Brief Description of the Histological, Cytological and Functional Aspects of the Ovary

  • María Luisa EscobarEmail author
  • Gerardo H. Vázquez-Nin
  • Olga M. Echeverría


The ovary is formed by three main compartments: superficial epithelium, cortex and medulla. The superficial epithelium is constituted by one layer of cubic cells. The cortex is a wide peripheral zone containing the follicles, the functional and structural unit of the ovary, and a stroma formed by compact connective tissue. Every follicle is formed by one oocyte surrounded by follicular cells, also called granulosa cells, and a basal lamina surrounding them. The medulla is the central region of the ovary formed by connective tissue with numerous blood vessels. As the follicles develop they change their size, morphology and physiology. Primordial follicles are formed by the oocyte surrounded by flat follicular cells. Primary ­follicles are characterized by the initiation of follicular growth. Secondary follicles are characterized by two or more layers of granulosa cells and no antrum. The early antral follicles are characterized by the formation and progressive growth of a cavity, due to the accumulation of a fluid. Once the antrum is formed the follicle goes through several stages: (a) basal growth, (b) selection and (c) dominance. The process of follicular growth is controlled by extra-ovarian and intra-ovarian factors and the importance of each of these factors depends on the stage of follicle development. Extra-ovarian factors regulate growth of antral and preovulatory follicles, while intra-ovarian factors regulate growth of preantral and early antral follicles. The ovary is not only involved in sexual reproduction, but also has great influence on the entire hormonal functioning during development of the organism. The ovary is the site of the highest synthesis and secretion of progesterone and estrogen in mammals and gives rise to cyclical fluctuations in the levels of these hormones in the blood. Before ovulation, granulosa cells mature to form the corpus luteum, which is responsible for the secretion of progesterone and estrogen.


Luteinizing Hormone Granulosa Cell Follicle Stimulate Hormone Corpus Luteum Connective Tissue Growth Factor 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.

List of Abbreviations


Luteinizing hormone


Follicle stimulant hormone


Transforming growth factor beta


Epidermal growth factor


  1. Adhikari D, Liu K (2009) Molecular mechanisms underlying the activation of mammalian primordial follicles. Endocr Rev 30:438–464PubMedCrossRefGoogle Scholar
  2. Albertini DF, Combelles CM, Benecchi E, Carabatsos MJ (2001) Cellular basis for paracrine regulation of ovarian follicle development. Reproduction 121:647–653PubMedCrossRefGoogle Scholar
  3. Armstrong DT, Goff AK, Dorrington JH (1979) Regulation of follicular estrogen biosynthesis. In: Midgley AR Jr, Sadler WA Jr (eds) Ovarian Follicular Development and Function. Raven, New YorkGoogle Scholar
  4. Attisano L, Wrana JL (1998) Mads and Smads in TGF beta signalling. Curr Opin Cell Biol 10:188–194PubMedCrossRefGoogle Scholar
  5. Bassett DL (1943) The changes in the vascular pattern of the ovary of the albino rat during the estrous cycle. Am J Anat 73:252–292CrossRefGoogle Scholar
  6. Bergh C, Olsson JH, Selleskog U, Hillens ST (1993) Steroid production in cultured thecal cell obtained from human ovarian follicles. Hum Reprod 8:519–524PubMedGoogle Scholar
  7. Bomsel H, Gougeon A, Thebault A et al (1979) Healthy and atretic human follicles in the preovulatory phase: differences in evolution of follicular morphology and steroid content of the follicular fluid. J Clin Endocrinol Metab 48:686–694Google Scholar
  8. Brand T, Schneider MD (1996) Transforming growth factor-b signal transduction. Circ Res 78:173–179PubMedGoogle Scholar
  9. Campbell BK, Scaramuzzi RJ, Webb R (1995) Control of follicle development and selection in sheep and cattle. J Reprod Fertil Suppl 49:335–350PubMedGoogle Scholar
  10. Cate RL, Mattaliano RJ, Hession C et al (1986) Isolation of the bovine and human genes for Müllerian inhibiting substance and expression of the human gene in animal cells. Cell 45:685–698PubMedCrossRefGoogle Scholar
  11. Chang H, Brown CW, Matzuk MM (2002) Genetic analysis of the mammalian transforming growth factor-β superfamily. Endocr Rev 23:787–823PubMedCrossRefGoogle Scholar
  12. Conneely OM, Mulac-Jericevic B, Lydon JP (2003) Progesterone-dependent regulation of female reproductive activity by two distinct progesterone receptor isoforms. Steroids 68:771–778PubMedCrossRefGoogle Scholar
  13. Dennefors BL, Hamberger L, Nilsson L (1983) Influence of human chorionic gonadotropin in vivo on steroid formation and gonadotropin responsiveness of isolated human preovulatory follicular cells. Fertil Sterll 39:56–61Google Scholar
  14. Driancourt MA, Reynaud K, Cortvrindt R, Smitz J (2000) Roles of Kit and Kit Ligand in ovarian function. Rev Reprod 5:143–152PubMedCrossRefGoogle Scholar
  15. Drummond AE, Dyson M, Le Tan M et al (2003) Ovarian follicle populations of the rat express TGF-ß signaling pathways. Mol Cell Endocrinol 202:53–57PubMedGoogle Scholar
  16. Dube JL, Wang P, Elvin J et al (1998) The bone morphogenic protein 15 gene is X-linked and expressed in oocytes. Mol Endocrinol 12:1809–1917PubMedCrossRefGoogle Scholar
  17. Durlinger A, Kramer P, Karels B et al (1999) Control of primordial follicle recruitment by anti-Müllerian hormone in the mouse ovary. Endocrinol 140:5789–5796Google Scholar
  18. Fair T, Hulshof SCJ, Hyttel P et al (1997a) Oocyte ultrastructure in bovine primordial to early tertiary follicles. Anat Embryol 195:327–336PubMedCrossRefGoogle Scholar
  19. Fair T, Hulshof SCJ, Hyttel P et al (1997b) Nucleus ultrastructure and transcriptional activity of bovine oocytes in preantral and early antral follicles. Mol Reprod Dev 46:208–215PubMedCrossRefGoogle Scholar
  20. Findlay JK (1993) An updata on the roles of inhibin, activin, and follistatin as local regulators of folliculogenesis. Boi Reprod 48:15–23CrossRefGoogle Scholar
  21. Franchi LL (1960) Electron microscopy of oocyte-follicle relationships in the rat ovary. J Biophys Biochem Cytol 7:397–398PubMedCrossRefGoogle Scholar
  22. Garrido C, Saule S, Gospodarowicz D (1993) Transcriptional regulation of vascular endothelial growth factor gene expression in bovine granulosa cells. Growth Factors 8:109–117PubMedCrossRefGoogle Scholar
  23. Goddard I, Hendrick JC, Bnahmed M, Morera AM (1995) Transforming growth factor b receptor expression in cultured porcine granulosa cells. Mol Cell Endocrinol 115:207–213PubMedCrossRefGoogle Scholar
  24. Gougeon A (1984) Influence of cyclic variations in gonadotrophin and steroid hormones on ­follicular growth in the human ovary. In: de Brux I. Gautrav JP (eds) Clinical Pathology of the Endocrine Ovary. MTP Press, LancasterGoogle Scholar
  25. Graham JD, Clarke CL (1997) Physiological action of progesterone in target tissues. Endocr Rev 18(4):502–19PubMedCrossRefGoogle Scholar
  26. Harlow ChR, Davidson L, Burns KH et al (2002) FSH and TGF-b supefamily members regulate granulosa cell connective tissue growth factor gene expression in vitro and in vivo. Endocrinology 143(9):3316–3325PubMedCrossRefGoogle Scholar
  27. Hayashi M, McGee EA, Min G et al (1999) Recombinant growth differentiation factor-9 (GDF9) enhances growth and differentiation of cultured early ovarian follicles. Endocrinology 140:1236–1244PubMedCrossRefGoogle Scholar
  28. Hirshfield AN (1991) Development of follicles in the mammalian ovary. Int Rev Cytol 124:43–101PubMedCrossRefGoogle Scholar
  29. Huang EJ, Manova K, Packer AI et al (1993) The murine Steel Panda mutation affects kit ligand expression and growth of early ovarian follicles. Develop Biol 157:100–109PubMedCrossRefGoogle Scholar
  30. Ingraham HA, Hirokawa Y, Roberts LM et al (2000) Autocrine and paracrine hormone signaling in reproduction. Recent Prog Horm Res 55:30–38Google Scholar
  31. Ireland JJ, Martin TL, Ireland JLH, Aulerich RJ (1992) Immunoneutralization of inhibin suppress reproduction in female mink. Biol Reprod 47:746–750PubMedCrossRefGoogle Scholar
  32. Jaatinen TA, Penttila TL, Kaipia A et al (1994) Expression of inhibin a, bA y bB Messenger ribonucleic acids in the normal ovary and in polycystic ovarian syndrome. J Endocrin 143:127–137CrossRefGoogle Scholar
  33. Jordan AW III, Caffrey JL, Niswender GD (1978) Catecholamine induced stimulation of progesterone and adenosine 39, 59-monophosphate production by dispersed ovine luteal cells. Endocrinology 103:385–392PubMedCrossRefGoogle Scholar
  34. Katz D, Niederberger C, Slaughter GR, Cooney AJ (1997) Characterization of germ cell-specific expression of the orphan nuclear receptor, germ cell nuclear factor. Endocrinology 138:4364–4372PubMedCrossRefGoogle Scholar
  35. Keri RA, Nilson JH (1996) A steroidogenic factor-1 binding site is required for activity of the luteinizing hormone b subunit promoter in gonadotropes of transgenic mice. J Biol Chem 271:10782–10785PubMedCrossRefGoogle Scholar
  36. Klinger FG, De Felici M (2002) In vitro development of growing oocytes from fetal mouse oocytes: stage-specific regulation by stem cell factor and granulosa cells. Develop Biol 244:85–95PubMedCrossRefGoogle Scholar
  37. Knight PG, Glister C (2003) Local roles of TGFbeta superfamily members in the control of ovarian follicle development. Anim Reprod Sci 78:15–183CrossRefGoogle Scholar
  38. Levallet J, Koskimies P, Rahman N, Huhtaniemi I (2001) The promoter of murine follicle-stimulating hormone receptor: functional characterization and regulation by transcription factor steroidogenic factor 1. Mol Endocrinol 15:80–92PubMedCrossRefGoogle Scholar
  39. Lydon JP, DeMayo FJ, Funk CR et al (1995) Mice lacking progesterone receptor exhibit pleiotropic reproductive abnormalities. Genes Dev 9:2266–2278PubMedCrossRefGoogle Scholar
  40. Magoffin DA (2005) Ovarian theca cell. Int J Biochem Cel Biol 37:1344–1349CrossRefGoogle Scholar
  41. Manova K, Nocka K, Besmer P, Bachvarova RF (1990) Gonadal expression of c-kit encoded at the W locus of the mouse. Development 110:1057–1069PubMedGoogle Scholar
  42. Massague J (1992) Receptors for the TGF-b family. Cell 69:1067–1070PubMedCrossRefGoogle Scholar
  43. Massague J, Chen YG (2000) Controling TGF-b signaling. Genes Dev 14:627–644PubMedGoogle Scholar
  44. Massague J, Attisano L, Wrana JL (1994) The TGF-b family and its composite receptors. Trends Cell Biol 4:172–178PubMedCrossRefGoogle Scholar
  45. McGee EA, Hsueh AJW (2000) Initial and cyclic recruitment of ovarian follicles. Endocr Rev 21(2):200–214PubMedCrossRefGoogle Scholar
  46. McGrath SA, Esquela AF, Lee SJ (1995) Oocyte specific expression of growth/differentiation factor-9. Mol Endocrinol 9:131–136PubMedCrossRefGoogle Scholar
  47. Miro F, Smyth CD, Whitelaw PF et al (1995) Regulation of 3b-hydroxysteroid dehydrogenase5/4-isomerase and cholesterol side-chain cleavage cytochrome P450 by activin in rat granulosa cells. Endocrinology 136:3247–3252PubMedCrossRefGoogle Scholar
  48. Montro B, Bernstein A (1993) Dynamic changes in ovarian c-kit and Steel expression during the estrous reproductive cycle. Dev Dyn 197:69–79CrossRefGoogle Scholar
  49. Moore GPM, Lintern-Moore S, Peters H, Faber M (1974) RNA synthesis in the mouse oocyte. J Cell Biol 60:416–422PubMedCrossRefGoogle Scholar
  50. Morita Y, Tilly JL (1999) Oocyte apoptosis: like sand through an hourglass. Dev Biol 213:1–17PubMedCrossRefGoogle Scholar
  51. Motta PM, Makabe S, Naguro T, Correr S (1994) Oocyte follicle cells association during development of human ovarian follicle. A study by high resolution scanning and transmission electron microscopy. Arch Histol Cytol 57:369–394PubMedCrossRefGoogle Scholar
  52. Natraj U, Richards JS (1993) Hormonal regulation, localization, and functional activity of the progesterone receptor in granulosa cells of rat preovulatory follicles. Endocrinology 133:761–769PubMedCrossRefGoogle Scholar
  53. Nilsson EE, Kezele P, Skinner MK (2002) Leukemia inhibitory factor (LIF) promotes the primordial to primary follicle transition in rat ovaries. Mol Cell Endocrinol 188:65–73PubMedCrossRefGoogle Scholar
  54. Norman A, Litwack G (1987) Hormones. Academy Press, Inc., LondonGoogle Scholar
  55. Otsuka F, Yao Z, Lee T et al (2000) Bone morphogenetic protein-15. Identification of target cells and biological functions. J Biol Chem 275:39523–39528PubMedCrossRefGoogle Scholar
  56. Park JY, Su YQ, Ariga M et al (2004) EGF-like growth factors as mediators of LH action in the ovulatory follicle. Science 303(5658):682–684PubMedCrossRefGoogle Scholar
  57. Picton H, Briggs D, Gosden R (1998) The molecular basis of oocyte growth and development. Mol Cell Endocrinol 145:27–37PubMedCrossRefGoogle Scholar
  58. Rajkovic A, Pangas SA, Ballow D et al (2004) Nobox deficiency disrupts early folliculogenesis and oocyte-specific gene expression. Science 305:1157–1159PubMedCrossRefGoogle Scholar
  59. Rankin T, Familari M, Lee E et al (1996) Mice homozygous for an insertional mutation in the Zp3 gene lack a zona pellucida and are infertile. Development 122:2903–2910PubMedGoogle Scholar
  60. Richards JS (1980a) Maturation of ovarian follicles: actions and interactions of pituitary and ­ovarian hormones on follicular cell differentiation. Physiol Rev 60(1):51–89PubMedGoogle Scholar
  61. Richards JS (2001) Perspective: the ovarian follicle: a perspective in 2001. Endocrinology 142(6):2184–2193PubMedCrossRefGoogle Scholar
  62. Robertson DM, Klein R, deVos FL et al (1987) The isolation of polypeptides with FSH suppressing activity from bovine follicular fluid which are structurally different to inhibin. Biochem Biophys Res Commun 149:744–749PubMedCrossRefGoogle Scholar
  63. Rugglu M, Speed R, Taggart M et al (1997) The mouse Dazla gene encodes a cytoplasmic protein essential for gametogenesis. Nature 389:73–77CrossRefGoogle Scholar
  64. Shimasaki S, Moore RK, Otsuka F, Erickson GF (2004) The bone morphogenetic protein system in mammalian reproduction. Endocr Rev 25:72–101PubMedCrossRefGoogle Scholar
  65. Shimonaka M, Inouye S, Shimasaki S, Ling N (1991) Follistatin binds to both activin and inhibin through the common subunit. Endocrinology 128(6):3313–15PubMedCrossRefGoogle Scholar
  66. Sotelo JR, Porter K (1959) An electron microscope study of rat ovum. J Biophys Biochem Cytol 5:327–342PubMedCrossRefGoogle Scholar
  67. Soyal SM, Amleh A, Dean J (2000) Fig α a germ cell-specific transcription factor required for ovarian follicle development. Development 127:4645–4655PubMedGoogle Scholar
  68. Symonds D, Tomic D, Borgeest C et al (2003) Smad3 regulates proliferation of the mouse ovarian surface epithelium. Anat Rec 273A:681–686CrossRefGoogle Scholar
  69. Teixeira J, He WW, Shah PC et al (1996) Developmental expression of a candidate Müllerian inhibiting substance type II receptor. Endocrinology 137:160–165PubMedCrossRefGoogle Scholar
  70. Tilly JL (2001) Commuting the death sentence: how oocytes strive to survive. Nat Rev Mol Cell Biol 2:838–848PubMedCrossRefGoogle Scholar
  71. Tisdall DJ, Fidler AE, Smith P et al (1999) Stem cell factor and c-kit gene expression and protein localization in the sheep ovary during fetal development. J Reprod Fertil 116:277–291PubMedCrossRefGoogle Scholar
  72. Tomic D, Brodie SG, Deng C et al (2002a) Smad3 may regulate follicular growth in the mouse ovary. Biol Reprod 66:917–923PubMedCrossRefGoogle Scholar
  73. Tomic D, Miller KP, Kenny HA et al (2004a) Ovarian follicle development requires Smad3. Mol Endocrinol 18(9):2224–40PubMedCrossRefGoogle Scholar
  74. Tremblay JJ, Viger RS (2001) GATA factors differentially activate multiple gonadal promoters through conserved GATA regulatory elements. Endocrinology 142:977–986PubMedCrossRefGoogle Scholar
  75. Ueno S, Kuroda T, Maclaughlin DT et al (1989) Müllerian inhibiting substance in the adult rat ovary during various stages of the estrous cycle. Endocrinology 125:1060–1066PubMedCrossRefGoogle Scholar
  76. van den Hurk R, Bevers MM, Dieleman SJ (1999) Folliculogenesis and oocyte development in mammals (livestock animals). In: Joy KP, Krishna A, Haldar C (eds) Comparative endocrinology and reproduction. Narosa Publishing House, New DelhiGoogle Scholar
  77. van den Hurk R, Abir R, Telfer EE, Bevers MM (2000) Primate and bovine immature oocytes and follicles as sources of fertilizable oocytes. Hum Reprod 6:457–474CrossRefGoogle Scholar
  78. Vitt UA, Hayashi M, Klein C, Hsueh AJ (2000) Growth differentiation factor-9 stimulates proliferation but suppresses the follicle stimulating hormone-induced differentiation of cultured granulosa cells from small antral and preovulatory rat follicles. Biol Reprod 62:370–377PubMedCrossRefGoogle Scholar
  79. Wandji SA, Gadsby JE, Barber JA, Hammond JM (2000) Messenger ribonucleic acids for MAC25 and connective tissue growth factor (CTGF) are inversely regulated during folliculogenesis and early luteogenesis. Endocrinology 141:2648–2657PubMedCrossRefGoogle Scholar
  80. Wang QF, Tilly KL, Tilly JL et al (1996) Activin inhibits basal an androgen-stimulated ­proliferation and induces apoptosis in the human prostatic cancer cell line, LNCaP. Endocrinology 137:5476–5483PubMedCrossRefGoogle Scholar
  81. Webb R, Campbell BK, Garverick HA et al (1999) Molecular mechanisms regulating follicular recruitment and selection. J Reprod Fertil Suppl 54:33–48PubMedGoogle Scholar
  82. Xu J, Oakley J, McGee EA (2002) Stage-specific expression of Smad2 and Smad3 during folliculogenesis. Biol Reprod 66:1571–1578PubMedCrossRefGoogle Scholar
  83. Zimmerman CM, Padgett RW (2000) Transforming growth factor-β signalling mediators and modulators. Gene 249:17–30PubMedCrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media B.V. 2011

Authors and Affiliations

  • María Luisa Escobar
    • 1
    Email author
  • Gerardo H. Vázquez-Nin
    • 1
  • Olga M. Echeverría
    • 1
  1. 1.Laboratory of Electron Microscopy, Department of Cell Biology, Faculty of SciencesNational University of Mexico (UNAM)Mexico CityMexico

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