Difference between revisions of "Endocrine - Gonad Development"

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{{Header}}
 
==Introduction==
 
==Introduction==
[[File:XXhpgaxis.gif|thumb|Female HPG axis]]
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[[File:XXhpgaxis.jpg|thumb|Female HPG axis]]
Note this section of notes refers to the development of the gonad as an endocrine organ. A detailed description of the gonad development is covered in [[Ovary Development]] and [[Testis Development]].
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The term gonad refers to both the female {{ovary}} and the male {{testis}}, both have their own detailed topic pages. This section of notes refers only to the development of the gonad as an endocrine organ.  
  
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Embryonically, initial endocrine development of the testis is required for development of both the internal genital tract and the external genitalia.
  
{{Template:Endocrine Links}} | [[2009_Lecture_16|Lecture - Genital Development]] | [http://embryology.med.unsw.edu.au/Notes/endocrine15.htm original page]
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Postnatally, the gonads are part of an integrated '''H'''ypothalamus - '''P'''ituitary - '''G'''onad (HPG) axis. Post-puberty, this axis is the key regulator of the female reproductive cycle, the {{menstrual cycle}}.
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{{Endocrine Links}}
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<br>
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{{Menstrual_Links}}
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<br>
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{{Genital Links}}
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<br>
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== Some Recent Findings ==
 +
[[File:Model male androsterone synthesis.jpg|alt=Model male second trimester androsterone synthesis|thumb|Model male second trimester androsterone synthesis{{#pmid:30763313|PMID30763313}}]]
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{|
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|-bgcolor="F5FAFF"
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|
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* '''Transcriptional activity of oestrogen receptors in the course of embryo development'''{{#pmid:30012715|PMID30012715}} "Oestrogens are well-known proliferation and differentiation factors that play an essential role in the correct development of sex-related organs and behaviour in mammals. With the use of the ERE-Luc reporter mouse model, we show herein that throughout mouse development, oestrogen receptors (ERs) are active starting from day 12 post conception. Most interestingly, we show that prenatal luciferase expression in each organ is proportionally different in relation to the germ layer of the origin. The luciferase content is highest in ectoderm-derived organs (such as brain and skin) and is lowest in endoderm-derived organs (such as liver, lung, thymus and intestine). Consistent with the testosterone surge occurring in male mice at the end of pregnancy, in the first 2 days after birth, we observed a significant increase in the luciferase content in several organs, including the liver, bone, gonads and hindbrain. The results of the present study show a widespread transcriptional activity of ERs in developing embryos, pointing to the potential contribution of these receptors in the development of non-reproductive as well as reproductive organs. Consequently, the findings reported here might be relevant in explaining the significant differences in male and female physiopathology reported by a growing number of studies and may underline the necessity for more systematic analyses aimed at the identification of the prenatal effects of drugs interfering with ER signalling, such as aromatase inhibitors or endocrine disrupter chemicals."
 +
 
 +
* '''Regulation of seminiferous tubule-associated stem Leydig cells in adult rat testes'''{{#pmid:26929346|PMID26929346}} "Testicular Leydig cells are the primary source of testosterone in males. Adult Leydig cells have been shown to arise from stem cells present in the neonatal testis. Once established, adult Leydig cells turn over only slowly during adult life, but when these cells are eliminated experimentally from the adult testis, new Leydig cells rapidly reappear. As in the neonatal testis, stem cells in the adult testis are presumed to be the source of the new Leydig cells. ... The proliferation of the stem Leydig cells was stimulated by paracrine factors including Desert hedgehog (DHH), basic fibroblast growth factor (FGF2), platelet-derived growth factor (PDGF), and activin. Suppression of proliferation occurred with transforming growth factor β (TGF-β). The differentiation of the stem cells was regulated positively by DHH, lithium- induced signaling, and activin, and negatively by TGF-β, PDGFBB, and FGF2. DHH functioned as a commitment factor, inducing the transition of stem cells to the progenitor stage and thus into the Leydig cell lineage."
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|}
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{| class="wikitable mw-collapsible mw-collapsed"
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! More recent papers &nbsp;
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|-
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| [[File:Mark_Hill.jpg|90px|left]] {{Most_Recent_Refs}}
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Search term: [http://www.ncbi.nlm.nih.gov/pubmed/?term=Endocrine+Gonad+Development ''Endocrine Gonad Development''] | [http://www.ncbi.nlm.nih.gov/pubmed/?term=Testosterone+Embryology ''Testosterone Embryology''] | [http://www.ncbi.nlm.nih.gov/pubmed/?term=Granulosa+Cell+Embryology ''Granulosa Cell Embryology''] | [http://www.ncbi.nlm.nih.gov/pubmed/?term=Leydig+Cell+Embryology ''Leydig Cell Embryology''] | [http://www.ncbi.nlm.nih.gov/pubmed/?term=Interstitial+Cell+Embryology ''Interstitial Cell Embryology'']
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|}
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{| class="wikitable mw-collapsible mw-collapsed"
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! Older papers &nbsp;
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|-
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| {{Older papers}}
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* '''Fetal Testosterone (FT) Influences Sexually Dimorphic Gray Matter in the Human Brain'''{{#pmid:22238103|PMID22238103}} "These results bridge a long-standing gap between human and nonhuman species by showing that fetal testosterone acts as an organizing mechanism for the development of regional sexual dimorphism in the human brain." [[#Male Hormone Levels|Male Hormone Levels]] | [[Neural System Development]]
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|}
  
 
HPG Axis - [http://www.ncbi.nlm.nih.gov/bookshelf/br.fcgi?book=endocrin&part=A972&rendertype=box&id=A1057 Endocrinology - Simplified diagram of the actions of gonadotrophins]
 
HPG Axis - [http://www.ncbi.nlm.nih.gov/bookshelf/br.fcgi?book=endocrin&part=A972&rendertype=box&id=A1057 Endocrinology - Simplified diagram of the actions of gonadotrophins]
  
 
===Gonad Development===
 
===Gonad Development===
 
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[[File:Infant_ovary.jpg|thumb|Infant Ovary]]
 
* mesoderm - mesothelium and underlying mesenchyme, primordial germ cells  
 
* mesoderm - mesothelium and underlying mesenchyme, primordial germ cells  
 
* Gonadal ridge - mesothelium thickening, medial mesonephros
 
* Gonadal ridge - mesothelium thickening, medial mesonephros
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[[File:Steroidogenesis.png|600px|Steroidogenesis]]
 
[[File:Steroidogenesis.png|600px|Steroidogenesis]]
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==Testosterone==
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Testicular Leydig cells (interstitial cells) are the main source of testosterone in males.
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[[File:Male_testosterone_and_AMH_level_graph.jpg|600px]]
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Human Male Testosterone and Anti-Müllerian Hormone (AMH) relative levels{{#pmid:16544032|PMID16544032}}
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===Leydig Cells===
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[[File:Leydig cells stained for LHCGR1.jpg|400px]]
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Leydig cells stained for LHCGR1{{#pmid:18433292|PMID18433292}}
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===Androgen and Digit ratio (2D:4D)===
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[[File:Finger_length_ratio_-_2D4D.jpg|thumb|Androgen and Digit ratio (2D:4D]]
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The ratio of 2nd and 4th finger (D, digit) length. This ratio has been suggested to relate to high fetal [[T#testosterone|testosterone]] concentration (males have lower 2D:4D than females) and has been shown for several species.{{#pmid:16504142|PMID16504142}} Although a study in mice has not shown the same correlation.{{#pmid:19495421|PMID19495421}} There have been some suggestions that the ratio may also be an indicator of various neurological abnormalities.
 +
 +
'''To measure (2D:4D)''' - using your right hand palm up, measure the index finger (2) and ring finger (4) length from palm to tip. Dividing the index finger by the ring finger gives the 2D:4D ratio, average women ratio is 1, average men is 0.98.
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==Anti-Mullerian Hormone==
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{{AMH}}
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==Estrogen==
 +
{|
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| The estradiol ({{estrogen}}, oestrogen) hormone is a steroid sex hormone expressed in both male and female.
 +
 +
estrogenic activity in human placental extracts was due to the presence of at least three compounds: estriol, estrone, and 17β-estradiol.
 +
 +
In the female, this hormone together with progesterone regulate changes that occur each menstrual cycle. During female development the fetal adrenal gland cortex synthesises DHEA (and DHEA/S), an oestrogen precursor (see image [[:File:Fetal adrenal gland steroidogenesis.jpg|Fetal adrenal gland steroidogenesis]]), converted by the {{placenta}} into estrogen compounds; estriol, estrone, and 17β-estradiol. During {{puberty}}, ovarian estrogen production is responsible for development of the secondary feminine sex characteristics.
 +
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In the male, [[:File:Testis histology 2.jpg|Leydig cells]] produce estrogen into the rete testis fluid at variable levels in different species. During male embryonic development exposure to high levels of estrogen can lead to genital abnormalities.
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| [[File:Estradiol.jpg|alt=estrogen|300px]]
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Estradiol
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|}
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==Progesterone==
 +
 +
{|
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| The progesterone (progestin) hormone is produced by the granulosa cells of the ovarian follicles at different levels during the {{menstrual cycle}} and at high levels by the luteal cells (P4) of the {{corpus luteum}}.
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In 1934 progesterone (progestin) C<sub>21</sub>H<sub>30</sub>O<sub>2</sub> was first isolated from the {{corpus luteum}} and its structure reported by four separate groups of researchers.<ref name=Butenandt1934>Butenandt A. Neure Ergebnisse auf dem Gebiet der Sexualhormone. Wien Klin Wochenschr. 1934;47:936.</ref><ref name=Slotta1934>Slotta KH, Ruschig H, Fels E. Ȕber der Hormon aus dem Corpus-luteum. Ber Chem Ges. 1934;67:1270.</ref><ref name=Hartmann1934>Hartmann M, Wettstein A. Ein krystallisiertes Hormon aus Corpus-luteum. Helv Chim Acta. 1934;17:878.</ref>{{#pmid:17817057|PMID17817057}}
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| [[File:Progesterone.jpg|alt=Progesterone molecular structure|300px]]
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Progesterone molecular structure
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|}
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{{Genital Links}}
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==Adult Histology==
  
 
== References ==
 
== References ==
Line 39: Line 129:
 
===Search PubMed===
 
===Search PubMed===
  
Search April 2010
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'''Search Pubmed:''' [http://www.ncbi.nlm.nih.gov/sites/entrez?db=pubmed&cmd=search&term=endocrine%20gonad%20development endocrine gonad development]
* Endocrine Development - All (14277) Review (4620) Free Full Text (3140)
 
  
'''Search Pubmed:''' [http://www.ncbi.nlm.nih.gov/sites/entrez?db=pubmed&cmd=search&term=pineal%20development pineal development]
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==External Links==
  
==Additional Images==
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{{External Links}}
  
===Adult Histology===
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* '''Andrology Australia''' - [http://www.andrologyaustralia.org/docs/clinical-summary-guide04_May2010.pdf Androgen deficiency PDF] | [http://www.andrologyaustralia.org/docs/clinical-summary-guide02_May2010.pdf Childhood and adolescence. Examination of male genitals and secondary sexual characteristics PDF]
<gallery>
 
File:Pineal_histology_001.jpg|Pineal (high power)
 
</gallery>
 
  
==Terms==
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==Additional Images==
  
{{Template:Glossary}}
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{{Glossary}}
  
{{Template:Footer}}
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{{Footer}}
  
 
[[Category:Endocrine]] [[Category:Gonad]]
 
[[Category:Endocrine]] [[Category:Gonad]]

Latest revision as of 13:22, 20 May 2019

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Introduction

Female HPG axis

The term gonad refers to both the female ovary and the male testis, both have their own detailed topic pages. This section of notes refers only to the development of the gonad as an endocrine organ.

Embryonically, initial endocrine development of the testis is required for development of both the internal genital tract and the external genitalia.

Postnatally, the gonads are part of an integrated Hypothalamus - Pituitary - Gonad (HPG) axis. Post-puberty, this axis is the key regulator of the female reproductive cycle, the menstrual cycle.


Endocrine Links: Introduction | BGD Lecture | Science Lecture | Lecture Movie | pineal | hypothalamus‎ | pituitary | thyroid | parathyroid | thymus | pancreas | adrenal | endocrine gonad‎ | endocrine placenta | other tissues | Stage 22 | endocrine abnormalities | Hormones | Category:Endocrine
Historic Embryology - Endocrine  
1903 Islets of Langerhans | 1904 interstitial Cells | 1908 Pancreas Different Species | 1908 Pituitary | 1908 Pituitary histology | 1911 Rathke's pouch | 1912 Suprarenal Bodies | 1914 Suprarenal Organs | 1915 Pharynx | 1916 Thyroid | 1918 Rabbit Hypophysis | 1920 Adrenal | 1935 Mammalian Hypophysis | 1926 Human Hypophysis | 1927 Hypophyseal fossa | 1932 Pineal Gland and Cysts | 1935 Hypophysis | 1937 Pineal | 1938 Parathyroid | 1940 Adrenal | 1941 Thyroid | 1950 Thyroid Parathyroid Thymus | 1957 Adrenal


Menstrual Cycle Links: Introduction | menstrual histology | ovary | corpus luteum | oocyte | uterus | Uterine Gland | estrous cycle | pregnancy test
Historic Embryology - Menstrual 
1839 Corpus Luteum Structure | 1851 Corpus Luteum | 1933 Pap Smear | 1937 Corpus Luteum Hormone | 1942 Human Reproduction Hormones | 1951 Corpus Luteum | 1969 Ultrastructure of Development and Regression | 1969 Ultrastructure during Pregnancy


Genital Links: genital | Lecture - Medicine | Lecture - Science | Lecture Movie | Medicine - Practical | primordial germ cell | meiosis | Female | X | ovary | corpus luteum |oocyte | uterus | vagina | reproductive cycles | menstrual cycle | Male | Y | SRY | testis | spermatozoa | ductus deferens | penis | prostate | endocrine gonad‎ | Genital Movies | genital abnormalities | Assisted Reproductive Technology | puberty | Category:Genital
Historic Embryology - Genital 
1901 Urinogenital Tract | 1902 The Uro-Genital System | 1904 Ovary and Testis | 1904 Leydig Cells | 1904 Hymen | 1905 Testis vascular | 1909 Prostate | 1912 Prostate | 1912 Urinogenital Organ Development | 1914 External Genitalia | 1914 Female | 1915 Cowper’s and Bartholin’s Glands | 1920 Wolffian tubules | 1921 Urogenital Development | 1921 External Genital | 1927 Female Foetus 15 cm | 1932 Postnatal Ovary | 1935 Prepuce | 1935 Wolffian Duct | 1942 Sex Cords | 1943 Testes Descent | 1953 Germ Cells | Historic Embryology Papers | Historic Disclaimer


Some Recent Findings

Model male second trimester androsterone synthesis
Model male second trimester androsterone synthesis[1]
  • Transcriptional activity of oestrogen receptors in the course of embryo development[2] "Oestrogens are well-known proliferation and differentiation factors that play an essential role in the correct development of sex-related organs and behaviour in mammals. With the use of the ERE-Luc reporter mouse model, we show herein that throughout mouse development, oestrogen receptors (ERs) are active starting from day 12 post conception. Most interestingly, we show that prenatal luciferase expression in each organ is proportionally different in relation to the germ layer of the origin. The luciferase content is highest in ectoderm-derived organs (such as brain and skin) and is lowest in endoderm-derived organs (such as liver, lung, thymus and intestine). Consistent with the testosterone surge occurring in male mice at the end of pregnancy, in the first 2 days after birth, we observed a significant increase in the luciferase content in several organs, including the liver, bone, gonads and hindbrain. The results of the present study show a widespread transcriptional activity of ERs in developing embryos, pointing to the potential contribution of these receptors in the development of non-reproductive as well as reproductive organs. Consequently, the findings reported here might be relevant in explaining the significant differences in male and female physiopathology reported by a growing number of studies and may underline the necessity for more systematic analyses aimed at the identification of the prenatal effects of drugs interfering with ER signalling, such as aromatase inhibitors or endocrine disrupter chemicals."
  • Regulation of seminiferous tubule-associated stem Leydig cells in adult rat testes[3] "Testicular Leydig cells are the primary source of testosterone in males. Adult Leydig cells have been shown to arise from stem cells present in the neonatal testis. Once established, adult Leydig cells turn over only slowly during adult life, but when these cells are eliminated experimentally from the adult testis, new Leydig cells rapidly reappear. As in the neonatal testis, stem cells in the adult testis are presumed to be the source of the new Leydig cells. ... The proliferation of the stem Leydig cells was stimulated by paracrine factors including Desert hedgehog (DHH), basic fibroblast growth factor (FGF2), platelet-derived growth factor (PDGF), and activin. Suppression of proliferation occurred with transforming growth factor β (TGF-β). The differentiation of the stem cells was regulated positively by DHH, lithium- induced signaling, and activin, and negatively by TGF-β, PDGFBB, and FGF2. DHH functioned as a commitment factor, inducing the transition of stem cells to the progenitor stage and thus into the Leydig cell lineage."
More recent papers  
Mark Hill.jpg
PubMed logo.gif

This table allows an automated computer search of the external PubMed database using the listed "Search term" text link.

  • This search now requires a manual link as the original PubMed extension has been disabled.
  • The displayed list of references do not reflect any editorial selection of material based on content or relevance.
  • References also appear on this list based upon the date of the actual page viewing.


References listed on the rest of the content page and the associated discussion page (listed under the publication year sub-headings) do include some editorial selection based upon both relevance and availability.

More? References | Discussion Page | Journal Searches | 2019 References

Search term: Endocrine Gonad Development | Testosterone Embryology | Granulosa Cell Embryology | Leydig Cell Embryology | Interstitial Cell Embryology

Older papers  
These papers originally appeared in the Some Recent Findings table, but as that list grew in length have now been shuffled down to this collapsible table.

See also the Discussion Page for other references listed by year and References on this current page.

  • Fetal Testosterone (FT) Influences Sexually Dimorphic Gray Matter in the Human Brain[4] "These results bridge a long-standing gap between human and nonhuman species by showing that fetal testosterone acts as an organizing mechanism for the development of regional sexual dimorphism in the human brain." Male Hormone Levels | Neural System Development

HPG Axis - Endocrinology - Simplified diagram of the actions of gonadotrophins

Gonad Development

Infant Ovary
  • mesoderm - mesothelium and underlying mesenchyme, primordial germ cells
  • Gonadal ridge - mesothelium thickening, medial mesonephros
  • Primordial Germ cells - yolk sac, to mesentery of hindgut, to genital ridge of developing kidney

Differentiation

  • testis-determining factor (TDF) from Y chromosome: presence (testes), absence (ovaries)

Testis

  • 8 Weeks, mesenchyme, interstitial cells (of Leydig) secrete testosterone, androstenedione
  • 8 to 12 Weeks - hCG stimulates testosterone production
  • Sustentacular cells - produce anti-mullerian hormone to puberty

Ovary

  • X chromosome genes regulate ovary development

Steroidogenesis

Steroidogenesis

Testosterone

Testicular Leydig cells (interstitial cells) are the main source of testosterone in males.

Male testosterone and AMH level graph.jpg

Human Male Testosterone and Anti-Müllerian Hormone (AMH) relative levels[5]

Leydig Cells

Leydig cells stained for LHCGR1.jpg

Leydig cells stained for LHCGR1[6]

Androgen and Digit ratio (2D:4D)

Androgen and Digit ratio (2D:4D

The ratio of 2nd and 4th finger (D, digit) length. This ratio has been suggested to relate to high fetal testosterone concentration (males have lower 2D:4D than females) and has been shown for several species.[7] Although a study in mice has not shown the same correlation.[8] There have been some suggestions that the ratio may also be an indicator of various neurological abnormalities.

To measure (2D:4D) - using your right hand palm up, measure the index finger (2) and ring finger (4) length from palm to tip. Dividing the index finger by the ring finger gives the 2D:4D ratio, average women ratio is 1, average men is 0.98.

Anti-Mullerian Hormone

AMH

Estrogen

The estradiol (estrogen, oestrogen) hormone is a steroid sex hormone expressed in both male and female.

estrogenic activity in human placental extracts was due to the presence of at least three compounds: estriol, estrone, and 17β-estradiol.

In the female, this hormone together with progesterone regulate changes that occur each menstrual cycle. During female development the fetal adrenal gland cortex synthesises DHEA (and DHEA/S), an oestrogen precursor (see image Fetal adrenal gland steroidogenesis), converted by the placenta into estrogen compounds; estriol, estrone, and 17β-estradiol. During puberty, ovarian estrogen production is responsible for development of the secondary feminine sex characteristics.

In the male, Leydig cells produce estrogen into the rete testis fluid at variable levels in different species. During male embryonic development exposure to high levels of estrogen can lead to genital abnormalities.

estrogen

Estradiol

Progesterone

The progesterone (progestin) hormone is produced by the granulosa cells of the ovarian follicles at different levels during the menstrual cycle and at high levels by the luteal cells (P4) of the corpus luteum.


In 1934 progesterone (progestin) C21H30O2 was first isolated from the corpus luteum and its structure reported by four separate groups of researchers.[9][10][11][12]

Progesterone molecular structure

Progesterone molecular structure


Genital Links: genital | Lecture - Medicine | Lecture - Science | Lecture Movie | Medicine - Practical | primordial germ cell | meiosis | Female | X | ovary | corpus luteum |oocyte | uterus | vagina | reproductive cycles | menstrual cycle | Male | Y | SRY | testis | spermatozoa | ductus deferens | penis | prostate | endocrine gonad‎ | Genital Movies | genital abnormalities | Assisted Reproductive Technology | puberty | Category:Genital
Historic Embryology - Genital 
1901 Urinogenital Tract | 1902 The Uro-Genital System | 1904 Ovary and Testis | 1904 Leydig Cells | 1904 Hymen | 1905 Testis vascular | 1909 Prostate | 1912 Prostate | 1912 Urinogenital Organ Development | 1914 External Genitalia | 1914 Female | 1915 Cowper’s and Bartholin’s Glands | 1920 Wolffian tubules | 1921 Urogenital Development | 1921 External Genital | 1927 Female Foetus 15 cm | 1932 Postnatal Ovary | 1935 Prepuce | 1935 Wolffian Duct | 1942 Sex Cords | 1943 Testes Descent | 1953 Germ Cells | Historic Embryology Papers | Historic Disclaimer

Adult Histology

References

  1. O'Shaughnessy PJ, Antignac JP, Le Bizec B, Morvan ML, Svechnikov K, Söder O, Savchuk I, Monteiro A, Soffientini U, Johnston ZC, Bellingham M, Hough D, Walker N, Filis P & Fowler PA. (2019). Alternative (backdoor) androgen production and masculinization in the human fetus. PLoS Biol. , 17, e3000002. PMID: 30763313 DOI.
  2. Della Torre S, Rando G, Meda C, Ciana P, Ottobrini L & Maggi A. (2018). Transcriptional activity of oestrogen receptors in the course of embryo development. J. Endocrinol. , 238, 165-176. PMID: 30012715 DOI.
  3. Li X, Wang Z, Jiang Z, Guo J, Zhang Y, Li C, Chung J, Folmer J, Liu J, Lian Q, Ge R, Zirkin BR & Chen H. (2016). Regulation of seminiferous tubule-associated stem Leydig cells in adult rat testes. Proc. Natl. Acad. Sci. U.S.A. , 113, 2666-71. PMID: 26929346 DOI.
  4. Lombardo MV, Ashwin E, Auyeung B, Chakrabarti B, Taylor K, Hackett G, Bullmore ET & Baron-Cohen S. (2012). Fetal testosterone influences sexually dimorphic gray matter in the human brain. J. Neurosci. , 32, 674-80. PMID: 22238103 DOI.
  5. Rey R. (2005). Anti-Müllerian hormone in disorders of sex determination and differentiation. Arq Bras Endocrinol Metabol , 49, 26-36. PMID: 16544032 DOI.
  6. Kossack N, Simoni M, Richter-Unruh A, Themmen AP & Gromoll J. (2008). Mutations in a novel, cryptic exon of the luteinizing hormone/chorionic gonadotropin receptor gene cause male pseudohermaphroditism. PLoS Med. , 5, e88. PMID: 18433292 DOI.
  7. McIntyre MH. (2006). The use of digit ratios as markers for perinatal androgen action. Reprod. Biol. Endocrinol. , 4, 10. PMID: 16504142 DOI.
  8. Yan RH, Bunning M, Wahlsten D & Hurd PL. (2009). Digit ratio (2Dratio4D) differences between 20 strains of inbred mice. PLoS ONE , 4, e5801. PMID: 19495421 DOI.
  9. Butenandt A. Neure Ergebnisse auf dem Gebiet der Sexualhormone. Wien Klin Wochenschr. 1934;47:936.
  10. Slotta KH, Ruschig H, Fels E. Ȕber der Hormon aus dem Corpus-luteum. Ber Chem Ges. 1934;67:1270.
  11. Hartmann M, Wettstein A. Ein krystallisiertes Hormon aus Corpus-luteum. Helv Chim Acta. 1934;17:878.
  12. Allen WM & Wintersteiner O. (1934). CRYSTALLINE PROGESTIN. Science , 80, 190-1. PMID: 17817057 DOI.


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Cite this page: Hill, M.A. (2019, July 18) Embryology Endocrine - Gonad Development. Retrieved from https://embryology.med.unsw.edu.au/embryology/index.php/Endocrine_-_Gonad_Development

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