ischemia, the higher the chance that the testicle will be lost. There is a 90% chance to save the testicle if de-torsion surgery is performed within six hours of testicular torsion onset.[32]
- Testicular rupture is severe trauma affecting the tunica albuginea.[33]
- Penetrating injuries to the scrotum may cause castration, or physical separation or destruction of the testes, possibly along with part or all of the penis, which results in total sterility if the testicles are not reattached quickly. In an effort to avoid severe infection, ample application of saline and bacitracin help remove debris and foreign objects from the wound.[34]
- Jockstraps support and protect the testicles.
Diseases and conditions
Testicular disease | |
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Specialty | Urology, Reproductive medicine |
- To improve the chances of catching cases of neoplasms, or other health issues early, regular testicular self-examinationis recommended.
- Varicocele, swollen vein(s) from the testes, usually affecting the left side, the testis usually being normal.[35][36]
- Hydrocele testis is swelling around testes caused by accumulation of clear liquid within a membranous sac, the testis usually being normal. It is the most common cause of scrotal swelling.[37]
- Spermatocele is a retention cyst of a tubule of the rete testis or the head of the epididymis distended with barely watery fluid that contains spermatozoa.[36]
- Endocrine disorderscan also affect the size and function of the testis.
- Certain inherited conditions involving mutations in key developmental genes also impair testicular descent, resulting in abdominal or inguinal testes, which remain nonfunctional and may become cancerous.Müllerian ducts. Both excess and deficient levels of estrogens can disrupt spermatogenesis and cause infertility.[39]
- Orchitis is inflammation of the testicles
- Epididymitis is a painful inflammation of the epididymis or epididymides, frequently caused by bacterial infection but sometimes of unknown origin.
- Anorchiais the absence of one or both testicles.
- Cryptorchidism, or "undescended testicles", is when the testicle does not descend into the scrotum of an infant boy.[30]
- Testicular enlargement is an unspecific sign of various testicular diseases, and can be defined as a testicular size of more than 5 cm (long axis) × 3 cm (short axis).[42]
- Blue balls is a condition concerning temporary fluid congestion in the testicles and prostate region, caused by prolonged sexual arousal.
Scientists are working on developing lab-grown testicles that might help infertile men in the future.[43]
Effects of exogenous hormones
To some extent, it is possible to change testicular size. Short of direct injury or subjecting them to adverse conditions, e.g., higher temperature than they are normally accustomed to, they can be shrunk by competing against their intrinsic hormonal function through the use of externally administered
Stimulation of testicular functions via gonadotropic-like hormones may enlarge their size. Testes may shrink or atrophy during hormone replacement therapy or through chemical castration.
In all cases, the loss in testes volume corresponds with a loss of spermatogenesis.
Society and culture
The testicles of
In the Middle Ages, men who wanted a boy sometimes had their left testicle removed. This was because people believed that the right testicle made "boy" sperm and the left made "girl" sperm.[45] As early as 330 BC, Aristotle prescribed the ligation (tying off) of the left testicle in men wishing to have boys.[46]
Etymology and slang
One theory about the etymology of the word testis is based on Roman law. The original Latin word testis, "witness", was used in the firmly established legal principle "Testis unus, testis nullus" (one witness [equals] no witness), meaning that testimony by any one person in court was to be disregarded unless corroborated by the testimony of at least another. This led to the common practice of producing two witnesses, bribed to testify the same way in cases of lawsuits with ulterior motives. Since such witnesses always came in pairs, the meaning was accordingly extended, often in the diminutive (testiculus, testiculi).[citation needed]
Another theory says that testis is influenced by a loan translation, from Greek parastatēs "defender (in law), supporter" that is "two glands side by side".[47]
There are multiple slang terms for the testes. They may be referred to as "balls". Frequently, "nuts" (sometimes intentionally misspelled as "nutz") are also a slang term for the testes due to the geometric resemblance. One variant of the term includes "Deez Nuts", which was used for a satirical political candidate in 2016.
In
Other animals
External appearance
In seasonal breeders, the weight of the testes often increases during the breeding season.[48] The testicles of a dromedary camel are 7–10 cm (2.8–3.9 in) long, 4.5 cm (1.8 in) deep and 5 cm (2.0 in) in width. The right testicle is often smaller than the left.[49]
In
Location
Internal
The basal condition for mammals is to have internal testes.
External
Boreoeutherian land mammals, the large group of mammals that includes humans, have externalized testes.[64] Their testes function best at temperatures lower than their core body temperature. Their testes are located outside of the body and are suspended by the spermatic cord within the scrotum.
There are several hypotheses as to why most boreotherian mammals have external testes that operate best at a temperature that is slightly less than the core body temperature. One view is that it is stuck with enzymes evolved in a colder temperature due to external testes evolving for different reasons. Another view is that the lower temperature of the testes simply is more efficient for sperm production.
The classic hypothesis is that cooler temperature of the testes allows for more efficient fertile spermatogenesis. There are no possible enzymes operating at normal core body temperature that are as efficient as the ones evolved.
Early mammals had lower body temperatures and thus their testes worked efficiently within their body. However, boreotherian mammals may have higher body temperatures than the other mammals and had to develop external testes to keep them cool. One argument is that mammals with internal testes, such as the monotremes, armadillos, sloths, elephants, and rhinoceroses, have a lower core body temperatures than those mammals with external testes.[citation needed]
Researchers have wondered why birds, despite having very high core body temperatures, have internal testes and did not evolve external testes.
Some mammals with seasonal breeding cycles keep their testes internal until the breeding season. After that, their testes descend and increase in size and become external.[66]
The ancestor of the boreoeutherian mammals may have been a small mammal that required very large testes for sperm competition and thus had to place its testes outside the body.[67] This might have led to enzymes involved in spermatogenesis, spermatogenic DNA polymerase beta and recombinase activities evolving a unique temperature optimum that is slightly less than core body temperature. When the boreoeutherian mammals diversified into forms that were larger or did not require intense sperm competition, they still produced enzymes that operated best at cooler temperatures and had to keep their testes outside the body. This position is made less parsimonious because the kangaroo, a non-boreoeutherian mammal, has external testicles. Separately from boreotherian mammals, the ancestors of kangaroos might have also been subject to heavy sperm competition and thus developed external testes; however, kangaroo external testes are suggestive of a possible adaptive function for external testes in large animals.
One argument for the evolution of external testes is that it protects the testes from abdominal cavity pressure changes caused by jumping and galloping.[68]
Mild, transient scrotal heat stress causes DNA damage, reduced fertility and abnormal embryonic development in mice.[69] DNA strand breaks were found in spermatocytes recovered from testicles subjected to 40 °C or 42 °C for 30 minutes.[69] These findings suggest that the external location of the testicles provides the adaptive benefit of protecting spermatogenic cells from heat-induced DNA damage that could otherwise lead to infertility and germline mutation.
Size
The relative size of the testes is often influenced by mating systems.[70] Testicular size as a proportion of body weight varies widely. In the mammalian kingdom, there is a tendency for testicular size to correspond with multiple mates (e.g., harems, polygamy). Production of testicular output sperm and spermatic fluid is also larger in polygamous animals, possibly a spermatogenic competition for survival. The testes of the right whale are likely to be the largest of any animal, each weighing around 500 kg (1,100 lb).[71]
Among the Hominidae, gorillas have little female promiscuity and sperm competition and the testes are small compared to body weight (0.03%). Chimpanzees have high promiscuity and large testes compared to body weight (0.3%). Human testicular size falls between these extremes (0.08%).[72]
Testis weight also varies in seasonal breeders like red foxes,[73] golden jackals,[74] and coyotes.[48]
Internal structure
Gallery
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Testicle
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Testicle
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Testicle hanging on cremaster muscle. These are two healthy testicles. Heat causes them to descend, allowing cooling.
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A healthy scrotum containing normal size testes. The scrotum is in tight condition. The image also shows the texture.
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Testicle of a cat: 1: Extremitas capitata, 2: Extremitas caudata, 3: Margo epididymalis, 4: Margo liber, 5: Mesorchium, 6: Epididymis, 7: testicular artery and vene, 8: Ductus deferens
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Testis surface
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Testis cross section
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The right testis, exposed by laying open the tunica vaginalis.
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Microscopic view of rabbit testis 100×
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Testicle
See also
- Anorchia
- Bollocks
- Cryptorchidism (cryptorchismus)
- Ejaculation
- Eunuchs
- Gelding
- List of homologues of the human reproductive system
- Neutering
- Perineum
- Polyorchidism
- Sterilization (surgical procedure), vasectomy
- Testicular nubbin
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