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From Wikipedia, the free encyclopedia

The ovary is an organ found in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary (from Latin ovarium, meaning 'egg, nut') found on the left and right sides of the body. The ovaries also secrete hormones that play a role in the menstrual cycle and fertility. The ovary progresses through many stages beginning in the prenatal period through menopause. It is also an endocrine gland because of the various hormones that it secretes.[1]

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  • ✪ The ovarian cycle | Reproductive system physiology | NCLEX-RN | Khan Academy
  • ✪ Structure or Histology of ovary (Female reproductive system / Human reproduction)
  • ✪ Germ cell ovarian tumors - causes, symptoms, diagnosis, treatment, pathology
  • ✪ Dermoid Cyst of Ovary - Laparoscopic Removal Without Spillage
  • ✪ Laparoscopic Surgery for Left Ovarian Dermoid Cyst

Transcription

- We're gonna talk about the ovarian cycle. The ovaries are two structures in a female's reproductive system that produce her eggs. Each month her eggs go through a maturation process called the ovarian cycle, and that cycle creates a secondary oocyte than can be then fertilized by a sperm to result in a pregnancy. The ovarian cycle is also responsible for what we commonly know as the menstrual cycle. Basically, the primary oocytes that are destined to be ovulated will develop in the ovaries, complete meiosis one just before ovulation, and then they'll be ejected out of the ovary as a secondary oocyte to be picked up by the fimbriae and swept into the uterine tube to hope for fertilization. So let's start from the beginning. Inside the ovaries, eggs develop in structures called follicles, these purple circles here. And they start off as primordial follicles. And so what a follicle is- I'll just blow that up for you- It's one primary oocyte, so an egg cell, surrounded by a layer of cells called granulosa cells. And the granulosa cells develop and become more numerous as the follicle matures. Now the granulosa cells also secrete a few hormones. Estrogen, a little progesterone and some inhibin, and we'll talk about the functions of those a little bit later on. So let's put a timeline on this. Now the ovarian cycle lasts 28 days. This is day zero here at the primordial follicle, where we're going counter-clockwise. All the way over here, this is day 13. Here, where the secondary oocyte gets ejected, or ovulated, that's day 14. And then the rest of the time spent getting back to the primordial follicle stage are days 15 through 28. So now you have an idea of about how long this all takes. So you remember when we said that the granulosa cells produce hormones? Well, as the follicles develop over the first 13 days, and you can see the changes between the one here and the one here. It's got a lot more purple cells around here. Those are granulosa cells. So the number of granulosa cells goes up, and since they produce hormones, what do you think happens to the hormone levels in the blood? They go up. So that's sort of just a general point. So keep that in mind, but first we'll jump back to these. We know these are primordial follicles here. The next stage of development are these guys here, and these are called primary follicles. And in the primary follicles, the layers of granulosa cells and the oocyte, the egg, start to be separated by this other layer that starts to form between them. That's called the zona pellucida, and I'll draw it here in light blue. And even though the egg I've drawn in blue, there's still a layer of zona pellucida, even though the egg is originally drawn in blue because I wanted to draw the egg in blue. There's still a layer of zone pellucida around it. Now even though the zona pellucida is there separating the granulosa cells from the actual egg, the granulosa cells can still nourish the egg through gap junctions that go through the zona pellucida and into the egg. Gap junctions are just little passageways from one cell to another cell where they can exchange nutrients or other signals. And actually, through those gap junctions, the granulosa cells send through little chemicals that keep those primary oocytes stuck at that meiosis one stage, 'cause you remember at this point all of these primary oocytes are stuck in meiotic arrest. They're not dividing and reducing their chromosome copy number. So as we develop from our primordial to our primary to our next follicle here, called our pre-antral follicle, and you'll see why it's called that in a minute, the granulosa cells are actually starting to divide and become a lot greater in number. You can see that there's a pretty big difference in granulosa cell number from our primary follicles to our pre-antral follicle here. And remember the granulosa cells are shaded in in purple here. So while the granulosa cells are proliferating, this wall on the outside of the follicle called the theca starts to form. Theca cells have receptors for luteinizing hormone from the anterior pituitary, and when luteinizing hormone, or LH, binds these theca cells, they produce a hormone called androstenedione. And when the thecas get androstenedione, they give it to the granulosa cells, who then convert it to estrogen and release it into the blood. So the blood estrogen levels start to go up at this point. And so that's what these red and blue bits running down the middle of the ovary are, blood vessels, arteries and veins. And if they look a little bit weird to you, or unusual, that's just because they're cut in cross-section as well. Now you might be wondering what an antral refers to, like what you see in the pre-antral follicle and this early antral follicle here. It actually refers to the antrum, which will be formed in the next step. This space here is called an antrum. And the antrum is just basically fluid that's being produced by the granulosa cells. And it's that antrum and the fluid in the antrum that pushes against the edges of the follicle and causes it to expand. Now just so you're aware, during this ovarian cycle, multiple follicles are actually forming. It's not just this one pre-antral follicle, and then this one early antral follicle, and this one mature follicle. You're getting a lot of these happening at one time. But only one of the biggest ones is the one that eventually gets ovulated, because you only ovulate one egg every 28 days. And that one that gets ovulated is called the dominant follicle. So let's just say that what we're seeing here is an example of the dominant follicle's development. Because the rest of the ones that were developing along this pathway sort of degenerate and die off in a process called atresia. So I'll write that at the bottom here. And atresia just means to degenerate. So another note. In the ones that undergo atresia, both the follicle and the eggs they contain die off. And that means that a woman loses anywhere between 15 to 25 eggs per menstrual cycle to atresia, while only one gets ovulated. So you can kind of imagine how you go from two to four million eggs when you were born to having zero after about 35-ish years of ovulation. It's not just that one egg you lose by ovulation. You lose quite a few. So anyway, back to the development of the dominant follicle. It enlarges mostly due to the expanding antrum, as I mentioned earlier. And granulosa cells actually start to form this bit of a mound here that protrudes into the middle of the antrum. This mound of granulosa cells is called the cumulus oophorus. As part of the development of the dominant follicle, the cumulus oophorus and the egg sort of separate together from the wall of the follicle and float around in the middle of the antrum, like a little island. And the follicle increases in size. So the actual follicle is increasing in size as it gets filled with more and more fluid from the granulosa cells. And the granulosa cells are just producing fluid as a by-product of their metabolism and creation of hormones. Eventually this dominant follicle, which at this point is called the mature follicle, it starts to balloon out the side of the ovary, kind of like this. Just starts to push out against the edge of the ovary. And then because the edge of the ovary and the wall of the mature follicle are in such close proximity, enzymes within the follicle break down that common wall between them, and the egg pops out onto the surface of the ovary, because now this wall is broken down. And by the way, an enzyme is a protein that carries out a specific task. The task here is to break down the wall between the mature follicle and the ovary, and that happens on day 14. So it takes day zero to 13 of build up to get to this event. When this happens, some women feel a little bit of pelvic pain. And actually sometimes, by chance, two or more follicles reach maturity, and they all pop out. And that's how you get twins or triplets or quadruplets or octuplets, when they all pop out and get fertilized by different sperm each. Because they're all subsequently swept up into the uterine tubes where sperm can fertilize them. So now you have the egg out here, but what about the old follicle it was in? The follicle actually collapses a little and transforms into a structure called the corpus luteum. And in this transformation the granulosa cells get a lot bigger and start to produce more estrogen, progesterone and that other hormone, inhibin, that we mentioned before. Just briefly, inhibin lowers the amount of FSH, follicle stimulating hormone, that comes from the anterior pituitary. And it does that because follicle stimulating hormone actually propagates this whole process of follicle maturation, as you can imagine from the name. So if you didn't know this before, these are the exact follicles that follicle stimulating hormone refers to. At least in a female. Anyway, if the egg doesn't get fertilized, then the corpus luteum reaches a maximum size in about 10 days. So that's about day 25, which it's probably sitting at in this diagram. And then it degenerates by apoptosis. That's a process that cells use to sort of self-destruct and die off. And here I'm abbreviating corpus luteum as CL, just so you know what I mean. But if the egg is fertilized, i.e., it travels into the uterine tubes and gets fertilized by a sperm, then the corpus luteum persists, I mean it keeps living, because we want it to keep producing estrogen and progesterone. That's because estrogen and progesterone prepare the inner lining of the uterus, that's called the endometrium, for implantation, which would be really handy since we have a fertilized egg now that needs to develop. And that's where it does it, by implanting in the endometrium of the uterus. So just a final note. Ovulation doesn't happen forever. At about age 50 to 51, females undergo something called menopause. First menstrual cycles become less and less regular. In other words, they don't happen every 28 days like they do when you're under the age of 50. And then ultimately, they stop happening entirely. And that cessation of ovulation is called menopause. The main cause of menopause is sometimes referred to as ovarian failure. Basically the ovaries lose the ability to respond to signalling hormones from the brain called gonadotropins. And we know these as LH and FSH. And this happens because most, or all of the follicles and eggs have already gone through that process that we talked about called atresia. In other words, they've degenerated.

Contents

Structure

The ovaries are considered the female gonads.[2] Each ovary is whitish in color and located alongside the lateral wall of the uterus in a region called the ovarian fossa. 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. This area is about 4 cm x 3 cm x 2 cm in size.[3][4] The ovaries are surrounded by a capsule, and have an outer cortex and an inner medulla.[4]

Usually, ovulation occurs in one of the two ovaries releasing an egg each menstrual cycle; however, if there was a case where one ovary was absent or dysfunctional then the other ovary would continue providing eggs to be released without any changes in cycle length or frequency.[medical citation needed]

The side of the ovary closest to the fallopian tube is connected to it by infundibulopelvic ligament,[3] and the other side points downwards attached to the uterus via the ovarian ligament.

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 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 that covers the ovary is known as the mesovarium.[4]

The ovarian pedicle is made up part of the fallopian tube, mesovarium, ovarian ligament, and ovarian blood vessels.[5]

Microanatomy

The surface of the ovaries is covered with membrane consisting of a lining of simple cuboidal-to-columnar shaped mesothelium.[6]

The outermost layer is called the germinal epithelium.

The outer layer is the ovarian cortex, consisting of ovarian follicles and stroma in between them. Included in the follicles are the cumulus oophorus, membrana granulosa (and the granulosa cells inside it), corona radiata, zona pellucida, and primary oocyte. Theca of follicle, antrum and liquor folliculi are also contained in the follicle. Also in the cortex is the corpus luteum derived from the follicles. The innermost layer is the ovarian medulla.[7] It can be hard to distinguish between the cortex and medulla, but follicles are usually not found in the medulla.

Follicular cells flat epithelial cells that originate from surface epithelium covering the ovary, are surrounded by Granulosa cells - that have changed from flat to cuboidal and proliferated to produce a stratified epithelium

Other

The ovary also contains blood vessels and lymphatics.[9]

Function

At puberty, the ovary begins to secrete increasing levels of hormones. Secondary sex characteristics begin to develop in response to the hormones. The ability to produce eggs and reproduce develops. The ovary changes structure and function beginning at puberty.[1]

Gamete production

The ovaries are the site of production and periodical release of egg cells, the female gametes. In the ovaries, the developing egg cells (or oocytes) 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.[10]:833

When the oocyte finishes its maturation in the ovary, a surge of luteinizing hormone secreted by the pituitary gland stimulates the release of the oocyte through the rupture of the follicle, a process called ovulation.[11] The follicle remains functional and reorganizes into a corpus luteum, which secretes progesterone in order to prepare the uterus for an eventual implantation of the embryo.[10]:839

Hormone secretion

At maturity, ovaries secrete estrogen, testosterone,[12][13] inhibin, and progesterone.[14][15][1] In women, fifty percent of testosterone is produced by the ovaries and adrenal glands and released directly into the blood stream.[16] Estrogen is responsible for the appearance of secondary sex characteristics for females at puberty 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 changes in the endometrium.[medical citation needed]

Ovarian aging

As women age, they experience a decline in reproductive performance leading to menopause. 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,[17] 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.[medical citation needed]

Women with an inherited mutation in the DNA repair gene BRCA1 undergo menopause prematurely,[18] 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.[19] 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.[20] (see Meiosis and Origin and function of meiosis). Homologous recombinational repair is especially promoted during meiosis. Titus et al.[19] 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.

Clinical significance

Ovarian diseases can be classified as endocrine disorders or as a disorders of the reproductive system.[medical citation needed]

If the egg fails to release from the follicle in the ovary an ovarian cyst may form. Small ovarian cysts are common in healthy women. Some women have more follicles than usual (polycystic ovary syndrome), which inhibits the follicles to grow normally and this will cause cycle irregularities.

Notes Ref(s)
Ovarian neoplasms
Germ cell tumor Seen most often in young women or adolescent girls
Other germ cell tumors are: Endodermal sinus tumor and teratoma,
[21]
Ovarian cancer includes ovarian epithelial cancer [22][23][24]
Luteoma [25]
Ovaritis syn. oophoritis [15]
Ovarian remnant syndrome [15]
Hypogonadism
Hyperthecosis
Ovarian torsion
Ovarian apoplexy (rupture)
Premature ovarian failure
Anovulation
Follicular cyst of ovary
Corpus luteum cyst
Theca lutein cyst
Chocolate cyst
Ovarian germ cell tumors benign [26]
Dysgerminoma
Choriocarcinoma
Yolk sac tumor
Teratoma
Ovarian serous cystadenoma
Serous cystadenocarcinoma
Mucinous cystadenoma
Mucinous cystadenocarcinoma
Brenner tumor
Granulosa cell tumor
Krukenberg tumor

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,[27] for example in hematologic malignancies or breast cancer.[28] 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),[28] where it starts to produce new eggs, allowing normal conception to take place.[29] A study of 60 procedures concluded that ovarian tissue harvesting appears to be safe.[28] The ovarian tissue may also be transplanted into mice that are immunocompromised (SCID mice) to avoid graft rejection, and tissue can be harvested later when mature follicles have developed.[30]

Other animals

Ovary of a marine fish and its parasite, the nematode Philometra fasciati

Birds have only one functional ovary (the left), while the other remains vestigial. Ovaries in females are analogous to testes in males, in that they are both gonads and endocrine glands. Ovaries of some kind are found in the female reproductive system of many animals that employ sexual reproduction, including invertebrates. However, they develop in a very different way in most invertebrates than they do in vertebrates, and are not truly homologous.[31]

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 monotremes, the egg is relatively large, filling the follicle, and distorting the shape of the ovary at maturity.[31]

Amphibians and reptiles have no ovarian medulla; the central part of the ovary is a hollow, lymph-filled space.[32]

The ovary of teleosts is also often hollow, but in this case, the eggs are shed into the cavity, which opens into the oviduct.[31] Certain nematodes of the genus Philometra 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.[33] Although most normal female vertebrates have two ovaries, this is not the case in all species. In most birds and in platypuses, the right ovary never matures, so that only the left is functional. (Exceptions include the kiwi and some, but not all raptors, in which both ovaries persist.[34][35]) 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.[31]

Additional images

See also

References

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  2. ^ "Dorlands Medical Dictionary". www.mercksource.com. Retrieved 2017-11-20.
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  5. ^ Baskett, Thomas F.; Calder, Andrew A.; Arulkumaran, Sabaratnam (2014). Munro Kerr's Operative Obstetrics E-Book. Elsevier Health Sciences. p. 268. ISBN 9780702052484.
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  20. ^ Harris Bernstein, Carol Bernstein and Richard E. Michod (2011). Meiosis as an Evolutionary Adaptation for DNA Repair. Chapter 19 in DNA Repair. Inna Kruman editor. InTech Open Publisher. DOI: 10.5772/25117 http://www.intechopen.com/books/dna-repair/meiosis-as-an-evolutionary-adaptation-for-dna-repair
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  25. ^ "MeSH Browser". meshb.nlm.nih.gov. Retrieved 2017-12-01.
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  29. ^ Livebirth after orthotopic transplantation of cryopreserved ovarian tissue The Lancet, Sep 24, 2004
  30. ^ Lan C, Xiao W, Xiao-Hui D, Chun-Yan H, Hong-Ling Y (December 2008). "Tissue culture before transplantation of frozen-thawed human fetal ovarian tissue into immunodeficient mice". Fertil. Steril. 93 (3): 913–9. doi:10.1016/j.fertnstert.2008.10.020. PMID 19108826.
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  32. ^ "Animal reproductive system - Accessory glands".
  33. ^ Moravec, František; Justine, Jean-Lou (2014). "Philometrids (Nematoda: Philometridae) in carangid and serranid fishes off New Caledonia, including three new species". Parasite. 21: 21. doi:10.1051/parasite/2014022. ISSN 1776-1042. PMC 4023622. PMID 24836940. open access
  34. ^ Fitzpatrick, F. L. (1934). "Unilateral and bilateral ovaries in raptorial birds". The Wilson Bulletin. 46 (1): 19–22.
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Bibliography

  • Venes, Donald (2013). Taber's cyclopedic medical dictionary. Philadelphia: F.A. Davis. ISBN 9780803629790.

External links

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