Ovarian Capsule
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The ovary is an organ in the female reproductive system that produces an
ovum The egg cell, or ovum (plural ova), is the female reproductive cell, or gamete, in most anisogamous organisms (organisms that reproduce sexually with a larger, female gamete and a smaller, male one). The term is used when the female gamete is ...
. When released, this travels down the
fallopian tube The fallopian tubes, also known as uterine tubes, oviducts or salpinges (singular salpinx), are paired tubes in the human female that stretch from the uterus to the ovaries. The fallopian tubes are part of the female reproductive system. In ot ...
into the
uterus The uterus (from Latin ''uterus'', plural ''uteri'') or womb () is the organ in the reproductive system of most female mammals, including humans that accommodates the embryonic and fetal development of one or more embryos until birth. The uter ...
, where it may become fertilized by a
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, whi ...
. There is an ovary () found on each side of the body. The ovaries also secrete
hormone A hormone (from the Greek participle , "setting in motion") is a class of signaling molecules in multicellular organisms that are sent to distant organs by complex biological processes to regulate physiology and behavior. Hormones are required ...
s that play a role in the
menstrual cycle The menstrual cycle is a series of natural changes in hormone production and the structures of the uterus and ovaries of the female reproductive system that make pregnancy possible. The ovarian cycle controls the production and release of eggs a ...
and
fertility Fertility is the capability to produce offspring through reproduction following the onset of sexual maturity. The fertility rate is the average number of children born by a female during her lifetime and is quantified demographically. Fertili ...
. The ovary progresses through many stages beginning in the prenatal period through
menopause Menopause, also known as the climacteric, is the time in women's lives when menstrual periods stop permanently, and they are no longer able to bear children. Menopause usually occurs between the age of 47 and 54. Medical professionals often d ...
. It is also an
endocrine gland Endocrine glands are ductless glands of the endocrine system that secrete their products, hormones, directly into the blood. The major glands of the endocrine system include the pineal gland, pituitary gland, pancreas, ovaries, testes, thyroid ...
because of the various
hormone A hormone (from the Greek participle , "setting in motion") is a class of signaling molecules in multicellular organisms that are sent to distant organs by complex biological processes to regulate physiology and behavior. Hormones are required ...
s that it secretes.


Structure

The ovaries are considered the female gonads. Each ovary is whitish in color and located alongside the lateral wall of the
uterus The uterus (from Latin ''uterus'', plural ''uteri'') or womb () is the organ in the reproductive system of most female mammals, including humans that accommodates the embryonic and fetal development of one or more embryos until birth. The uter ...
in a region called the
ovarian fossa The ovarian fossa is a shallow depression on the lateral wall of the pelvis, where in the ovary lies. This ovarian fossa has the following boundaries: * superiorly: by the external iliac artery and vein * anteriorly and inferiorly: by the broad ...
. The ovarian fossa is the region that is bounded by the external iliac artery and in front of the ureter and the
internal iliac artery The internal iliac artery (formerly known as the hypogastric artery) is the main artery of the pelvis. Structure The internal iliac artery supplies the walls and viscera of the pelvis, the buttock, the reproductive organs, and the medial compart ...
. This area is about 4 cm x 3 cm x 2 cm in size.Daftary, Shirish; Chakravarti, Sudip (2011). Manual of Obstetrics, 3rd Edition. Elsevier. pp. 1-16. . The ovaries are surrounded by a capsule, and have an outer cortex and an inner medulla. The capsule is of dense connective tissue and is known as the tunica albuginea. Usually, ovulation occurs in one of the two ovaries releasing an egg each menstrual cycle. The side of the ovary closest to the
fallopian tube The fallopian tubes, also known as uterine tubes, oviducts or salpinges (singular salpinx), are paired tubes in the human female that stretch from the uterus to the ovaries. The fallopian tubes are part of the female reproductive system. In ot ...
is connected to it by
infundibulopelvic ligament The suspensory ligament of the ovary, also infundibulopelvic ligament (commonly abbreviated IP ligament or simply IP), is a fold of peritoneum that extends out from the ovary to the wall of the pelvis. Some sources consider it a part of the broad ...
, and the other side points downwards attached to the uterus via the
ovarian ligament The ovarian ligament (also called the utero-ovarian ligament or proper ovarian ligament) is a fibrous ligament that connects the ovary to the lateral surface of the uterus. Structure The ovarian ligament is composed of muscular and fibrous tissue; ...
. Other structures and tissues of the ovaries include the hilum.


Ligaments

The ovaries lie within the peritoneal cavity, on either side of the uterus, to which they are attached via a fibrous cord called the
ovarian ligament The ovarian ligament (also called the utero-ovarian ligament or proper ovarian ligament) is a fibrous ligament that connects the ovary to the lateral surface of the uterus. Structure The ovarian ligament is composed of muscular and fibrous tissue; ...
. The ovaries are uncovered in the peritoneal cavity but are tethered to the body wall via the suspensory ligament of the ovary which is a posterior extension of the broad ligament of the uterus. The part of the
broad ligament of the uterus The broad ligament of the uterus is the wide fold of peritoneum that connects the sides of the uterus to the walls and floor of the pelvis. Structure Subdivisions Contents The contents of the broad ligament include the following: * Reprodu ...
that covers the ovary is known as the
mesovarium The mesovarium is the portion of the broad ligament of the uterus that suspends the ovaries. The ovary is not covered by the mesovarium; rather, it is covered by germinal epithelium. At first the mesonephros and genital ridge are suspended by a c ...
. The ovarian pedicle is made up part of the
fallopian tube The fallopian tubes, also known as uterine tubes, oviducts or salpinges (singular salpinx), are paired tubes in the human female that stretch from the uterus to the ovaries. The fallopian tubes are part of the female reproductive system. In ot ...
,
mesovarium The mesovarium is the portion of the broad ligament of the uterus that suspends the ovaries. The ovary is not covered by the mesovarium; rather, it is covered by germinal epithelium. At first the mesonephros and genital ridge are suspended by a c ...
, ovarian ligament, and ovarian blood vessels.


Microanatomy

The surface of the ovaries is covered with a membrane consisting of a lining of
simple cuboidal Simple cuboidal epithelium is a type of epithelium that consists of a single layer of cuboidal (cube-like) cells which have large, spherical and central nuclei. Simple cuboidal epithelium is found on the surface of ovaries, the lining of nephro ...
-to-columnar shaped
mesothelium The mesothelium is a membrane composed of simple squamous epithelial cells of mesodermal origin, which forms the lining of several body cavities: the pleura (pleural cavity around the lungs), peritoneum (abdominopelvic cavity including the mesente ...
, called the germinal epithelium. The outer layer is the
ovarian cortex The ovarian cortex is the outer portion of the ovary. The ovarian follicles are located within the ovarian cortex. The ovarian cortex is made up of connective tissue. Ovarian cortex tissue transplant has been performed to treat infertility I ...
, consisting of
ovarian follicles An ovarian follicle is a roughly spheroid cellular aggregation set found in the ovaries. It secretes hormones that influence stages of the menstrual cycle. At the time of puberty, women have approximately 200,000 to 300,000 follicles, each with th ...
and stroma in between them. Included in the follicles are the
cumulus oophorus The cumulus oophorus, (discus proligerus), is a cluster of cells that surround the oocyte both in the ovarian follicle and after ovulation. In the antral follicle, it may be regarded as an extension of the membrana granulosa. The innermost layer ...
,
membrana granulosa The larger ovarian follicles An ovarian follicle is a roughly spheroid cellular aggregation set found in the ovaries. It secretes hormones that influence stages of the menstrual cycle. At the time of puberty, women have approximately 200,000 to ...
(and the
granulosa cells A granulosa cell or follicular cell is a somatic cell of the sex cord that is closely associated with the developing female gamete (called an oocyte or egg) in the ovary of mammals. Structure and function In the primordial ovarian follicle, and l ...
inside it),
corona radiata In neuroanatomy, the corona radiata is a white matter sheet that continues inferiorly as the internal capsule and superiorly as the centrum semiovale. This sheet of both ascending and descending axons carries most of the neural traffic from and ...
, zona pellucida, and primary oocyte. Theca of follicle, antrum and
liquor folliculi Follicular fluid is a liquid which fills the follicular antrum and surrounds the ovum in an ovarian follicle. This fluid is rich in hyaluronic acid, and is used in a modified intracytoplasmic sperm injection (ICSI) called physiological ICSI In ...
are also contained in the follicle. Also in the cortex is the
corpus luteum The corpus luteum (Latin for "yellow body"; plural corpora lutea) is a temporary endocrine structure in female ovaries involved in the production of relatively high levels of progesterone, and moderate levels of estradiol, and inhibin A. It is t ...
derived from the follicles. The innermost layer is the
ovarian medulla The medulla of ovary (or Zona vasculosa of Waldeyer) is a highly vascular stroma in the center of the ovary. It forms from embryonic mesenchyme and contains blood vessels, lymphatic vessels, and nerves. This stroma forms the tissue of the hilum ...
. It can be hard to distinguish between the cortex and medulla, but follicles are usually not found in the medulla. Follicular cells are flat epithelial cells that originate from surface epithelium covering the ovary. They are surrounded by granulosa cells that have changed from flat to cuboidal and proliferated to produce a stratified epithelium. The ovary also contains
blood vessel The blood vessels are the components of the circulatory system that transport blood throughout the human body. These vessels transport blood cells, nutrients, and oxygen to the tissues of the body. They also take waste and carbon dioxide away ...
s and lymphatics.


Function

At
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 girl, the testes in a boy. ...
, the ovary begins to secrete increasing levels of hormones. Secondary sex characteristics begin to develop in response to the hormones. The ovary changes structure and function beginning at puberty. Since the ovaries are able to regulate hormones, they also play an important role in pregnancy and fertility. When egg cells (oocytes) are released from the Fallopian tube, a variety of feedback mechanisms stimulate the endocrine system which cause hormone levels to change. These feedback mechanisms are controlled by the hypothalamus and pituitary glands. Messages or signals from the hypothalamus are sent to the pituitary gland. In turn, the pituitary gland releases hormones to the ovaries. From this signaling, the ovaries release their own hormones.


Gamete production

The ovaries are the site of production and periodical release of
egg cell The egg cell, or ovum (plural ova), is the female reproductive cell, or gamete, in most anisogamous organisms (organisms that reproduce sexually with a larger, female gamete and a smaller, male one). The term is used when the female gamete is ...
s, the female gametes. In the ovaries, the developing egg cells (or
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 ...
s) mature in the fluid-filled follicles. Typically, only one oocyte develops at a time, but others can also mature simultaneously. Follicles are composed of different types and number of cells according to the stage of their maturation, and their size is indicative of the stage of oocyte development. When an oocyte completes its maturation in the ovary, a surge of luteinizing hormone is secreted by the
pituitary gland In vertebrate anatomy, the pituitary gland, or hypophysis, is an endocrine gland, about the size of a chickpea and weighing, on average, in humans. It is a protrusion off the bottom of the hypothalamus at the base of the brain. The ...
which stimulates the release of the oocyte through the rupture of the follicle, a process called ovulation. The follicle remains functional and reorganizes into a
corpus luteum The corpus luteum (Latin for "yellow body"; plural corpora lutea) is a temporary endocrine structure in female ovaries involved in the production of relatively high levels of progesterone, and moderate levels of estradiol, and inhibin A. It is t ...
, which secretes
progesterone Progesterone (P4) is an endogenous steroid and progestogen sex hormone involved in the menstrual cycle, pregnancy, and embryogenesis of humans and other species. It belongs to a group of steroid hormones called the progestogens and is the m ...
in order to prepare the
uterus The uterus (from Latin ''uterus'', plural ''uteri'') or womb () is the organ in the reproductive system of most female mammals, including humans that accommodates the embryonic and fetal development of one or more embryos until birth. The uter ...
for an eventual implantation of the embryo.


Hormone secretion

At maturity, ovaries secrete
estrogen Estrogen or oestrogen is a category of sex hormone responsible for the development and regulation of the female reproductive system and secondary sex characteristics. There are three major endogenous estrogens that have estrogenic hormonal acti ...
, androgen,
inhibin Activin and inhibin are two closely related protein complexes that have almost directly opposite biological effects. Identified in 1986, activin enhances FSH biosynthesis and secretion, and participates in the regulation of the menstrual cy ...
, and
progestogen Progestogens, also sometimes written progestagens or gestagens, are a class of natural or synthetic steroid hormones that bind to and activate the progesterone receptors (PR). Progesterone is the major and most important progestogen in the body. ...
. In women before menopause, 50% of testosterone is produced by the ovaries and released directly into the blood stream. The other 50% of testosterone in the blood stream is made from conversion of the adrenal pre-androgens ( DHEA and androstenedione) to testosterone in other parts of the body. Estrogen is responsible for the appearance of secondary sex characteristics for females at
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 girl, the testes in a boy. ...
and for the maturation and maintenance of the reproductive organs in their mature functional state. Progesterone prepares the uterus for pregnancy, and the mammary glands for lactation. Progesterone functions with estrogen by promoting
menstrual cycle The menstrual cycle is a series of natural changes in hormone production and the structures of the uterus and ovaries of the female reproductive system that make pregnancy possible. The ovarian cycle controls the production and release of eggs a ...
changes in the
endometrium The endometrium is the inner epithelial layer, along with its mucous membrane, of the mammalian uterus. It has a basal layer and a functional layer: the basal layer contains stem cells which regenerate the functional layer. The functional laye ...
.


Ovarian aging

As women age, they experience a decline in reproductive performance leading to
menopause Menopause, also known as the climacteric, is the time in women's lives when menstrual periods stop permanently, and they are no longer able to bear children. Menopause usually occurs between the age of 47 and 54. Medical professionals often d ...
. This decline is tied to a decline in the number of ovarian follicles. Although about 1 million oocytes are present at birth in the human ovary, only about 500 (about 0.05%) of these ovulate, and the rest are wasted. The decline in ovarian reserve appears to occur at a constantly increasing rate with age, and leads to nearly complete exhaustion of the reserve by about age 52. As ovarian reserve and fertility decline with age, there is also a parallel increase in pregnancy failure and meiotic errors resulting in chromosomally abnormal conceptions. The ovarian reserve and fertility perform optimally around 20–30 years of age. Around 45 years of age, the menstrual cycle begins to change and the follicle pool decreases significantly. The events that lead to ovarian aging remain unclear. The variability of aging could include environmental factors, lifestyle habits or genetic factors. Women with an inherited mutation in the DNA repair gene BRCA1 undergo menopause prematurely, suggesting that naturally occurring DNA damages in oocytes are repaired less efficiently in these women, and this inefficiency leads to early reproductive failure. The BRCA1 protein plays a key role in a type of DNA repair termed homologous recombinational repair that is the only known cellular process that can accurately repair DNA double-strand breaks. Titus et al. showed that DNA double-strand breaks accumulate with age in humans and mice in primordial follicles. Primordial follicles contain oocytes that are at an intermediate (prophase I) stage of meiosis. Meiosis is the general process in eukaryotic organisms by which germ cells are formed, and it is likely an adaptation for removing DNA damages, especially double-strand breaks, from germ line DNA (see
Meiosis Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately resu ...
and Origin and function of meiosis). Homologous recombinational repair is especially promoted during meiosis. Titus et al. also found that expression of 4 key genes necessary for homologous recombinational repair of DNA double-strand breaks (BRCA1, MRE11, RAD51 and ATM) decline with age in the oocytes of humans and mice. They hypothesized that DNA double-strand break repair is vital for the maintenance of oocyte reserve and that a decline in efficiency of repair with age plays a key role in ovarian aging. A study identified 290 genetic determinants of ovarian ageing, also found that
DNA damage DNA repair is a collection of processes by which a cell identifies and corrects damage to the DNA molecules that encode its genome. In human cells, both normal metabolic activities and environmental factors such as radiation can cause DNA da ...
response processes are implicated and suggests that possible effects of extending fertility in women would improve bone health, reduce risk of type 2 diabetes and increase the risk of hormone-sensitive cancers. A variety of testing methods can be used in order to determine fertility based on maternal age. Many of these tests measure levels of hormones FSH, and GnrH. Methods such as measuring AMH (anti-mullerian) hormone levels, and AFC (antral follicule count) can predict ovarian aging. AMH levels serve as an indicator of ovarian aging since the quality of ovarian follicles can be determined.


Clinical significance

Ovarian disease Ovarian diseases refer to diseases or disorders of the ovary. These can be classified as endocrine disorders or as a disorders of the reproductive system. If the egg fails to release from the follicle in the ovary an ovarian cyst may form. Smal ...
s can be classified as
endocrine disorder Endocrine diseases are disorders of the endocrine system. The branch of medicine associated with endocrine disorders is known as endocrinology. Types of disease Broadly speaking, endocrine disorders may be subdivided into three groups: # Endocrin ...
s or as a disorders of the
reproductive system The reproductive system of an organism, also known as the genital system, is the biological system made up of all the anatomical organs involved in sexual reproduction. Many non-living substances such as fluids, hormones, and pheromones are als ...
. If the egg fails to release from the follicle in the ovary an
ovarian cyst An ovarian cyst is a fluid-filled sac within the ovary. Often they cause no symptoms. Occasionally they may produce bloating, lower abdominal pain, or lower back pain. The majority of cysts are harmless. If the cyst either breaks open or causes ...
may form. Small ovarian cysts are common in healthy women. Some women have more follicles than usual (
polycystic ovary syndrome Polycystic ovary syndrome, or PCOS, is the most common endocrine disorder in women of reproductive age. The syndrome is named after the characteristic cysts which may form on the ovaries, though it is important to note that this is a sign and no ...
), which inhibits the follicles to grow normally and this will cause cycle irregularities.


Society and culture


Cryopreservation

Cryopreservation of ovarian tissue, often called ''ovarian tissue cryopreservation'', is of interest to women who want to preserve their reproductive function beyond the natural limit, or whose reproductive potential is threatened by cancer therapy, for example in hematologic malignancies or breast cancer. The procedure is to take a part of the ovary and carry out slow freezing before storing it in liquid nitrogen whilst therapy is undertaken. Tissue can then be thawed and implanted near the fallopian, either orthotopic (on the natural location) or heterotopic (on the abdominal wall), where it starts to produce new eggs, allowing normal conception to take place. A study of 60 procedures concluded that ovarian tissue harvesting appears to be safe. The ovarian tissue may also be transplanted into mice that are immunocompromised (
SCID mice The severe combined immunodeficiency (SCID) is a severe immunodeficiency genetic disorder that is characterized by the complete inability of the adaptive immune system to mount, coordinate, and sustain an appropriate immune response, usually due t ...
) to avoid graft rejection, and tissue can be harvested later when mature follicles have developed.


History

In former centuries, medical authors, for example
Galen Aelius Galenus or Claudius Galenus ( el, Κλαύδιος Γαληνός; September 129 – c. AD 216), often Anglicized as Galen () or Galen of Pergamon, was a Greek physician, surgeon and philosopher in the Roman Empire. Considered to be one of ...
, referred to a woman's ovaries as "female testes".


Other animals

Birds have only one functional ovary (the left), while the other remains vestigial. Ovaries in females are analogous to
testes 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 testoster ...
in males, in that they are both
gonad A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces sper ...
s and
endocrine gland Endocrine glands are ductless glands of the endocrine system that secrete their products, hormones, directly into the blood. The major glands of the endocrine system include the pineal gland, pituitary gland, pancreas, ovaries, testes, thyroid ...
s. Ovaries of some kind are found in the female reproductive system of many animals that employ
sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote tha ...
, including invertebrates. However, they develop in a very different way in most invertebrates than they do in vertebrates, and are not truly homologous. Many of the features found in human ovaries are common to all vertebrates, including the presence of follicular cells, tunica albuginea, and so on. However, many species produce a far greater number of eggs during their lifetime than do humans, so that, in fish and amphibians, there may be hundreds, or even millions of fertile eggs present in the ovary at any given time. In these species, fresh eggs may be developing from the germinal epithelium throughout life. Corpora lutea are found only in mammals, and in some elasmobranch fish; in other species, the remnants of the follicle are quickly resorbed by the ovary. In birds, reptiles, and
monotreme Monotremes () are prototherian mammals of the order Monotremata. They are one of the three groups of living mammals, along with placentals (Eutheria), and marsupials (Metatheria). Monotremes are typified by structural differences in their brain ...
s, the egg is relatively large, filling the follicle, and distorting the shape of the ovary at maturity. Amphibians and reptiles have no ovarian medulla; the central part of the ovary is a hollow,
lymph Lymph (from Latin, , meaning "water") is the fluid that flows through the lymphatic system, a system composed of lymph vessels (channels) and intervening lymph nodes whose function, like the venous system, is to return fluid from the tissues to ...
-filled space. The ovary of teleosts is also often hollow, but in this case, the eggs are shed into the cavity, which opens into the oviduct. Certain
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
s of the genus ''
Philometra ''Philometra'' is a genus of nematodes, which are parasites of marine and freshwater fishes. The genus was erected by Oronzio Gabriele Costa in 1845. Species in this genus are worldwide. They parasitize the body cavities, tissues and ovaries of ...
'' are parasitic in the ovary of marine fishes and can be spectacular, with females as long as 40 cm, coiled in the ovary of a fish half this length. Although most normal female vertebrates have two ovaries, this is not the case in all species. In most birds and in
platypus The platypus (''Ornithorhynchus anatinus''), sometimes referred to as the duck-billed platypus, is a semiaquatic, egg-laying mammal Endemic (ecology), endemic to Eastern states of Australia, eastern Australia, including Tasmania. The platypu ...
es, the right ovary never matures, so that only the left is functional. (Exceptions include the
kiwi Kiwi most commonly refers to: * Kiwi (bird), a flightless bird native to New Zealand * Kiwi (nickname), a nickname for New Zealanders * Kiwifruit, an edible berry * Kiwi dollar or New Zealand dollar, a unit of currency Kiwi or KIWI may also refe ...
and some, but not all raptors, in which both ovaries persist.) In some elasmobranchs, only the right ovary develops fully. In the primitive jawless fish, and some teleosts, there is only one ovary, formed by the fusion of the paired organs in the embryo.


Additional images

File:Slide1DDD.JPG, Left Ovary File:Ovaries.jpg, Ovaries


See also

*
Ovarian reserve Ovarian reserve is a term that is used to determine the capacity of the ovary to provide egg cells that are capable of fertilization resulting in a healthy and successful pregnancy. With advanced maternal age the number of egg cell that can be succe ...
* Folliculogenesis * Oophorectomy * Ovarian drilling *
Artificial ovary An artificial ovary is a potential fertility preservation treatment that aims to mimic the function of the natural ovary. Conventional fertility preservation for females involves oocyte cryopreservation or ovarian tissue cryopreservation. However ...
* Ovary culture


References


Bibliography

*


External links


From the American Medical Association


{{Authority control Organs (anatomy) Endocrine system Pelvis Human female reproductive system Women's health Gynaecology