C. Glidewell-kenney, Nonclassical estrogen receptor ?? signaling mediates negative feedback in the female mouse reproductive axis, Proceedings of the National Academy of Sciences, vol.104, issue.19, pp.8173-8177, 2007.
DOI : 10.1073/pnas.0611514104

G. Olmos, Synaptic remodeling in the rat arcuate nucleus during the estrous cycle, Neuroscience, vol.32, issue.3, pp.663-667, 1989.
DOI : 10.1016/0306-4522(89)90288-1

M. Langub, B. Maley, and R. Watson, Estrous cycle-associated axosomatic synaptic plasticity upon estrogen receptive neurons in the rat preoptic area, Brain Research, vol.641, issue.2, pp.303-310, 1994.
DOI : 10.1016/0006-8993(94)90159-7

A. Parducz, Synaptic remodeling induced by gonadal hormones: Neuronal plasticity as a mediator of neuroendocrine and behavioral responses to steroids, Neuroscience, vol.138, issue.3, pp.977-985, 2006.
DOI : 10.1016/j.neuroscience.2005.07.008

R. Cohen and D. Pfaff, Ultrastructure of neurons in the ventromedial nucleus or the hypothalamus in ovariectomized rats with or without estrogen treatment, Cell and Tissue Research, vol.217, issue.3, pp.451-470, 1981.
DOI : 10.1007/BF00219357

E. Gould, C. Woolley, M. Frankfurt, and B. Mcewen, Gonadal steroids regulate dendritic spine density in hippocampal pyramidal cells in adulthood, J Neurosci, vol.10, pp.1286-1291, 1990.

C. Woolley, E. Gould, M. Frankfurt, and B. Mcewen, Naturally occurring fluctuation in dendritic spine density on adult hippocampal pyramidal neurons, J Neurosci, vol.10, pp.4035-4039, 1990.

F. Naftolin, Estrogen-Induced Hypothalamic Synaptic Plasticity and Pituitary Sensitization in the Control of the Estrogen-Induced Gonadotrophin Surge, Reproductive Sciences, vol.14, issue.2, pp.101-116, 2007.
DOI : 10.1177/1933719107301059

H. Mukai, Rapid modulation of long-term depression and spinogenesis via synaptic estrogen receptors in hippocampal principal neurons, Journal of Neurochemistry, vol.17, issue.4, pp.950-967, 2007.
DOI : 10.1111/j.1471-4159.2006.04264.x

URL : https://hal.archives-ouvertes.fr/hal-00190993

H. Carlisle and M. Kennedy, Spine architecture and synaptic plasticity, Trends in Neurosciences, vol.28, issue.4, pp.182-187, 2005.
DOI : 10.1016/j.tins.2005.01.008

A. Mazumdar and R. Kumar, Estrogen regulation of Pak1 and FKHR pathways in breast cancer cells, FEBS Letters, vol.276, issue.1-3, pp.6-10, 2003.
DOI : 10.1016/S0014-5793(02)03846-2

T. Tsakiridis, C. Taha, S. Grinstein, and A. Klip, Insulin Activates a p21-activated Kinase in Muscle Cells via Phosphatidylinositol 3-Kinase, Journal of Biological Chemistry, vol.271, issue.33, pp.19664-19667, 1996.
DOI : 10.1074/jbc.271.33.19664

M. Nikolic, The Pak1 Kinase: An Important Regulator of Neuronal Morphology and Function in the Developing Forebrain, Molecular Neurobiology, vol.281, issue.2-3, pp.187-202, 2008.
DOI : 10.1007/s12035-008-8032-1

S. Rayala and R. Kumar, Sliding p21-activated kinase 1 to nucleus impacts tamoxifen sensitivity, Biomedicine & Pharmacotherapy, vol.61, issue.7, pp.408-411, 2007.
DOI : 10.1016/j.biopha.2007.05.006

E. Manser, PAK Kinases Are Directly Coupled to the PIX Family of Nucleotide Exchange Factors, Molecular Cell, vol.1, issue.2, pp.183-192, 1998.
DOI : 10.1016/S1097-2765(00)80019-2

L. Zhao, Role of p21-activated kinase pathway defects in the cognitive deficits of Alzheimer disease, Nature Neuroscience, vol.152, issue.2, pp.234-242, 2006.
DOI : 10.1523/JNEUROSCI.2860-04.2004

P. Penzes, Rapid Induction of Dendritic Spine Morphogenesis by trans-Synaptic EphrinB-EphB Receptor Activation of the Rho-GEF Kalirin, Neuron, vol.37, issue.2, pp.263-274, 2003.
DOI : 10.1016/S0896-6273(02)01168-6

H. Maruta, H. He, A. Tikoo, and M. Nur-e-kamal, Cytoskeletal Tumor Suppressors That Block Oncogenic RAS Signaling, Annals of the New York Academy of Sciences, vol.8, issue.1 ANTICANCER MO, pp.48-57, 1999.
DOI : 10.1016/S0968-0004(97)01097-9

A. Obermeier, PAK promotes morphological changes by acting upstream of Rac, The EMBO Journal, vol.17, issue.15, pp.4328-4339, 1998.
DOI : 10.1093/emboj/17.15.4328

T. Nheu, PAK Is Essential for RAS-Induced Upregulation of Cyclin D1 During the G1 to S Transition, Cell Cycle, vol.3, issue.1, pp.71-74, 2004.
DOI : 10.4161/cc.3.1.593

P. Wender, The design, synthesis, and evaluation of molecules that enable or enhance cellular uptake: Peptoid molecular transporters, Proceedings of the National Academy of Sciences, vol.97, issue.24, pp.13003-13008, 2000.
DOI : 10.1073/pnas.97.24.13003

M. Aarts, Treatment of Ischemic Brain Damage by Perturbing NMDA Receptor- PSD-95 Protein Interactions, Science, vol.298, issue.5594, pp.846-850, 2002.
DOI : 10.1126/science.1072873

G. Cao, In vivo delivery of a Bcl-xL fusion protein containing the TAT protein transduction domain protects against ischemic brain injury and neuronal apoptosis, J Neurosci, vol.22, pp.5423-5431, 2002.

H. Kusuhara and Y. Sugiyama, Efflux transport systems for drugs at the blood???brain barrier and blood???cerebrospinal fluid barrier (Part 2), Drug Discovery Today, vol.6, issue.4, pp.206-212, 2001.
DOI : 10.1016/S1359-6446(00)01643-3

J. Arreguin-arevalo and T. Nett, A Nongenomic Action of Estradiol as the Mechanism Underlying the Acute Suppression of Secretion of Luteinizing Hormone in Ovariectomized Ewes1, Biology of Reproduction, vol.74, issue.1, pp.202-208, 2006.
DOI : 10.1095/biolreprod.105.044685

M. Kelly, O. Ronnekleiv, and R. Eskay, Identification of estrogen-responsive LHRH Neurons in the guinea pig hypothalamus, Brain Research Bulletin, vol.12, issue.4, pp.399-407, 1984.
DOI : 10.1016/0361-9230(84)90112-6

I. Abraham, Estrogen receptor beta mediates rapid estrogen actions on gonadotropin-releasing hormone neurons in vivo, J Neurosci, vol.23, pp.5771-5777, 2003.

J. Qiu, Rapid signaling of estrogen in hypothalamic neurons involves a novel G-protein-coupled estrogen receptor that activates protein kinase C, J Neurosci, vol.23, pp.9529-9540, 2003.

F. Strobl, C. Gilmore, and J. Levine, Castration Induces Luteinizing Hormone (LH) Secretion in Hypophysectomized Pituitary-Grafted Rats Receiving Pulsatile LH-Releasing Hormone Infusions*, Endocrinology, vol.124, issue.3, pp.1140-1144, 1989.
DOI : 10.1210/endo-124-3-1140

C. Glidewell-kenney, Estrogen Receptor ?? Signaling Pathways Differentially Regulate Gonadotropin Subunit Gene Expression and Serum Follicle-Stimulating Hormone in the Female Mouse, Endocrinology, vol.149, issue.8, pp.4168-4176, 2008.
DOI : 10.1210/en.2007-1807

M. Nikolic, The p35/Cdk5 kinase is a neuron-specific Rac effector that inhibits Pak1 activity, Nature, vol.395, pp.194-198, 1998.

D. Thiel, Cell Cycle-Regulated Phosphorylation of p21-Activated Kinase 1, Current Biology, vol.12, issue.14, pp.1227-1232, 2002.
DOI : 10.1016/S0960-9822(02)00931-4

L. Sundberg-smith, J. Doherty, C. Mack, and J. Taylor, Adhesion Stimulates Direct PAK1/ERK2 Association and Leads to ERK-dependent PAK1 Thr212 Phosphorylation, Journal of Biological Chemistry, vol.280, issue.3, pp.2055-2064, 2005.
DOI : 10.1074/jbc.M406013200

L. Adam, Heregulin Regulates Cytoskeletal Reorganization and Cell Migration through the p21-activated Kinase-1 via Phosphatidylinositol-3 Kinase, Journal of Biological Chemistry, vol.273, issue.43, pp.28238-28246, 1998.
DOI : 10.1074/jbc.273.43.28238

S. Kato, Activation of the Estrogen Receptor Through Phosphorylation by Mitogen-Activated Protein Kinase, Science, vol.270, issue.5241, pp.1491-1494, 1995.
DOI : 10.1126/science.270.5241.1491

M. Sun, Phosphatidylinositol-3-OH Kinase (PI3K)/AKT2, activated in breast cancer, regulates and is induced by estrogen receptor alpha (ERalpha) via interaction between ERalpha and PI3K, Cancer Res, vol.61, pp.5985-5991, 2001.

I. Abraham, M. Todman, K. Korach, and A. Herbison, Roles for Classical Estrogen Receptors in Rapid Estrogen Actions on Intracellular Signaling in Mouse Brain, Endocrinology, vol.145, issue.7, pp.3055-3061, 2004.
DOI : 10.1210/en.2003-1676

G. Murakami, Comparison between basal and apical dendritic spines in estrogen-induced rapid spinogenesis of CA1 principal neurons in the adult hippocampus, Biochemical and Biophysical Research Communications, vol.351, issue.2, pp.553-558, 2006.
DOI : 10.1016/j.bbrc.2006.10.066

D. Srivastava, Rapid enhancement of two-step wiring plasticity by estrogen and NMDA receptor activity, Proceedings of the National Academy of Sciences, vol.105, issue.38, pp.14650-14655, 2008.
DOI : 10.1073/pnas.0801581105

K. Hayashi, T. Ohshima, M. Hashimoto, and K. Mikoshiba, Pak1 regulates dendritic branching and spine formation, Developmental Neurobiology, vol.18, issue.5, pp.655-669, 2007.
DOI : 10.1002/dneu.20363

Y. Meng, Abnormal Spine Morphology and Enhanced LTP in LIMK-1 Knockout Mice, Neuron, vol.35, issue.1, pp.121-133, 2002.
DOI : 10.1016/S0896-6273(02)00758-4

B. Boda, The Mental Retardation Protein PAK3 Contributes to Synapse Formation and Plasticity in Hippocampus, Journal of Neuroscience, vol.24, issue.48, pp.10816-10825, 2004.
DOI : 10.1523/JNEUROSCI.2931-04.2004

K. Tolias, The Rac1-GEF Tiam1 Couples the NMDA Receptor to the Activity-Dependent Development of Dendritic Arbors and Spines, Neuron, vol.45, issue.4, pp.525-538, 2005.
DOI : 10.1016/j.neuron.2005.01.024

L. Zhao and R. Brinton, Estrogen receptor ?? and ?? differentially regulate intracellular Ca2+ dynamics leading to ERK phosphorylation and estrogen neuroprotection in hippocampal neurons, Brain Research, vol.1172, pp.48-59, 2007.
DOI : 10.1016/j.brainres.2007.06.092

E. Hrabovszky, Estrogen receptor-beta immunoreactivity in luteinizing hormone-releasing hormone neurons of the rat brain, Endocrinology, vol.142, pp.3261-3264, 2001.

C. Christian and S. Moenter, Estradiol Induces Diurnal Shifts in GABA Transmission to Gonadotropin-Releasing Hormone Neurons to Provide a Neural Signal for Ovulation, Journal of Neuroscience, vol.27, issue.8, pp.1913-1921, 2007.
DOI : 10.1523/JNEUROSCI.4738-06.2007

T. Milner, Ultrastructural evidence that hippocampal alpha estrogen receptors are located at extranuclear sites, The Journal of Comparative Neurology, vol.389, issue.3, pp.355-371, 2001.
DOI : 10.1002/1096-9861(20010115)429:3<355::AID-CNE1>3.0.CO;2-#

S. Hart, M. Snyder, T. Smejkalova, and C. Woolley, Estrogen Mobilizes a Subset of Estrogen Receptor-??-Immunoreactive Vesicles in Inhibitory Presynaptic Boutons in Hippocampal CA1, Journal of Neuroscience, vol.27, issue.8, pp.2102-2111, 2007.
DOI : 10.1523/JNEUROSCI.5436-06.2007

J. Blaustein, M. Lehman, J. Turcotte, and G. Greene, Estrogen receptors in dendrites and axon terminals in the guinea pig hypothalamus, Endocrinology, vol.131, pp.281-290, 1992.

N. Romano, Nonclassical Estrogen Modulation of Presynaptic GABA Terminals Modulates Calcium Dynamics in Gonadotropin-Releasing Hormone Neurons, Endocrinology, vol.149, issue.11, pp.5335-5344, 2008.
DOI : 10.1210/en.2008-0424

M. Mcdevitt, Estrogen Response Element-Independent Estrogen Receptor (ER)-?? Signaling Does Not Rescue Sexual Behavior but Restores Normal Testosterone Secretion in Male ER?? Knockout Mice, Endocrinology, vol.148, issue.11, pp.5288-5294, 2007.
DOI : 10.1210/en.2007-0673

URL : https://hal.archives-ouvertes.fr/hal-00192063

S. Dupont, Effect of single and compound knockouts of estrogen receptors alpha (ERalpha) and beta (ERbeta) on mouse reproductive phenotypes, Development, vol.127, pp.4277-4291, 2000.

G. Gu, Hormonal regulation of CREB phosphorylation in the anteroventral periventricular nucleus, J Neurosci, vol.16, pp.3035-3044, 1996.

P. Mellon, Immortalization of hypothalamic GnRH by genetically targeted tumorigenesis, Neuron, vol.5, issue.1, pp.1-10, 1990.
DOI : 10.1016/0896-6273(90)90028-E

M. Kelly, R. Moss, and C. Dudley, Differential sensitivity of preoptic-septal neurons to microelectrophoressed estrogen during the estrous cycle, Brain Research, vol.114, issue.1, pp.152-157, 1976.
DOI : 10.1016/0006-8993(76)91017-9

C. Christian, C. Glidewell-kenney, J. Jameson, and S. Moenter, Classical Estrogen Receptor ?? Signaling Mediates Negative and Positive Feedback on Gonadotropin-Releasing Hormone Neuron Firing, Endocrinology, vol.149, issue.11, pp.5328-5334, 2008.
DOI : 10.1210/en.2008-0520

URL : http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2584581