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A testicle or testis ( testes) is the
gonad A gonad, sex gland, or reproductive gland is a Heterocrine gland, mixed gland and sex organ that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gon ...
in all male
bilateria Bilateria () is a large clade of animals characterised by bilateral symmetry during embryonic development. This means their body plans are laid around a longitudinal axis with a front (or "head") and a rear (or "tail") end, as well as a left†...
ns, including humans, and is homologous to the
ovary The ovary () is a gonad in the female reproductive system that produces ova; when released, an ovum travels through the fallopian tube/ oviduct into the uterus. There is an ovary on the left and the right side of the body. The ovaries are end ...
in females. Its primary functions are the production of
sperm Sperm (: sperm or sperms) is the male reproductive Cell (biology), cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm ...
and the secretion of
androgens An androgen (from Greek ''andr-'', the stem of the word meaning ) is any natural or synthetic steroid hormone that regulates the development and maintenance of male characteristics in vertebrates by binding to androgen receptors. This includes ...
, primarily
testosterone Testosterone is the primary male sex hormone and androgen in Male, males. In humans, testosterone plays a key role in the development of Male reproductive system, male reproductive tissues such as testicles and prostate, as well as promoting se ...
. The release of testosterone is regulated by
luteinizing hormone Luteinizing hormone (LH, also known as luteinising hormone, lutropin and sometimes lutrophin) is a hormone produced by gonadotropic cells in the anterior pituitary gland. The production of LH is regulated by gonadotropin-releasing hormone (G ...
(LH) from the
anterior pituitary gland The anterior pituitary (also called the adenohypophysis or pars anterior) is a major organ of the endocrine system. The anterior pituitary is the glandular, anterior lobe that together with the posterior pituitary (or neurohypophysis) makes up ...
. Sperm production is controlled by
follicle-stimulating hormone Follicle-stimulating hormone (FSH) is a gonadotropin, a glycoprotein polypeptide hormone. FSH is synthesized and secreted by the gonadotropic cells of the anterior pituitary gland and regulates the development, growth, puberty, pubertal maturat ...
(FSH) from the anterior pituitary gland and by testosterone produced within the gonads.


Structure


Appearance

Males have two testicles of similar size contained within the
scrotum In most terrestrial mammals, the scrotum (: scrotums or scrota; possibly from Latin ''scortum'', meaning "hide" or "skin") or scrotal sac is a part of the external male genitalia located at the base of the penis. It consists of a sac of skin ...
, which is an extension of the
abdominal wall In anatomy, the abdominal wall represents the boundaries of the abdominal cavity. The abdominal wall is split into the anterolateral and posterior walls. There is a common set of layers covering and forming all the walls: the deepest being the vi ...
. 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. For 85% of men, the right testis hangs lower than the left one.


Measurement and volume

The volume of the testicle can be estimated by palpating it and comparing it to
ellipsoid An ellipsoid is a surface that can be obtained from a sphere by deforming it by means of directional Scaling (geometry), scalings, or more generally, of an affine transformation. An ellipsoid is a quadric surface;  that is, a Surface (mathemat ...
s (an
orchidometer An orchidometer (or orchiometer) is a medical instrument used to measure the volume of the testicles. Instrument The orchidometer was introduced in 1966 by Swiss pediatric endocrinologist Andrea Prader of the University of Zurich.
Page 16 Page most commonly refers to: * Page (paper), one side of a leaf of paper, as in a book Page, PAGE, pages, or paging may also refer to: Roles * Page (assistance occupation), a professional occupation * Page (servant), traditionally a young mal ...
in: ...
) of known sizes. Another method is to use calipers, a ruler, or an
ultrasound Ultrasound is sound with frequency, frequencies greater than 20 Hertz, kilohertz. This frequency is the approximate upper audible hearing range, limit of human hearing in healthy young adults. The physical principles of acoustic waves apply ...
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: :Volume = \frac \cdot \pi \cdot \frac \cdot \frac \cdot \frac :\approx length \cdot width \cdot depth \cdot 0.52 However, the most accurate calculation of actual testicular volume is gained from the formula: :\approx length \cdot width \cdot depth \cdot 0.71 An average adult testicle measures up to . The
Tanner scale The Tanner scale (also known as the Tanner stages or sexual maturity rating (SMR)) is a scale of physical development as pre-pubescent children transition into adolescence, and then adulthood. The scale defines physical measurements of devel ...
, 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). The number of spermatozoa an adult human male produces is directly proportional to testicular volume, as larger testicles contain more seminiferous tubules and
Sertoli cells Sertoli cells are a type of sustentacular "nurse" cell found in human testes which contribute to the process of spermatogenesis (the production of sperm) as a structural component of the seminiferous tubules. They are activated by follicle-sti ...
as a result. As such, men with larger testicles produce on average more sperm cells in each
ejaculate Ejaculation is the discharge of semen (the ''ejaculate''; normally containing sperm) from the penis through the urethra. It is the final stage and natural objective of male sexual stimulation, and an essential component of natural concepti ...
, as testicular volume is positively correlated with semen profiles.


Internal structure


Duct system

The testes are covered by a tough fibrous shell called the tunica albuginea. Under the tunica albuginea, the testes contain very fine-coiled tubes called
seminiferous tubule Seminiferous tubules are located within the testicles, and are the specific location of meiosis, and the subsequent creation of male gametes, namely spermatozoa. Structure The epithelium of the tubule consists of a type of sustentacular cells k ...
s. The tubules are lined with a layer of cells (
germ cell A germ cell is any cell that gives rise to the gametes of an organism that reproduces sexually. In many animals, the germ cells originate in the primitive streak and migrate via the gut of an embryo to the developing gonads. There, they unde ...
s) that develop from
puberty Puberty is the process of physical changes through which a child's body matures into an adult body capable of sexual reproduction. It is initiated by hormonal signals from the brain to the gonads: the ovaries in a female, the testicles i ...
through old age into
sperm Sperm (: sperm or sperms) is the male reproductive Cell (biology), cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm ...
cells Cell most often refers to: * Cell (biology), the functional basic unit of life * Cellphone, a phone connected to a cellular network * Clandestine cell, a penetration-resistant form of a secret or outlawed organization * Electrochemical cell, a d ...
(also known as
spermatozoa A spermatozoon (; also spelled spermatozoön; : spermatozoa; ) is a motile sperm cell (biology), cell produced by male animals relying on internal fertilization. A spermatozoon is a moving form of the ploidy, haploid cell (biology), cell that is ...
or male
gamete A gamete ( ) is a Ploidy#Haploid and monoploid, haploid cell that fuses with another haploid cell during fertilization in organisms that Sexual reproduction, reproduce sexually. Gametes are an organism's reproductive cells, also referred to as s ...
s). The developing sperm travel through the seminiferous tubules to the
rete testis The rete testis ( ; : retia testes) is an anastomosis, anastomosing network of delicate tubules located in the hilum of the testicle (mediastinum testis) that carries spermatozoon, sperm from the seminiferous tubules to the efferent ducts. It is t ...
located in the
mediastinum testis The mediastinum testis is a thick yet incomplete septum at the posterior part of the testis formed by the tunica albuginea of testis projecting into the testis at its posterior aspect where the testis is not lined by the serous membrane to allow fo ...
, to the
efferent ducts The efferent ducts (also efferent ductules, ductuli efferentes, ductus efferentes, or vasa efferentia) connect the rete testis with the initial section of the epididymis.Hess 2018 There are two basic designs for efferent ductule structure: * a ...
, and then to the
epididymis The epididymis (; : epididymides or ) is an elongated tubular genital organ attached to the posterior side of each one of the two male reproductive glands, the testicles. It is a single, narrow, tightly coiled tube in adult humans, in length; ...
where newly created sperm cells mature (
spermatogenesis Spermatogenesis is the process by which haploid spermatozoa develop from germ cells in the seminiferous tubules of the testicle. This process starts with the Mitosis, mitotic division of the stem cells located close to the basement membrane of ...
). The sperm move into the
vas deferens The vas deferens (: vasa deferentia), ductus deferens (: ductūs deferentes), or sperm duct is part of the male reproductive system of many vertebrates. In mammals, spermatozoa are produced in the seminiferous tubules and flow into the epididyma ...
, and are eventually expelled through the
urethra The urethra (: urethras or urethrae) is the tube that connects the urinary bladder to the urinary meatus, through which Placentalia, placental mammals Urination, urinate and Ejaculation, ejaculate. The external urethral sphincter is a striated ...
and out of the urethral orifice through muscular contractions.


Primary cell types

Within the seminiferous tubules, the germ cells develop into
spermatogonia A spermatogonium (plural: ''spermatogonia'') is an undifferentiated male germ cell. Spermatogonia undergo spermatogenesis to form mature spermatozoa in the seminiferous tubules of the testicles. There are three subtypes of spermatogonia in human ...
,
spermatocyte Spermatocytes are a type of male gametocyte in animals. They derive from immature germ cells called spermatogonia. They are found in the testis, in a structure known as the seminiferous tubules. There are two types of spermatocytes, primary and s ...
s, spermatids and spermatozoa through the process of spermatogenesis. The gametes contain DNA for fertilization of an ovum. Sertoli cellsthe true epithelium of the seminiferous epithelium, critical for the support of germ cell development into spermatozoa. Sertoli cells secrete inhibin. Peritubular myoid cells surround the seminiferous tubules. Between tubules (interstitial cells) exist Leydig cellscells localized between seminiferous tubules that produce and secrete
testosterone Testosterone is the primary male sex hormone and androgen in Male, males. In humans, testosterone plays a key role in the development of Male reproductive system, male reproductive tissues such as testicles and prostate, as well as promoting se ...
and other androgens important for
puberty Puberty is the process of physical changes through which a child's body matures into an adult body capable of sexual reproduction. It is initiated by hormonal signals from the brain to the gonads: the ovaries in a female, the testicles i ...
(including secondary sexual characteristics like facial hair), Human sexual activity, sexual behavior, and libido. Sertoli cells support spermatogenesis. Testosterone controls testicular volume. Immature Leydig cells and interstitial macrophages and epithelial cells are also present.


Blood supply and lymphatic drainage

The testis has three sources of arterial blood supply: the testicular artery, the cremasteric artery, and the artery to the ductus deferens. Blood supply and lymphatic drainage of the testes and scrotum are distinct: * 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). * 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). 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. * Lymphatic drainage of the testes follows the testicular arteries back to the paraaortic lymph nodes, while lymph from the scrotum drains to the inguinal lymph nodes.


Layers

Many anatomical features of the adult testis reflect its developmental origin in the abdomen. The layers of tissue enclosing each testicle are derived from the layers of the anterior
abdominal wall In anatomy, the abdominal wall represents the boundaries of the abdominal cavity. The abdominal wall is split into the anterolateral and posterior walls. There is a common set of layers covering and forming all the walls: the deepest being the vi ...
. The cremasteric muscle arises from the internal oblique muscle.


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. 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. The function of the blood–testis barrier may be to prevent an auto-immune reaction. Mature sperm (and their antigens) emerge significantly after immune tolerance is set in infancy. Since sperm are antigenically different from self-tissue, a male animal can react immunologically to his own sperm. The male can make antibodies against them. Injection of sperm antigens causes inflammation of the testis (auto-immune orchitis) and reduced fertility. The blood–testis barrier may reduce the likelihood that sperm proteins will induce an immune response.


Temperature regulation and responses

Carl Richard Moore in 1926 proposed that testicles were external due to
spermatogenesis Spermatogenesis is the process by which haploid spermatozoa develop from germ cells in the seminiferous tubules of the testicle. This process starts with the Mitosis, mitotic division of the stem cells located close to the basement membrane of ...
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 Â°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. The temperature of the testes is maintained at 34.4 Â°C, a little below body temperature, as temperatures above 36.7 Â°C impede spermatogenesis. There are a number of mechanisms to maintain the testes at the optimum temperature. The cremasteric muscle covers the testicles and the spermatic cord. 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. When cooling is required, the cremasteric muscle relaxes and the testicles lower away from the warm body and are able to cool. Contraction also occurs in response to Stress (biology), physical stress, such as blunt trauma; the testicles withdraw and the scrotum shrinks very close to the body in an effort to protect them. The cremasteric reflex will reflexively raise the testicles. The testicles can also be lifted voluntarily using the pubococcygeus muscle, which partially activates related muscles.


Gene and protein expression

The human genome includes approximately 20,000 protein coding genes: 80% of these Gene expression, genes are expressed in adult testes. The testes have the highest fraction of tissue type-specific genes compared to other organs and tissues. About 1000 of them are highly specific for the testes, and about 2,200 show an elevated pattern of expression. A majority of these genes encode for proteins that are expressed in the seminiferous tubules and have functions related to spermatogenesis. Sperm cells express proteins that result in the development of flagella; these same proteins are expressed in the female in cells lining the fallopian tube and cause the development of cilia. Sperm cell flagella and fallopian tube cilia are homologous structures. The testis-specific proteins that show the highest level of expression are protamines.


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 ovaries or testes. In humans, starting at about week 4, the gonadal rudiments are present within the intermediate mesoderm adjacent to the developing kidneys. At about week 6, sex cords develop within the forming testes. These are made up of early Sertoli cells that surround and nurture the germ cells that migrate into the gonads shortly before sex determination begins. In males, the sex-specific gene SRY that is found on the Y chromosome initiates sex determination by downstream regulation of sex-determining factors (such as GATA4, SOX9 and Anti-MĂĽllerian hormone, AMH), which lead to development of the male phenotype, including directing development of the early bipotential gonad toward the male path of development. Testes follow the path of descent, from high in the posterior fetal abdomen to the inguinal ring and beyond to the inguinal canal and into the
scrotum In most terrestrial mammals, the scrotum (: scrotums or scrota; possibly from Latin ''scortum'', meaning "hide" or "skin") or scrotal sac is a part of the external male genitalia located at the base of the penis. It consists of a sac of skin ...
. In most cases (97% full-term, 70% preterm), both testes have descended by birth. In most other cases, only one testis fails to descend. This is called cryptorchidism. In most cases of cryptorchidism, the issue will mostly resolve itself within the first half year of life. However, if the testes do not descend far enough into the scrotum, surgical anchoring in the scrotum is required due to risks of infertility and testicular cancer. The testes grow in response to the start of spermatogenesis. Size depends on lytic function, sperm production (amount of spermatogenesis present in testis), interstitial fluid, and Sertoli cell fluid production. The testicles are fully descended before the male reaches puberty.


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. 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. * Testicular rupture is severe trauma affecting the tunica albuginea. * 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 (medicine), saline and bacitracin help remove debris and foreign objects from the wound. * Jockstraps support and protect the testicles.


Diseases and conditions

* To improve the chances of identifying cases of testicular cancer, neoplasms, and other health issues early, regular testicular self-examination is recommended. * Varicocele, swollen vein(s) from the testes, usually affecting the left side, the testis usually being normal. * 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. * Spermatocele is a retention cyst of a tubule of the
rete testis The rete testis ( ; : retia testes) is an anastomosis, anastomosing network of delicate tubules located in the hilum of the testicle (mediastinum testis) that carries spermatozoon, sperm from the seminiferous tubules to the efferent ducts. It is t ...
or the head of the
epididymis The epididymis (; : epididymides or ) is an elongated tubular genital organ attached to the posterior side of each one of the two male reproductive glands, the testicles. It is a single, narrow, tightly coiled tube in adult humans, in length; ...
distended with barely watery fluid that contains spermatozoa. * Endocrine disorders can 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. Other genetic conditions can result in the loss of the Wolffian ducts and allow for the persistence of MĂĽllerian ducts. Both excess and deficient levels of estrogens can disrupt
spermatogenesis Spermatogenesis is the process by which haploid spermatozoa develop from germ cells in the seminiferous tubules of the testicle. This process starts with the Mitosis, mitotic division of the stem cells located close to the basement membrane of ...
and cause infertility. * Bell-clapper deformity is a deformity in which the testicle is not attached to the scrotal walls, and can rotate freely on the spermatic cord within the tunica vaginalis. Those with Bell-clapper are at a higher risk of testicular torsion. * 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. * Anorchia is 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. *Testicular enlargement is an sensitivity and specificity, unspecific sign of various testicular diseases, and can be defined as a testicular size of more than 5 cm (Anatomical terms of location#Axes, long axis) Ă— 3 cm (short axis). *Blue balls is a condition concerning temporary fluid congestion in the testicles and prostate region, caused by prolonged sexual arousal. Testicular prostheses are available to mimic the appearance and feel of one or both testicles, when absent as from injury or as treatment in association to gender dysphoria. There have also been some instances of their implantation in dogs. Scientists are working on developing lab-grown testicles that might help infertile men in the future.


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 steroidal hormones. Steroids taken for muscle enhancement (especially anabolic steroids) often have the undesired side effect of testicular shrinkage. Stimulation of testicular functions via gonadotropin, gonadotropic-like hormones may enlarge their size. Testes may shrink or atrophy during Hormone therapy, 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 Calf (animal), calves, Sheep, lambs, roosters, Turkey as food, turkeys, and other animals are eaten in many parts of the world, often under euphemistic culinary names. Testicles are a by-product of the castration of young animals raised for meat, so they might have been a late-spring seasonal specialty. In modern times, they are generally frozen and available year-round. 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. As early as 330 BC, Aristotle prescribed the ligation (tying off) of the left testicle in men wishing to have boys.


Etymology and slang

One theory about the etymology of the word ''testis'' is based on Roman law. The original Latin word , was used in the firmly established legal principle "" (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 (). Another theory says that ''testis'' is influenced by a loan translation, from Greek that is "two glands side by side". There are multiple slang terms for the testes. They may be referred to as "balls". Frequently, "Nut (fruit), 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 Deez Nuts (satirist), satirical political candidate in 2016. In Spanish language, Spanish, the term is used, which is Spanish for eggs.


Other animals


External appearance

In seasonal breeders, the weight of the testes often increases during the breeding season. The Dromedary#Testicles, testicles of a dromedary camel are long, deep and in width. The right testicle is often smaller than the left. In sharks, the testicle on the right side is usually larger. In many bird and mammal species, the left may be larger. Fish_anatomy#Testes, Fish usually have two testes of a similar size. The primitive jawless fish have only a single testis, located in the midline of the body, although this forms from the fusion of paired structures in the embryo.


Location


Internal

The basal condition for mammals is to have internal testes. The testes of monotremes, xenarthrans, and afrotherians remain within the abdomen (testicondy). There are also some marsupials with external testes and boreoeutherian mammals with internal testes, such as the rhinoceros.Schaffer, N. E., et al.
Ultrasonography of the reproductive anatomy in the Sumatran rhinoceros (Dicerorhinus sumatrensis)
." Journal of Zoo and Wildlife Medicine (1994): 337-348.
Cetaceans such as whales and dolphins also have internal testes. As external testes would increase drag in the water, they have internal testes, which are kept cool by special circulatory systems that cool the arterial blood going to the testes by placing the arteries near veins bringing cooled venous blood from the skin. In odobenids and phocids, the location of the testes is para-abdominal, though otariids have scrotal testes.


External

Boreoeutherian land mammals, the large group of mammals that includes humans, have externalized testes. 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. Researchers have wondered why birds, despite having very high core body temperatures, have internal testes and did not evolve external testes.BIOLOGY OF REPRODUCTION 56, 1570–1575 (1997)- Determination of Testis Temperature Rhythms and Effects of Constant Light on Testicular Function in the Domestic Fowl (Gallus domesticus)
It was once theorized that birds used their air sacs to cool the testes internally, but later studies revealed that birds' testes are able to function at core body temperature. 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. 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. 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 descended testes in mammals, evolution of external testes is that it protects the testes from abdominal cavity pressure changes caused by jumping and galloping. Mild, transient scrotal heat stress causes DNA damage, infertility, reduced fertility and abnormal embryonic development in mice. DNA strand breaks were found in spermatocytes recovered from testicles subjected to 40 Â°C or 42 Â°C for 30 minutes. 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 Monogamy in animals#Testis size, relative size of the testes is often influenced by mating systems. 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 Sperm competition, 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). 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%). Testis weight also varies in seasonal breeders like red foxes, golden jackals, and coyotes.


Internal structure

Amphibians and most fish do not possess seminiferous tubules. Instead, the sperm are produced in spherical structures called ''sperm ampullae''. These are seasonal structures, releasing their contents during the breeding season, and then being reabsorbed by the body. Before the next breeding season, new sperm ampullae begin to form and ripen. The ampullae are otherwise essentially identical to the seminiferous tubules in higher vertebrates, including the same range of cell types.


Gallery

File:Slide4aaaaaa.JPG, Testicle File:Slide2CC.JPG, Testicle Image:Hanging testicles.JPG, Testicle hanging on cremaster muscle. These are two healthy testicles. Heat causes them to descend, allowing cooling. Image:Human_Scrotum.JPG, A healthy
scrotum In most terrestrial mammals, the scrotum (: scrotums or scrota; possibly from Latin ''scortum'', meaning "hide" or "skin") or scrotal sac is a part of the external male genitalia located at the base of the penis. It consists of a sac of skin ...
containing normal size testes. The scrotum is in tight condition. The image also shows the texture. Image:Testicle-cat.jpg, 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 Image:Illu testis surface.jpg, Testis surface Image:Illu testis cross section.jpg, Testis cross section Image:Gray1148.png, The right testis, exposed by laying open the tunica vaginalis. Image:Rabbitttestis100x2.jpg, Microscopic view of rabbit testis 100Ă— File:Slide8eee.JPG, Testicle


See also

* Anorchia * Anther * Antheridium * Bollocks * Cryptorchidism (cryptorchismus) * Ejaculation * Eunuchs * Gelding * List of homologues of the human reproductive system * Neutering * Perineum * Polyorchidism * Sterilization (medicine), Sterilization (surgical procedure), vasectomy * Testicular nubbin


General and cited references

*


Citations


External links

* wikt:Thesaurus:testicles, List of synonyms and slang words for testicles in many languages at wikt:Wiktionary:Thesaurus, Wiktionary's Thesaurus
Testis at the Human Protein Atlas
{{Authority control Testicle, Endocrine system Glands Human male reproductive system Mammal male reproductive system Men's health Scrotum Sex organs