Ovary Development

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Human ovary with corpus luteum.
Adult human ovary viewed by endoscopy

The female gonad is the ovary and is closely associated with female internal genital (reproductive) tract development. In humans, these laterally paired organs lie within the peritoneal cavity. Genes such as Wnt-4 and DAX-1 necessary for initiation of female pathway ovary development, female gonad is not considered a default process.

Initial gonad development in females and males is virtually identical with germ cells migrating into an indifferent gonad. In females with XX, the ovary then begins to develop and the subsequent structure and timecourse of germ cell then differs between males and females. In the ovary oocytes proliferate prior to birth and arrest in meiosis 1.

Links: Menstrual Cycle | X Chromosome | Category:Ovary

Genital Links: Introduction | Lecture - Medicine | Lecture - Science | Online Practical | Primordial Germ Cell | Meiosis | Female | Ovary | Oocyte | Uterus | Vagina | Male | Testis | Spermatozoa | Prostate | Genital Movies | Abnormalities | Assisted Reproductive Technology | Puberty | Category:Genital
Historic Embryology: 1912 Urinogenital Organ Development | 1921 Urogenital Development | 1921 External Genital Development | Historic Disclaimer

Some Recent Findings

  • Retransplantation of cryopreserved ovarian tissue: the first live birth in Germany[1] "Cryopreserved ovarian tissue can be retransplanted to restore fertility after radiation or chemotherapy. To date, 15 live births after retransplantation have been reported worldwide. We report the first pregnancy and the first live birth after retransplantation in Germany. This was the first live birth after retransplantation of cryopreserved ovarian tissue in Germany and also the first case with histological confirmation that the oocyte from which the patient conceived could only have come from the retransplanted tissue."
  • Mammalian ovary differentiation - A focus on female meiosis[2] "Over the past 50 years, the ovary development has been subject of fewer studies as compare to the male pathway. Nevertheless due to the advancement of genetics, mouse ES cells and the development of genetic models, studies of ovarian differentiation was boosted. This review emphasizes some of new progresses in the research field of the mammalian ovary differentiation that have occurred in recent years with focuses of the period around prophase I of meiosis and of recent roles of small non-RNAs in the ovarian gene expression."
  • Human RSPO1/R-spondin1 is expressed during early ovary development and augments β-catenin signaling[3] "Human testis development starts from around 42 days post conception with a transient wave of SRY expression followed by up-regulation of testis specific genes and a distinct set of morphological, paracrine and endocrine events. Although anatomical changes in the ovary are less marked, a distinct sub-set of ovary specific genes are also expressed during this time. The furin-domain containing peptide R-spondin1 (RSPO1) has recently emerged as an important regulator of ovary development through up-regulation of the WNT/β-catenin pathway to oppose testis formation. Here, we show that RSPO1 is upregulated in the ovary but not in the testis during critical early stages of gonad development in humans (between 6-9 weeks post conception), whereas the expression of the related genes WNT4 and CTNNB1 (encoding β catenin) is not significantly different between these tissues. Furthermore, reduced R-spondin1 function in the ovotestis of an individual (46,XX) with a RSPO1 mutation leads to reduced β-catenin protein and WNT4 mRNA levels, consistent with down regulation of ovarian pathways"

Human Ovary Timeline

Fetal gonad retinoid receptor expression[4]

Approximate Timeline of human development listed below.

24 days - intermediate mesoderm, pronephros primordium

28 days - mesonephros and mesonephric duct

35 days - uteric bud, metanephros, urogenital ridge

42 days - cloacal divison, gonadal primordium (indifferent)

49 days - paramesonephric duct, gonadal differentiation

56 days - paramesonephric duct fusion (female)

100 days - primary follicles (ovary)


Human ovary non-growing follicle model

The 2 human ovaries gradually lose follicles both before and after puberty (the beginning of ovulation); beginning with about several million before birth, maximum number at birth, 300-400,000 by puberty and finally by late 40’s have only a few follicles left. Recent studies suggest that the original calculations of ovary follicle numbers at birth were over-estimates and the actual figure should be about 2.5 million.[5] The number of antral follicles detected within the ovary also decreases with increasing materal age.

In humans, a primodial follicle take about 150 days to develop into a preantral follicle (primary) and another 120 days to form an antral follicle (secondary). A number of antral follicles will then "compete" for 14-15 days to become the dominant follicle, which will undergo ovulation.

Infant Ovary

Histology of the female infant ovary Human infant ovary follicle 01.jpg
Overview Follicle

This image shows a region (see inset) of the infant ovary cortex.

There are a large number of developing oocytes which will eventually form a dense primordial germ layer at the ovary periphery.

Later stages of follicle development are completely absent and will begin to only appear just prior to puberty.

Postnatal Oogenesis

There is a dogma in mammalian development that new oocyte and follicle production does not occur during postnatal life. There is substantial data that shows human ovarian changes postnatally are loss by apoptosis of prenatal oocytes. A research group (Tilly JL, Johnson J. 2004, 2007) has recently published experiments using mice, showing potentially other sources/sites (bone marrow) of oocyte (putative germ cell) generation. They recently stated that the argument should be based upon "experimental approaches than simply an absence of evidence, especially from gene expression analyses". Several other research groups (Eggan K etal. 2004 and Veitia etal. 2007) have argued against these findings.

Adult Follicle Structure

Secondary (Antral) Follicle Structure

A follicle usually contains a single oocyte (egg, ovum, female gamete) and a series of supporting cells and a single fluid-filled space in layers surrounding this cell. The 3 layers below are arranged in layers outward from the oocyte.

Granulosa Cells

  • A specific cell type that proliferates in association with the oocyte within the developing follicles of the ovary. These cells form the follicle stratum granulosa and are also given specific names based upon their position within the follicle.
  • With development of the antral follicle, there are two populations of granulosa cells with distinct characteristics and functions: mural granulosa cells and cumulus cells.[6]
    • mural granulosa cells - an endocrine role by producing steroid hormones and various other ligands
    • cumulus cells - play a support role for oocyte development

Alternate Histological Terms

    • The membrana granulosa cells sit on the follicular basal lamina and line the antrum as a stratified epitelium. Following ovulation, these granulosa cells contribute to corpus luteum.
    • The cumulus oophorus is a column of granulosa cells that attaches the oocyte to the follicle wall. At ovulation, this column of cells is broken or separates to release the oocyte from its follicle attachment.
    • The corona radiata are the granulosa cells that directly surround the oocyte, and are released along with it at ovulation. Following ovulation, the corona radiata provide physical protection to the oocyte and are the initial structural barrier that spermatazoa must penetrate during fertilization.

Follicular Fluid

  • The antrum is a fluid-filled space in the secondary (antral) follicle
  • At ovulation, fluid is released along with the oocyte
  • Thought to "carry" the oocyte out of the follicle (like a boat on a wave)
  • Aids entry into the uterine tube

Theca Interna

(Greek, thek = box) The ovarian follicle endocrine cells forming the inner layer of the theca folliculi surrounding the developing follicle within the ovary. This vascularized layer of cells respond to leutenizing hormone (LH) synthesizing and secreting androgens (androstendione) transported to glomerulosa cells which process initially into testosterone and then by aromatase into estrogen (estradiol). Theca cells do not begin hormonal functions until puberty.

Theca Externa

(Greek, thek = box) The ovarian follicle stromal cells forming the outer layer of the theca folliculi surrounding the developing follicle within the ovary. Consisting of connective tissue cells, smooth muscle and collagen fibers.

Follicle Classification

There are several different nomenclatures for the stages of follicle maturation. It probably does not matter which naming system you use, as long as you are consistent and use the same set of terminology for all stages. Early stages of follicle development appear to be gonadotropin (Gn) independent and with development become gonadotropin "sensitive" and then "dependent" . (UK spelling is gonadotrophin).

Follicle development stages and the relationship to gonadotropin (Gn)[7]
Human ovary follicle development

Primordial Follicle

Alternate nomenclature: small follicle or type 1, 2, 3 (25cells)

Primary Follicle

Alternate nomenclature: preantral follicle or type 4 (26-100 cells), type 5 (101-300 cells)

Secondary Follicle

Alternate nomenclature: small and large antral follicle or type 6 (3001-500 cells), type 7 (501-1000 cells)

Preovulatory Follicle

Alternate nomenclature: Graafian follicle or type 8 (>1000 cells)


At any one time the majority of follicles are destined not to complete maturation and at any stage (from type 4-7) degeneration of the follicle can occur. Cells die by apoptosis.

Follicle Factors

Ovarian developmental genes[8]

There are both external endocrine factors and internal follicle factors that can influence the development and atresia of ovarian follicles.

External Factors

Leutenizing Hormone (LH)

  • from the anterior pituitary
  • stimulate the theca interna to synthesize and secrete androgens (androstendione) transported to granulosa cells
  • granulosa cells process initially into testosterone and then by aromatase into estrogen (estradiol)

Follicle-stimulating hormone (FSH)

  • from the anterior pituitary
  • initiates follicle growth through the granulosa cells
  • involved in selecting the most advanced (sensitive) follicle to proceed to ovulation

Internal Factors

Oocyte Factors

  • Growth Differentiation Factor-9 (GDF-9) - involved in the differentiation of theca cells during this early stage of follicular development OMIM 601918
  • Bone morphogenetic protein 15 (BMP15)
  • Fibroblast growth factor 8B (FGF8B)

Granulosal Factor(s)

  • stimulates the recruitment of theca cells from cortical stromal cells

Thecal Factor(s)

  • appear to be several inhibitors of apoptotic cell death
  • Epidermal growth factor (EGF)
  • Transforming growth factor alpha (TGF-α)
  • keratinocyte growth factor (KGF)
  • hepatocyte growth factor (HGF)
  • Bone morphogenetic protein 7 (BMP-7) also known as osteogenic protein-1 or OP-1

Ovary Growth

Table below is from historic data (1912) from measurement of histological materials.[9]

Table showing the Growth of the Ovary
Greatest diam.
of head
Right Ovary Left Ovary Comparison between
Breadth Length Breadth Length Length Breadth
R. L. R. L.
50.0 42.6 0.9 1.9 0.9 2.5 .. +
125.0 123.0 1.2 5.9 1.5 4.1 + .. .. +
138.0 115.0 1.6 5.0 2.0 5.0 .. +
156.0 131.0 1.9 7.2 2.0 7.1 + .. .. +
173.0 163.5 3.0 9.0 ... ... .. .. .. ..
190.0 175.0 2.9 7.7 2.1 7.8 .. + + ..
223.0 162.0 2.9 10.5 3.0 9.1 + .. .. +
235.0 190.0 4.2 10.0 3.8 12.0 .. + + ..
260.0 213.0 3.6 11.1 4.0 11.4 .. + .. +
272.0 213.0 3.0 10.0 3.5 9.2 + .. .. +
305.0 238.0 3.0 9.9 3.9 10.9 .. + .. +
347.0 ... 3.5 10.8 4.9 8.5 + .. .. +
355.0 273.0 4.0 14.0 5.2 9.9 + .. .. +
386.0 324.0 5.1 11.5 3.0 9.9 + .. + ..
402.0 301.0 5.05 10.5 3.0 12.0 .. + + ..
3 weeks ... 5.0 17.0 5.0 14.0 + ..
6 weeks ... 7.5 15.0 7.0 14.7 + .. + ..
6 weeks ... 7.0 18.0 8.0 17.0 + .. .. +
10 weeks ... ... 14.0 ... 16.0 + ..
2 months ... 6.0 14.5 4.0 13.0 + .. + ..
3 months ... 6.0 15.5 5.0 14.7 + .. + ..
7 months ... 5.9 15.5 4.5 18.1 .. + + ..
15 months ... 9.0 18.0 9.0 19.5 + ..
I.75 years ... 7.0 20.0 8.5 15.0 + .. .. +
4 years ... 10.0 27.0 12.7 23.2 + .. .. +
5.5 years ... 11.1 29.0 9.1 26.1 + .. + ..
14 years ... 11.9 26.5 12.0 29.5 .. + .. +
The measurements are all given in millimeters, breech length is measured along the nape and the back.
    Links: 1912 Ovary Growth | Table Ovary Growth Table | Collapsible Table | Ovary Development

Postnatal Growth

Human ovary postnatal growth.jpg

Human ovary postnatal volume growth[10]

Corpus Luteum

Human ovary with corpus luteum (white ring).

(Latin, corpus = body, luteum = yellow) The remains of ovarian follicle formed after ovulation that acts as an endocrine organ (produce progesterone and oestrogens) supporting pregnancy and preventing menstruation (loss of the endometrial lining). Formed during the luteal phase (secretory phase) of the menstrual cycle by proliferation of both follicular granulosa cells (granulosa lutein cells) and thecal cells (theca lutein cells), which together interact to produce progesterone and oestrogens.

Peak luteal function during the menstrual cycle, determined by maximum luteal area, progesterone concentration and estradiol concentration, is observed about 6 days following ovulation. [11]

If fertilization and pregnancy does not occur, the corpus luteum degenerates to form the corpus albicans.


  • Regnier de Graaf (1641 – 1673) was the first observer in the ovary of a cow as a yellow structure, the yellow colour was caused by accumulation of steroidal hormones.
  • Ludwig Fraenkel (1870 - 1951) first identified the endocrine function of the corpus luteum.[12]

Embryo Virtual Slide

Human Ovary and Corpus Luteum

Human ovary - corpus luteum 01.jpg

 ‎‎Mobile | Desktop | Original

Ovary | Embryo Slides
Corpus Luteum Links: anatomy overview| histology overview | low power label | high power label | low power | high power | corpus albicans | theca and granulosa lutein cells | Ovary Development | Menstrual Cycle

Corpus Albicans

Human ovary with corpus albicans (white arrow).

Ovary histology 003.jpg

Corpus albicans histology

(corpora albicantia) (Latin, corpus = body, albicans = whitish) The histological structure formed by luteolysis of the corpus luteum in the ovary. If implantation does not occur and the hormone hCG is not released the corpus luteum degenerates and the structure is white, not yellow, because of the absence of steroid hormone synthesis/accumulation.

Corpus Luteum Links: anatomy overview| histology overview | low power label | high power label | low power | high power | corpus albicans | theca and granulosa lutein cells | Ovary Development | Menstrual Cycle

Histology Images

Ovary histology: Tunica Albuginea x20 | Tunica albuginea, Germinal epithelium x40 | Primary follicle, primordial follicle, oocyte, x40 | Secondary follicle, cumulus oophorus, zona pelucida, granulosa cells, oocyte x20 | Corpus luteum, theca lutein cells, granulosa lutein cells, Loupe | Corpus luteum, theca lutein cells, granulosa lutein cells, x10 | Corpus luteum, theca lutein cells, granulosa lutein cells, x40 | Corpus albicans, primary follicle, primordial follicle, granulosa cells, oocyte x20 | Menstrual Cycle | Ovary Development


See also Genital System - Abnormalities

International Classification of Diseases

E28 Ovarian dysfunction

  • Excl.: isolated gonadotropin deficiency (E23.0); postprocedural ovarian failure (E89.4)

E28.0 Estrogen excess

  • Use additional external cause code (Chapter XX), if desired, to identify drug, if drug-induced.

E28.1 Androgen excess

  • Hypersecretion of ovarian androgens
  • Use additional external cause code (Chapter XX), if desired, to identify drug, if drug-induced.

E28.2 Polycystic ovarian syndrome

  • Sclerocystic ovary syndrome
  • Stein-Leventhal syndrome

E28.3 Primary ovarian failure

  • Decreased oestrogen
  • Premature menopause NOS
  • Resistant ovary syndrome
  • Excl.: menopausal and female climacteric states (N95.1); pure gonadal dysgenesis (Q99.1); Turner syndrome (Q96.-)

E28.8 Other ovarian dysfunction

  • Ovarian hyperfunction NOS

E28.9 Ovarian dysfunction, unspecified

Polycystic Ovary Syndrome

Mouse ovary normal and polycystic ovary syndrome model.[13]

International Classification of Diseases - E28.2 Polycystic ovarian syndrome.

Polycystic ovary syndrome (PCOS) or Stein-Leventhal syndrome (1930s researchers) clinical term for a metabolic hormone syndrome leading to anovulation and with many other symptoms (hyperandrogenism, insulin resistance) and is one of the most common forms of female infertility, see review.[14] Using the European Society for Human Reproduction and Embryology/American Society for Reproductive Medicine criteria, about 15% - 20% of women suffer from this disease. Ovarian cysts arise through incomplete follicular development or failure of ovulation (anovulation). A range of drugs has been used to induce ovulation in these women and more recently in vitro maturation (IVM) has also been suggested as a possible future technique.

In December 2012, the NIH held a workshop on "Evidence-based Methodology on Polycystic Ovary Syndrome". PCOS is a common disorder affecting 5 million women of reproductive-age in the United States. Symptoms include: irregular or no menstrual periods in women of reproductive age (ovulatory dysfunction), acne, weight gain, excess hair growth on the face and body (hirsutism), thinning scalp hair, ovarian cysts (polycystic ovarian morphology) and mental health problems.

One key workshop finding was that the name "Polycystic Ovary Syndrome" is inappropriate for defining the condition and should be renamed to reflect the complex metabolic, hypothalamic, pituitary, ovarian, and adrenal interactions that characterize the syndrome and their reproductive implications.

  • Androgen Excess + Ovulatory Dysfunction
  • Androgen Excess + Polycystic Ovarian Morphology
  • Ovulatory Dysfunction + Polycystic Ovarian Morphology
  • Androgen Excess + Ovulatory Dysfunction + Polycystic Ovarian Morphology

Recurrent pregnancy loss also occurs in about 50% of total pregnancies with polycystic ovary syndrome.[15]

A recent study in the Han Chinese population[16] identified associations between PCOS and three genetic loci: 2p16.3, 2p21 and 9q33.3.

Links: Menstrual Cycle | Genital Abnormalities | Endocrine Abnormalities | 2012 NIH Workshop on PCOS | 2011 Australia Guideline assessment and management PCOS | OMIM 184700]

Ovarian Cancer

Ovarian cancer is a major cause of death in the postnatal female population. A recent mouse study has identified a population of cancer-prone stem cells located at the hilum region of the ovary that are prone to epithelial ovarian cancer.[17]

Luteoma of Pregnancy

Luteoma of pregnancy is a rare nonneoplastic tumor-like mass of the ovary that emerges during pregnancy and regresses spontaneously after delivery. Luteomas can be hormonally active producing androgens that can result in maternal and fetal hirsutism and virilization.

Image: Dr Ed Uthman (Houston, Texas) - other pathology images


Links: Hematoxylin and Eosin | Histology Stains


  1. Andreas Müller, Katja Keller, Jennifer Wacker, Ralf Dittrich, Gudrun Keck, Markus Montag, Hans Van der Ven, David Wachter, Matthias W Beckmann, Wolfgang Distler Retransplantation of cryopreserved ovarian tissue: the first live birth in Germany. Dtsch Arztebl Int: 2012, 109(1-2);8-13 PMID:22282711
  2. Adrienne Baillet, Béatrice Mandon-Pepin Mammalian ovary differentiation - A focus on female meiosis. Mol Cell Endocrinol: 2011; PMID:21964319
  3. Sara Tomaselli, Francesca Megiorni, Lin Lin, Maria Cristina Mazzilli, Dianne Gerrelli, Silvia Majore, Paola Grammatico, John C Achermann Human RSPO1/R-spondin1 is expressed during early ovary development and augments β-catenin signaling. PLoS ONE: 2011, 6(1);e16366 PMID:21297984
  4. Andrew J Childs, Gillian Cowan, Hazel L Kinnell, Richard A Anderson, Philippa T K Saunders Retinoic Acid signalling and the control of meiotic entry in the human fetal gonad. PLoS ONE: 2011, 6(6);e20249 PMID:21674038
  5. W Hamish B Wallace, Thomas W Kelsey Human ovarian reserve from conception to the menopause. PLoS ONE: 2010, 5(1);e8772 PMID:20111701
  6. Maj A Hultén, Suketu Patel, Jon Jonasson, Erik Iwarsson On the origin of the maternal age effect in trisomy 21 Down syndrome: the Oocyte Mosaicism Selection model. Reproduction: 2010, 139(1);1-9 PMID:19755486
  7. Makoto Orisaka, Kimihisa Tajima, Benjamin K Tsang, Fumikazu Kotsuji Oocyte-granulosa-theca cell interactions during preantral follicular development. J Ovarian Res: 2009, 2(1);9 PMID:19589134 | J Ovarian Res.
  8. José Elias Garcia-Ortiz, Emanuele Pelosi, Shakib Omari, Timur Nedorezov, Yulan Piao, Jesse Karmazin, Manuela Uda, Antonio Cao, Steve W Cole, Antonino Forabosco, David Schlessinger, Chris Ottolenghi Foxl2 functions in sex determination and histogenesis throughout mouse ovary development. BMC Dev. Biol.: 2009, 9();36 PMID:19538736 | PMC2711087 | BMC Dev Biol.
  9. Keibel, F. and Mall, F.P. Manual of Human Embryology II. J. B. Lippincott Company, Philadelphia (1912) Chapter XIX. The Development of the Urinogenital Organs - Development of the Reproductive Glands and their Duct
  10. V V Khadilkar, A V Khadilkar, A S Kinare, H S Tapasvi, S S Deshpande, G B Maskati Ovarian and uterine ultrasonography in healthy girls between birth to 18 years. Indian Pediatr: 2006, 43(7);625-30 PMID:16891683
  11. A R Baerwald, G P Adams, R A Pierson Form and function of the corpus luteum during the human menstrual cycle. Ultrasound Obstet Gynecol: 2005, 25(5);498-507 PMID:15846762
  12. H H Simmer The first experiments to demonstrate an endocrine function of the corpus luteum. On the occasion of the 100th birthday of Ludwig Fraenkel (1870-1951). Sudhoffs Arch: 1971, 55(4);392-417 PMID:4261581
  13. Aylin Yaba, Necdet Demir The mechanism of mTOR (Mammalian Target Of Rapamycin) in a mouse model of polycystic ovary syndrome (PCOS). J Ovarian Res: 2012, 5(1);38 PMID:23185989 | PMC3538528 | J Ovarian Res.
  14. Susan M Sirmans, Kristen A Pate Epidemiology, diagnosis, and management of polycystic ovary syndrome. Clin Epidemiol: 2013, 6();1-13 PMID:24379699
  15. Pratip Chakraborty, S K Goswami, Shweta Rajani, Sunita Sharma, Syed N Kabir, Baidyanath Chakravarty, Kuladip Jana Recurrent pregnancy loss in polycystic ovary syndrome: role of hyperhomocysteinemia and insulin resistance. PLoS ONE: 2013, 8(5);e64446 PMID:23700477
  16. Zi-Jiang Chen, Han Zhao, Lin He, Yuhua Shi, Yingying Qin, Yongyong Shi, Zhiqiang Li, Li You, Junli Zhao, Jiayin Liu, Xiaoyan Liang, Xiaoming Zhao, Junzhao Zhao, Yingpu Sun, Bo Zhang, Hong Jiang, Dongni Zhao, Yuehong Bian, Xuan Gao, Ling Geng, Yiran Li, Dongyi Zhu, Xiuqin Sun, Jin-E Xu, Cuifang Hao, Chun-E Ren, Yajie Zhang, Shiling Chen, Wei Zhang, Aijun Yang, Junhao Yan, Yuan Li, Jinlong Ma, Yueran Zhao Genome-wide association study identifies susceptibility loci for polycystic ovary syndrome on chromosome 2p16.3, 2p21 and 9q33.3. Nat. Genet.: 2011, 43(1);55-9 PMID:21151128
  17. Andrea Flesken-Nikitin, Chang-Il Hwang, Chieh-Yang Cheng, Tatyana V Michurina, Grigori Enikolopov, Alexander Yu Nikitin Ovarian surface epithelium at the junction area contains a cancer-prone stem cell niche. Nature: 2013, 495(7440);241-5 PMID:23467088


  • Complementary pathways in mammalian female sex determination. Nef S, Vassalli JD. J Biol. 2009 Sep 2;8(8):74. Review. PMID: 19735582
  • On the origin of the maternal age effect in trisomy 21 Down syndrome: the Oocyte Mosaicism Selection model. Hultén MA, Patel S, Jonasson J, Iwarsson E. Reproduction. 2010 Jan;139(1):1-9. Epub . Review. PMID: 19755486


  • The forkhead transcription factor FOXL2 is expressed in somatic cells of the human ovary prior to follicle formation. Duffin K, Bayne RA, Childs AJ, Collins C, Anderson RA. Mol Hum Reprod. 2009 Dec;15(12):771-7. Epub 2009 Aug 25. PMID: 19706741 | Mol Hum Reprod.

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Cite this page: Hill, M.A. (2014) Embryology Ovary Development. Retrieved April 24, 2014, from http://embryology.med.unsw.edu.au/embryology/index.php?title=Ovary_Development

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