臺灣博碩士論文加值系統

真骨魚 (telesot fish)的腦部有著其獨特的神經內分泌調控機制；其神經生成的過程與扮演著神經前驅細胞的放射性神經膠細胞連結著，而芳香酶的濃度也比哺乳動物高出一百到一千倍。目前已確定其腦部可以產生類固醇，並且在發育的過程中扮演著關鍵的角色。在真骨魚性腺分化過程中，腦部發育的早期受到許多神經內分泌因子及受體的調控，而這些調控的機制還有許多未被了解的地方。在這篇論文中，我們以點帶石斑魚 (Epinephelus coioides)為動物模型來研究上述的機制。點帶石斑魚是一種雌性先熟雌雄同体的魚類；在生命週期的前六年為雌性生殖而在約第七年的時候開始轉換為雄性生殖
我們假設初期點帶石斑魚的腦部可以產生一定量的神經性雌二醇，因而調控下游訊息傳遞以及神經性類固醇的表現，進而影響性腺分化。我們首先選殖及分析star (steroidogenic acute regulatory protein，類固醇生成急性調控蛋白) 、cyp11a1 (cytochrome P450 side chain cleavage，細胞色素P450支鏈裂解酵素)、hsd3b1 (3β-hydroxysteroid dehydrogenase，3β-羥類固醇去氫酶)、cyp17a1 (cytochrome P450c17，細胞色素P450c17)、arcyp19a1b (cytochrome P450 aromatase，细胞色素P450芳香化酶)這些在顳腦中與神經性雌激素生合成有關的主要基因的表現，同時也利用即時定量聚合酶連鎖反應觀察不同發育年齡(孵化後九十至一百八十天)時前腦、中腦、下視丘及後腦中雌激素受體 (α、β1、β2)、雌激素細胞膜受體 (GPR30)以及雄激素受體。結果指出除了上述除了雄激素受體以外的基因在性腺分化時均有顯著表現上升的情形. 而細胞增生活性指標pcna (proliferating cell nuclear antigen， 細胞增生核抗原)的表現在孵化後一百一十至一百五十天的也觀察到表現量上升。
另外，我們也發現相較於其他的時間點，在孵化後一百一十及一百二十天時前腦內的有著較高濃度的芳香酶以及雌二醇，更進一步的，我們觀察到在前腦及下視丘中、雌二醇可提高cyp17a1及cyp19a1b的表現，而在中腦中則是可提高hsd3b1的表現，至於cyp11a1則是在所有觀察的腦區中均無變化。而在中腦腦室、前視區及間腦腦室內利用外源表現的雌二醇也同樣的可以提高Cyp19a1b的表現，證明了雌二醇確實可調控類固醇激素基因。原位雜交的分析結果則顯示不管是在前腦、下視丘或是後腦，cyp11a1、hsd3b1及cyp17a1 均擁有相似的表現量，而在小腦中，hsd3b1則有著較高的表現，在放射性神經膠細胞中則發現cyp19a1b的基因與蛋白質量表現均有提高。綜合以上的結果，我們認為cyp19a1b只存在放射性神經膠細胞中。Cyp19a1b與PCNA的免疫螢光組織染色結果則顯示絕大部分增生的細胞均為放射性神經膠細胞。最後，結合了Cyp19a1b免疫螢光染色以及cyp11a1、hsd3b1和cyp17a1螢光原位雜合的結果則顯示cyp11a1, hsd3b1 and cyp17a1的mRNA存在於有著Cyp19a1b活性的放射性神經膠細胞中，代表cyp11a1, hsd3b1 and cyp17a1 mRNA來自於放射性神經膠細胞。
我們同時也分析了早期腦部中sf-1以及 dax-1這兩種孤兒核受體(orphan nuclear receptors)的表現與孵化後不同天數至性腺分化時促類固醇生成酶(star、cyp11a1、hsd3b1、cyp17a1與cyp19a1b)以及雌激素受體(ERα、ERβ1、與ERβ2)基因表現程度的關聯性。定量聚合酶連鎖反應顯示sf-1 mRNA與上述性腺分化促類固醇生成酶有顯著的正相關，而dax-1則剛好相反，代表這些孤兒核受體確實在腦中確實有其功能性。而在不同的發育時期時，sf-1以及 dax-1會在下視丘中表現，另外，在下視丘腹內側細胞中利用原位雜交發現star、 sf-1 與dax-1的大量表現，而在活體中作用雌二醇之後，雖然並未發現dax-1 的表現有所變化，但star 與 sf-1 的表現量確實有明顯的上升。最後，實驗數據也表示在早期腦部發育時，精催產素(arginine vasotocin) 的產生神經內分泌功能可能是透過神經類固醇與雌激素的訊息傳遞來執行。綜合以上，我們的結果證明了點帶石斑魚腦中擁有著所有類固醇生成基因、類固醇生成因子、固醇類性荷爾蒙受體、其他固醇以及芳香酶的活性。更進一步的，我們的結果證明了上述分子的基因從孵化後腦部發育早期就開始活化並轉錄，並在整個需要依賴性荷爾蒙的腦部發育過程中有著重要的功能,同時，研究結果也證明了點帶石斑魚的腦中有著產生所需要雌二醇的機制，因此擁有著在性腺分化時調控腦部發育的能力，而也正因為如此，點帶石斑魚的腦部發育的高峰期恰巧位於性腺發育的時候。

Since the brain of telesot fish exhibits a significant degree of neuroendocrine regulation, notably an intense neurogenic process linked to the persistence of radial glial cells acting as neural progenitors and remarkably high activity of brain aromatase, which is 100-1000 times higher than mammals, it has become clear that the brain is a true steroidogenic organ and important key player during development. In teleost, the neuroendocrine regulation of early brain development during gonadal sex differentiation is a complex phenomena regulated by various neuroendocrine factors and receptors during a critical period of development, which remains largely unknown. In the present study, we selected orange-spotted grouper Epinephelus coioides as a primary animal model to investigate the molecular mechanism underlying early brain development during gonadal sex differentiation. Groupers are protogynous hermaphrodite and functional females for the first six years of life and start to sex change around the age of seven years.
In the present study, we hypothesized that the grouper brain has the capacity to produce a significant amount of neural estrogen (E2) in the early brain, and this locally produced E2 could auto-regulate the functional peak of neurosteroidogenesis and estrogen signaling in the brain, which may have an association with the occurrence of gonadal sex differentiation in this species. To test this hypothesis, we first molecular cloned and analyzed the temporal expression patterns of key genes involved in neural estrogen biosynthesis pathway includes star (steroidogenic acute regulatory protein), cyp11a1 (cytochrome P450 side chain cleavage), hsd3b1 (3β-hydroxysteroid dehydrogenase), cyp17a1 (cytochrome P450c17) and cyp19a1b (cytochrome P450 aromatase), and sex steroid nuclear receptors for estrogen (ERα, ERβ1 and ERβ2), androgen (AR) and the plasma membrane-associated estrogen receptor (GPR30) were also studied at different developmental ages (from 90 to 180-dah: days after hatching) in different brain regions (forebrain, midbrain, hypothalamus and hindbrain) using quantitative real-time PCR (q-PCR) and RT-PCR analyses. The results indicated that the mRNA expressions of all key genes and estrogen receptors except for AR were significantly increased in the brain during the period of gonadal sex differentiation. Moreover, pcna transcripts (a marker for cell proliferation activity) were higher in the early brain at 110–150 dah.
Brain aromatase activity and E2 concentration levels, but not testosterone (T), were higher in the forebrain at 110-dah and 120-dah, respectively compared to other ages. Further, exogenous E2 upregulated cyp17a1 and cyp19a1b transcripts in the forebrain and hypothalamus, however, E2 did not alter cyp11a1 and hsd3b1 transcripts in any brain regions except hsd3b1 expression in the midbrain studied. Exogenous E2 also upregulated Cyp19a1b immunoreactivity in the telencephalic ventricle, preoptic area and diencephalic ventricle compared to their control brain, suggesting E2 regulation on steroidogenic genes. Furthermore, in situ hybridization (ISH) analysis revealed that cyp11a1, hsd3b1 and cyp17a1 transcripts were localized from the forebrain, hypothalamus and hindbrain with an overall similar expression pattern; at the same time, hsd3b1 transcript was higher in the cerebellum compared to other genes. High density in the cyp19a1b/Cyp19a1b expression was detected in radial glial cells. This is possibly concluded that the expression of grouper cyp19a1b/Cyp19a1b is restricted to radial glial cells. Besides, dual-labeled fluorescence immunohistochemistry (FIHC) of Cyp19a1b and PCNA further show that most of the proliferative cells were corresponded to radial glial cells. Moreover, by combining Cyp19a1b FIHC with florescence ISH (FISH) of cyp11a1, hsd3b1 and cyp17a1, we showed that sub-cellular localization of cyp11a1, hsd3b1 and cyp17a1 transcripts, in partial if not all, appeared to be in Cyp19a1b immunoreactive radial glial cell soma or process, suggesting mRNA export through radial glial cells.
Next, orphan nuclear receptors such as sf-1 and dax -1 expressions were studied in the early brain in relation to a wider suite of neurosteroidogenic enzyme genes (star, cyp11a1, hsd3b1, cyp17a1 and cyp19a1b) and estrogen receptors (ERα, ERβ1 and ERβ2) during different post hatching developmental ages and analyzed them against the timing of gonadal sex differentiation. q-PCR results showed that sf-1 mRNA expression was significantly increased together with other key steroidogenic genes during gonadal sex differentiation whereas dax-1 expression was decreased as opposite to sf-1, indicating functional interaction of these orphan nuclear receptors in the grouper brain. sf-1 and dax-1 transcripts were localized in the hypothalamus during different developmental ages by in situ hybridization analysis. Localization of star, sf-1 and dax-1 were highly detected in the cells of the ventromedial hypothalamus (VMH) and in vivo E2 treatment was significantly upregulated only star and sf-1 expressions in the VMH. However, there was no difference detected in the dax-1 expression. Furthermore, the present study also indicates that neurosteroids and estrogen signaling may mediate some of the neuroendocrine functions of AVT during the early brain development. Collectively, the present study demonstrated that the early grouper brain exhibits all necessary changes in the expression of steroidogenic genes, factors, sex steroid receptors, brain steroid concentration and brain aromatse activity. These data provide the evidence that these genes are functional during early post-hatching development in the brain, and have a distinct contribution to different aspects of sex hormone-dependent brain development. Overall, this study demonstrated that the orange-spotted grouper brain possesses all required mechanisms for the local production of E2 which possibly regulates early brain development during gonadal sex differentiation. Therefore, a peak time of development in the early brain is suggested to occur during gonadal sex differentiation in the grouper.

Abstract………………………………………………………… i
Declaration…………………………………………………………… ix
Acknowledgement…………………………………………………… x
Abbreviations………………………………………………………… xiii
Background of the study…………………………………………… 2
Aim of this research work……………………………………… 8
What is thesis about? …………………………………………… 12
Experimental fish…………………………………………………… 18
Materials and method………………………………………………… 18
Samples collection and experimental design………………… 19

Chapter 1: Literature review…………………………………… 20
1.1. Orange spotted-grouper…………………………… 21
1.2. Sex differentiation………………………………… 23
1.3. Types of gonadal sex differentiation in teleosts 29
1.4. Biosynthesis of neurosteroids in the brain………………………. 33
1.5. P450 aromatase (cyp19)……………………………………… 39
1.6. Radial glial cells acting as a neural progenitor
in teleosts brain 55
1.7. PCNA is a marker for cell proliferation activity… 56
1.8. Comparison of developmental changes and
localization of neurosteroidogenic enzyme genes
in response to gonadal differentiation in teleosts
and other vertebrate species………… 58
1.9. Physiological importance of steroidogenic
enzyme genes………61
1.10. Sex steroids and their receptors………………………65
1.11. Steroidogenic factor 1 (Sf-1) is a master regulator
of endocrine gland development and steroidogenesis…78
1.12. Dax-1 (Nr0b1) is a negative co-regulator of many
genes involving in the steroidogenic enzyme
pathway…………………………... 85
1.13. Neuropeptide and their possible role in developing
brain...….. 88
1.14. References……………………………………………………… 91
Chapter 2: Neurosteroidogenic enzymes and their
regulation in the Early Brain of the
protogynous grouper Epinephelus coioides
during gonadal sex differentiation………… 121
2.1. Abstract…………………………………………………………… 122
2.2. Introduction………………………………………………………. 123
2.3. Materials and methods…………………………………………… 131
2.4. Results…………………………………………………………….. 145
2.5. Discussion.………………………………………………………… 171
2.6. Conclusion.……………………………………………………… 178
2.7. Acknowledgments………………………………………………… 181
2.8. References…………………………………………………………. 181

Chapter 3: Developmental expression of genes involved
in neural estrogen biosynthesis and signaling
in the brain of orange-spotted grouper,
Epinephelus coioides during gonadal sex
differentiation……………………………………189
3.1. Abstract………………………………………………………… 190
3.2. Introduction………………………………………………………. 191
3.3. Materials and methods………………………………………… 196
3.4. Results……………………………………………………………... 209
3.5. Discussion………………………………………………………… 219
3.6. Conclusion.……………………………………………… 228
3.7. Acknowledgments………………………………………………... 229
3.8. References………………………………………………………… 229

Chapter 4: Increase in estrogen signaling in the early
brain of female grouper and male black porgy
during gonadal sex differentiation…………… 237
4.1. Abstract…………………………………………………………… 238
4.2. Introduction………………………………………………………. 239
4.3. A peak of estrogen signaling in the early brain
of black porgy and grouper during gonadal sex
differentiation………………. 242
4.4. Aromatse expressed in radial glial cells of brain
of teleosts…… 245
4.5. Sex differences in the aromatase expression in
teleosts brain…. 248
4.6. Expression of steroidogenic enzymes in the brain of
black porgy and grouper fish………………………………251
4.7. Conclusion…………………………………………………………253
4.8. References..……………………………………………………….. 253

Chapter 5: sf-1, dax-1 and steroidogenic genes in the
early brain of orange-spotted grouper
Epinephelus coioides during gonadal sex
differentiation……………………………... 259
5.1. Abstract…………………………………………………………… 260
5.2. Introduction………………………………………………………. 261
5.3. Material and methods……………………………………………. 266
5.4. Results……………………………………………………………... 275
5.5. Discussion……………………………………………… 281
5.6. Conclusion……………………………………………... 296
5.7. References………………………………………………………… 297

Chapter 6: Integrated functions of neuropeptide,
neurosteroidogenic enzymes and estrogen
receptors in the early brain of orange-
spotted grouper (Epinephelus coioides):
Potential for functional interactions
during gonadal sex differentiation…………… 304
6.1. Abstract…………………………………………………………… 305
6.2. Introduction……………………………………………………… 306
6.3. Material and methods………………………………………… 312
6.4. Results…………………………………………………………… 320
6.5. Discussion………………………………………………………… 330
6.6. Conclusion………………………………………………………… 346
6.7. References………………………………………………………… 347
7. Conclusion and perspectives………………………………… 355

Acher, R., 1996. Molecular evolution of fish neurohypophysial hormones: Neutral and selective evolutionary mechanisms. Gen. Comp. Endocrinol. 102, 157–172.
Agate, R.J., Perlman, W.R., Arnold, A.P., 2002. Cloning and expression of zebra finch (Taeniopygia guttata) steroidogenic factor 1: overlap with hypothalamic but not with telencephalic aromatase. Biol. Reprod. 66, 1127–1133.
Anderson, T., Forrester, J., 2001. Administration of estradiol to barramundi, Lates calcarifer, induces protandrous sex change, in: H.I.Th. Goos, R.K. Rastogi, H. Vaudry, R. Pierantoni (Eds). Perspective in Comparative Endocrinology, Unity and Diversity, Monduzzi Editiors, Bologna. pp. 155-164.
Andersson, S., Berman, D.M., Jenkins, E.P., Russell, D.W., 1991. Deletion of steroid 5α-reductase 2 gene in male pseudohermaphroditism. Nature 354, 159–161.
Arnold, A.P., Xu, J., Grisham, W., Chen, X., Kim, Y.H., Itoh, Y., 2004. Minireview: Sex chromosomes and brain sexual differentiation. Endocrinology 145, 1057–1062.
Aubrey Milunsky; Jeff Milunsky., 2010. Genetic Disorders and the Fetus: Diagnosis, Prevention and Treatment. John Wiley and Sons. pp. 600–. ISBN 978-1-4051-9087-9.
B. Adolf, P. Chapouton, C.S. Lam, S. Topp, B. Tannhauser, U. Strahle, M. Gotz, L. Bally-Cuif, Conserved and acquired features of adult neurogenesis in the zebrafish telencephalon, Dev. Biol. 295 (2006) 278–293.
Ball, G.F., Balthazart, J., 2004. Hormonal regulation of brain circuits mediating male sexual behavior in birds. Physiol. Behav. 83, 329–346.
Balment, R.J., Lu, W., Weybourne, E., Warne, J.M., 2006. Arginine vasotocin a key hormone in fish physiology and behaviour: a review with insights from mammalian models. Gen. Comp. Endocrinol. 147, 9–16.
Balthazart, J., Ball, G.F., 1998. New insights into the regulation and function of brain estrogen synthase (aromatase), Trends Neurosci. 21, 243–249.
Balthazart, J., Ball, G.F., 2006. Is brain estradiol a hormone or neurotransmitter? Trends Neurosci. 29, 241–249.
Barha CK, Lieblich SE, Galea LA. 2009. Different forms of oestrogen rapidly up-regulate cell proliferation in the dentate gyrus of adult female rats. J Neuroendo-crinol 21:155–166.
Baroiller, J.-F., Guiguen, Y., Fostier, A., 1999. Endocrine and environmental aspects of sex differentiation in fish. Cell Mol. Life Sci. 55, 910–931.
Bhandari, R.K., Alam, M.A., Soyano, K., Nakamura, M., 2006. Induction of female-to-male sex change in the honeycomb grouper (Epinephelus merra) by 11-ketotestosterone treatments. Zool. Sci. 23, 65–69.
Bhandari, R.K., Komuro, H., Nakamura, S., Higa, M., Nakamura, M., 2003. Gonadal restructuring and correlative steroid hormone profiles during natural sex change in protogynous honeycomb grouper (Epinephelus merra). Zool. Sci. 20, 1399–1404.
Birnbaumer, M., 2000. Vasopressin receptors. Trends Endocrinol. Metab. 11, 406–410.
Blazquez, M., Gonzalez, A., Papadaki, M., Mylonas, C., Piferrer, F., 2008. Sex-related changes in estrogen receptors and aromatase gene expression and enzymatic activity during early development and sex differentiation in the European sea bass (Dicentrarchus labrax). Gen. Comp. Endocrinol. 158, 95–101.
Blazquez, M., Piferrer, F., 2004. Cloning, sequence analysis, tissue distribution, and sex-specific expression of the neural form of P450 aromatase in juvenile sea bass (Dicentrarchus labrax). Mol. Cell. Endocrinol. 219, 83–94.
Bokenkamp, D., Jungblut, P., Thole, H.H., 1994. The C-terminal half of the porcine estradiol receptor contains no post-translational modification: determination of the primary structure. Mol. Cell Endocrinol. 104, 163–172.
Borg, B., 1994. Androgens in teleost fishes. Comp. Biochem. Physiol. 109 C, 219–245.
Borg, B., Andersson, E., Mayer, I., JG., L., 1989. Aromatase activity in the brain of the three-spined stickleback, Gasterosteus aculeatus. III. Effects of castration under different conditions and of replacement with different androgens. Exp. Biol. 48, 149–152.
Borg, B., Timmers, R., Lambert, J., 1987. Aromatase activity in the brain of the three-spined stickleback, Gasterosteus aculeatus. I. Distribution and effects of season and photoperiod. Exp. Biol. 47, 63–68.
Brailoiu, E., Dun, S.L., Brailoiu, G.C., Mizuo, K., Sklar, L.A., 2007. Distribution and characterization of estrogen receptor G protein-coupled receptor 30 in the rat central nervous system. J. Endocrinol. 193, 311–21.
Brueggemeier, R.W., 2004. Aromatase inhibitors: new endocrine treatment of breast cancer. Semin. Reprod. Med. 22, 31–43.
Bruzzone, F., Do Rego, J.L., Luu-The, V., Pelletier, G., Vallarino, M., Vaudry, H., 2010. Immunohistochemical localization and biological activity of 3β-hydroxysteroid dehydrogenase and 5α-reductase in the brain of the frog, Rana esculenta, during development. J. Chem. Neuroanat. 39, 35–50.
Bryan, M.B., Scott, A.P., Li, W., 2007. The sea lamprey (Petromyzon marinus) has a receptor for androstenedione. Biol. Reprod. 77, 688–696.
Callard, G.V., Petro, Z., Ryan, K.J., 1978. Phylogenetic distribution of aromatase and other androgen-converting enzymes in the central nervous system. Endocrinology 103, 2283–2290.
Capel, B., 1998. Sex in the 90s: SRY and the switch to the male pathway. Annu. Rev. Physiol. 60, 497–523.
Carreau, S., Lambard, S., Delalande, C., Denis-Galeraud, I., Bilinska, B., Bourguiba, S., 2003. Aromatase expression and role of estrogens in male gonad: a review. Reprod. Biol. Endocrinol. 1, 35.
Chang, C., Lin, B., 1998. Estradiol-17β stimulates aromatase activity and reversible sex change in protandrous black porgy, Acanthopagrus schlegeli. J. Exp. Zool. 280, 165–173.
Chang, X.T., Kobayashi, T., Sentilkumaran, B., Kobayashi-Kajiura, H., Sudhakumari, C.C., Nagahama, Y., 2005. Two types of aromatase with different encoding genes, tissue distribution and developmental expression in Nile tilapia (Oreochromis niloticus). Gen. Comp. Endocrinol. 141, 101–115.
Chang, Y.T., Kappy, M.S., Iwamoto, K., Wang, J., Yang, X., Pang, S., 1993. Mutations in the type II 3β-hydroxysteroid dehydrogenase gene in a patient with classic salt-wasting 3-HSD deficiency congenital adrenal hyperplasia. Pediatr. Res. 34, 698–700.
Chaube, R., Singh, R.K., Joy, K.P., 2012. Estrogen regulation of brain vasotocin secretion in the catfish Heteropneustes fossilis: an interaction with catecholaminergic system. Gen. Comp. Endocrinol. 175, 206–213.
Chen, H., Zhang, Y., Li, S., Lin, M., Shi, Y., Sang, Q., Liu, M., Zhang, H., Lu, D., Meng, Z., Liu, X., Lin, H., 2011. Molecular cloning, characterization and expression profiles of three estrogen receptors in protogynous hermaphroditic orange-spotted grouper (Epinephelus coioides). Gen. Comp. Endocrinol. 172, 371–381.
Chiang, E.F., Yan, Y.L., Guiguen, Y., Postlethwait, J., Chung, B., 2001. Two Cyp19 (P450 aromatase) genes on duplicated zebrafish chromosomes are expressed in ovary or brain, Mol. Biol. Evol. 18, 542–550.
Choi, C.Y., Habibi, H.R., 2003. Molecular cloning of estrogen receptor alpha and expression pattern of estrogen receptor subtypes in male and female goldfish. Mol. Cell Endocrinol. 204, 169–177.
Choi, C.Y., Habibi, H.R., 2005. Molecular cloning and tissue distribution of SF-1-related orphan receptors during sexual maturation in female goldfish. Biotechnol. Lett. 27, 1283–1290.
Choi, J.Y., Park, J.G., Jeong, H.B., Lee, Y.D., Takemura, A., Kim, S.J., 2005. Molecular cloning of cytochrome P450 aromatases in the protogynous wrasse, Halichoeres tenuispinis, Comp. Biochem. Physiol. B: Biochem. Mol. Biol. 141, 49–59.
Cole, K.S., Noakes, D.L.G., 1997. Gonadal development and sexual allocation in mangrove killifish, Rivulus marmoratus (Pisces: Atherinomorpha). Copeia 1997, 596–600.
Compagnone, N.A., Bulfone, A., Rubenstein, J.L., Mellon, S.H., 1995. Expression of the steroidogenic enzyme P450scc in the central and peripheral nervous systems during rodent embryogenesis. Endocrinology 136, 2689–2696.
Compagnone, N.A., Mellon, S.H., 2000. Neurosteroids: biosynthesis and function of these novel neuromodulators. Front. Neuroendocrinol. 21, 1–56.
Conley, A.J., Bird, I.M., 1997. Minireview: the role of cytochrome P450 17α-hydroxylase and 3β-hydroxysteroid dehydrogenase in the integration of gonadal and adrenal steroidogenesis via the Δ5 and Δ4 pathways of steroidogenesis in mammals. Biol. Reprod. 56, 789–799.
Conte, F.A., Grumbach, M.M., Ito, Y., 1994. A syndrome of female pseudo-hermaphrodism, hypergonadotropic hypogonadism, and multicystic ovaries associated with missense mutations in the gene encoding aro-matase (P450arom). J. Clin. Endocrinol. Metab. 78, 1287–1292.
Corley, D.R., Li, X., Lei, Z.M., Rao, C.V., 2000. Potential regulation of GnRH gene by a steroidogenic factor-1-like protein. Mol. Hum. Reprod. 6, 671–676.
Cornett, L.E., Kirby, J.D., Vizcarra, J.A., Ellison, J.C., Thrash, j., Mayeux, P.R., Crew, M.D., Jones, S.M., Ali, N., Baeyens, D.A., 2003. Molecular cloning and functional characterization of a vasotocin receptor sub-type expressed in the pituitary gland of the domestic chicken (Gallus domesticus): avian homolog of the mammalian V1b-vasopressin recep-tor. Regul. Peptides 110, 231–239.
Corpéchot, C., Robel, P., Axelson, M., Sjövall, J., Baulieu, E.E., 1981. Characterization and measurement of dehydroepiandrosterone sulfate in rat brain, Proc. Natl. Acad. Sci. USA 78, 4704–4707.
Corpéchot, C., Synguelakis, M., Talha, S., Axelson, M., Sjövall, J., Vihko, R., Baulieu, E.E., Robel. P., 1983. Pregnenolone and its sulfate ester in the rat brain, Brain Res. 270, 119–125.
Couse, J.F., Korach, K.S., 1999. Estrogen receptor null mice: What have we learned and where will they lead us? Endocr. Rev. 20, 358–417.
Cross, E., Roselli, C.E., 1999. 17β-Estradiol rapidly facilitates chemoinvestigation and mounting in castrated male rats. Am. J. Physiol. Regul. Integr. Comp. Physiol. 276, R1346–R1350.
Dalla Valle, L., Lunardi, L., Colombo, L., Belvedere, P., 2002. European sea bass (Dicentrarchus labrax L.) cytochrome P450arom: cDNA cloning, expression and genomic organization. J. Steroid Biochem. Mol. Biol. 80, 25–34.
Darlison, M.G., Richter, D., 1999. Multiple genes for neuropeptides and their recep-tors: co-evolution and physiology. Trends Neurosci. 22, 81–88.
de Keyzer, Y., Lenne, F., Auzan, C., Jegou, S., Rene, P., Vaudry, H., Khun, J-M., Luton, J-P., Clauser, E., Bertagna, X., 1996. The pituitary V3 vasopressin receptor and the corticotroph phenotype in ectopic ACTH syndrome. J. Clin. Invest. 97, 1311–1318.
de Vries, G.J., So dersten, P., 2009. Sex differences in the brain: the relation between structure and function. Horm. Behav. 55, 589–596.
Deladoëy, J., Flück, C., Bex, M., 1999. Aromatase deficiency caused by a novel P450arom gene mutation: impact of absent estrogen production on serum gonadotropin concentration in a boy. J. Clin. Endocrinol. Metab. 84, 4050–4054.
Devlin, R.H., Nagahama, Y., 2002. Sex determination and sex differentiation in fish: an overview of genetic, physiological, and environmental influences. Aquaculture 208, 191–364.
Diotel, N., Le Page, Y., Mouriec, K., Tong, S.K., Pellegrini, E., Vaillant, C., Anglade, I., Brion, F., Pakdel, F., Chung, B.C., Kah, O., 2010. Aromatase in the brain of teleost fish: expression, regulation and putative functions. Front. Neuroendocrinol. 31, 172–192.
Diotel, N., Do Rego, J.L., Anglade, I., Vaillant, C., Pellegrini, E., Gueguen, M.M., Mironov, S,
Vaudry, H., Kah, O., 2011. Activity and expression of steroidogenic enzymes in the brain of adult zebrafish. Eur. J. Neurosci. 34, 45–56.
Do Rego, J.L., Seong, J.Y., Burel, D., Leprince, J., Luu-The, V., Tsutsui, K., Tonon, M.C., Pelletier, G., Vaudry, H., 2009. Neurosteroid biosynthesis: enzymatic pathways and neuroendocrine regulation by neurotransmitters and neuropeptides. Front. Neuroendocrinol. 3, 259–301.
Do Rego, J.L., Tremblay, Y., Luu-The, V., Repetto, E., Castel, H., Vallarino, M., Bélanger, A., Pelletier, G., Vaudry, H., 2007. Immunohistochemical localization and biological activity of the steroidogenic enzyme cytochrome P450 17α-hydroxylase/C17, 20-lyase (P450C17) in the frog brain and pituitary. J. Neurochem. 100, 251–268.
Do-Rego, J.L., Acharjee, S., Seong, J.Y., Galas, L., Alexandre, D., Bizet, P., Burlet, A., Kwon, H.B., Luu-The, V., Pelletier, G., Vaudry, H., 2006. Vasotocin and mesotocin stimulate the biosynthesis of neurosteroids in the frog brain. J. Neurosci. 26, 6749–6760.
Dulac, C., Kimchi, T., 2007. Neural mechanisms underlying sex specific behaviors in vertebrates. Curr. Opin. Neurobiol. 17, 675–683.
Duval, D.L., Nelson, S.E., Clay, C.M., 1997. A binding site for steroidogenic factor-1 is part of a complex enhancer that mediates expression of the murine gonadotropin-releasing hormone receptor gene. Biol. Reprod. 56, 160–168.
Leonardi, E., Girlando, S., Serio, G., Mauri, F.A., Perrone, G., Scampini, S., Dalla Palma, P.,
Barbareschi, M.,1992. PCNA and Ki67 expression in breast carcinoma: correlations with clinical and biological variables. J. Clin. Pathol. 45, 416–419.
Ekstrom, P., Johnsson, C.M., Ohlin, L.M., 2001. Ventricular proliferation zones in the brain of an adult teleost fish and their relation to neuromeres and migration (secondary matrix) zones. J. Comp. Neurol. 436, 92–110.
Essers, J., Theil, A.F., Baldeyron, C., van Cappellen, W.A., Houtsmuller, A.B., Kanaar, R., Vermeulen, W., 2005. Nuclear dynamics of PCNA in DNA replication and repair, Mol. Cell. Biol. 25, 9350–9359.
Ezaz, M.T., Harvey, S.C., Boonphakdee, C., Teale, A.J., McAndrew, B.J., Penman, D.J., 2004. Isolation and physical mapping of sex-linked AFLP markers in Nile tilapia, Oreochromis niloticus L. Mar. Biotechnol. 6, 435–445.
Felip, A., Young, W.P., Wheeler, P.A., Thorgaard, G.H., 2005. An AFLP based approach for the identification of sex-linked markers in rainbow trout, Oncorhynchus mykiss. Aquaculture 247:35–43.
Filby, A.L., Tyler, C.R., 2005. Molecular characterization of estrogen receptors 1, 2a, and 2b and their tissue and ontogenic expression profiles in fathead minnow (Pimephales promelas). Biol Reprod. 73, 648–662.
Fleming, A., Wibbels, T., Skipper, J.K., Crews, D., 1999. Developmental expression of steroidogenic factor 1 in a turtle with temperature-dependent sex determination. Gen. Comp. Endocrinol. 116, 336–346.
Foran, C.M., Bass, A.H., 1999. Preoptic GnRH and AVT: axes for sexual plasticity in teleost fish. Gen. Comp. Endocrinol. 116, 141–152.
Galien, R., Garcia T., 1997. Estrogen receptor impairs interleukin-6 expression by preventing protein binding on the NF-kB site. Nucleic Acids Res. 25, 2424–2429.
Garcia-Segura LM, Azcoitia I, DonCarlos LL. 2001. Neuroprotection by estradiol. Prog Neurobiol 63:29–60.
Garcia-Segura LM, Wozniak A, Azicoita I, Rodriques JR, Hutchinson RE, Hutchinson JB., 1999. Aromatase expression by astrocytes after brain injury: implications for local estrogen formation in brain repair. Neuroscience 89:567–578.
Gelinas, D., Pitoc, G.A., Callard, G.V., 1998. Isolation of a goldfish brain cytochrome P450 aromatase cDNA: mRNA expression during the seasonal cycle and after steroid treatment.
Mol. Cell. Endocrinol. 138, 81–93.
Godwin, J., Sawby, R., Warner, R.R., Crews, D., Grober, M.S., 2000. Hypothalamic arginine vasotocin mRNA abundance variation across sexes and with sex change in a coral reef fish. Brain Behav. Evol. 55, 77–84.
Gonzalez, A., Piferrer, F., 2003. Aromatase activity in the European sea bass (Dicentrarchus labrax L.) brain. Distribution and changes in relation to age, sex, and the annual reproductive cycle. Gen. Comp. Endocrinol. 132, 223–230.
Goodson, J.L., 2008. Nonapeptides and the evolutionary patterning of sociality. Prog. Brain Res. 170, 3–15.
Govoroun, M., McMeel, O.M., D'Cotta, H., Ricordel, M.-J., Smith, T., Fostier, A., Guiguen, Y., 2001. Steroid enzyme gene expressions during natural and androgen-induced gonadal differentiation in the rainbow trout, Oncorhynchus mykiss. J. Exp. Zool. 290, 558–566.
Greco, T.L., Payne, A.H., 1994. Ontogeny of expression of the genes for steroidogenic enzymes P450 side-chain cleavage, 3β-hydroxysteroid dehydrogenase, P450 17α-hydroxylase/C17-20 lyase, and P450 aromatase in fetal mouse gonads. Endocrinology 135, 262–268.
Green, S., Waher, P., Kumar, V., Kmst, A., Bornert, J.M, Argos, P. and Chambon, P., 1986. Human oestrogen receptor cDNA: sequence, expression and homology to v-erb-A. Nature (NSC) 320,134–139.
Greytak, S.R., Champlin, D., Callard, G.V., 2005. Isolation and characterization of two cytochrome P450 aromatase forms in killifish (Fundulus heteroclitus): differential expression in fish from polluted and unpolluted environments. Aquat. Toxicol. 71, 371–389.
Guennoun, R., Fiddes, R.J., Gouezou, M., Lombes, M., Baulieu, E.E., 1995. A key enzyme in the biosynthesis of neurosteroids, 3b-hydroxysteroid dehydrogenaser/Δ5 Δ4-isomerase (3β-HSD), is expressed in rat brain, Mol. Brain Res. 30, 287–300.
Guiguen, Y., Baroiller, J.-F., Ricordel, M.-J., Iseki, K., McMeel, O.M., Martin, S.A.M., Fostier, A., 1999. Involvement of estrogens in the process of sex differentiation in two fish species: the rainbow trout (Oncorhynchus mykiss) and a tilapia (Oreochromis niloticus). Mol. Reprod. Dev. 54, 154–162.
Halm, S., Kwon, J.Y., Rand-Weaver, M., Sumpter, J.P., Pounds, N., Hutchinson, T.H., Tyler,
C.R., 2003. Cloning and gene expression of P450 17α-hydroxylase,17,20-lyase cDNA in the gonads and brain of the fathead minnow Pimephales promelas. Gen. Comp. Endocrinol. 130, 256–66.
Halvorson, L.M., Kaiser, U.B., Chin, W.W., 1996. Stimulation of luteinizing hormone beta gene promoter activity by the orphan nuclear receptor, steroidogenic factor-1. J. Biol. Chem. 271, 6645–6650.

Hammer, F., Compagnone, N.A., Vigne, J.L., Bair, S.R., Mellon, S.H., 2004. Transcriptional
regulation of P450scc gene expression in the embryonic rodent nervous system. Endocrinology 145, 901–912.
Harbott, L.K., Burmeister, S.S., White, R.B., Vagell, M., Fernald, R.D., 2007. Androgen receptor in a cichlid fish, Astatotilapia burtoni: structure, localization, and expression levels. J. Comp. Neurol. 504, 57–73.
Hasunuma, I., Sakai, T., Nakada, T., Toyoda, F., Namiki, H., Kikuyama, S., 2007. Molec-ular cloning of three types of arginine vasotocin receptor in the newt, Cynops pyrrhogaster. Gen. Comp. Endocrinol. 151, 252–258.
Hatano, O., Takayama, K., Imai, T., Waterman, M.R., Takakusu, A., Omura, T., Morohashi, K., 1994. Sex-dependent expression of a transcription factor, Ad4BP, regulating steroidogenic P-450 genes in the gonads during prenatal andpostnatal rat development. Development 120, 2787–2797.
Hayashi, M., Sasaki, S., Tsuganezawa, H., Monkawa, T., Kitajima, W., Konishi, K., Fushimi, K., Marumo, F., Saruta, T., 1994. Expression and distribution of aquaporin of collecting duct are regulated by vasopressin V-2 receptor in rat kidney. J. Clin. Invest. 94, 1778–1783.
Hazell, G.G., Yao, S.T., Roper, J.A., Prossnitz, E.R., O'Carroll, A.M, Lolait, S.J., 2009. Localisation of GPR30, a novel G protein-coupled oestrogen receptor, suggests multiple functions in rodent brain and peripheral tissues. J. Endocrinol. 202, 223–236.
Henderson, R.G., Brown, A.E., Tobet, S.A., 1999. Sex differences in cell migration in the preoptic area/anterior hypothalamus of mice. J Neurobiol. 41, 252–266.
Hendry, C.I., Martin-Robichaud, D.J., Benfey, T.J., 2002. Gonadal sex differentiation in Atlantic halibut. J. Fish Biol. 60, 1431–1442.
Honda, S., Morohashi, K., Nomura, M., Takeya, H., Kitajima, M., Omura, T., 1993. Ad4BP regulating steroidogenic P-450 gene is a member of steroid hormone receptor superfamily.
J. Biol. Chem. 268, 7494–7502.
Hoogervorst, E.M., van Steeg, H., de Vries. A., 2005. Nucleotide excision repair- and p53-deficient mouse models in cancer research. Mutat. Res. 574, 3–21.
Hossain, M.S., Larsson, A., Scherbak, N., Olsson, P.E., Orban, L., 2008. Zebrafish androgen receptor: isolation, molecular, and biochemical characterization. Biol. Reprod. 78, 361–369.
Hrabovszky, E., Kallo, I., Hajszan, T., Shughrue, P.J., Merchenthaler, I., Liposits, Z., 1998. Expression of estrogen receptor-β mRNA in oxytocin and vasopressin neurons of the rat supraoptic and paraventricular nuclei. Endocrinology 139, 2600–2604.
Hu, M.C., Hsu, H.J., Guo, I.C., Chung, B.C., 2004. Function of CYP11A1 in animal models. Mol. Cell. Endocrinol. 215, 95–100.
Hu, M.C., Hsu, N.C., Pai, C.I., Wang, C.K., Chung, B., 2001. Functions of the upstream and proximal steroidogenic factor 1 (SF-1)-binding sites in the CYP11A1 promoter in basal transcription and hormonal response. Mol. Endocrinol. 15, 812–818.
Ijiri, S., Kazeto, Y., Lokman, P.M., Adachi, S., Yamauchi, K., 2003. Characterization of a cDNA encoding P-450 aromatase (CYP19) from Japanese eel ovary and its expression in ovarian follicles during induced ovarian development. Gen. Comp. Endocrinol. 130, 193–203.
Ikeda, Y., Luo, X., Abbud, R., Nilson, J.H., Parker, K.L., 1995. The nuclear receptor steroidogenic factor 1 is essential for the formation of the ventromedial hypothalamic nucleus. Mol. Endocrinol. 9, 478– 486.
Ikeda, Y., Takeda, Y., Shikayama, T., Mukai, T., Hisano, S., Morohashi, K.I., 2001. Comparative localization of Dax-1 and Ad4BP/SF-1 during development of the hypothalamic-pituitary-gonadal axis suggests their closely related and distinct functions. Dev. Dyn. 220, 363–376.
Ikeuchi, T., Todo, T., Kobayashi, T., Nagahama, Y., 1999. cDNA cloning of a novel androgen receptor subtype. J. Biol. Chem. 274, 25205–25209.
Jacobs, E.C., Arnold, A.P., Campagnoni, A.T., 1996. Zebra finch estrogen receptor cDNA: cloning and mRNA expression. J. Steroid Biochem. Mol. Biol. 59, 135–45.
Jefcoate, C., 2002. High-flux mitochondrial cholesterol trafficking, a specialized function of the adrenal cortex. J. Clin. Invest. 110, 881–890.
Jeyasuriya, P., Place, A.R., 1998. Embryonic brain-gonadal axis in temperature dependent sex
determination in reptiles: a role for P450 aromatase (CYP19). J. Exp. Zool. 281, 428–449.
Jurkevich, A., Berghman, L.R., Cornett, L.E., Kuenzel, W.J., 2005. Characterization and immunohistochemical visualization of the vasotocin VT2 receptor in the pituitary gland of the chicken, Gallus gallus. Gen. Comp. Endocrinol. 143, 82–91.
Kanda, H., Okubo, T., Omori, N., Niihara, H., Matsumoto, N., Yamada, K., Yoshimoto, S., Ito,
M., Yamashita, S., Shiba, T., Takamatsu, N., 2006. Transcriptional regulation of the rainbow trout CYP19a gene by FTZ-F1 homologue. J. Steroid Biochem. Mol. Biol. 99, 85–92.
Karube, M., Fernandino, J.I., Strobl-Mazzulla, P., Strüssmann, C.A., Yoshizaki, G., Somoza, G.M., Patiño, R., 2007. Characterization and expression profile of the ovarian cytochrome P-450 aromatase (cyp19A1) gene during thermolabile sex determination in pejerrey, Odontesthes bonariensis. J. Exp. Zool. Part. A: Ecol. Genet. Physiol. 307A, 625–636.
Kater C.E., Biglieri E.G., 1994. Disorders of steroid 17α-hydroxylase deficiency. Endocrinol. Metab. Clin. North. Am. 23:341-357.
Katsu Y, Lange A, Miyagawa S, Urushitani H, Tatarazako N, Kawashima Y, Tyler CR, Iguchi T. Cloning, expression and functional characterization of carp, Cyprinus carpio, estrogen receptorsand their differential activations by estrogens. J Appl Toxicol. 2011 Jul 1. doi: 10.1002/jat.1707.
Katsumata, N., Ohtake, M., Hojo, T., Ogawa, E., Hara, T., Sato, N., Tanaka, T., 2002. Compound heterozygous mutations in the cholesterol side-chain cleavage enzyme gene (CYP11A) cause congenital adrenal insufficiency in humans. J. Clin. Endocrinol. Metab. 87, 3808–3813.
Khan, I. A., and Thomas, P. (1999). GABA exerts stimulatory and inhibitory influences on gonadotropin II secretion in the Atlantic croaker (Micropogonias undulatus). Neuroendocrinology 69, 261–268.
Khan, I. A., Thomas, P., 1992. Stimulatory effects of serotonin on maturational gonadotropin release in the Atlantic croaker, Micropogonias undulatus. Gen. Comp. Endocrinol. 88, 388–396.
Kimoto, T., Tsurugizawa, T., Ohta, Y., Makino, J., Tamura, H., Hojo, Y., Takata, N., Kawato,
S., 2001. Neurosteroid synthesis by cytochrome p450-containing systems localized in the rat brain hippocampal neurons: N-methyl-D-aspartate and calcium-dependent synthesis. Endocrinology 142, 3578–3589.
Kishida,M., Callard, G.V., 2001. Distinct cytochrome P450 aromatase isoforms in zebrafish
(Danio rerio) brain and ovary are differentially programmed and estrogen regulated during early development. Endocrinology 142, 740–750.
Kitano, T., Takamune, K., Kobayashi, T., Nagahama, Y., Abe, S.-I., 1999. Suppression of P450 aromatase gene expression in sex-reversed males produced by rearing genetically female larvae at a high water temperature during a period of sex differentiation in the Japanese flounder (Paralichthys olivaceus). J. Mol. Endocrinol. 23, 167–176.
Kitano, T., Takamune, K., Nagahama, Y., Abe, S.I., 2000. Aromatase inhibitor and 17α-methyltestosterone cause sex-reversal from genetical females to phenotypic males and suppression of P450 aromatase gene expression in Japanese flounder (Paralichthys olivaceus). Mol. Reprod. Dev. 56, 1–5.
Kline, R.J., O'Connell, L.A., Hofmann, H.A., Holt, G.J., Khan, I.A., 2011. The distribution of an AVT V1a receptor in the brain of a sex changing fish, Epinephelus adscensionis. J. Chem. Neuroanat. 42, 72–88.
Kohchi, C., Ukena, K., Tsutsui, K., 1998. Age- and region-specific expressions of the messenger RNAs encoding for steroidogenic enzymes p450scc, P450c17 and 3β-HSD in the postnatal rat brain. Brain Res. 801, 233–238.
Kohchi, C., Ukena, K., Tsutsui, K., 1998. Age- and region-specific expressions of the messenger RNAs encoding for steroidogenic enzymes P450scc, P450c17 and 3β-HSD in the postnatal rat brain. Brain. Res. 801, 233–238.
Köhler, T., Pröls, F., Brand-Saberi, B., 2005. PCNA in situ hybridization: a novel and reliable tool for detection of dynamic changes in proliferative activity. Histochem. Cell. Biol. 123, 315–327.
Koopman, P., 1999. Sry and Sox9: mammalian testis-determining genes. Cell. Mol. Life Sci. 55, 839–856.
Koopman, P., 2001. Sry, Sox9 and mammalian sex determination. EXS 91, 25–56.
Kovacs, B., Egedi, S., Bartfai, R., Orban, L., 2000. Male-specific DNA markers from African
catfish, Clarias gariepinus. Genetica. 110, 267–276.
Kroon, F., Liley, N., 2000. The role of steroid hormones in protogynous sex change in black-eye goby, Coryphoptrus nicholsi (Telesotei: Gobidae). Gen. Comp. Endocrinol. 118, 273–283.
Kuiper, G.G., Carlsson, B., Grandien, K., Enmark, E., Haggblad, J., 1997. Comparison of the ligand binding specificity and transcript tissue distribution of estrogen receptors α and β. Endocrinology 138, 863–70.
Kuiper, G.G., Enmark, E., Peho-Huikko, M., Nilsson, S., Gustafsson, J., 1996. Cloning of a novel estrogen receptor expressed in rat prostate and ovary. Proc. Nati. Acad. Sci. USA 93. 5925–5930.
Kumar, V., Green, S., Stack, G., Berry, M., Jin, J.R., Chambon, P., 1987. Functional domains of the human estrogen receptor. Cell 51, 941–951.
Kwon, J.Y., McAndrew, B.J., Penman, D.J., 2001. Cloning of brain aromatase gene and expression of brain and ovarian aromatase genes during sexual differentiation in genetic male and female Nile tilapia, Oreochromis niloticus, Mol. Reprod. Dev. 59, 359–370.
Lala, D.S., Rice, D.A., Parker, K.L., 1992. Steroidogenic factor I, a key regulator of steroidogenic enzyme expression, is the mouse homolog of fushi tarazu-factor I. Mol. Endocrinol. 6, 1249–1258.
Lance, V.A., 2008. Is regulation of aromatase expression in reptiles the key to understanding temperature-dependent sex determination? J. Exp. Zool. 309A, 314–322.
Le Goascogne, C., Robel, P., Gouézou, M., Sananès, N., Baulieu, E.E., Waterman, M., 1987. Neurosteroids: cytochrome P-450scc in rat brain. Science 237, 1212–1215.
Le Goascogne, C., Robel, P., Gouézou, M., Sananès, N., Baulieu, E.E., Waterman, M., 1987. Neurosteroids: cytochrome P-450scc in rat brain. Science 237, 1212–1215.
Lee, Y.D., Rho, S., Chang, Y.J., Baek, H.J., An, C.M., 1996. Sex differentiation of the rockfish, Sebastes schlegeli. J. Korean Fish Soc. 29, 44–50.
Lee, Y.H., Wu, G.C., Du, J.L., Chang, C.F., 2004. Estradiol-17β induced a reversible sex change
in the fingerlings of protandrous black porgy, Acanthopagrus schlegeli Bleeker: the
possible roles of luteinizing hormone in sex change. Biol Reprod. 71, 1270–1278.
Leers-Sucheta, S., Morohashi, K., Mason, J.I., Melner, M.H., 1997. Synergistic activation of the human type II 3β-hydroxysteroid dehydrogenase/Δ5-Δ4 isomerase promoter by the transcription factor steroidogenic factor-1/adrenal 4-binding protein and phorbol ester. J. Biol. Chem. 272, 7960–7967.
Lema, S.C., 2010. Identification of multiple vasotocin receptor cDNAs in teleost fish: sequences, phylogenetic analysis, sites of expression, and regulation in the hypothalamus and gill in response to hyperosmotic challenge. Mol. Cell. Endocrinol. 321, 215–230.
Lema, S.C., Nevitt, G.A., 2004. Variation in vasotocin immunoreactivity in the brain of recently isolated populations of a death valley pupfish, Cyprinodon nevadensis. Gen. Comp. Endocrinol. 135, 300–309.
Liang, J.H., Sankai, T., Yoshida, T., Cho, F., Yoshikawa, Y., 1999. Localization of immunoreactive testosterone and 3β-hydroxysteroid dehydrogenase/Δ5-Δ4 isomerase in cynomolgus monkey (Macaca fascicularis) testes during postnatal development. J. Med. Primatol. 28, 62–66.
Lolait, S.J., O’Carrol, A-M., McBride, O.W., Konig, M., Morel, A., Brown-stein, M.J., 1992. Cloning and characterization of a vasopressin V2 receptor and possible link to nephrogenic diabetes insipidus. Nature 357, 336–339.
Loomis, A.K., Thomas, P., 1999. Binding characteristics of estrogen receptor (ER) in Atlantic croaker (Micropogonias undulatus) testis: different affinity for estrogens and xenobiotics from that of hepatic ER. Biol. Reprod. 61, 51–60.
Lorence, M.C., Murry, B.A., Trant J,M., Mason, J.I., 1990. Human 3β-hydroxysteroid dehydrogenase/Δ 5-4 isomerase from placenta: expression in nonsteroidogenic cells of a protein that catalyzes the dehydrogenation/isomerization of C21 and C19 steroids. Endocrinology 126, 2493–2498.
Luckenbach, J.A., Early, L.W., Rowe, A.H., Borski, R.J., Daniels, H.V., Godwin, J., 2005. Aromatase cytochrome P450: cloning, intron variation and ontogeny of gene expression in southern flounder (Paralichthys lethostigma). J. Exp. Zool. 303A, 643–656.
Lynch, J.P., Lala, D.S., Peluso, J.J., Luo, W., Parker, K.L., White, B.A., 1993. Steroidogenic
factor 1, an orphan nuclear receptor, regulates the expression of the rat aromatase gene in
gonadal tissues. Mol. Endocrinol. 7, 776–786.
Maack, G., Segner, H., 2003. Morphological development of the gonads in zebrafish. J. Fish Biol. 62, 895–906.
Marsh, K.E., Creutz, L.M., Hawkins, M.B., Godwin, J., 2006. Aromatase immunoreactivity in the bluehead wrasse brain. Thalassoma bifasciatum: immunolocalization and co-regionalization with arginine vasotocin and tyrosine hydroxylase. Brain Res. 1126, 91–101.
Martínez, P., Bouza, C., Hermida, M., Fernández, J., Toro, M.A., Vera, M., Pardo, B., Millán, A., Fernández, C., Vilas, R., Viñas, A., Sánchez, L., Felip, A., Piferrer, F., Ferreiro, I., Cabaleiro, S., 2009. Identification of the major sex-determining region of turbot (Scophthalmus maximus). Genetics 183, 1443–1452.
Martinez-Cerdeno, V., Noctor, S.C., Kriegstein, A.R., 2006. Estradiol stimulates progenitor cell division in the ventricular and subventricular zones of the embryonic neocortex, Eur. J. Neurosci. 24, 3475–3488.
Martin-Robichaud, D.J., 1993. Sexual differentiation and hormonal feminization of lumpfish (Cyclopterus lumpus Linnaeus). MSc Thesis, University of New Brunswick (Fredericton).
Martins, R.S., Deloffre, L.A., Mylonas, C.C., Power, D.M., Canário, A.V., 2007. Developmental expression of DAX1 in the European sea bass, Dicentrarchus labrax: lack of evidence for sexual dimorphism during sex differentiation. Reprod. Biol. Endocrinol. 30, 5─19.
Matsuda, M., Nagahama, Y., Shinomiya, A., Sato, T., Matsuda, C., Kobayashi, T., Morrey, C.E., Shibata, N., Asakawa, S., Shimizu, N., Hori, H., Hamaguchi, S., Sakaizumi, M., 2002. DMY is a Y-specific DM-domain gene required for male development in the medaka fish. Nature 417, 559–563.
Matsunaga, M., Ukena, K., Baulieu, E.E., Tsutsui, K., 2004. 7α-hydroxypregnenolone acts as a neuronal activator to stimulate locomotor activity of breeding newts by means of the dopaminergic system. Proc. Natl. Acad. Sci. USA 101, 17282–17287.
Matsunaga, M., Ukena, K., Tsutsui, K., 2001. Expression and localization of cytochrome P450 17α-hydroxylase/c17,20-lyase in the avian brain. Brain Res. 899, 112–122.
Matsuoka, M.P., van Nes, S., Andersen, Ø., Benfey, T.J., Reith, M., 2006. Real-time PCR
analysis of ovary- and brain-type aromatase gene expression during Atlantic halibut (Hippoglossus hippoglossus) development. Comp. Biochem. Physiol. B: Biochem. Mol. Biol. 144, 128–35.
Matthews, J., Gustafsson, J.A., 2003. Estrogen signaling: a subtle balance between ERα and ERβ. Mol. Interv. 3, 281–292.
Maxwell, B.L., McDonnell, D.P., Conneely, O.M., Schulz, T.Z., Greene, G.L., O'Malley, B.W., 1987. Structural organization and regulation of the chicken estrogen receptor. Mol Endocrinol. 1, 25–35.
McEwen, B.S., 1991. Steroid hormones are multifunctional messengers in the brain. Trends Endocrinol. Metab. 2, 62– 67.
Mellon, S.H., 1994. Neurosteroids: Biochemistry, modes of action and clinical relevance. J. Clin. Endocrinol. Metab. 78, 1003–1008.
Mellon, S.H., Deschepper, C.F., 1993. Neurosteroid biosynthesis: Genes for adrenal steroidogenic enzymes are expressed in the brain. Brain Res 629, 283–292.
Mensah-Nyagan, A.G., Feuilloley, M., Do-Rego, J.L., Marcual, A., Lange, C, Tonon, M.C., Pelletier, G., Vaudry, H., 1996. Localization of 17b-hydroxysteroid dehydrogenase and characterization of testosterone in the brain of the male frog. Proc. Natl. Acad. Sci. USA 93, 1423–1428.
Mensah-Nyagan, A.G., Feuilloley, M., Dupont, E., Do-Rego, J.-L., Leboulenger, F., Pelletier, G., Vaudry, H., 1994. Immunocytochemical localization and biological activity of 3β-hydroxysteroid dehydrogenase in the central nervous system of the frog, J. Neurosci. 14, 306–7318.
Menuet, A., Pellegrini, E., Brion, F., Gueguen, M.M., Anglade, I., Pakdel, F., Kah, O., 2005. Expression and estrogen-dependent regulation of the zebrafish brain aromatase gene. J. Comp. Neurol. 485, 304–320.
Menuet, A., Pellegrini, E., Anglade, I., Blaise, O., Laudet, V., Kah, O., Pakdel, F., 2002. Molecular characterization of three estrogen forms in zebrafish: binding characteristics, transactivation properties, and tissue distributions. Biol. Reprod. 66, 1881–1892.
Menuet, A., Anglade, I., Le Guevel, R., Pellegrini, E., Pakdel, F., Kah, O., 2003. Distribution of aromatase mRNA and protein in the brain and pituitary of female rainbow trout:
Comparison with estrogen receptor alpha. J. Comp. Neurol. 462, 180–193.
Mesiano, S., Coulter, C.L., Jaffe, R.B., 1993. Localization of cytochrome P450 cholesterol side-chain cleavage, cytochrome P450 17α-hydroxylase/17,20-lyase, and 3β-hydroxysteroid dehydrogenase isomerase steroidogenic enzymes in human and rhesus monkey fetal adrenal glands: reappraisal of functional zonation. J. Clin. Endocrinol. Metab. 77, 1184–1189.
Micevych, P.E., Chaban, V., Ogi, J., Lakhter, A., Lu, J.K.H., Sinchak, K., 2007. Estra-diol stimulates progesterone synthesis in hypothalamic astrocyte cultures. Endocrinology 148, 782–789.
Miura, S., Nakamura, S., Kobayashi, Y., Piferrer, F., Nakamura, M., 2008. Differentiation of ambisexual gonads and immunohistochemical localization of P450 cholesterol side-chain cleavage enzyme during gonadal sex differentiation in the protandrous anemonefish, Amphiprion clarkia. Comp. Biochem. Physiol B. Biochem. Mol. Biol. 1491, 29–37.
Moore, F.L., Wood, R.E., Boyd, S.K., 1992. Sex steroid and vasotocin interact in a female amphibian (Taricha granulosa) to elicit female-like egg-laying behavior of male-like courtship behavior. Horm. Behav. 26, 156–166.
Moore, F.L., Richardson, C., Lowry, C.A., 2000. Sexual dimorphism in numbers of vasotocin-immunoreactive neurons in brain areas associated with reproductive behaviors in the roughskin newt. Gen. Comp. Endocrinol. 117, 281─98.
Morris, J.A., Jordan, C.L., Breedlove, S.M., 2004. Sexual differentiation of the vertebrate nervous system. Nat. Neurosci. 7, 1034–1039.
Mouriec K, Pellegrini E, Anglade I, Menuet A, Adrio F, Thieulant ML, Pakdel F, Kah O. 2008.
Synthesis of estrogens in progenitor cells of adult fish brain: evolutive novelty or exaggeration of a more general mechanism implicating estrogens in neurogenesis? Brain Res Bull 75:274–280.
Mouriec, K., Lareyre, J.J., Tong, S.K., Le Page, Y., Vaillant, C., Pellegrini, E., Pakdel, F., Chung, B.C., Kah, O., Anglade, I., 2009. Early regulation of brain aromatase (cyp19a1b)
estrogen receptors during zebrafish development. Dev. Dyn. 238, 2641–2651.
Mullis, P.E., Yoshimura, N., Kuhlmann, B., 1997. Aromatase deficiency in a female who is
compound heterozygote for two new point mutations in the P450arom gene: impact of estrogens on hypergonadotropic hy-pogonadism, multicystic ovaries, and bone densitometry in childhood. J. Clin. Endocrinol. Metab. 82, 1739–1745.
Munchrath, L.A., Hofmann, H.A., 2010. Distribution of sex steroid hormone receptors in the brain of an African cichlid fish, Astatotilapia burtoni. J. Comp. Neurol. 518, 3302–3326.
Nagarajan, G., Tsai, Y.J., Chen, C.Y., Chang, C.F., 2011. Developmental expression of genes involved in neural estrogen biosynthesis and signaling in the brain of the orange-spotted grouper Epinephelus coioides during gonadal sex differentiation, J. Steroid Biochem. Mol. Biol. 127, 155–166.
Nakamura, M., Hourigan, T.F., Yamauchi, K., Nagahama, Y., 1989. Histological and ultrastructural evidence for the role of gonadal steroid hormones in sex change in the protogynous wrasse, Thalassoma duperrey. Environ. Biol. Fish 24, 117–136.
Nakamura, M., Kobayashi, T., Chang, X.T., Nagahama, Y., 1998. Gonadal sex differentiation in teleost fish. J. Exp. Zool. 281, 362–372.
Nanda, I., Kondo, M., Hornung, U., Asakawa, S., Winkler, C., Shimizu, A., Shan, Z., Haaf, T., Shimizu, N., Shima, A., Schmid, M., Schartl, M., 2002. A duplicated copy of DMRT1 in the sex-determining region of the Y chromosome of the medaka, Oryzias latipes. PNAS 99, 11778–11783.
Nelson, D.R., Kamataki, T., Waxman, D.J., Guengerich, F.P., Estabrook, R.W., 1993. The P450 superfamily: update on new sequences, gene mapping, accession numbers, early trivial names of enzymes, and nomenclature. DNA Cell Biol.12, 1–51.
Nelson, D.R., Koymans, L., Kamataki, T., Stegeman, J.J., Feyereisen, R., 1996. P450 superfamily: update on new sequences, gene mapping, accession numbers and nomenclature. Pharmacogenetics 6, 1–42.
Nocillado, J.N., Elizur, A., Avitan, A., Carrick, F., Levavi-Sivan, B., 2007. Cytochrome P450 aromatase in grey mullet: cDNA and promoter isolation; brain, pituitary and ovarian expression during puberty. Mol. Cell. Endocrinol. 263, 65–78.
Noctor, S.C., Flint, A.C., Weissman, T.A., Dammerman, R.S., Kriegstein, A.R., 2001. Neurons derived from radial glial cells establish radial units in neocortex. Nature 409, 714–720.
Nomura, M., McKenna, E., Korach, K.S., Pfaff, D.W., Ogawa, S., 2002. Estrogen receptor-β regulates transcript levels for oxytocin and arginine vasopressin in the hypothalamic paraventricular nucleus of male mice, Brain Res. Mol. Brain Res. 109, 84–94.
Okubo, K., Takeuchi, A., Chaube, R., Paul-Prasanth, B., Kanda, S., Oka, Y., Nagahama, Y., 2011. Sex differences in aromatase gene expression in the medaka brain. J. Neuroendocrinol. 23, 412–23.
Olivier, B., Chan. J.S., Snoeren, E.M., Olivier, J.D., Veening, J.G., Vinkers, C.H., Waldinger, M.D., Oosting, R.S., 2011. Differences in sexual behaviour in male and female rodents: role of serotonin. Curr. Top Behav. Neurosci. 8, 15─36.
Osawa, Y., Yoshida, N., Franckowiak, M., Kitawaki, J., 1987. Immunoaffinity purification of aromatase cytochrome P450 from human placental microsomes, metabolic switching from aromatization to 1β and 2β-monohydroxylation, and recognition of aromatase isoenzymes. Steroids 50, 11–28.
Paech, K., Webb, P., Kuiper, G.G., Nilsson, S., Gustafsson, J., Kushner, P.J., Scanlan, T.S., 1997. Differential ligand activation of estrogen receptors ERα and ERβ at AP1 sites. Science 277, 1508–1510.
Pakdel, F., Le Guellec, C., Vaillant, C., Le Roux, M.G., Valotaire, Y., 1989. Identification and estrogen induction of two estrogen receptors (ER) messenger ribonucleic acids in the rainbow trout liver: sequence homology with other ERs. Mol. Endocrinol. 3, 44–51.
Pandian, T.S., Sheela, S., 1995. Hormonal induction of sex reversal in fish. Aquaculture 138,1-22
Pang, Y., Dong, J., Thomas, P., 2008. Estrogen signaling characteristics of Atlantic croaker G protein-coupled receptor 30 (GPR30) and evidence it is involved in maintenance of oocyte meiotic arrest. Endocrinology 149, 3410–3426.
Pang, Y., Thomas, P., 2010. Role of G protein-coupled estrogen receptor 1, GPER, in inhibition of oocyte maturation by endogenous estrogens in zebrafish. Dev. Biol. 342, 194–206.
Panzica, G.C., Castagna, C., Viglietti-Panzica, C., Russo, C., Tlemcani, O., Balthazart, J., 1998. Organizational effects of estrogens on brain vasotocin and sexual behavior in quail. J. Neurobiol. 37, 684–699.
Parker, K.L., Schimmer, B.P., 1997. Steroidogenic factor 1: a key determinant of endocrine
development and function. Endocr. Rev. 18, 361–377.
Pasmanik, M., Callard, G.V., 1985. Aromatase and 5α-reductase in the teleost brain, spinal cord, and pituitary gland. Gen Comp Endocrinol 60, 244–251.
Pasqualini, J.R., Gelly, C., Nguyen, B.L., Vella, C., 1989. Importance of estrogen sulfates in breast cancer. J. Steroid Biochem.34, 155–63.
Payne, A.H., Hales, D.B., 2004. Overview of steroidogenic enzymes in the pathway from cholesterol to active steroid hormones. Endocr. Rev. 25, 947–970.
Pellegrini, E., Mouriec, K., Anglade, I., Menuet, A., Le Page, Y., Gueguen, M.M., Marmignon, M.H., Brion, F., Pakdel, F., Kah, O., 2007. Identification of aromatase-positive radial glial cells as progenitor cells in the ventricular layer of the forebrain in zebrafish. J. Comp. Neurol. 501, 150–167.
Pellegrini, E., Menue,t A., Lethimonier, C., Adrio, F., Gueguen, M.M., Tascon, C., Anglade, I., Pakdel, F., Kah, O., 2005. Relationships between aromatase and estrogen receptors in the brain of teleost fish. Gen. Comp. Endocrinol. 142, 60–66.
Peterson, R.S., Lee, D.W., Fernando, G., Schlinger, B.A., 2004. Radial glia expresses aromatase in the injured zebrafinch brain. J. Comp. Neurol. 475, 261–269.
Pietras, R.J., Szego, C.M., 1977. Specific binding sites for oestrogen at the outer surfaces of
isolated endometrial cells. Nature 265, 69–72.
Pilon, N., Behdjani, R., Daneau, I., Lussier, J.G., Silversides, D.W., 1998. Porcine steroidogenic factor-1 gene (pSF-1) expression and analysis of embryonic pig gonads during sexual differentiation. Endocrinology 139, 3803–3812.
Place, A.R., Lang, J., Gavasso, S., Jeyasuriya, P., 2001. Expression of P450arom in Malachemys terrapin and Chelydra serpentina: a tale of two sites. J. Exp. Zool. 290, 673–690.
Plassart-Schiess, E., Baulieu, E.E., 2001. Neurosteroids: recent findings. Brain Res. Brain Res. Rev. 37, 133–140.
Prossnitz, E.R., Arterburn, J.B., Smith, H.O., Oprea, T.I., Sklar, L.A., Hathaway, H.J., 2008. Estrogen signaling through the transmembrane G protein-coupled receptor GPR30. Annu.
Rev. Physiol. 70, 165–190.
Quigley, C.A., De Bellis, A., Marschke, K.B., el-Awady, M.K., Wilson, E.M., French, F.S.,
1995. Androgen receptor defects: historical, clinical, and molecular perspectives. Endocr. Rev. 16, 271–321.
Ramallo, M.R., Grober, M., Cánepa, M.M., Morandini, L., Pandolfi, M., 2012. Arginine-vasotocin expression and participation in reproduction and social behavior in males of the cichlid fish Cichlasoma dimerus. Gen. Comp. Endocrinol. 179, 221–231.
Rasheeda, M.K., Sridevi, P., Senthilkumaran, B., 2010. Cytochrome P450 aromatases: Impact on gonadal development, recrudescence and effect of hCG in the catfish, Clarias gariepinus. Gen. Comp. Endocrinol. 167, 234–245.
Reinboth, R., Brusle´ Sicard, S., 1997. Histological and ultrastructural studies on the effects of hCG on sex inversion in the protogynous teleost Coris julis. J. Fish Biol. 51, 738–749.
Remage-Healey, L., Bass, A.H., 2004. Rapid, hierarchical modulation of vocal patterning by steroid hormones. J. Neurosci. 24, 5892–5900.
Rhe´aume, E., Simard, J., Morel, Y., Mebarki, F., Zachmann, M., Forest, M.G., New, M.I.,
Labrie, F., 1992. Congenital adrenal hyperplasia due to point mutations in the type II 3β-hydroxysteroid dehydrogenase gene. Nat. Genet. 1, 239–245.
Sadovsky, Y., Crawford PA, Woodson K.G., Polish, J.A., Clements, M.A., Tourtellotte, L.M., Simburger, K., Milbrandt, J., 1995. Mice deficient in the or-phan receptor steroidogenic factor 1 lack adrenal glands and gonads but express P450 side-chain-cleavage enzyme in the placenta and have normal embryonic serum levels of corticosteroids. Proc Natl Acad Sci USA 92, 10939–10943.
Sakamoto, H., Ukena, K., Tsutsui, K., 2001. Activity and localization of 3β-hydroxysteroid dehydrogenase/Δ5–Δ4-isomerase in the zebrafish central nervous system. J. Comp. Neurol. 439, 291–305.
Sakurai, N., Maruo, K., Haraguchi, S., Uno, Y., Oshima, Y., Tsutsui, K., Matsuda, Y., Do Rego, J.L., Pelletier, G., Vaudry, H., Nakamura, M., 2008. Immunohistochemical detection and biological activities of CYP17 (P450c17) in the indifferent gonad of the frog Rana rugosa. J. Steroid. Biochem. Mol. Biol. 112, 5–12.
Sanne, J.L., Krueger, K.E., 1995. Expression of cytochrome P450 side-chain cleavage enzyme
and 3β-hydroxysteroid dehydrogenase in the rat central nervous system: a study by
polymerase chain reaction and in situ hybridization. J. Neurochem. 65, 528–536.
Saville, B., Wormke, M., Wang, F., Nguyen, T., Enmark, E., Kuiper, G., Gustafsson, J.A., Safe, S., 2000. Ligand-, cell-, and estrogen receptor subtype (α/β)-dependent activation at GC-rich (Sp1) promoter elements. J. Biol. Chem. 275, 5379–5387.
Semsar, K., Kandel, F.L.M., Godwin, J., 2001. Manipulations of the AVT system shift social status and related courtship and aggressive behavior in the bluehead wrasse. Horm. Behav. 40, 21–31.
Shao, C.W., Chen, S.L., Scheuring, C.F., Xu, J.Y., Sha, Z.X., Dong, X.L., Zhang, H.B., 2010. Construction of two BAC libraries from half-smooth tongue sole Cynoglossus semilaevis and identification of clones containing candidate sex-determination genes. Mar. Biotechnol. 12, 558–568.
Shen, W.H., Moore, C.C., Ikeda, Y., Parker, K.L., Ingraham, H.A., 1994. Nuclear receptor
steroidogenic factor 1 regulates the mullerian inhibiting substance gene: a link to the sex determination cascade. Cell 77, 651–661.
Shinoda, K., Lei, H., Yoshii, H., Nomura, M., Nagano, M., Shiba, H., Sasaki, H., Osawa, Y., Ninomiya, Y., Niwa, O., Morohashi, K., Li, E., 1995. Developmental defects of the ventromedial hypothalamic nucleus and pituitary gonadotroph in the Ftz-F1 disrupted mice. Dev. Dyn. 204, 22–29.
Shozu, M., Sebastian, S., Takayama, K., 2003. Estrogen excess associated with novel gain-of-function mutations affecting the aromatase gene. N. Engl. J. Med. 348, 1855–1865.
Simard J., Moisan A.M., Morel Y., 2002. "Congenital adrenal hyperplasia due to 3β-hydroxysteroid dehydrogenase/Δ5-Δ4 isomerase deficiency". Semin. Reprod. Med. 20 (3): 255–76.
Simard, J., Sanchez, R., Durocher, F., Rhe´aume, E., Turgeon, C., Labrie, Y., Luu-The, V., Mebarki, F., Morel, Y., Launoit, Y., Labrie, F., 1995. Structure-function relationships and molecular genetics of the 3β-hydroxysteroid dehydrogenase gene family. J. Steroid Biochem. Mol. Biol. 55, 489–505.
Simerly, R.B., 2002. Wired for reproduction: organization and development of sexually dimorphic circuits in the mammalian forebrain. Annu. Rev. Neurosci. 25, 507–536.
Simpson, E.R., Mahendroo, M.S., Means, G.D., Kilgore, M.W., Hinshelwood, M.M., Graham-Lorence, S., Amarneh, B., Ito, Y., Fisher, C.R., Michael, M.D., Mendelson, C.R., Bulun, S.E., 1994. Aromatase cytochrome P450, the enzyme responsible for estrogen biosynthesis. Endocr. Rev. 15, 342–355.
Sinchak, K., Mills, R.H., Tao, L., LaPolt, P., Lu, J.K., Micevych, P., 2003. Estrogen induces de novo progesterone synthesis in astrocytes. Dev. Neurosci. 25, 343–348.
Socorro, S., Power, D.M., Olsson, P.E., Canario, A.V., 2000. Two estrogen receptors expressed in the teleost fish, Sparus aurata: cDNA cloning, characterization and tissue distribution. J. Endocrinol. 166, 293–306.
Sperry, T.S., Thomas, P., 1999a. Identification of two nuclear androgen receptors in kelp bass
(Paralabrax clathratus) and their binding affinities for xenobiotics: comparison with Atlantic croaker (Micropogonias undulatus) androgen receptors. Biol. Reprod. 61, 1152–1161.
Sperry, T.S., Thomas, P., 1999b. Characterization of two nuclear androgen receptors in Atlantic croaker: comparison of their biochemical properties and binding specificities. Endocrinology 140, 1602–1611.
Sridevi, P., Chaitanya, R.K., Dutta-Gupta, A., Senthilkumaran, B., 2012. FTZ-F1 and FOXL2 up-regulate catfish brain aromatase gene transcription by specific binding to the promoter motifs. Biochim. Biophys. Acta. 1819, 57–66.
St. Mary, C.M., 1998. Characteristic gonad structure in the gobiid genus Lythrypnus with comparisons to other hermaphroditic gobies. Copeia 1998, 720–724.
Strobl-Mazzulla, P.H., Lethimonier, C., Gueguen, M.M., Karube, M., Fernandino, J.I., Yoshizaki, G., Patiño, R., Strüssmann, C.A., Kah, O., Somoza, G.M., 2008. Brain aromatase (Cyp19A2) and estrogen receptors, in larvae and adult pejerrey fish Odontesthes bonariensis: Neuroanatomical and functional relations. Gen. Comp. Endocrinol. 158, 191–201.
Sugawara, T., Holt, J.A., Kiriakidou, M., Strauss 3rd, J.F., 1996. Steroidogenic factor 1-
dependent promoter activity of the human steroidogenic acute regulatory protein (StAR) gene. Biochemistry 35, 9052–9059.
Sugita, J., Takase, M., Nakamura, M., 2001. Expression of Dax-1 during gonadal development
of the frog. Gene 280, 67–74.
Swain, A., Lovell-Badge, R., 1997. A molecular approach to sex determination in mammals. Acta Paediatr. 423, 46–49.
Tajima, T., Fujieda, K., Kouda, N., Nakae, J., Miller, W.L., 2001. Heterozygous mutation in the cholesterol side chain cleavage enzyme (p450scc) gene in a patient with 46,XY sex reversal and adrenal insufficiency. J. Clin. Endocrinol. Metab. 86, 3820–3825.
Takeo, J., Yamashita, S., 1999. Two distinct isoforms of cDNA. Encoding rainbow trout androgen receptors. J. Biochem. 274, 5674–5680.
Tan, F.L., Lolait, S.J., Brownstein, M.J., Saito, N., MacLeod, V., Baeyens, D.A., Mayeux, P.R., Jones, S.M., Cornett, L.E., 2000. Molecular cloning and functional characterization of a vasotocin receptor subtype that is expressed in the shell gland and brain of the domestic chicken. Biol. Reprod. 62, 8–15.
Tan, N.S., Lam, T.J., Ding, J.L., 1996. The first contiguous estrogen receptor gene from a fish, Oreochromis aureus: evidence for multiple transcripts. Mol. Cell Endocrinol. 120, 177–192.
Tanaka, M., Fukada, S., Matsuyamma, M., Nagahama Y., 1995. Structure and promoter analysis of the cytochrome P450 aromatase gene of the teleost fish, Medaka Oryzias latipes. J. Biochem. 177, 719–725.
Tang, B., Hu, W., Hao, J., Zhu, Z., 2010. Developmental expression of steroidogenic factor-1, cyp19a1a and cyp19a1b from common carp (Cyprinus carpio). Gen. Comp. Endocrinol. 167, 408–416.
Tchoudakova, A., Callard, G.V., 1998. Identification of multiple CYP19 genes encoding different cytochrome P450 aro-matase isozymes in brain and ovary. Endocrinology 139, 2179–2189.
Tchoudakova, A., Kishida, M., Wood, E., Callard, G.V., 2001. Promoter characteristics of two
cyp19 genes differentially expressed in the brain and ovary of teleost fish. J. Steroid. Biochem. Mol. Biol. 78, 427–439.
Thibonnier, M., Auzan, C., Madhun, Z., Wilkins, P., Berti-Mattera, L., Clauser, E., 1994.
Molecular cloning, sequencing, and functional expression of a cDNA encoding the human V1a vasopressin receptor. J. Biol. Chem. 269, 3304–3310.
Tobet, S.A., Hanna, I.K., 1997. Ontogeny of sex differences in the mammalian hypothalamus and preoptic area. Cell Mol. Neurobiol. 17, 565–601.
Todo, T., Ikeuchi, T., Kobayashi, T., Nagahama, Y., 1999. Fish androgen receptor: cDNAcloning, steroid activationof transcription in trans-fectedmammalian cells, and tissue mRNA levels. Biochem. Biophys. Res. Commun. 254, 378–383.
Todo, T., Adachi, S., Yamauchi, K., 1996. Molecular cloning and characterization of Japanese eel estrogen receptor cDNA. Mol. Cell Endocrinol. 119, 37–45.
Toft, D., Gorski, J., 1966. A receptor molecule for estrogens: isolation from the rat uterus and preliminary characterization. Proc. Natl. Acad. Sci. USA 55, 1574–1581.
Tomy, S., Wu, G.C., Huang, H.R., Chang, C.F., 2009. Age-dependent differential expression of genes involved in steroid sig-nalling pathway in the brain of protandrous black porgy, Acanthopagrus schlegeli. Dev. Neurobiol. 69, 299–313.
Tomy, S., Wu, G.C., Huang, H.R., Dufour, S., Chang, C.F., 2007. Developmental expression of key steroidogenic enzymes in the brain of protandrous black porgy fish, Acanthopagrus schlegeli. J. Neuroendocrinol. 19, 643–655.
Tong, S.K., Chiang, E.F.L., Hsiao, P.H., Chung, B.C., 2001. Phylogeny, expression and enzyme activity of zebrafish cyp19 (P450 aromatase) genes. J. Steroid Biochem. Mol. Biol. 79, 299–303.
Tong, S.K., Chung, B.C., 2003. Analysis of zebrafish cyp19 promoters. J. Steroid Biochem. Mol. Biol. 86, 381–386.
Touhata, K., Kinoshita, M., Toyohara, H., Sakaguchi, M., 1998. Sequence and expression of a cDNA encoding the red seabream estrogen receptor. Fisheries Science 64,131–135.
Trant, J.M., Gavasso, S., Ackers, J., Chung, B.-C., Place, A.R., 2001. Developmental expression
of cytochrome P450 aromatase genes (CYP19a and CYP19b) in zebrafish fry (Danio rerio). J. Exp. Zool. 290, 475–483.
Tremblay, G.B., Tremblay, A., Copeland, N.G., Gilbert, D.J., Jenkins, N.A., Labrie, F., Giguere, V., 1997. Cloning, chromosomal localization, and functional analysis of the murine
estrogen receptor β. Mol. Endocrinol. 11,353–365.
Tsutsui, K., Ukena, K., Takase, M., Kohchi, C., Lea, R.W., 1999. Neurosteroid biosynthesis in vertebrate brains. Comp. Biochem. Physiol. C. Pharmacol. Toxicol. Endocrinol. 124, 121–129.
Tzukerman, M.T., Esty, A., Santiso-Mere, D., Danielian, P., Parker, M.G., Stein, R.B., Pike, J.W., McDonnell, D.P., 1994. Human estrogen receptor transactivational capacity is determined by both cellular and promoter context and mediated by two functionally distinct intramolecular regions. Mol. Endocrinol. 8, 21–30.
Ukena, K., Kohchi, C., Tsutsui, K., 1999. Expression and activity of 3β-hydroxysteroid dehydrogenase/Δ5-Δ4-isomerase in the rat Purkinje neuron during neonatal life. Endocrinology 140, 805–813.
Ukena, K., Usui, M., Kohchi, C., Tsutsui, K., 1998. Cytochrome P450 side-chain cleavage enzyme in the cerebellar Purkinje neuron and its neonatal change in rats. Endocrinology 139, 137–147.
Urushitani, H., Nakai, M., Inanaga, H., Shimohigashi, Y., Shimizu, A., Katsu, Y., Iguchi, T., 2003. Cloning and characterization of estrogen receptor α in mummichog, Fundulus heteroclitus. Mol. Cell Endocrinol. 203, 41–50.
Valle, L.D., Ramina, A., Vianello, S., Belvedere, P., Colombo, L., 2002. Cloning of two mRNA variants of brain aromatase cytochrome P450 in rainbow trout (Oncorhynchus mykiss Walbaum), J. Steroid Biochem. Mol. Biol. 82, 19–32.
van Nes, S., Andersen, Ø., 2006. Temperature effects on sex determination and ontogenetic gene expression of the aromatases cyp19a and cyp19b, and the estrogen receptors esr1 and esr2 in Atlantic halibut (Hippoglossus hippoglossus). Mol. Reprod. Dev. 73, 1481–1490.
Vizziano, D., Randuineau, G., Baron, D., Cauty, C., Guiguen, Y., 2007. Characterization of early molecular sex differenti-ation in rainbow trout, Oncorhynchus mykiss. Dev. Dyn. 236, 2198–2206.
Vizziano-Cantonnet, D., Anglade, I., Pellegrini, E., Gueguen, M.M., Fostier, A., Guiguen, Y., Kah, O., 2011. Sexual dimorphism in the brain aromatase expression and activity, and in the central expression of other steroidogenic enzymes during the period of sex differentiation in monosex rainbow trout populations. Gen. Comp. Endocrinol. 170, 346–355.
von Hofsten, J., Olsson, P.E., 2005. Zebrafish sex determination and differentiation: involvement of FTZ-F1 genes. Reprod. Biol. Endocrinol. 3, 63.
Watanabe, M., Tanaka, M., Kobayashi, D., Yoshiura, Y., Oba, Y., Nagahama, Y., 1999. Medaka (Oryzias latipes) FTZ-F1 potentially regulates the transcription of P450 aromatase in ovarian follicles: cDNA cloning and functional characterization. Mol. Cell Endocrinol. 149, 221–228.
Wehrenberg, U., Prange-Kiel, J., Rune, G.M., 2001. Steroidogenic factor-1 expression in marmoset and rat hippocampus: co-localization with StAR and aromatase. J. Neurochem. 76, 1879–1886.
Wu, G.C, Tomy, S., Nakamura, M., Chang, C.F., 2008. Dual roles of cyp19a1a in gonadal sex differentiation and development in the protandrous black porgy, Acanthopagrus schlegeli. Biol. Reprod. 79, 1111–1120.
Wu, M.V., Manoli, D.S., Fraser, E.J., Coats, J.K., Tollkuhn, J., Honda, S., Harada, N., Shah, N.M., 2009. Estrogen masculinizes neural pathways and sex-specific behaviors. Cell 139, 61–72.
Yamamoto, T., 1958. Artificial induction of functional sex-reversal in genotypic females of the medaka (Oryzias latipes). J. Exp. Zool. 137, 227–260.
Yamamoto, T., 1968. Effects of 17-α-hydroxyprogesterone and androstenedione upon sex differentiation in the medaka, Oryzias latipes. Gen. Comp. Endocrinol. 10, 8–13.
Yamamoto, T., 1969. Sex differentiation. In: Hoar, WS, Randall, DJ (Eds.), Fish Physiology, vol. 3. Academic Press, New York, pp. 117–175.
Yanase T, Simpson ER, Waterman MR., 1991. 17α-hydroxylase/17,20 lyase deficiency: from clinical investigation to molecular definition. Endocr. Rev. 12:91–108.
Yeh, S., Tsai, M.Y., Xu, Q., Mu, X.M., Lardy, H., Huang, K.E., Lin, H., Yeh, S.D., Altuwaijri, S., Zhou, X., Xing, L., Boyce, B.F., 2002. Generation and characterization of androgen receptor knockout (ARKO) mice: an in vivo model for the study of androgen functions in selective tissues. Proc. Natl. Acad. Sci. USA 99, 13498–13503.
Yoshiura, Y., Senthilkumaran, B., Watanabe, M., Oba, Y., Kobayashi, T., Nagahama, Y., 2003. Synergistic expression of Ad4BP/SF-1 and cytochrome P-450 aromatase (ovarian type) in the ovary of Nile tilapia, Oreochromis niloticus, during vitellogenesis suggests transcriptional interaction. Biol. Reprod. 68, 1545–1553.
Zhang, Y., Zhang, W., Yang, H., Zhou, W., Hu, C., Zhang, L., 2008. Two cytochrome P450 aromatase genes in the hermaphrodite ricefield eel, Monopterus albus: mRNA expression during ovarian development and sex change, J. Endocrinol. 199, 317–331.
Zhang, Y., Zhang, W., Zhang, L., Zhu, T., Tian, J., Li, X., Lin, H., 2004. Two distinct cytochrome P450 aromatases in the orange-spotted grouper (Epinephelus coioides): cDNA cloning and differential mRNA expression, J. Steroid Biochem. Mol. Biol. 92, 39–50.
Zhao, J., Mak, P., Tchoudakova, A., Callard, G., Chen, S., 2001. Different catalytic properties and inhibitor responses of the goldfish brain and ovary aromatase isozymes. Gen. Comp. Endocrinol. 123, 180–191.
Zhu, P., Zhang, Y., Zhuo, Q., Lu, D., Huang, J., Liu, X., Lin, H., 2008. Discovery of four estrogen receptors and their expression profiles during testis recrudescence in male Spinibarbus denticulatus. Gen. Comp. Endocrinol. 156, 265–276.
Zupanc, G.K., 2001. Adult neurogenesis and neuronal regeneration in the central nervous system of teleost fish. Brain Behav. Evol. 58, 250–275.