The epididymis (; plural: epididymides or ) is a tube that connects a

testicle A testicle or testis (plural testes) is the male reproductive gland or gonad in all bilaterians, including humans. It is homologous to the female ovary. The functions of the testes are to produce both sperm and androgens, primarily testoste ...

to a

vas deferens The vas deferens or ductus deferens is part of the male reproductive system of many vertebrates. The ducts transport sperm from the epididymis to the ejaculatory ducts in anticipation of ejaculation. The vas deferens is a partially coiled tube ...

in the

male reproductive system The male reproductive system consists of a number of sex organs that play a role in the process of human reproduction. These organs are located on the outside of the body and within the pelvis. The main male sex organs are the penis and the te ...

. It is a single, narrow, tightly-coiled tube in adult humans, in length. It serves as an interconnection between the multiple efferent ducts at the rear of a testicle (proximally), and the vas deferens (distally).

Anatomy

The epididymis is situated posterior and somewhat lateral to the testis. The epididymis is invested completely by the tunica vaginalis (which is continuous with the tunica vaginalis covering the testis). The epididymis can be divided into three main regions: * The head ( la, caput). The head of the epididymis receives

spermatozoa A spermatozoon (; also spelled spermatozoön; ; ) is a motile sperm cell, or moving form of the haploid cell that is the male gamete. A spermatozoon joins an ovum to form a zygote. (A zygote is a single cell, with a complete set of chromos ...

via the efferent ducts of the mediastinium of the

testis A testicle or testis (plural testes) is the male reproductive gland or gonad in all bilaterians, including humans. It is homologous to the female ovary. The functions of the testes are to produce both sperm and androgens, primarily testostero ...

at the superior pole of the testis. The head is characterized

histological Histology, also known as microscopic anatomy or microanatomy, is the branch of biology which studies the microscopic anatomy of biological tissues. Histology is the microscopic counterpart to gross anatomy, which looks at larger structures vis ...

ly by a thick

epithelium Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellul ...

with long stereocilia (described below) and a little

smooth muscle Smooth muscle is an involuntary non- striated muscle, so-called because it has no sarcomeres and therefore no striations (''bands'' or ''stripes''). It is divided into two subgroups, single-unit and multiunit smooth muscle. Within single-unit ...

. It is involved in absorbing fluid to make the

sperm Sperm is the male reproductive 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 with a tail known as a flagellum, ...

more concentrated. The concentration of the sperm here is dilute. * The body ( la, corpus, links=no). This has an intermediate epithelium and smooth muscle thickness. * The tail ( la, cauda, links=no). This has the thinnest epithelium of the three regions and the greatest quantity of smooth muscle. The tail is distally continuous with (the convoluted portion of) the ductus deferens (s. vas deferens).

Histology

The epididymis is covered by a two layered pseudostratified epithelium. The epithelium is separated by a

basement membrane The basement membrane is a thin, pliable sheet-like type of extracellular matrix that provides cell and tissue support and acts as a platform for complex signalling. The basement membrane sits between epithelial tissues including mesothelium an ...

from the

connective tissue Connective tissue is one of the four primary types of animal tissue, along with epithelial tissue, muscle tissue, and nervous tissue. It develops from the mesenchyme derived from the mesoderm the middle embryonic germ layer. Connective tiss ...

wall which has smooth muscle cells. The major cell types in the epithelium are: * Principal cells: columnar cells that, with the basal cells, form the majority of the epithelium. In the caput (head) region these cells have long stereocilia that are tuft-like extensions that project into the lumen. The stereocilia are much shorter in the cauda (tail) segment. They also secrete

carnitine Carnitine is a quaternary ammonium compound involved in metabolism in most mammals, plants, and some bacteria. In support of energy metabolism, carnitine transports long-chain fatty acids into mitochondria to be oxidized for energy production, an ...

sialic acid Sialic acids are a class of alpha-keto acid sugars with a nine-carbon backbone. The term "sialic acid" (from the Greek for saliva, - ''síalon'') was first introduced by Swedish biochemist Gunnar Blix in 1952. The most common member of this ...

, glycoproteins, and glycerylphosphorylcholine into the lumen. * Basal cells: shorter, pyramid-shaped cells which contact the basal lamina but taper off before their apical surfaces reach the lumen. These are thought to be undifferentiated precursors of principal cells. * Apical cells: predominantly found in the head region * Clear cells: predominant in the tail region * Intraepithelial lymphocytes: distributed throughout the tissue. * Intraepithelial macrophages

Stereocilia

The stereocilia of the epididymis are long cytoplasmic projections that have an actin filament backbone. These filaments have been visualized at high resolution using fluorescent phalloidin that binds to actin filaments. The stereocilia in the epididymis are non-motile. These membrane extensions increase the surface area of the cell, allowing for greater absorption and secretion. It has been shown that epithelial sodium channel

ENaC The epithelial sodium channel (ENaC), (also known as amiloride-sensitive sodium channel) is a membrane-bound ion channel that is selectively permeable to sodium ions (). It is assembled as a heterotrimer composed of three homologous subunits α ...

that allows the flow of Na⁺ ions into the cell is localized on stereocilia. Because sperm are initially non-motile as they leave the seminiferous tubules, large volumes of fluid are secreted to propel them to the epididymis. The core function of the stereocilia is to resorb 90% of this fluid as the spermatozoa start to become motile. This absorption creates a fluid current that moves the immobile sperm from the seminiferous tubules to the epididymis. Spermatozoa do not reach full motility until they reach the

vagina In mammals, the vagina is the elastic, muscular part of the female genital tract. In humans, it extends from the vestibule to the cervix. The outer vaginal opening is normally partly covered by a thin layer of mucosal tissue called the hymen ...

, where the alkaline pH is neutralized by acidic vaginal fluids.

Development

In the

embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...

, the epididymis develops from tissue that once formed the mesonephros, a primitive

kidney The kidneys are two reddish-brown bean-shaped organs found in vertebrates. They are located on the left and right in the retroperitoneal space, and in adult humans are about in length. They receive blood from the paired renal arteries; blo ...

found in many aquatic vertebrates. Persistence of the cranial end of the mesonephric duct will leave behind a remnant called the ''

appendix of the epididymis The appendix of the epididymis (or pedunculated hydatid) is a small stalked appendage (sometimes duplicated) on the head of the epididymis. It is usually regarded as a detached efferent duct. This structure is derived from the Wolffian duct (Meso ...

''. In addition, some mesonephric tubules can persist as the '' paradidymis'', a small body caudal to the efferent ductules. A Gartner's duct is a homologous remnant in the female.

Function

Role in storage of sperm and ejaculant

Spermatozoa formed in the testis enter the caput epididymis, progress to the corpus, and finally reach the cauda region, where they are stored. Sperm entering the caput epididymis are incomplete—they lack the ability to swim forward (motility) and to

fertilize Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Pro ...

an egg. Epididymal transit takes 2 to 6 days in humans and 10–13 in rodents. During their transit in the epididymis, sperm undergo maturation processes necessary for them to acquire motility and fertility. Final maturation (''

capacitation Capacitation is the penultimate step in the maturation of mammalian spermatozoa and is required to render them competent to fertilize an oocyte. This step is a biochemical event; the sperm move normally and look mature prior to capacitation. ''In ...

'') is completed in the female reproductive tract. The epididymis secretes immobilin, a large glycoprotein that is responsible for the creating of the viscoelastic luminal environment that serves to mechanically immobilize spermatozoa until

ejaculation Ejaculation is the discharge of semen (the ''ejaculate''; normally containing sperm) from the male reproductory tract as a result of an orgasm. It is the final stage and natural objective of male sexual stimulation, and an essential compo ...

. Immobilin is predominantly secreted into the proximal caput epididymis prior to the acquisition of the potential for sperm motility. During emission, sperm flow from the ''cauda epididymis'' (which functions as a storage reservoir) into the vas deferens where they are propelled by the

peristaltic Peristalsis ( , ) is a radially symmetrical contraction and relaxation of muscles that propagate in a wave down a tube, in an anterograde direction. Peristalsis is progression of coordinated contraction of involuntary circular muscles, whic ...

action of

muscle Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of mus ...

layers in the wall of the

, and are mixed with the diluting fluids of the

prostate The prostate is both an accessory gland of the male reproductive system and a muscle-driven mechanical switch between urination and ejaculation. It is found only in some mammals. It differs between species anatomically, chemically, and phys ...

seminal vesicle The seminal vesicles (also called vesicular glands, or seminal glands) are a pair of two convoluted tubular glands that lie behind the urinary bladder of some male mammals. They secrete fluid that partly composes the semen. The vesicles are 5� ...

s, and other accessory glands prior to ejaculation (forming

semen Semen, also known as seminal fluid, is an organic bodily fluid created to contain spermatozoa. It is secreted by the gonads (sexual glands) and other sexual organs of male or hermaphroditic animals and can fertilize the female ovum. Sem ...

). Contrary to popular belief, sperm are capable of causing a pregnancy even without ever travelling through the epididymis. This has been proven in two cases in the United States in the 1980s where a couple of men's vasa deferentia were directly surgically attached to their efferent ducts and these men both subsequently impregnated their partners within the next couple of years. This has also been proven in a similar case in Western Europe in the early 1990s.

Antioxidant defenses

During their transit through the epididymis, the

undergo a series of transformations in preparation for their ultimate task of fertilizing the

oocyte An oocyte (, ), oöcyte, or ovocyte is a female gametocyte or germ cell involved in reproduction. In other words, it is an immature ovum, or egg cell. An oocyte is produced in a female fetus in the ovary during female gametogenesis. The female ...

. In order to protect the spermatozoa during their transit through the epididymis, the epididymal

produces a variety of

antioxidant Antioxidants are compounds that inhibit oxidation, a chemical reaction that can produce free radicals. This can lead to polymerization and other chain reactions. They are frequently added to industrial products, such as fuels and lubrica ...

proteins that help protect the spermatozoa from

oxidative damage Oxidative stress reflects an imbalance between the systemic manifestation of reactive oxygen species and a biological system's ability to readily detoxify the reactive intermediates or to repair the resulting damage. Disturbances in the normal r ...

O'Flaherty C. Orchestrating the antioxidant defenses in the epididymis. Andrology. 2019 Sep;7(5):662-668. doi: 10.1111/andr.12630. Epub 2019 May 1. PMID: 31044545. The antioxidant proteins produced include

catalase Catalase is a common enzyme found in nearly all living organisms exposed to oxygen (such as bacteria, plants, and animals) which catalyzes the decomposition of hydrogen peroxide to water and oxygen. It is a very important enzyme in protecting t ...

, glutathione peroxidases, glutathione-S-transferases,

peroxiredoxin Peroxiredoxins (Prxs, ; HGNC root symbol ''PRDX'') are a ubiquitous family of antioxidant enzymes that also control cytokine-induced peroxide levels and thereby mediate signal transduction in mammalian cells. The family members in humans are P ...

superoxide dismutase Superoxide dismutase (SOD, ) is an enzyme that alternately catalyzes the dismutation (or partitioning) of the superoxide () radical into ordinary molecular oxygen (O2) and hydrogen peroxide (). Superoxide is produced as a by-product of oxygen ...

thioredoxin reductase Thioredoxin reductases (TR, TrxR) () are enzymes that reduce thioredoxin (Trx). Two classes of thioredoxin reductase have been identified: one class in bacteria and some eukaryotes and one in animals. In bacteria TrxR also catalyzes the reduction ...

and thioredoxins. Deficiencies in the availability of these antioxidant proteins reduces sperm quality by affecting a variety of the proteins necessary for the motility needed to fertilize oocytes. Reduced antioxidant activity also causes increased oxidative damage to the sperm DNA.

Clinical significance

Inflammation

An inflammation of the epididymis is called epididymitis. It is much more common than testicular inflammation, termed

orchitis Orchitis is inflammation of the testes. It can also involve swelling, pains and frequent infection, particularly of the epididymis, as in epididymitis. The term is from the Ancient Greek ὄρχις meaning "testicle"; same root as ''orchid''. S ...

Surgical removal

Epididymotomy is the placing of an incision into the epididymis and is sometimes considered as a treatment option for acute suppurating epididymitis. Epididymectomy is the surgical removal of the epididymis sometimes performed for post-vasectomy pain syndrome and for refractory cases of epididymitis. Epididymectomy is also performed for

sterilization Sterilization may refer to: * Sterilization (microbiology), killing or inactivation of micro-organisms * Soil steam sterilization, a farming technique that sterilizes soil with steam in open fields or greenhouses * Sterilization (medicine) rende ...

on some

male Male (symbol: ♂) is the sex of an organism that produces the gamete (sex cell) known as sperm, which fuses with the larger female gamete, or ovum, in the process of fertilization. A male organism cannot reproduce sexually without access to ...

animals Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...

of livestock species so they can be used to detect estrus in females ready for artificially insemination.

Non-human animals

The epididymis is present in all male reptiles, birds, and mammals. In reptiles, there is an additional canal between the testis and the head of the epididymis and which receives the various efferent ducts. This is, however, absent in all birds and mammals.

Gallery

Image:Illu repdt male.jpg, Human male reproductive system Image:Illu testis 1b.jpg, Testis Image:Mesorchium.svg, Schematic drawing: cross-section through a testicle Image:Epididymis high mag.jpg,

Micrograph A micrograph or photomicrograph is a photograph or digital image taken through a microscope or similar device to show a magnified image of an object. This is opposed to a macrograph or photomacrograph, an image which is also taken on a m ...

of epididymis -

H&E stain Hematoxylin and eosin stain ( or haematoxylin and eosin stain or hematoxylin-eosin stain; often abbreviated as H&E stain or HE stain) is one of the principal tissue stains used in histology. It is the most widely used stain in medical diag ...

Image:Epididymis.JPG, Micrograph File:Slide4UMR.JPG, Deep dissection of epididymis

Notes

External links

* {{Authority control Scrotum