B. Adolf, P. Chapouton, C. S. Lam, S. Topp, B. Tannhauser et al., Conserved and acquired features of adult neurogenesis in the zebrafish telencephalon, Dev Biol, vol.295, pp.278-293, 2006.

A. Alvarez-buylla and D. A. Lim, For the long run: maintaining germinal niches in the adult brain, Neuron, vol.41, pp.683-686, 2004.

A. Alvarez-buylla, D. R. Buskirk, and F. Nottebohm, Monoclonal antibody reveals radial glia in adult avian brain, J Comp Neurol, vol.264, pp.159-170, 1987.

I. Anglade, A. M. Zandbergen, and O. Kah, Origin of the pituitary innervation in the goldfish, Cell Tissue Res, vol.273, pp.345-355, 1993.

M. Arochena, R. Anadon, and S. M. Diaz-regueira, Development of vimentin and glial fibrillary acidic protein immunoreactivities in the brain of gray mullet (Chelon labrosus), an advanced teleost, J Comp Neurol, vol.469, pp.413-436, 2004.

I. Azcoitia, A. Sierra, and L. M. Garcia-segura, Localization of estrogen receptor beta-immunoreactivity in astrocytes of the adult rat brain, Glia, vol.26, pp.260-267, 1999.

I. Azcoitia, A. Sierra, S. Veiga, and L. M. Garcia-segura, Aromatase expression by reactive astroglia is neuroprotective, Ann N Y Acad Sci, vol.1007, pp.298-305, 2003.

I. Azcoitia, A. Sierra, S. Veiga, and L. M. Garcia-segura, Brain steroidogenesis: emerging therapeutic strategies to prevent neurodegeneration, 2005.

, J Neural Transm, vol.112, pp.171-176

J. Balthazart and G. F. Ball, New insights into the regulation and function of brain estrogen synthase (aromatase), Trends Neurosci, vol.21, pp.243-249, 1998.

P. I. Bang, W. F. Sewell, and J. J. Malicki, Morphology and cell type heterogeneities of the inner ear epithelia in adult and juvenile zebrafish (Danio rerio), J Comp Neurol, vol.438, pp.173-190, 2001.

C. Behl, Oestrogen as a neuroprotective hormone, Nat Rev Neurosci, vol.3, pp.433-442, 2002.

C. Behl, Oxidative stress in Alzheimer's disease: implications for prevention and therapy, Subcell Biochem, vol.38, pp.65-78, 2005.

M. S. Brainard and A. J. Doupe, Postlearning consolidation of birdsong: stabilizing effects of age and anterior forebrain lesions, J Neurosci, vol.21, pp.2501-2517, 2001.

A. Carleton, C. Rochefort, J. Morante-oria, D. Desmaisons, J. D. Vincent et al., New insights into the regulation and function of brain estrogen synthase (aromatase), Trends Neurosci, vol.21, issue.6, pp.243-249, 1998.

P. L. Bardet, B. Horard, M. Robinson-rechavi, V. Laudet, and J. M. Vanacker, Characterization of oestrogen receptors in zebrafish (Danio rerio), J Mol Endocrinol, vol.28, issue.3, pp.153-163, 2002.

M. Blazquez and F. Piferrer, 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, vol.219, issue.1-2, pp.83-94, 2004.

G. V. Callard and A. Tchoudakova, Evolutionary and functional significance of two CYP19 genes differentially expressed in brain and ovary of goldfish, J Steroid Biochem Mol Biol, vol.61, issue.3-6, pp.387-392, 1997.

G. V. Callard, A. V. Tchoudakova, M. Kishida, and W. E. , Differential tissue distribution, developmental programming, estrogen regulation and promoter characteristics of cyp19 genes in teleost fish, J Steroid Biochem Mol Biol, vol.79, issue.1-5, pp.305-314, 2001.

E. G. De-jesus and T. Hirano, Changes in whole body concentrations of cortisol, thyroid hormones, and sex steroids during early development of the chum salmon, Oncorhynchus keta, Gen Comp Endocrinol, vol.85, issue.1, pp.55-61, 1992.

A. L. Filby and C. R. Tyler, Molecular characterization of estrogen receptors 1, 2a, and 2b and their tissue and ontogenic expression profiles in fathead minnow (Pimephales promelas), Biol Reprod, vol.73, issue.4, pp.648-662, 2005.

A. L. Filby and C. R. Tyler, Appropriate 'housekeeping' genes for use in expression profiling the effects of environmental estrogens in fish, BMC Mol Biol, vol.8, p.10, 2007.

G. Flouriot, F. Pakdel, and Y. Valotaire, Transcriptional and post-transcriptional regulation of rainbow trout estrogen receptor and vitellogenin gene expression, Mol Cell Endocrinol, vol.124, issue.1-2, pp.173-183, 1996.

P. M. Forlano, D. L. Deitcher, D. A. Myers, and A. H. Bass, Anatomical distribution and cellular basis for high levels of aromatase activity in the brain of teleost fish: aromatase enzyme and mRNA expression identify glia as source, J Neurosci, vol.21, issue.22, pp.8943-8955, 2001.

D. Gelinas, G. A. Pitoc, and G. V. Callard, Isolation of a goldfish brain cytochrome P450 aromatase cDNA: mRNA expression during the seasonal cycle and after steroid treatment, Mol Cell Endocrinol, vol.138, issue.1-2, pp.81-93, 1998.

S. Halm, N. Pounds, S. Maddix, M. Rand-weaver, J. P. Sumpter et al., Exposure to exogenous 17beta-oestradiol disrupts p450aromB mRNA expression in the brain and gonad of adult fathead minnows, 2002.

, Aquat Toxicol, vol.60, issue.3-4, pp.285-299

M. B. Hawkins, J. W. Thornton, D. Crews, J. K. Skipper, A. Dotte et al., Identification of a third distinct estrogen receptor and reclassification of estrogen receptors in teleosts, Proc Natl Acad Sci U S A, vol.97, issue.20, pp.10751-10756, 2000.

G. A. Hines, L. R. Boots, T. Wibbels, and S. A. Watts, Steroid levels and steroid metabolism in relation to early gonadal development in the tilapia Oreochromis niloticus (Teleostei: cyprinoidei), Gen Comp Endocrinol, vol.114, issue.2, pp.235-248, 1999.

E. Kallivretaki, R. I. Eggen, S. C. Neuhauss, O. Kah, and H. Segner, The zebrafish, brainspecific, aromatase cyp19a2 is neither expressed nor distributed in a sexually dimorphic manner during sexual differentiation, Dev Dyn, vol.236, issue.11, pp.3155-3166, 2007.
URL : https://hal.archives-ouvertes.fr/hal-00180993

C. B. Kimmel, W. W. Ballard, S. R. Kimmel, B. Ullmann, and T. F. Schilling, Stages of embryonic development of the zebrafish, Dev Dyn, vol.203, issue.3, pp.253-310, 1995.

M. Kishida, M. Mclellan, J. A. Miranda, and G. V. Callard, Estrogen and xenoestrogens upregulate the brain aromatase isoform (P450aromB) and perturb markers of early development in zebrafish (Danio rerio), Comp Biochem Physiol B Biochem Mol Biol, vol.129, issue.2-3, pp.261-268, 2001.

C. S. Lassiter, B. Kelley, and E. Linney, Genomic structure and embryonic expression of estrogen receptor beta a (ERbetaa) in zebrafish (Danio rerio), Gene, vol.299, issue.1-2, pp.141-151, 2002.

L. Page, Y. Menuet, A. Kah, O. Pakdel, and F. , Characterization of a cis-acting element involved in cell-specific expression of the zebrafish brain aromatase gene, Mol, 2008.
URL : https://hal.archives-ouvertes.fr/hal-00316100

L. Page, Y. Scholze, M. Kah, O. Pakdel, and F. , Assessment of xenoestrogens using three distinct estrogen receptors and the zebrafish brain aromatase gene in a highly responsive glial cell system, Environ Health Perspect, vol.114, issue.5, pp.752-758, 2006.
URL : https://hal.archives-ouvertes.fr/hal-00068400

K. E. Marsh, L. M. Creutz, M. B. Hawkins, and J. Godwin, Aromatase immunoreactivity in the bluehead wrasse brain, Thalassoma bifasciatum: immunolocalization and coregionalization with arginine vasotocin and tyrosine hydroxylase, Brain Res, vol.1126, issue.1, pp.91-101, 2006.

V. Martinez-cerdeno, S. C. Noctor, and A. R. Kriegstein, Estradiol stimulates progenitor cell division in the ventricular and subventricular zones of the embryonic neocortex, Eur J Neurosci, vol.24, issue.12, pp.3475-3488, 2006.

A. Menuet, I. Anglade, L. Guevel, R. Pellegrini, E. Pakdel et al., Distribution of aromatase mRNA and protein in the brain and pituitary of female rainbow trout: Comparison with estrogen receptor alpha, J Comp Neurol, vol.462, issue.2, pp.180-193, 2003.
URL : https://hal.archives-ouvertes.fr/hal-00295588

A. Menuet, L. Page, Y. Torres, O. Kern, L. Kah et al., Analysis of the estrogen regulation of the zebrafish estrogen receptor (ER) reveals distinct effects of ERalpha, ERbeta1 and ERbeta2, J Mol Endocrinol, vol.32, issue.3, pp.975-986, 2004.
URL : https://hal.archives-ouvertes.fr/hal-00295170

A. Menuet, E. Pellegrini, I. Anglade, O. Blaise, V. Laudet et al., Molecular characterization of three estrogen receptor forms in zebrafish: binding characteristics, transactivation properties, and tissue distributions, Biol Reprod, vol.66, issue.6, pp.1881-1892, 2002.

A. Menuet, E. Pellegrini, F. Brion, M. M. Gueguen, I. Anglade et al., Expression and estrogen-dependent regulation of the zebrafish brain aromatase gene, J Comp Neurol, vol.485, issue.4, pp.304-320, 2005.
URL : https://hal.archives-ouvertes.fr/hal-00294769

F. Pakdel, S. Feon, L. Gac, F. , L. Menn et al., In vivo estrogen induction of hepatic estrogen receptor mRNA and correlation with vitellogenin mRNA in rainbow trout, Mol Cell Endocrinol, vol.75, issue.3, pp.205-212, 1991.
URL : https://hal.archives-ouvertes.fr/hal-02715541

M. Pasmanik and G. V. Callard, Aromatase and 5 alpha-reductase in the teleost brain, spinal cord, and pituitary gland, Gen Comp Endocrinol, vol.60, issue.2, pp.244-251, 1985.

E. Pellegrini, K. Mouriec, I. Anglade, A. Menuet, L. Page et al., Identification of aromatase-positive radial glial cells as progenitor cells in the ventricular layer of the forebrain in zebrafish, J Comp Neurol, vol.501, issue.1, pp.150-167, 2007.
URL : https://hal.archives-ouvertes.fr/hal-00096389

R. S. Peterson, D. W. Lee, G. Fernando, and B. A. Schlinger, Radial glia express aromatase in the injured zebra finch brain, J Comp Neurol, vol.475, issue.2, pp.261-269, 2004.

S. Rozen and H. Skaletsky, Primer3 on the WWW for general users and for biologist programmers, Methods in molecular biology, vol.132, pp.365-386, 2000.

T. Sabo-attwood, K. J. Kroll, and N. D. Denslow, Differential expression of largemouth bass (Micropterus salmoides) estrogen receptor isotypes alpha, beta, and gamma by estradiol, Mol Cell Endocrinol, vol.218, issue.1-2, pp.107-118, 2004.

S. J. Sawyer, K. A. Gerstner, and G. V. Callard, Real-time PCR analysis of cytochrome P450 aromatase expression in zebrafish: gene specific tissue distribution, sex differences, developmental programming, and estrogen regulation, Gen Comp Endocrinol, vol.147, issue.2, pp.108-117, 2006.

D. Shin, C. H. Shin, J. Tucker, E. A. Ober, F. Rentzsch et al., Bmp and Fgf signaling are essential for liver specification in zebrafish, Development, vol.134, issue.11, pp.2041-2050, 2007.

D. Steinke, S. Hoegg, H. Brinkmann, and A. Meyer, Three rounds (1R/2R/3R) of genome duplications and the evolution of the glycolytic pathway in vertebrates, BMC Biol, vol.4, p.16, 2006.

P. H. Strobl-mazzulla, C. Lethimonier, M. M. Gueguen, M. Karube, J. I. Fernandino et al., Brain aromatase (Cyp19A2) and estrogen receptors, in larvae and adult pejerrey fish Odontesthes bonariensis: Neuroanatomical and functional relations, Gen Comp Endocrinol, vol.158, issue.2, pp.191-201, 2008.
URL : https://hal.archives-ouvertes.fr/hal-00319804

P. H. Strobl-mazzulla, N. P. Moncaut, G. C. Lopez, L. A. Miranda, A. V. Canario et al., Brain aromatase from pejerrey fish (Odontesthes bonariensis): cDNA cloning, tissue expression, and immunohistochemical localization, Gen Comp Endocrinol, vol.143, issue.1, pp.21-32, 2005.

J. R. Tata and D. F. Smith, Vitellogenesis: a versatile model for hormonal regulation of gene expression, Recent progress in hormone research, vol.35, pp.47-95, 1979.

A. Tchoudakova and G. V. Callard, Identification of multiple CYP19 genes encoding different cytochrome P450 aromatase isozymes in brain and ovary, Endocrinology, vol.139, issue.4, pp.2179-2189, 1998.

A. Tingaud-sequeira, M. Andre, J. Forgue, C. Barthe, and P. J. Babin, Expression patterns of three estrogen receptor genes during zebrafish (Danio rerio) development: evidence for high expression in neuromasts, Gene Expr Patterns, vol.4, issue.5, pp.561-568, 2004.
URL : https://hal.archives-ouvertes.fr/hal-02673320

S. K. Tong and B. C. Chung, Analysis of zebrafish cyp19 promoters, J Steroid Biochem Mol Biol, vol.86, issue.3-5, pp.381-386, 2003.

C. D. Toran-allerand, Minireview: A plethora of estrogen receptors in the brain: where will it end?, Endocrinology, vol.145, issue.3, pp.1069-1074, 2004.

J. M. Trant, S. Gavasso, J. Ackers, B. C. Chung, and A. R. Place, Developmental expression of cytochrome P450 aromatase genes (CYP19a and CYP19b) in zebrafish fry (Danio rerio), J Exp Zool, vol.290, issue.5, pp.475-483, 2001.

C. L. Tsai, S. L. Chang, L. H. Wang, and T. Y. Chao, Temperature influences the ontogenetic expression of aromatase and oestrogen receptor mRNA in the developing tilapia (Oreochromis mossambicus) brain, J Neuroendocrinol, vol.15, issue.1, pp.97-102, 2003.

D. Uchida, M. Yamashita, T. Kitano, and T. Iguchi, An aromatase inhibitor or high water temperature induce oocyte apoptosis and depletion of P450 aromatase activity in the gonads of genetic female zebrafish during sex-reversal, Comp Biochem Physiol A Mol Integr Physiol, vol.137, issue.1, pp.11-20, 2004.

L. Wang, S. Andersson, M. Warner, and J. A. Gustafsson, Estrogen receptor (ER)beta knockout mice reveal a role for ERbeta in migration of cortical neurons in the developing brain, Proc Natl Acad Sci U S A, vol.100, issue.2, pp.703-708, 2003.

L. H. Wang and C. L. Tsai, Photoperiod influences the ontogenetic expression of aromatase and estrogen receptor alpha in the developing tilapia brain, Gen Comp Endocrinol, vol.145, issue.1, pp.62-66, 2006.

J. Balthazart and G. F. Ball, New insights into the regulation and function of brain estrogen synthase (aromatase), Trends Neurosci, vol.21, pp.243-249, 1998.

C. Behl, Estrogen can protect neurons: modes of action, J Steroid Biochem Mol Biol, vol.83, pp.195-197, 2002.

K. Cheshenko, F. Brion, Y. Le-page, N. Hinfray, F. Pakdel et al., Expression of zebra fish aromatase cyp19a and cyp19b genes in response to the ligands of estrogen receptor and aryl hydrocarbon receptor, Toxicol Sci, vol.96, pp.255-267, 2007.
URL : https://hal.archives-ouvertes.fr/hal-00140538

E. F. Chiang, Y. L. Yan, S. K. Tong, P. H. Hsiao, Y. Guiguen et al., Characterization of duplicated zebrafish cyp19 genes, J Exp Zool, vol.290, pp.709-714, 2001.
URL : https://hal.archives-ouvertes.fr/hal-02683253

A. Conley and M. Hinshelwood, Mammalian aromatases, Reproduction, vol.121, pp.685-695, 2001.

P. De-waal, D. Wang, W. Nijenhuis, R. Schulz, and J. Bogerd, Functional characterization and expression analysis of the androgen receptor in zebrafish, 2008.

P. P. De-waal, D. S. Wang, W. A. Nijenhuis, R. W. Schulz, and J. Bogerd, Functional characterization and expression analysis of the androgen receptor in zebrafish (Danio rerio) testis, Reproduction, vol.136, pp.225-234, 2008.

P. M. Forlano, D. L. Deitcher, D. A. Myers, and A. H. Bass, Anatomical distribution and cellular basis for high levels of aromatase activity in the brain of teleost fish: aromatase enzyme and mRNA expression identify glia as source, J Neurosci, vol.21, pp.8943-8955, 2001.

L. M. Garcia-segura, F. Naftolin, J. B. Hutchison, I. Azcoitia, and J. A. Chowen, , 1999.

, Role of astroglia in estrogen regulation of synaptic plasticity and brain repair, J Neurobiol, vol.40, pp.574-584

L. M. Garcia-segura, S. Veiga, A. Sierra, R. C. Melcangi, A. et al., , 2003.

, Aromatase: a neuroprotective enzyme, Prog Neurobiol, vol.71, pp.31-41

D. Gelinas, G. A. Pitoc, and G. V. Callard, Isolation of a goldfish brain cytochrome P450 aromatase cDNA: mRNA expression during the seasonal cycle and after steroid treatment, Mol Cell Endocrinol, vol.138, pp.81-93, 1998.

A. Gonzalez and F. Piferrer, 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, vol.132, pp.223-230, 2003.

M. Gotz, E. Hartfuss, and P. Malatesta, Radial glial cells as neuronal precursors: a new perspective on the correlation of morphology and lineage restriction in the developing cerebral cortex of mice, Brain Res Bull, vol.57, pp.777-788, 2002.

R. J. Handa, T. R. Pak, A. E. Kudwa, T. D. Lund, and L. Hinds, An alternate pathway for androgen regulation of brain function: activation of estrogen receptor beta by the metabolite of dihydrotestosterone, 5alpha-androstane-3beta,17beta-diol, Horm Behav, vol.53, pp.741-752, 2008.

M. B. Hawkins, J. W. Thornton, D. Crews, J. K. Skipper, A. Dotte et al., , 2000.

, Identification of a third distinct estrogen receptor and reclassification of estrogen receptors in teleosts, Proc Natl Acad Sci U S A, vol.97, pp.10751-10756

M. Kishida and G. V. Callard, Distinct cytochrome P450 aromatase isoforms in zebrafish (Danio rerio) brain and ovary are differentially programmed and estrogen regulated during early development, Endocrinology, vol.142, pp.740-750, 2001.

G. G. Kuiper, P. J. Shughrue, I. Merchenthaler, and J. A. Gustafsson, The estrogen receptor beta subtype: a novel mediator of estrogen action in neuroendocrine systems, 1998.

, Front Neuroendocrinol, vol.19, pp.253-286

J. Y. Kwon, B. J. Mcandrew, and D. J. Penman, 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, vol.59, pp.359-370, 2001.

C. S. Lassiter and E. Linney, Embryonic expression and steroid regulation of brain aromatase cyp19a1b in zebrafish (Danio rerio), Zebrafish, vol.4, pp.49-57, 2007.

L. Page, Y. Menuet, A. Kah, O. Pakdel, and F. , Characterization of a cis-acting element involved in cell-specific expression of the zebrafish brain aromatase gene, Mol Reprod Dev, 2008.
URL : https://hal.archives-ouvertes.fr/hal-00316100

L. Page, Y. Scholze, M. Kah, O. Pakdel, and F. , Assessment of xenoestrogens using three distinct estrogen receptors and the zebrafish brain aromatase gene in a highly responsive glial cell system, Environ Health Perspect, vol.114, pp.752-758, 2006.
URL : https://hal.archives-ouvertes.fr/hal-00068400

E. D. Lephart, A review of brain aromatase cytochrome P450, Brain Res Brain Res Rev, vol.22, pp.1-26, 1996.

C. Lethimonier, G. Flouriot, Y. Valotaire, O. Kah, and B. Ducouret, Transcriptional interference between glucocorticoid receptor and estradiol receptor mediates the inhibitory effect of cortisol on fish vitellogenesis, Biol Reprod, vol.62, pp.1763-1771, 2000.

V. Martinez-cerdeno, S. C. Noctor, and A. R. Kriegstein, Estradiol stimulates progenitor cell division in the ventricular and subventricular zones of the embryonic neocortex, Eur J Neurosci, vol.24, pp.3475-3488, 2006.

I. Mayer, B. Borg, I. Berglund, and J. G. Lambert, Effects of castration and androgen treatment on aromatase activity in the brain of mature male Atlantic salmon (Salmo salar L.) parr, Gen Comp Endocrinol, vol.82, pp.86-92, 1991.

M. M. Mccarthy, S. K. Amateau, and J. A. Mong, Steroid modulation of astrocytes in the neonatal brain: implications for adult reproductive function, Biol Reprod, vol.67, pp.691-698, 2002.

A. Menuet, I. Anglade, R. Le-guevel, E. Pellegrini, F. Pakdel et al., , 2003.

, Distribution of aromatase mRNA and protein in the brain and pituitary of female rainbow trout: Comparison with estrogen receptor alpha, J Comp Neurol, vol.462, pp.180-193

A. Menuet, E. Pellegrini, I. Anglade, O. Blaise, V. Laudet et al., , 2002.

, Molecular characterization of three estrogen receptor forms in zebrafish: binding characteristics, transactivation properties, and tissue distributions, Biol Reprod, vol.66, pp.1881-1892

A. Menuet, E. Pellegrini, F. Brion, M. M. Gueguen, I. Anglade et al., Expression and estrogen-dependent regulation of the zebrafish brain aromatase gene, J Comp Neurol, vol.485, pp.304-320, 2005.
URL : https://hal.archives-ouvertes.fr/hal-00294769

Y. Merot, R. Metivier, G. Penot, D. Manu, C. Saligaut et al., The relative contribution exerted by AF-1 and AF-2 transactivation functions in estrogen receptor alpha transcriptional activity depends upon the differentiation stage of the cell, J Biol Chem, vol.279, pp.26184-26191, 2004.
URL : https://hal.archives-ouvertes.fr/hal-00295059

K. Mouriec, E. Pellegrini, I. Anglade, A. Menuet, F. Adrio et al., 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, vol.75, pp.274-280, 2008.
URL : https://hal.archives-ouvertes.fr/hal-00266593

F. Naftolin, Brain aromatization of androgens, J Reprod Med, vol.39, pp.257-261, 1994.

S. C. Noctor, A. C. Flint, T. A. Weissman, W. S. Wong, B. K. Clinton et al., Dividing precursor cells of the embryonic cortical ventricular zone have morphological and molecular characteristics of radial glia, J Neurosci, vol.22, pp.3161-3173, 2002.

T. R. Pak, W. C. Chung, L. R. Hinds, and R. J. Handa, Estrogen receptor-beta mediates dihydrotestosterone-induced stimulation of the arginine vasopressin promoter in neuronal cells, Endocrinology, vol.148, pp.3371-3382, 2007.

T. R. Pak, W. C. Chung, T. D. Lund, L. R. Hinds, C. M. Clay et al., , 2005.

, The androgen metabolite, 5alpha-androstane-3beta, 17beta-diol, is a potent modulator of estrogen receptor-beta1-mediated gene transcription in neuronal cells, Endocrinology, vol.146, pp.147-155

M. Pasmanik and G. V. Callard, Aromatase and 5 alpha-reductase in the teleost brain, spinal cord, and pituitary gland, Gen Comp Endocrinol, vol.60, pp.244-251, 1985.

E. Pellegrini, A. Menuet, C. Lethimonier, F. Adrio, M. M. Gueguen et al., Relationships between aromatase and estrogen receptors in the brain of teleost fish, Gen Comp Endocrinol, vol.142, pp.60-66, 2005.
URL : https://hal.archives-ouvertes.fr/hal-00294797

E. Pellegrini, K. Mouriec, I. Anglade, A. Menuet, Y. Le-page et al., Identification of aromatasepositive radial glial cells as progenitor cells in the ventricular layer of the forebrain in zebrafish, J Comp Neurol, vol.501, pp.150-167, 2007.
URL : https://hal.archives-ouvertes.fr/hal-00096389

R. S. Peterson, G. Fernando, L. Day, T. A. Allen, J. D. Chapleau et al., Aromatase expression and cell proliferation following injury of the adult zebra finch hippocampus, Dev Neurobiol, vol.67, pp.1867-1878, 2007.

R. S. Peterson, D. W. Lee, G. Fernando, and B. A. Schlinger, Radial glia express aromatase in the injured zebra finch brain, J Comp Neurol, vol.475, pp.261-269, 2004.

P. Rakic, Neuronal migration and contact guidance in the primate telencephalon, 1978.

, Postgrad Med J 54 Suppl, vol.1, pp.25-40

C. E. Roselli and J. A. Resko, Aromatase activity in the rat brain: hormonal regulation and sex differences, J Steroid Biochem Mol Biol, vol.44, pp.499-508, 1993.

C. F. Roselli, Brain aromatase: roles in reproduction and neuroprotection, J Steroid Biochem Mol Biol, vol.106, pp.143-150, 2007.

T. D. Schmittgen and K. J. Livak, Analyzing real-time PCR data by the comparative C(T) method, Nat Protoc, vol.3, pp.1101-1108, 2008.

S. Y. Shu, G. Ju, and L. Z. Fan, The glucose oxidase-DAB-nickel method in peroxidase histochemistry of the nervous system, Neurosci Lett, vol.85, pp.169-171, 1988.

D. Steinke, S. Hoegg, H. Brinkmann, and A. Meyer, Three rounds (1R/2R/3R) of genome duplications and the evolution of the glycolytic pathway in vertebrates, BMC Biol, vol.4, p.16, 2006.

A. Tchoudakova, M. Kishida, E. Wood, and G. V. Callard, Promoter characteristics of two cyp19 genes differentially expressed in the brain and ovary of teleost fish, J Steroid Biochem Mol Biol, vol.78, pp.427-439, 2001.

S. K. Tong, C. , and B. C. , Analysis of zebrafish cyp19 promoters, J Steroid Biochem Mol Biol, vol.86, pp.381-386, 2003.

J. M. Trant, S. Gavasso, J. Ackers, B. C. Chung, and A. R. Place, Developmental expression of cytochrome P450 aromatase genes (CYP19a and CYP19b) in zebrafish fry (Danio rerio), J Exp Zool, vol.290, pp.475-483, 2001.

J. G. Yague, E. Lavaque, J. Carretero, I. Azcoitia, and L. M. Garcia-segura, , 2004.

, Aromatase, the enzyme responsible for estrogen biosynthesis, is expressed by human and rat glioblastomas, Neurosci Lett, vol.368, pp.279-284

C. Zhao, R. Fujinaga, M. Tanaka, A. Yanai, K. Nakahama et al., , 2007.

, Region-specific expression and sex-steroidal regulation on aromatase and its mRNA in the male rat brain: immunohistochemical and in situ hybridization analyses, J Comp Neurol, vol.500, pp.557-573

I. H. Zwain, Y. , and S. S. , Neurosteroidogenesis in astrocytes, oligodendrocytes, and neurons of cerebral cortex of rat brain, Endocrinology, vol.140, pp.3843-3852, 1999.

I. H. Zwain, S. S. Yen, and C. Y. Cheng, Astrocytes cultured in vitro produce estradiol-17beta and express aromatase cytochrome P-450 (P-450 AROM) mRNA, 1997.

, Biochim Biophys Acta, vol.1334, pp.338-348

, Aromatase, the estrogen-synthesizing enzyme, is strongly expressed in radial glial cells of the adult fish brain

, Emerging new functions of estrogens in embryonic, adult or reparative neurogenesis

, What triggers aromatase expression in the brain of fish?

. .. Conclusions,

. .. Acknowledgements,

. .. References,

B. Adolf, P. Chapouton, C. S. Lam, S. Topp, B. Tannhaüser et al., Conserved and acquired features of adult neurogenesis in the zebrafish telencephalon, Dev. Biol, vol.295, pp.278-293, 2006.

J. Altman, Are new neurons formed in the brains of adult mammals?, Science, vol.135, pp.1127-1128, 1962.

A. Alvarez-buylla, D. R. Buskirk, and F. Nottebohm, Monoclonal antibody reveals radial glia in adult avian brain, J. Comp. Neurol, vol.264, pp.159-170, 1987.

A. Alvarez-buylla, B. Seri, and F. Doetsch, Identification of neural stem cells in the adult vertebrate brain, Brain Res. Bull, vol.57, pp.751-758, 2002.

T. E. Anthony, C. Klein, G. Fishell, and N. Heintz, Radial glia serve as neuronal progenitors in all regions of the central nervous system, Neuron, vol.41, pp.881-890, 2004.

T. E. Anthony, H. A. Mason, T. Gridley, G. Fishell, and N. Heintz, Brain lipidbinding protein is a direct target of Notch signaling in radial glial cells, Genes Dev, vol.19, pp.1028-1033, 2005.

J. Balthazart and G. F. Ball, New insights into the regulation and function of brain estrogen synthase (aromatase), Trends Neurosci, vol.21, pp.243-249, 1998.

R. Brandstätter and K. , Brain growth patterns in four European cyprinid fish species (Cyprinidae, Teleostei): roach (Rutilus rutilus), bream (Abramis brama), common carp (Cyprinus carpio) and sabre carp (Pelecus cultratus), Brain Behav. Evol, vol.35, pp.195-211, 1990.

R. Brandstätter and K. , Life history of roach, Rutilus rutilus (Cyprinidae, Teleostei). A qualitative and quantitative study on the development of sensory brain areas, Brain Behav. Evol, vol.34, pp.35-42, 1989.

D. W. Brann, K. Dhandapani, C. Wakade, V. B. Mahesh, and M. M. Khan, Neurotrophic and neuroprotective actions of estrogen: basic mechanisms and clinical implications, Steroids, vol.72, pp.381-405, 2007.

S. E. Bulun, K. Takayama, T. Suzuki, H. Sasano, B. Yilmaz et al., Organization of the human aromatase p450 (CYP19) gene, Semin. Reprod. Med, vol.22, pp.5-9, 2004.

G. V. Callard, Z. Petro, and K. J. Ryan, Phylogenetic distribution of aromatase and other androgen-converting enzymes in the central nervous system, Endocrinology, vol.103, pp.2283-2290, 1978.

G. V. Callard, A. V. Tchoudakova, M. Kishida, and E. Wood, Differential tissue distribution, developmental programming, estrogen regulation and promoter characteristics of cyp19 genes in teleost fish, J. Steroid Biochem

, Mol. Biol, vol.79, pp.305-314, 2001.

K. Cheshenko, F. Pakdel, H. Segner, O. Kah, and R. I. Eggen, Interference of endocrine disrupting chemicals with aromatase CYP19 expression or activity, and consequences for reproduction of teleost fish, Gen. Comp. Endocrinol, 2007.
URL : https://hal.archives-ouvertes.fr/hal-00195359

E. F. Chiang, Y. L. Yan, S. K. Tong, P. H. Hsiao, Y. Guiguen et al., Characterization of duplicated zebrafish cyp19 genes, J. Exp. Zool, vol.290, pp.709-714, 2001.
URL : https://hal.archives-ouvertes.fr/hal-02683253

P. Ekström, C. M. Johnsson, and L. M. Ohlin, Ventricular proliferation zones in the brain of an adult teleost fish and their relation to neuromeres and migration (secondary matrix) zones, J. Comp. Neurol, vol.436, pp.92-110, 2001.

L. Fester, V. Ribeiro-gouveia, J. Prange-kiel, C. Schassen, M. Bottner et al., Proliferation and apoptosis of hippocampal granule cells require local oestrogen synthesis, J. Neurochem, vol.97, pp.1136-1144, 2006.

P. M. Forlano, D. L. Deitcher, D. A. Myers, and A. H. Bass, Anatomical distribution and cellular basis for high levels of aromatase activity in the brain of teleost fish: aromatase enzyme and mRNA expression identify glia as source, J. Neurosci, vol.21, pp.8943-8955, 2001.

P. M. Forlano, B. A. Schlinger, and A. H. Bass, Brain aromatase: new lessons from non-mammalian model systems, Front. Neuroendocrinol, vol.27, pp.247-274, 2006.

C. D. Fowler, F. Johnson, and Z. Wang, Estrogen regulation of cell proliferation and distribution of estrogen receptor-alpha in the brains of adult female prairie and meadow voles, J. Comp. Neurol, vol.489, pp.166-179, 2005.

L. A. Galea, M. D. Spritzer, J. M. Barker, and J. L. Pawluski, Gonadal hormone modulation of hippocampal neurogenesis in the adult, Hippocampus, vol.16, pp.225-232, 2006.

L. M. Garcia-segura, I. Azcoitia, and L. L. Doncarlos, Neuroprotection by estradiol, Prog. Neurobiol, vol.63, pp.29-60, 2001.

L. M. Garcia-segura, S. Veiga, A. Sierra, R. C. Melcangi, and I. Azcoitia, Aromatase: a neuroprotective enzyme, Prog. Neurobiol, vol.71, pp.31-41, 2003.

A. Gonzalez and F. Piferrer, Characterization of aromatase activity in the sea bass: effects of temperature and different catalytic properties of brain and ovarian homogenates and microsomes, J. Exp. Zool, vol.293, pp.500-510, 2002.

M. Götz, Glial cells generate neurons-master control within CNS regions: developmental perspectives on neural stem cells, Neuroscientist, vol.9, pp.379-397, 2003.

M. Götz and Y. A. Barde, Radial glial cells defined and major intermediates between embryonic stem cells and CNS neurons, Neuron, vol.46, pp.369-372, 2005.

H. Grandel, J. Kaslin, J. Ganz, I. Wenzel, and M. Brand, Neural stem cells and neurogenesis in the adult zebrafish brain: origin, proliferation dynamics, migration and cell fate, Dev. Biol, vol.295, pp.263-277, 2006.

Y. Hojo, T. A. Hattori, T. Enami, A. Furukawa, K. Suzuki et al., Adult male rat hippocampus synthesizes estradiol from pregnenolone by cytochromes P45017alpha and P450 aromatase localized in neurons, Proc. Natl. Acad. Sci. U.S.A, vol.101, pp.865-870, 2004.

R. Jagasia, H. Song, F. H. Gage, and D. C. Lie, New regulators in adult neurogenesis and their potential role for repair, vol.12, pp.400-405, 2006.

K. Jin, X. Wang, L. Xie, X. O. Mao, W. Zhu et al., Evidence for stroke-induced neurogenesis in the human brain, Proc. Natl. Acad. Sci. U.S.A, vol.103, pp.13198-13202, 2006.

M. S. Kaplan and J. W. Hinds, Neurogenesis in the adult rat: electron microscopic analysis of light radioautographs, Science, vol.197, pp.1092-1094, 1977.

Y. Kazeto, S. Ijiri, A. R. Place, Y. Zohar, and J. M. Trant, The 5 -flanking regions of CYP19A1 and CYP19A2 in zebrafish, Biochem. Biophys. Res. Commun, vol.288, pp.503-508, 2001.

Y. Kazeto, A. R. Place, and J. M. Trant, Effects of endocrine disrupting chemicals on the expression of CYP19 genes in zebrafish (Danio rerio) juveniles, Aquat. Toxicol, vol.69, pp.25-34, 2004.

W. Kirsche, On postembryonic matrix zones in the brain of various vertebrates and their relationship to the study of the brain structure, Z. Mikrosk. Anat. Forsch, vol.77, pp.313-406, 1967.

M. Kishida, M. Mclellan, J. A. Miranda, and G. V. Callard, Estrogen and xenoestrogens up-regulate the brain aromatase isoform (P450aromB) and perturb markers of early development in zebrafish (Danio rerio), Comp. Biochem. Physiol. B: Biochem. Mol. Biol, vol.129, pp.261-268, 2001.

D. Kranz and W. Richter, Autoradiographic studies on the localization of the matrix zones of the diencephalon of young and adult Lebistes reticulatus (Teleostae), Z. Mikrosk. Anat. Forsch, vol.82, pp.42-66, 1970.

D. Kranz and W. Richter, Autoradiographic studies on the synthesis of DNA in the cerebellum and medulla oblongata of teleosts of various ages, Z. Mikrosk. Anat. Forsch, vol.82, pp.264-292, 1970.

Y. L. Page, M. Scholze, O. Kah, and F. Pakdel, Assessment of xenoestrogens using three distinct estrogen receptors and the zebrafish brain aromatase gene in a highly responsive glial cell system, Environ. Health Perspect, vol.114, pp.752-758, 2006.
URL : https://hal.archives-ouvertes.fr/hal-00068400

R. R. Leker, F. Soldner, I. Velasco, D. K. Gavin, A. Androutsellis-theotokis et al., Long-lasting regeneration after ischemia in the cerebral cortex, Stroke, vol.38, pp.153-161, 2007.

E. D. Lephart, A review of brain aromatase cytochrome P450, Brain Res. Brain Res. Rev, vol.22, pp.1-26, 1996.

B. W. Lindsey and V. Tropepe, A comparative framework for understanding the biological principles of adult neurogenesis, Prog. Neurobiol, vol.80, pp.281-307, 2006.

P. M. Lledo, M. Alonso, and M. S. Grubb, Adult neurogenesis and functional plasticity in neuronal circuits, Nat. Rev. Neurosci, vol.7, pp.179-193, 2006.
URL : https://hal.archives-ouvertes.fr/pasteur-00160881

S. E. London and B. A. Schlinger, Steroidogenic enzymes along the ventricular proliferative zone in the developing songbird brain, J. Comp. Neurol, vol.502, pp.507-521, 2007.

K. E. Marsh, L. M. Creutz, M. B. Hawkins, and J. Godwin, Aromatase immunoreactivity in the bluehead wrasse brain, Thalassoma bifasciatum: immunolocalization and co-regionalization with arginine vasotocin and tyrosine hydroxylase, Brain Res, vol.1126, pp.91-101, 2006.

V. Martinez-cerdeno, S. C. Noctor, and A. R. Kriegstein, Estradiol stimulates progenitor cell division in the ventricular and subventricular zones of the embryonic neocortex, Eur. J. Neurosci, vol.24, pp.3475-3488, 2006.

C. A. Mazzucco, S. E. Lieblich, B. I. Bingham, M. A. Williamson, V. Viau et al., Both estrogen receptor alpha and estrogen receptor beta agonists enhance cell proliferation in the dentate gyrus of adult female rats, Neuroscience, vol.141, pp.1793-1800, 2006.

M. M. Mccarthy, Molecular aspects of sexual differentiation of the rodent brain, Psychoneuroendocrinology, vol.19, pp.415-427, 1994.

A. Menuet, I. Anglade, R. L. Guevel, E. Pellegrini, F. Pakdel et al., Distribution of aromatase mRNA and protein in the brain and pituitary of female rainbow trout: comparison with estrogen receptor alpha, J. Comp. Neurol, vol.462, pp.180-193, 2003.
URL : https://hal.archives-ouvertes.fr/hal-00295588

A. Menuet, E. Pellegrini, I. Anglade, O. Blaise, V. Laudet et al., Molecular characterization of three estrogen receptor forms in zebrafish: binding characteristics, transactivation properties, and tissue distributions, Biol. Reprod, vol.66, pp.1881-1892, 2002.

A. Menuet, E. Pellegrini, F. Brion, M. M. Gueguen, I. Anglade et al., Expression and estrogen-dependent regulation of the zebrafish brain aromatase gene, J. Comp. Neurol, vol.485, pp.304-320, 2005.
URL : https://hal.archives-ouvertes.fr/hal-00294769

J. Ninkovic and M. Götz, Signaling in adult neurogenesis: from stem cell niche to neuronal networks, Curr. Opin. Neurobiol, 2007.

S. C. Noctor, A. C. Flint, T. A. Weissman, W. S. Wong, B. K. Clinton et al., Dividing precursor cells of the embryonic cortical ventricular zone have morphological and molecular characteristics of radial glia, J. Neurosci, vol.22, pp.3161-3173, 2002.

S. C. Noctor, V. Martinez-cerdeno, L. Ivic, and A. R. Kriegstein, Cortical neurons arise in symmetric and asymmetric division zones and migrate through specific phases, Nat. Neurosci, vol.7, pp.136-144, 2004.

M. Pasmanik and G. V. Callard, Aromatase and 5 alpha-reductase in the teleost brain, spinal cord, and pituitary gland, Gen. Comp. Endocrinol, vol.60, pp.244-251, 1985.

E. Pellegrini, A. Menuet, C. Lethimonier, F. Adrio, M. M. Gueguen et al., Relationships between aromatase and estrogen receptors in the brain of teleost fish, vol.142, pp.60-66, 2005.
URL : https://hal.archives-ouvertes.fr/hal-00294797

E. Pellegrini, K. Mouriec, I. Anglade, A. Menuet, Y. L. Page et al., Identification of aromatase-positive radial glial cells as progenitor cells in the ventricular layer of the forebrain in zebrafish, J. Comp. Neurol, vol.501, pp.150-167, 2007.
URL : https://hal.archives-ouvertes.fr/hal-00096389

R. S. Peterson, D. W. Lee, G. Fernando, and B. A. Schlinger, Radial glia express aromatase in the injured zebra finch brain, J. Comp. Neurol, vol.475, pp.261-269, 2004.

R. S. Peterson, C. J. Saldanha, and B. A. Schlinger, Rapid up-regulation of aromatase mRNA and protein following neural injury in the zebra finch (Taeniopygia guttata), J. Neuroendocrinol, vol.13, pp.317-323, 2001.

P. Rakic, Neuronal migration and contact guidance in the primate telencephalon, Postgrad. Med. J, vol.54, pp.25-40, 1978.

S. Rastogi and D. A. Liberles, Subfunctionalization of duplicated genes as a transition state to neofunctionalization, BMC Evol. Biol, vol.5, p.28, 2005.

F. Saravia, Y. Revsin, V. Lux-lantos, J. Beauquis, F. Homo-delarche et al., Oestradiol restores cell proliferation in dentate gyrus and subventricular zone of streptozotocin-diabetic mice, J. Neuroendocrinol, vol.16, pp.704-710, 2004.

F. E. Saravia, J. Beauquis, Y. Revsin, F. Homo-delarche, E. R. De-kloet et al., Hippocampal neuropathology of diabetes mellitus is relieved by estrogen treatment, Cell. Mol. Neurobiol, vol.26, pp.943-957, 2006.

P. H. Strobl-mazzulla, N. P. Moncaut, G. C. Lopez, L. A. Miranda, A. V. Canario et al., Brain aromatase from pejerrey fish (Odontesthes bonariensis): cDNA cloning, tissue expression, and immunohistochemical localization, Gen. Comp. Endocrinol, vol.143, pp.21-32, 2005.

S. Suzuki, C. M. Brown, and P. M. Wise, Mechanisms of neuroprotection by estrogen, Endocrine, vol.29, pp.209-215, 2006.

S. Suzuki, L. M. Gerhold, M. Bottner, S. W. Rau, C. Cruz et al., Estradiol enhances neurogenesis following ischemic stroke through estrogen receptors alpha and beta, J. Comp. Neurol, vol.500, pp.1064-1075, 2007.

A. Tchoudakova and G. V. Callard, Identification of multiple CYP19 genes encoding different cytochrome P450 aromatase isozymes in brain and ovary, Endocrinology, vol.139, pp.2179-2189, 1998.

R. J. Timmers, J. G. Lambert, J. Peute, H. G. Vullings, and P. G. Van-oordt, Localization of aromatase in the brain of the male African catfish, Clarias gariepinus (Burchell), by microdissection and biochemical identification, J. Comp. Neurol, vol.258, pp.368-377, 1987.

S. K. Tong and B. C. Chung, Analysis of zebrafish cyp19 promoters, J. Steroid Biochem. Mol. Biol, vol.86, pp.381-386, 2003.

C. D. Toran-allerand, Minireview: a plethora of estrogen receptors in the brain: where will it end?, Endocrinology, vol.145, pp.1069-1074, 2004.

C. Wiltrout, B. Lang, Y. Yan, R. J. Dempsey, and R. Vemuganti, Repairing brain after stroke: a review on post-ischemic neurogenesis, Neurochem. Int, 2007.

J. G. Yague, A. Munoz, P. De-monasterio-schrader, J. Defelipe, L. M. Garcia-segura et al., Aromatase expression in the human temporal cortex, Neuroscience, vol.138, pp.389-401, 2006.

C. Zhao, R. Fujinaga, M. Tanaka, A. Yanai, K. Nakahama et al., Region-specific expression and sex-steroidal regulation on aromatase and its mRNA in the male rat brain: immunohistochemical and in situ hybridization analyses, J. Comp. Neurol, vol.500, pp.557-573, 2007.

G. K. Zupanc, K. Hinsch, F. H. Gage, and . Mccarthy, Proliferation, migration, neuronal differentiation, and long-term survival of new cells in the adult zebrafish brain, contraire, chez les poissons, vol.488, pp.290-319, 2002.

(. Kallivretaki, Ces dernières données semblent donc suggérer que, comme chez les mammifères, l'établissement d'un dimorphisme sexuel cérébral pourrait impliquer l, 2001.

. Lee, De plus, le sexe phénotypique du poisson zèbre est réversible, en fonction de facteurs la gonade, 1993.

;. Nakanishi, . Stacey, ;. Kobayashi, and . Es-s, Par exemple, chez de nombreux poissons coralliens, la mort d'un mâle dominant vivant avec plusieurs femelles entraîne une, 1996.

S. E. Abdelgadir, J. A. Resko, S. R. Ojeda, E. D. Lephart, M. J. Mcphaul et al., Androgens regulate aromatase cytochrome P450 messenger ribonucleic acid in rat brain, Endocrinology, vol.135, pp.395-401, 1994.

B. Adolf, P. Chapouton, C. S. Lam, S. Topp, B. Tannhauser et al., Conserved and acquired features of adult neurogenesis in the zebrafish telencephalon, Dev Biol, vol.295, pp.278-293, 2006.

J. Akimoto, H. Itoh, T. Miwa, and K. Ikeda, Immunohistochemical study of glutamine synthetase expression in early glial development, Brain Res Dev Brain Res, vol.72, pp.9-14, 1993.

J. Altman, Autoradiographic and histological studies of postnatal neurogenesis. 3. Dating the time of production and onset of differentiation of cerebellar microneurons in rats, J Comp Neurol, vol.136, pp.269-293, 1969.

J. Altman and G. D. Das, Autoradiographic and histological evidence of postnatal hippocampal neurogenesis in rats, J Comp Neurol, vol.124, pp.319-335, 1965.

A. Alunni, S. Vaccari, S. Torcia, M. E. Meomartini, A. Nicotra et al., Characterization of glial fibrillary acidic protein and astroglial architecture in the brain of a continuously growing fish, the rainbow trout, Eur J Histochem, vol.49, pp.157-166, 2005.

A. Alvarez-buylla, Mechanism of neurogenesis in adult avian brain, Experientia, vol.46, pp.948-955, 1990.

A. Alvarez-buylla and J. M. Garcia-verdugo, Neurogenesis in adult subventricular zone, J Neurosci, vol.22, pp.629-634, 2002.

A. Alvarez-buylla and F. Nottebohm, Migration of young neurons in adult avian brain, Nature, vol.335, pp.353-354, 1988.

A. Alvarez-buylla, B. Seri, and F. Doetsch, Identification of neural stem cells in the adult vertebrate brain, Brain Res Bull, vol.57, pp.751-758, 2002.

A. Alvarez-buylla, M. Theelen, and F. Nottebohm, Mapping of radial glia and of a new cell type in adult canary brain, J Neurosci, vol.8, pp.2707-2712, 1988.

S. K. Amateau and M. M. Mccarthy, Sexual differentiation of astrocyte morphology in the developing rat preoptic area, J Neuroendocrinol, vol.14, pp.904-910, 2002.

J. H. Anolik, C. M. Klinge, C. L. Brolly, R. A. Bambara, and R. Hilf, Stability of the ligand-estrogen receptor interaction depends on estrogen response element flanking sequences and cellular factors, J Steroid Biochem Mol Biol, vol.59, pp.413-429, 1996.

J. H. Anolik, C. M. Klinge, R. Hilf, and R. A. Bambara, Cooperative binding of estrogen receptor to DNA depends on spacing of binding sites, flanking sequence, and ligand, Biochemistry, vol.34, pp.2511-2520, 1995.

T. E. Anthony, C. Klein, G. Fishell, and N. Heintz, Radial glia serve as neuronal progenitors in all regions of the central nervous system, Neuron, vol.41, pp.881-890, 2004.

E. S. Anton, M. A. Marchionni, K. F. Lee, and P. Rakic, Role of GGF/neuregulin signaling in interactions between migrating neurons and radial glia in the developing cerebral cortex, Development, vol.124, pp.3501-3510, 1997.

M. Arochena, R. Anadon, and S. M. Diaz-regueira, Development of vimentin and glial fibrillary acidic protein immunoreactivities in the brain of gray mullet (Chelon labrosus), an advanced teleost, J Comp Neurol, vol.469, pp.413-436, 2004.

I. Azcoitia, A. Sierra, and L. M. Garcia-segura, Localization of estrogen receptor beta-immunoreactivity in astrocytes of the adult rat brain, Glia, vol.26, pp.260-267, 1999.

I. Azcoitia, A. Sierra, S. Veiga, S. Honda, N. Harada et al., , 2001.

, Brain aromatase is neuroprotective, J Neurobiol, vol.47, pp.318-329

J. Bakker, S. Honda, N. Harada, and J. Balthazart, Restoration of male sexual behavior by adult exogenous estrogens in male aromatase knockout mice, Horm Behav, vol.46, pp.1-10, 2004.

J. Balthazart, M. Baillien, T. D. Charlier, C. A. Cornil, and G. F. Ball, Multiple mechanisms control brain aromatase activity at the genomic and non-genomic level, 2003.

, Steroid Biochem Mol Biol, vol.86, pp.367-379

J. Balthazart and G. F. Ball, New insights into the regulation and function of brain estrogen synthase (aromatase), Trends Neurosci, vol.21, pp.243-249, 1998.

J. Balthazart, L. Evrard, and C. Surlemont, Effects of the nonsteroidal inhibitor R76713 on testosterone-induced sexual behavior in the Japanese quail (Coturnix coturnix japonica), Horm Behav, vol.24, pp.510-531, 1990.

J. Balthazart, A. Foidart, C. Surlemont, N. Harada, and F. Naftolin, , 1992.

, Neuroanatomical specificity in the autoregulation of aromatase-immunoreactive neurons by androgens and estrogens: an immunocytochemical study, Brain Res, vol.574, pp.280-290

J. Balthazart, C. Surlemont, and N. Harada, Aromatase as a cellular marker of testosterone action in the preoptic area, Physiol Behav, vol.51, pp.395-409, 1992.

E. E. Baulieu, Neurosteroids: of the nervous system, by the nervous system, for the nervous system, Recent Prog Horm Res, vol.52, pp.1-32, 1997.

E. E. Baulieu and P. Robel, Neurosteroids: a new brain function?, J Steroid Biochem Mol Biol, vol.37, pp.395-403, 1990.

M. Beato, P. Herrlich, and G. Schutz, Steroid hormone receptors: many actors in search of a plot, Cell, vol.83, pp.851-857, 1995.

C. Behl, Oestrogen as a neuroprotective hormone, Nat Rev Neurosci, vol.3, pp.433-442, 2002.

M. Bentivoglio and P. Mazzarello, The history of radial glia, Brain Res Bull, vol.49, pp.305-315, 1999.

R. L. Bernardos, R. , and P. A. , GFAP transgenic zebrafish, Gene Expr Patterns, vol.6, pp.1007-1013, 2006.

M. Blazquez and F. Piferrer, Cloning, sequence analysis, tissue distribution, and sexspecific expression of the neural form of P450 aromatase in juvenile sea bass (Dicentrarchus labrax), Mol Cell Endocrinol, vol.219, pp.83-94, 2004.

W. Bourguet, P. Germain, and H. Gronemeyer, Nuclear receptor ligand-binding domains: three-dimensional structures, molecular interactions and pharmacological implications, Trends Pharmacol Sci, vol.21, pp.381-388, 2000.

S. Bourguiba, S. Lambard, and S. Carreau, Steroids control the aromatase gene expression in purified germ cells from the adult male rat, J Mol Endocrinol, vol.31, pp.83-94, 2003.
URL : https://hal.archives-ouvertes.fr/hal-02673267

E. Brailoiu, S. L. Dun, G. C. Brailoiu, K. Mizuo, L. A. Sklar et al., Distribution and characterization of estrogen receptor G proteincoupled receptor 30 in the rat central nervous system, J Endocrinol, vol.193, pp.311-321, 2007.

R. Brandstatter and K. Kotrschal, Life history of roach, Rutilus rutilus (Cyprinidae, Teleostei). A qualitative and quantitative study on the development of sensory brain areas, 1989.

, Brain Behav Evol, vol.34, pp.35-42

R. Brandstatter and K. Kotrschal, Brain growth patterns in four European cyprinid fish species (Cyprinidae, Teleostei): roach (Rutilus rutilus), bream (Abramis brama), common carp (Cyprinus carpio) and sabre carp (Pelecus cultratus), Brain Behav Evol, vol.35, pp.195-211, 1990.

K. Brannvall, L. Korhonen, and D. Lindholm, Estrogen-receptor-dependent regulation of neural stem cell proliferation and differentiation, Mol Cell Neurosci, vol.21, pp.512-520, 2002.

S. E. Bulun, S. Sebastian, K. Takayama, T. Suzuki, H. Sasano et al., The human CYP19 (aromatase P450) gene: update on physiologic roles and genomic organization of promoters, J Steroid Biochem Mol Biol, vol.86, pp.219-224, 2003.

S. Carreau, The testicular aromatase: from gene to physiological role, Reprod Biol, vol.2, pp.5-12, 2002.

R. E. Caton, K. E. Farnell, R. B. Chronister, L. E. White, . Jr et al., , 1980.

, Regeneration of the forebrain of the Japanese carp, Cyprinus carpio: a Golgi analysis, J Hirnforsch, vol.21, pp.257-263

X. Chang, T. Kobayashi, B. Senthilkumaran, H. Kobayashi-kajura, C. C. Sudhakumari et al., Two types of aromatase with different encoding genes, tissue distribution and developmental expression in Nile tilapia, 2005.

, Comp Endocrinol, vol.141, pp.101-115

A. Chenn and S. K. Mcconnell, Cleavage orientation and the asymmetric inheritance of Notch1 immunoreactivity in mammalian neurogenesis, Cell, vol.82, pp.631-641, 1995.

E. F. Chiang, Y. L. Yan, Y. Guiguen, J. Postlethwait, C. et al., Two Cyp19 (P450 aromatase) genes on duplicated zebrafish chromosomes are expressed in ovary or brain, Mol Biol Evol, vol.18, pp.542-550, 2001.
URL : https://hal.archives-ouvertes.fr/hal-02680091

C. Y. Choi and H. R. Habibi, Molecular cloning of estrogen receptor alpha and expression pattern of estrogen receptor subtypes in male and female goldfish, Mol Cell Endocrinol, vol.204, pp.169-177, 2003.

N. A. Compagnone and S. H. Mellon, Neurosteroids: biosynthesis and function of these novel neuromodulators, Front Neuroendocrinol, vol.21, pp.1-56, 2000.

A. Conley, J. Corbin, T. Smith, M. Hinshelwood, Z. Liu et al., Porcine aromatases: studies on tissue-specific, functionally distinct isozymes from a single gene?, J Steroid Biochem Mol Biol, vol.61, pp.407-413, 1997.

A. Conley and M. Hinshelwood, Mammalian aromatases, Reproduction, vol.121, pp.685-695, 2001.

R. A. Cornell and J. S. Eisen, Notch in the pathway: the roles of Notch signaling in neural crest development, Semin Cell Dev Biol, vol.16, pp.663-672, 2005.

J. F. Couse and K. S. Korach, Estrogen receptor null mice: what have we learned and where will they lead us?, Endocr Rev, vol.20, pp.358-417, 1999.

F. Doetsch, I. Caille, D. A. Lim, J. M. Garcia-verdugo, and A. Alvarez-buylla, , 1999.

, Subventricular zone astrocytes are neural stem cells in the adult mammalian brain, Cell, vol.97, pp.703-716

D. B. Dubal, H. Zhu, J. Yu, S. W. Rau, P. J. Shughrue et al., Estrogen receptor alpha, not beta, is a critical link in estradiolmediated protection against brain injury, Proc Natl Acad Sci U S A, vol.98, pp.1952-1957, 2001.

S. Dutta, J. E. Dietrich, M. Westerfield, and Z. M. Varga, Notch signaling regulates endocrine cell specification in the zebrafish anterior pituitary, Dev Biol, vol.319, pp.248-257, 2008.

P. Ekstrom, C. M. Johnsson, and L. M. Ohlin, Ventricular proliferation zones in the brain of an adult teleost fish and their relation to neuromeres and migration (secondary matrix) zones, J Comp Neurol, vol.436, pp.92-110, 2001.

F. Estay, A. Diaz, R. Pedrazza, C. , and N. , Oogenesis and plasma levels of sex steroids in cultured females of brown trout (Salmo trutta linnaeus, 1758) in Chile, 2003.

, Exp Zoolog A Comp Exp Biol, vol.298, pp.60-66

L. Feng and N. Heintz, Differentiating neurons activate transcription of the brain lipid-binding protein gene in radial glia through a novel regulatory element, Development, vol.121, pp.1719-1730, 1995.

E. J. Filardo, J. A. Quinn, A. R. Frackelton, . Jr, and K. I. Bland, Estrogen action via the G protein-coupled receptor, GPR30: stimulation of adenylyl cyclase and cAMPmediated attenuation of the epidermal growth factor receptor-to-MAPK signaling axis, 2002.

, Mol Endocrinol, vol.16, pp.70-84

S. K. Fisher and G. P. Lewis, Muller cell and neuronal remodeling in retinal detachment and reattachment and their potential consequences for visual recovery: a review and reconsideration of recent data, Vision Res, vol.43, pp.887-897, 2003.

P. M. Forlano and A. H. Bass, Steroid regulation of brain aromatase expression in glia: female preoptic and vocal motor nuclei, J Neurobiol, vol.65, pp.50-58, 2005.

P. M. Forlano, D. L. Deitcher, D. A. Myers, and A. H. Bass, Anatomical distribution and cellular basis for high levels of aromatase activity in the brain of teleost fish: aromatase enzyme and mRNA expression identify glia as source, J Neurosci, vol.21, pp.8943-8955, 2001.

E. M. Frojdo, J. Westerlund, and B. Isomaa, Culturing and characterization of astrocytes isolated from juvenile rainbow trout (Oncorhynchus mykiss), Comp Biochem Physiol A Mol Integr Physiol, vol.133, pp.17-28, 2002.

T. Funakoshi, A. Yanai, K. Shinoda, M. M. Kawano, and Y. Mizukami, G proteincoupled receptor 30 is an estrogen receptor in the plasma membrane, Biochem Biophys Res Commun, vol.346, pp.904-910, 2006.

A. Furukawa, A. Miyatake, T. Ohnishi, and Y. Ichikawa, Steroidogenic acute regulatory protein (StAR) transcripts constitutively expressed in the adult rat central nervous system: colocalization of StAR, cytochrome P-450SCC (CYP XIA1), and 3beta-hydroxysteroid dehydrogenase in the rat brain, J Neurochem, vol.71, pp.2231-2238, 1998.

L. Fusani and M. Gahr, Hormonal influence on song structure and organization: the role of estrogen, Neuroscience, vol.138, pp.939-946, 2006.

L. A. Galea, M. D. Spritzer, J. M. Barker, and J. L. Pawluski, Gonadal hormone modulation of hippocampal neurogenesis in the adult, Hippocampus, vol.16, pp.225-232, 2006.

D. Garcia-ovejero, S. Veiga, L. M. Garcia-segura, and L. L. Doncarlos, Glial expression of estrogen and androgen receptors after rat brain injury, J Comp Neurol, vol.450, pp.256-271, 2002.

L. M. Garcia-segura, F. Naftolin, J. B. Hutchison, I. Azcoitia, and J. A. Chowen, , 1999.

, Role of astroglia in estrogen regulation of synaptic plasticity and brain repair, J Neurobiol, vol.40, pp.574-584

L. M. Garcia-segura, A. Wozniak, I. Azcoitia, J. R. Rodriguez, R. E. Hutchison et al., Aromatase expression by astrocytes after brain injury: implications for local estrogen formation in brain repair, Neuroscience, vol.89, pp.567-578, 1999.

D. Gelinas, G. A. Pitoc, and G. V. Callard, Isolation of a goldfish brain cytochrome P450 aromatase cDNA: mRNA expression during the seasonal cycle and after steroid treatment, Mol Cell Endocrinol, vol.138, pp.81-93, 1998.

C. Genissel, J. Levallet, and S. Carreau, Regulation of cytochrome P450 aromatase gene expression in adult rat Leydig cells: comparison with estradiol production, J Endocrinol, vol.168, pp.95-105, 2001.

P. Germain, B. Staels, C. Dacquet, M. Spedding, and V. Laudet, Overview of nomenclature of nuclear receptors, Pharmacol Rev, vol.58, pp.685-704, 2006.
URL : https://hal.archives-ouvertes.fr/hal-02657483

S. A. Goldman and F. Nottebohm, Neuronal production, migration, and differentiation in a vocal control nucleus of the adult female canary brain, Proc Natl Acad Sci U S A, vol.80, pp.2390-2394, 1983.

S. A. Goldman, A. Zukhar, K. Barami, T. Mikawa, and D. Niedzwiecki, , 1996.

, Ependymal/subependymal zone cells of postnatal and adult songbird brain generate both neurons and nonneuronal siblings in vitro and in vivo, J Neurobiol, vol.30, pp.505-520

A. Gonzalez and F. Piferrer, Characterization of aromatase activity in the sea bass: effects of temperature and different catalytic properties of brain and ovarian homogenates and microsomes, J Exp Zool, vol.293, pp.500-510, 2002.

A. Gonzalez and F. Piferrer, 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, vol.132, pp.223-230, 2003.

M. Gotz and Y. A. Barde, Radial glial cells defined and major intermediates between embryonic stem cells and CNS neurons, Neuron, vol.46, pp.369-372, 2005.

M. Gotz and W. B. Huttner, The cell biology of neurogenesis, Nat Rev Mol Cell Biol, vol.6, pp.777-788, 2005.

M. Gotz, A. Stoykova, and P. Gruss, Pax6 controls radial glia differentiation in the cerebral cortex, Neuron, vol.21, pp.1031-1044, 1998.

L. G. Graddy, A. A. Kowalski, F. A. Simmen, S. L. Davis, W. W. Baumgartner et al., Multiple isoforms of porcine aromatase are encoded by three distinct genes, J Steroid Biochem Mol Biol, vol.73, pp.49-57, 2000.

J. M. Hall, D. P. Mcdonnell, and K. S. Korach, Allosteric regulation of estrogen receptor structure, function, and coactivator recruitment by different estrogen response elements, Mol Endocrinol, vol.16, pp.469-486, 2002.

S. Halm, J. Y. Kwon, M. Rand-weaver, J. P. Sumpter, N. Pounds et al., Cloning and gene expression of P450 17alpha-hydroxylase, 20-lyase cDNA in the gonads and brain of the fathead minnow Pimephales promelas, vol.17, 2003.

, Comp Endocrinol, vol.130, pp.256-266

S. Halm, N. Pounds, S. Maddix, M. Rand-weaver, J. P. Sumpter et al., Exposure to exogenous 17beta-oestradiol disrupts p450aromB mRNA expression in the brain and gonad of adult fathead minnows (Pimephales promelas), Aquat Toxicol, vol.60, pp.285-299, 2002.

E. Hartfuss, E. Forster, H. H. Bock, M. A. Hack, P. Leprince et al., Reelin signaling directly affects radial glia morphology and biochemical maturation, Development, vol.130, pp.4597-4609, 2003.

E. Hartfuss, R. Galli, N. Heins, and M. Gotz, Characterization of CNS precursor subtypes and radial glia, Dev Biol, vol.229, pp.15-30, 2001.

J. Hatakeyama and R. Kageyama, Notch1 expression is spatiotemporally correlated with neurogenesis and negatively regulated by Notch1-independent Hes genes in the developing nervous system, Cereb Cortex, vol.16, issue.1, pp.132-137, 2006.

M. E. Hatten, Central nervous system neuronal migration, Annu Rev Neurosci, vol.22, pp.511-539, 1999.

M. E. Hatten, New directions in neuronal migration, Science, vol.297, pp.1660-1663, 2002.

M. B. Hawkins, J. Godwin, D. Crews, T. , and P. , The distributions of the duplicate oestrogen receptors ER-beta a and ER-beta b in the forebrain of the Atlantic croaker (Micropogonias undulatus): evidence for subfunctionalization after gene duplication, Proc Biol Sci, vol.272, pp.633-641, 2005.

M. B. Hawkins, T. , and P. , The unusual binding properties of the third distinct teleost estrogen receptor subtype ERbetaa are accompanied by highly conserved amino acid changes in the ligand binding domain, Endocrinology, vol.145, pp.2968-2977, 2004.

J. P. Hobbs, P. L. Munday, and G. P. Jones, Social induction of maturation and sex determination in a coral reef fish, Proc Biol Sci, vol.271, pp.2109-2114, 2004.

Y. Hojo, T. A. Hattori, T. Enami, A. Furukawa, K. Suzuki et al., Adult male rat hippocampus synthesizes estradiol from pregnenolone by cytochromes P45017alpha and P450 aromatase localized in neurons, Proc Natl Acad Sci U S A, vol.101, pp.865-870, 2004.

J. B. Hutchison, C. Beyer, S. Green, and A. Wozniak, Brain formation of oestrogen in the mouse: sex dimorphism in aromatase development, J Steroid Biochem Mol Biol, vol.49, pp.407-415, 1994.

W. B. Huttner and Y. Kosodo, Symmetric versus asymmetric cell division during neurogenesis in the developing vertebrate central nervous system, Curr Opin Cell Biol, vol.17, pp.648-657, 2005.

J. S. Jacobi, C. Martin, G. Nava, M. C. Jeziorski, C. Clapp et al., 17-Beta-estradiol directly regulates the expression of adrenergic receptors and kisspeptin/GPR54 system in GT1-7 GnRH neurons, Neuroendocrinology, vol.86, pp.260-269, 2007.

E. Kallivretaki, R. I. Eggen, S. C. Neuhauss, O. Kah, and H. Segner, The zebrafish, brain-specific, aromatase cyp19a2 is neither expressed nor distributed in a sexually dimorphic manner during sexual differentiation, Dev Dyn, vol.236, pp.3155-3166, 2007.
URL : https://hal.archives-ouvertes.fr/hal-00180993

M. Kalman, Astroglial architecture of the carp (Cyprinus carpio) brain as revealed by immunohistochemical staining against glial fibrillary acidic protein (GFAP), Anat Embryol (Berl), vol.198, pp.409-433, 1998.

M. S. Kaplan, Neurogenesis in the 3-month-old rat visual cortex, J Comp Neurol, vol.195, pp.323-338, 1981.

Y. Kazeto, S. Ijiri, A. R. Place, Y. Zohar, and J. M. Trant, The 5'-flanking regions of CYP19A1 and CYP19A2 in zebrafish, Biochem Biophys Res Commun, vol.288, pp.503-508, 2001.

D. E. Kime and I. P. Dolben, Hormonal changes during induced ovulation of the carp, Cyprinus carpio, Gen Comp Endocrinol, vol.58, pp.137-149, 1985.

T. Kimoto, T. Tsurugizawa, Y. Ohta, J. Makino, H. Tamura et al., Neurosteroid synthesis by cytochrome p450-containing systems localized in the rat brain hippocampal neurons: N-methyl-D-aspartate and calciumdependent synthesis, Endocrinology, vol.142, pp.3578-3589, 2001.

M. Kishida and G. V. Callard, Distinct cytochrome P450 aromatase isoforms in zebrafish (Danio rerio) brain and ovary are differentially programmed and estrogen regulated during early development, Endocrinology, vol.142, pp.740-750, 2001.

L. Klein-hitpass, M. Schorpp, U. Wagner, and G. U. Ryffel, An estrogenresponsive element derived from the 5' flanking region of the Xenopus vitellogenin A2 gene functions in transfected human cells, Cell, vol.46, pp.1053-1061, 1986.

L. Klein-hitpass, C. Schwerk, S. Kahmann, and L. Vassen, Targets of activated steroid hormone receptors: basal transcription factors and receptor interacting proteins, 1998.

, Mol Med, vol.76, pp.490-496

C. M. Klinge, Estrogen receptor interaction with co-activators and co-repressors, Steroids, vol.65, pp.227-251, 2000.

C. M. Klinge, Estrogen receptor interaction with estrogen response elements, Nucleic Acids Res, vol.29, pp.2905-2919, 2001.

C. M. Klinge, F. V. Peale, . Jr, R. Hilf, R. A. Bambara et al., Cooperative estrogen receptor interaction with consensus or variant estrogen responsive elements in vitro, Cancer Res, vol.52, pp.1073-1081, 1992.

M. Kobayashi and T. Nakanishi, 11-ketotestosterone induces male-type sexual behavior and gonadotropin secretion in gynogenetic crucian carp, Carassius auratus langsdorfii, Gen Comp Endocrinol, vol.115, pp.178-187, 1999.

D. Kranz and W. Richter, , 1970.

, Z Mikrosk Anat Forsch, vol.82, pp.42-66

V. D. Kranz and W. Richter, , 1975.

, Z Alternsforsch, vol.30, pp.371-382

P. J. Kushner, D. A. Agard, G. L. Greene, T. S. Scanlan, A. K. Shiau et al., Estrogen receptor pathways to AP-1, J Steroid Biochem Mol Biol, vol.74, pp.311-317, 2000.

M. C. Langub, . Jr, R. E. Watson, and . Jr, Estrogen receptor-immunoreactive glia, endothelia, and ependyma in guinea pig preoptic area and median eminence: electron microscopy, Endocrinology, vol.130, pp.364-372, 1992.

P. M. Lansdorp, Immortal strands? Give me a break, Cell, vol.129, pp.1244-1247, 2007.

L. Page, Y. Menuet, A. Kah, O. Pakdel, and F. , Characterization of a cis-acting element involved in cell-specific expression of the zebrafish brain aromatase gene, Mol Reprod Dev, 2008.
URL : https://hal.archives-ouvertes.fr/hal-00316100

L. Page, Y. Scholze, M. Kah, O. Pakdel, and F. , Assessment of xenoestrogens using three distinct estrogen receptors and the zebrafish brain aromatase gene in a highly responsive glial cell system, Environ Health Perspect, vol.114, pp.752-758, 2006.
URL : https://hal.archives-ouvertes.fr/hal-00068400

W. K. Lee, K. W. Lee, E. J. Kwak, S. W. Yang, K. S. Yang et al., Effects of environmental endocrine disruptors on the sex differentiation in Korean rockfish, Sebastes schlegeli, Water Sci Technol, vol.47, pp.65-70, 2003.

E. D. Lephart, A review of brain aromatase cytochrome P450, Brain Res Brain Res Rev, vol.22, pp.1-26, 1996.

C. Y. Lin, V. B. Vega, J. S. Thomsen, T. Zhang, S. L. Kong et al., Whole-genome cartography of estrogen receptor alpha binding sites, PLoS Genet, vol.3, p.87, 2007.

M. Liu, P. D. Hurn, C. E. Roselli, A. , and N. J. , Role of P450 aromatase in sex-specific astrocytic cell death, J Cereb Blood Flow Metab, vol.27, pp.135-141, 2007.

S. E. London, D. A. Monks, J. Wade, and B. A. Schlinger, Widespread capacity for steroid synthesis in the avian brain and song system, Endocrinology, vol.147, pp.5975-5987, 2006.

S. E. London and B. A. Schlinger, Steroidogenic enzymes along the ventricular proliferative zone in the developing songbird brain, J Comp Neurol, vol.502, pp.507-521, 2007.

P. Malatesta, M. A. Hack, E. Hartfuss, H. Kettenmann, W. Klinkert et al., Neuronal or glial progeny: regional differences in radial glia fate, Neuron, vol.37, pp.751-764, 2003.

P. Malatesta, E. Hartfuss, and M. Gotz, Isolation of radial glial cells by fluorescentactivated cell sorting reveals a neuronal lineage, Development, vol.127, pp.5253-5263, 2000.

D. J. Mangelsdorf, C. Thummel, M. Beato, P. Herrlich, G. Schutz et al., The nuclear receptor superfamily: the second decade, Cell, vol.83, pp.835-839, 1995.

K. E. Marsh, L. M. Creutz, M. B. Hawkins, G. , and J. , Aromatase immunoreactivity in the bluehead wrasse brain, Thalassoma bifasciatum: immunolocalization and co-regionalization with arginine vasotocin and tyrosine hydroxylase, Brain Res, vol.1126, pp.91-101, 2006.

V. Martinez-cerdeno, S. C. Noctor, and A. R. Kriegstein, Estradiol stimulates progenitor cell division in the ventricular and subventricular zones of the embryonic neocortex, Eur J Neurosci, vol.24, pp.3475-3488, 2006.

M. Mathieu, A. G. Mensah-nyagan, M. Vallarino, J. L. Do-rego, D. Beaujean et al., Immunohistochemical localization of 3 beta-hydroxysteroid dehydrogenase and 5 alpha-reductase in the brain of the African lungfish Protopterus annectens, J Comp Neurol, vol.438, pp.123-135, 2001.

T. Matsumoto, S. Honda, and N. Harada, Neurological effects of aromatase deficiency in the mouse, J Steroid Biochem Mol Biol, vol.86, pp.357-365, 2003.

M. M. Mccarthy, S. K. Amateau, and J. A. Mong, Steroid modulation of astrocytes in the neonatal brain: implications for adult reproductive function, Biol Reprod, vol.67, pp.691-698, 2002.

L. D. Mccullough, K. Blizzard, E. R. Simpson, O. K. Oz, and P. D. Hurn, , 2003.

, Aromatase cytochrome P450 and extragonadal estrogen play a role in ischemic neuroprotection, J Neurosci, vol.23, pp.8701-8705

A. C. Melo and J. S. Ramsdell, Sexual dimorphism of brain aromatase activity in medaka: induction of a female phenotype by estradiol, Environ Health Perspect, vol.109, pp.257-264, 2001.

A. Menuet, I. Anglade, R. Le-guevel, E. Pellegrini, F. Pakdel et al., , 2003.

, Distribution of aromatase mRNA and protein in the brain and pituitary of female rainbow trout: Comparison with estrogen receptor alpha, J Comp Neurol, vol.462, pp.180-193

A. Menuet, E. Pellegrini, I. Anglade, O. Blaise, V. Laudet et al., , 2002.

, Molecular characterization of three estrogen receptor forms in zebrafish: binding characteristics, transactivation properties, and tissue distributions, Biol Reprod, vol.66, pp.1881-1892

A. Menuet, E. Pellegrini, F. Brion, M. M. Gueguen, I. Anglade et al., Expression and estrogen-dependent regulation of the zebrafish brain aromatase gene, J Comp Neurol, vol.485, pp.304-320, 2005.
URL : https://hal.archives-ouvertes.fr/hal-00294769

F. T. Merkle, A. D. Tramontin, J. M. Garcia-verdugo, and A. Alvarez-buylla, , 2004.

, Radial glia give rise to adult neural stem cells in the subventricular zone, Proc Natl Acad Sci U S A, vol.101, pp.17528-17532

T. Miyata, A. Kawaguchi, K. Saito, M. Kawano, T. Muto et al., , 2004.

, Asymmetric production of surface-dividing and non-surface-dividing cortical progenitor cells, Development, vol.131, pp.3133-3145

T. Mori, A. Buffo, and M. Gotz, The novel roles of glial cells revisited: the contribution of radial glia and astrocytes to neurogenesis, Curr Top Dev Biol, vol.69, pp.67-99, 2005.

S. Mosselman, J. Polman, and R. Dijkema, ER beta: identification and characterization of a novel human estrogen receptor, FEBS Lett, vol.392, pp.49-53, 1996.

T. Mueller and M. F. Wullimann, Anatomy of neurogenesis in the early zebrafish brain, Brain Res Dev Brain Res, vol.140, pp.137-155, 2003.

F. Naftolin, K. J. Ryan, and Z. Petro, Aromatization of androstenedione by the diencephalon, J Clin Endocrinol Metab, vol.33, pp.368-370, 1971.

S. C. Noctor, A. C. Flint, T. A. Weissman, R. S. Dammerman, and A. R. Kriegstein, Neurons derived from radial glial cells establish radial units in neocortex, Nature, vol.409, pp.714-720, 2001.

S. C. Noctor, A. C. Flint, T. A. Weissman, W. S. Wong, B. K. Clinton et al., Dividing precursor cells of the embryonic cortical ventricular zone have morphological and molecular characteristics of radial glia, J Neurosci, vol.22, pp.3161-3173, 2002.

S. C. Noctor, V. Martinez-cerdeno, L. Ivic, and A. R. Kriegstein, Cortical neurons arise in symmetric and asymmetric division zones and migrate through specific phases, Nat Neurosci, vol.7, pp.136-144, 2004.

S. C. Noctor, V. Martinez-cerdeno, and A. R. Kriegstein, Distinct behaviors of neural stem and progenitor cells underlie cortical neurogenesis, J Comp Neurol, vol.508, pp.28-44, 2008.

B. Onteniente, H. Kimura, and T. Maeda, Comparative study of the glial fibrillary acidic protein in vertebrates by PAP immunohistochemistry, J Comp Neurol, vol.215, pp.427-436, 1983.

B. K. Ormerod, T. T. Lee, and L. A. Galea, Estradiol initially enhances but subsequently suppresses (via adrenal steroids) granule cell proliferation in the dentate gyrus of adult female rats, J Neurobiol, vol.55, pp.247-260, 2003.

B. K. Ormerod, T. T. Lee, and L. A. Galea, Estradiol enhances neurogenesis in the dentate gyri of adult male meadow voles by increasing the survival of young granule neurons, Neuroscience, vol.128, pp.645-654, 2004.

Y. Pang, J. Dong, T. , and P. , Estrogen signaling characteristics of Atlantic croaker G protein-coupled receptor 30 (GPR30) and evidence it is involved in maintenance of oocyte meiotic arrest, Endocrinology, vol.149, pp.3410-3426, 2008.

M. Pasmanik and G. V. Callard, Aromatase and 5 alpha-reductase in the teleost brain, spinal cord, and pituitary gland, Gen Comp Endocrinol, vol.60, pp.244-251, 1985.

M. Pasmanik and G. V. Callard, Changes in brain aromatase and 5 alpha-reductase activities correlate significantly with seasonal reproductive cycles in goldfish (Carassius auratus), Endocrinology, vol.122, pp.1349-1356, 1988.

M. Pasmanik, B. A. Schlinger, and G. V. Callard, In vivo steroid regulation of aromatase and 5 alpha-reductase in goldfish brain and pituitary, Gen Comp Endocrinol, vol.71, pp.175-182, 1988.

G. A. Peters and S. A. Khan, Estrogen receptor domains E and F: role in dimerization and interaction with coactivator RIP-140, Mol Endocrinol, vol.13, pp.286-296, 1999.

R. S. Peterson, G. Fernando, L. Day, T. A. Allen, J. D. Chapleau et al., Aromatase expression and cell proliferation following injury of the adult zebra finch hippocampus, Dev Neurobiol, vol.67, pp.1867-1878, 2007.

R. S. Peterson, D. W. Lee, G. Fernando, and B. A. Schlinger, Radial glia express aromatase in the injured zebra finch brain, J Comp Neurol, vol.475, pp.261-269, 2004.

R. S. Peterson, C. J. Saldanha, and B. A. Schlinger, Rapid upregulation of aromatase mRNA and protein following neural injury in the zebra finch (Taeniopygia guttata), J Neuroendocrinol, vol.13, pp.317-323, 2001.

R. S. Peterson, L. Yarram, B. A. Schlinger, and C. J. Saldanha, Aromatase is presynaptic and sexually dimorphic in the adult zebra finch brain, Proc Biol Sci, vol.272, pp.2089-2096, 2005.

D. W. Pfaff, Autoradiographic localization of radioactivity in rat brain after injection of tritiated sex hormones, Science, vol.161, pp.1355-1356, 1968.

D. W. Pfaff, Autoradiographic localization of testosterone-3H in the female rat brain and estradiol-3H in the male rat brain, Experientia, vol.24, pp.958-959, 1968.

F. Piferrer, I. J. Baker, and E. M. Donaldson, Effects of natural, synthetic, aromatizable, and nonaromatizable androgens in inducing male sex differentiation in genotypic female chinook salmon (Oncorhynchus tshawytscha), Gen Comp Endocrinol, vol.91, pp.59-65, 1993.

A. Pontious, T. Kowalczyk, C. Englund, and R. F. Hevner, Role of intermediate progenitor cells in cerebral cortex development, Dev Neurosci, vol.30, pp.24-32, 2008.

C. S. Potten, G. Owen, and D. Booth, Intestinal stem cells protect their genome by selective segregation of template DNA strands, J Cell Sci, vol.115, pp.2381-2388, 2002.

P. Rakic, Elusive radial glial cells: historical and evolutionary perspective, Glia, vol.43, pp.19-32, 2003.

A. M. Reugels, B. Boggetti, N. Scheer, and J. A. Campos-ortega, Asymmetric localization of Numb:EGFP in dividing neuroepithelial cells during neurulation in Danio rerio, Dev Dyn, vol.235, pp.934-948, 2006.

C. M. Revankar, D. F. Cimino, L. A. Sklar, J. B. Arterburn, and E. R. Prossnitz, A transmembrane intracellular estrogen receptor mediates rapid cell signaling, Science, vol.307, pp.1625-1630, 2005.

M. Robinson-rechavi, H. Garcia, and V. Laudet, The nuclear receptor superfamily, J Cell Sci, vol.116, pp.585-586, 2003.
URL : https://hal.archives-ouvertes.fr/hal-02670443

M. Robinson-rechavi, O. Marchand, H. Escriva, P. L. Bardet, D. Zelus et al., Euteleost fish genomes are characterized by expansion of gene families, 2001.

, Genome Res, vol.11, pp.781-788

C. E. Roselli and J. A. Resko, Effects of gonadectomy and androgen treatment on aromatase activity in the fetal monkey brain, Biol Reprod, vol.35, pp.106-112, 1986.

C. E. Roselli and J. A. Resko, Sex differences in androgen-regulated expression of cytochrome P450 aromatase in the rat brain, J Steroid Biochem Mol Biol, vol.61, pp.365-374, 1997.

C. F. Roselli, Brain aromatase: roles in reproduction and neuroprotection, J Steroid Biochem Mol Biol, vol.106, pp.143-150, 2007.

D. J. Rossi, J. D. Brady, and C. Mohr, Astrocyte metabolism and signaling during brain ischemia, Nat Neurosci, vol.10, pp.1377-1386, 2007.

H. Sakamoto, K. Ukena, and K. Tsutsui, Activity and localization of 3beta-hydroxysteroid dehydrogenase/ Delta5-Delta4-isomerase in the zebrafish central nervous system, J Comp Neurol, vol.439, pp.291-305, 2001.

M. Sancho-tello, S. Valles, C. Montoliu, J. Renau-piqueras, and C. Guerri, , 1995.

, Developmental pattern of GFAP and vimentin gene expression in rat brain and in radial glial cultures, Glia, vol.15, pp.157-166

B. Saville, M. Wormke, F. Wang, T. Nguyen, E. Enmark et al., Ligand-, cell-, and estrogen receptor subtype (alpha/beta)-dependent activation at GC-rich (Sp1) promoter elements, J Biol Chem, vol.275, pp.5379-5387, 2000.

M. Schartl, A comparative view on sex determination in medaka, Mech Dev, vol.121, pp.639-645, 2004.

B. A. Schlinger, S. Amur-umarjee, P. Shen, A. T. Campagnoni, A. et al., Neuronal and non-neuronal aromatase in primary cultures of developing zebra finch telencephalon, J Neurosci, vol.14, pp.7541-7552, 1994.

B. A. Schlinger and S. E. London, Neurosteroids and the songbird model system, J Exp Zoolog A Comp Exp Biol, vol.305, pp.743-748, 2006.

J. Schnitzer, W. W. Franke, and M. Schachner, Immunocytochemical demonstration of vimentin in astrocytes and ependymal cells of developing and adult mouse nervous system, J Cell Biol, vol.90, pp.435-447, 1981.

J. W. Schwabe, L. Chapman, J. T. Finch, D. Rhodes, and D. Neuhaus, DNA recognition by the oestrogen receptor: from solution to the crystal, Structure, vol.1, pp.187-204, 1993.

A. Sharif, P. Legendre, V. Prevot, C. Allet, L. Romao et al., Transforming growth factor alpha promotes sequential conversion of mature astrocytes into neural progenitors and stem cells, Oncogene, vol.26, pp.2695-2706, 2007.

Q. Shen, W. Zhong, Y. N. Jan, and S. Temple, Asymmetric Numb distribution is critical for asymmetric cell division of mouse cerebral cortical stem cells and neuroblasts, Development, vol.129, pp.4843-4853, 2002.

P. J. Shughrue, M. V. Lane, and I. Merchenthaler, Comparative distribution of estrogen receptor-alpha and -beta mRNA in the rat central nervous system, J Comp Neurol, vol.388, pp.507-525, 1997.

P. J. Shughrue, M. V. Lane, P. J. Scrimo, and I. Merchenthaler, Comparative distribution of estrogen receptor-alpha (ER-alpha) and beta (ER-beta) mRNA in the rat pituitary, gonad, and reproductive tract, Steroids, vol.63, pp.498-504, 1998.

R. B. Simerly, M. C. Zee, J. W. Pendleton, D. B. Lubahn, and K. S. Korach, , 1997.

, Estrogen receptor-dependent sexual differentiation of dopaminergic neurons in the preoptic region of the mouse, Proc Natl Acad Sci U S A, vol.94, pp.14077-14082

N. Stacey and M. Kobayashi, Androgen induction of male sexual behaviors in female goldfish, Horm Behav, vol.30, pp.434-445, 1996.

P. H. Strobl-mazzulla, N. P. Moncaut, G. C. Lopez, L. A. Miranda, A. V. Canario et al., Brain aromatase from pejerrey fish (Odontesthes bonariensis): cDNA cloning, tissue expression, and immunohistochemical localization, Gen Comp Endocrinol, vol.143, pp.21-32, 2005.

A. Tchoudakova and G. V. Callard, Identification of multiple CYP19 genes encoding different cytochrome P450 aromatase isozymes in brain and ovary, Endocrinology, vol.139, pp.2179-2189, 1998.

R. J. Timmers, J. G. Lambert, J. Peute, H. G. Vullings, and P. G. Van-oordt, , 1987.

, Localization of aromatase in the brain of the male African catfish, Clarias gariepinus (Burchell), by microdissection and biochemical identification, J Comp Neurol, vol.258, pp.368-377

S. Tomy, G. C. Wu, H. R. Huang, S. Dufour, C. et al., Developmental expression of key steroidogenic enzymes in the brain of protandrous black porgy fish, Acanthopagrus schlegeli, J Neuroendocrinol, vol.19, pp.643-655, 2007.
URL : https://hal.archives-ouvertes.fr/hal-00180889

C. D. Toran-allerand, X. Guan, N. J. Maclusky, T. L. Horvath, S. Diano et al., ER-X: a novel, plasma membrane-associated, putative estrogen receptor that is regulated during development and after ischemic brain injury, J Neurosci, vol.22, pp.8391-8401, 2002.

C. D. Toran-allerand, M. Singh, G. Setalo, and . Jr, Novel mechanisms of estrogen action in the brain: new players in an old story, Front Neuroendocrinol, vol.20, pp.97-121, 1999.

J. M. Trant, S. Gavasso, J. Ackers, B. C. Chung, and A. R. Place, Developmental expression of cytochrome P450 aromatase genes (CYP19a and CYP19b) in zebrafish fry (Danio rerio), J Exp Zool, vol.290, pp.475-483, 2001.

V. L. Trudeau, R. E. Peter, and B. D. Sloley, Testosterone and estradiol potentiate the serum gonadotropin response to gonadotropin-releasing hormone in goldfish, Biol Reprod, vol.44, pp.951-960, 1991.

D. Uchida, M. Yamashita, T. Kitano, and T. Iguchi, An aromatase inhibitor or high water temperature induce oocyte apoptosis and depletion of P450 aromatase activity in the gonads of genetic female zebrafish during sex-reversal, Comp Biochem Physiol A Mol Integr Physiol, vol.137, pp.11-20, 2004.

M. Ueno, K. Katayama, H. Yamauchi, H. Nakayama, and K. Doi, Cell cycle progression is required for nuclear migration of neural progenitor cells, Brain Res, vol.1088, pp.57-67, 2006.

L. D. Valle, A. Ramina, S. Vianello, P. Belvedere, and L. Colombo, Cloning of two mRNA variants of brain aromatase cytochrome P450 in rainbow trout (Oncorhynchus mykiss Walbaum), J Steroid Biochem Mol Biol, vol.82, pp.19-32, 2002.

S. Van-nes, M. Moe, and O. Andersen, Molecular characterization and expression of two cyp19 (P450 aromatase) genes in embryos, larvae, and adults of Atlantic halibut (Hippoglossus hippoglossus), Mol Reprod Dev, vol.72, pp.437-449, 2005.

S. Veiga, I. Azcoitia, and L. M. Garcia-segura, Extragonadal synthesis of estradiol is protective against kainic acid excitotoxic damage to the hippocampus, Neuroreport, vol.16, pp.1599-1603, 2005.

D. Vizziano, G. Randuineau, D. Baron, C. Cauty, and Y. Guiguen, Characterization of early molecular sex differentiation in rainbow trout, Oncorhynchus mykiss, Dev Dyn, vol.236, pp.2198-2206, 2007.
URL : https://hal.archives-ouvertes.fr/hal-02656188

W. Volknandt, Vesicular release mechanisms in astrocytic signalling, Neurochem Int, vol.41, pp.301-306, 2002.

B. J. Walters and C. J. Saldanha, Glial aromatization increases the expression of bone morphogenetic protein-2 in the injured zebra finch brain, J Neurochem, vol.106, pp.216-223, 2008.

L. Wang, S. Andersson, M. Warner, and J. A. Gustafsson, Morphological abnormalities in the brains of estrogen receptor beta knockout mice, Proc Natl Acad Sci U S A, vol.98, pp.2792-2796, 2001.

L. Wang, S. Andersson, M. Warner, and J. A. Gustafsson, Estrogen receptor (ER)beta knockout mice reveal a role for ERbeta in migration of cortical neurons in the developing brain, Proc Natl Acad Sci U S A, vol.100, pp.703-708, 2003.

Y. Wang and W. Ge, Cloning of zebrafish ovarian P450c17 (CYP17, 17alpha-hydroxylase/17, 20-lyase) and characterization of its expression in gonadal and extragonadal tissues, Gen Comp Endocrinol, vol.135, pp.241-249, 2004.

Z. Wang, G. Benoit, J. Liu, S. Prasad, P. Aarnisalo et al., Structure and function of Nurr1 identifies a class of ligandindependent nuclear receptors, Nature, vol.423, pp.555-560, 2003.

J. T. Watson and E. Adkins-regan, Testosterone implanted in the preoptic area of male Japanese quail must be aromatized to activate copulation, Horm Behav, vol.23, pp.432-447, 1989.

U. Wehrenberg, J. Prange-kiel, and G. M. Rune, Steroidogenic factor-1 expression in marmoset and rat hippocampus: co-localization with StAR and aromatase, J Neurochem, vol.76, pp.1879-1886, 2001.

R. D. Wynne, S. Maas, and C. J. Saldanha, Molecular characterization of the injuryinduced aromatase transcript in the adult zebra finch brain, J Neurochem, vol.105, pp.1613-1624, 2008.

R. D. Wynne and C. J. Saldanha, Glial aromatization decreases neural injury in the zebra finch (Taeniopygia guttata): influence on apoptosis, J Neuroendocrinol, vol.16, pp.676-683, 2004.

R. D. Wynne, B. J. Walters, D. J. Bailey, and C. J. Saldanha, Inhibition of injuryinduced glial aromatase reveals a wave of secondary degeneration in the songbird brain, Glia, vol.56, pp.97-105, 2008.

Z. Xia, R. Patino, W. L. Gale, A. G. Maule, and L. D. Densmore, Cloning, in vitro expression, and novel phylogenetic classification of a channel catfish estrogen receptor, 1999.

, Gen Comp Endocrinol, vol.113, pp.360-368

F. Xiong, D. Liu, Y. Le-drean, H. P. Elsholtz, and C. L. Hew, Differential recruitment of steroid hormone response elements may dictate the expression of the pituitary gonadotropin II beta subunit gene during salmon maturation, Mol Endocrinol, vol.8, pp.782-793, 1994.

J. G. Yague, E. Lavaque, J. Carretero, I. Azcoitia, and L. M. Garcia-segura, , 2004.

, Aromatase, the enzyme responsible for estrogen biosynthesis, is expressed by human and rat glioblastomas, Neurosci Lett, vol.368, pp.279-284

J. G. Yague, A. Munoz, P. De-monasterio-schrader, J. Defelipe, L. M. Garcia-segura et al., Aromatase expression in the human temporal cortex, Neuroscience, vol.138, pp.389-401, 2006.

K. Yamada and M. Watanabe, Cytodifferentiation of Bergmann glia and its relationship with Purkinje cells, Anat Sci Int, vol.77, pp.94-108, 2002.

D. Zhang and V. L. Trudeau, Integration of membrane and nuclear estrogen receptor signaling, Comp Biochem Physiol A Mol Integr Physiol, vol.144, pp.306-315, 2006.

Z. Zhang, R. Kumar, R. J. Santen, and R. X. Song, The role of adapter protein Shc in estrogen non-genomic action, Steroids, vol.69, pp.523-529, 2004.

J. Zhao, P. Mak, A. Tchoudakova, G. Callard, C. et al., Different catalytic properties and inhibitor responses of the goldfish brain and ovary aromatase isozymes, 2001.

, Comp Endocrinol, vol.123, pp.180-191

B. Zikopoulos, M. Kentouri, and C. R. Dermon, Proliferation zones in the adult brain of a sequential hermaphrodite teleost species (Sparus aurata), Brain Behav Evol, vol.56, pp.310-322, 2000.

G. K. Zupanc, Neurogenesis and neuronal regeneration in the adult fish brain, J Comp Physiol A Neuroethol Sens Neural Behav Physiol, vol.192, pp.649-670, 2006.

G. K. Zupanc, C. , and S. C. , Potential role of radial glia in adult neurogenesis of teleost fish, Glia, vol.43, pp.77-86, 2003.

G. K. Zupanc and I. Horschke, Proliferation zones in the brain of adult gymnotiform fish: a quantitative mapping study, J Comp Neurol, vol.353, pp.213-233, 1995.

, Karen MOURIEC Régulation de l'aromatase B dans les cellules gliales radiaires dans le cerveau du poisson zèbre (Danio rerio) et rôles potentiels dans la neurogénèse adulte

, Résumé Chez les mammifères, la neurogénèse adulte est limitée, au contraire, chez les poissons téléostéens

, Mots clés: aromatase B, oestradiol, récepteurs aux oestrogènes, neurogénèse, cellules gliales

, Key words: aromatase B, estradiol, estrogen receptors, neurogenesis, radial glial cells