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Juliana I Candelaria Department of Animal Science, University of California Davis, Davis, California, USA

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Maria B Rabaglino Department of Applied Mathematics and Computer Science, Technical University of Denmark, Kongens Lyngby, Denmark

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Anna C Denicol Department of Animal Science, University of California Davis, Davis, California, USA

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Introduction Ovarian antral follicles require gonadotropin stimulation for development and ovulation whereas preantral follicles seem capable of development in the absence of follicle-stimulating hormone (FSH) and luteinizing hormone (LH

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Michael W Pankhurst Department of Anatomy, School of Biomedical Sciences, University of Otago, Dunedin, New Zealand

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Rebecca L Kelley School of Biosciences, University of Melbourne, Melbourne, Australia

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Rachel L Sanders Department of Anatomy, School of Biomedical Sciences, University of Otago, Dunedin, New Zealand

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Savana R Woodcock Department of Anatomy, School of Biomedical Sciences, University of Otago, Dunedin, New Zealand

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Dorothy E Oorschot Department of Anatomy, School of Biomedical Sciences, University of Otago, Dunedin, New Zealand
Brain Health Research Centre, University of Otago, Dunedin, New Zealand

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Nicola J Batchelor Department of Anatomy, School of Biomedical Sciences, University of Otago, Dunedin, New Zealand

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primordial follicles leads to transition to the primary stage. Subsequent progression through the preantral stages is facilitated by oocyte growth and proliferation of the associated granulosa and theca cells. The antral stages are characterized by the

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Marie M Devillers Sorbonne Paris Cité, Université Paris-Diderot, CNRS, INSERM, Biologie Fonctionnelle et Adaptative UMR 8251, Physiologie de l’Axe Gonadotrope U1133, Paris, France

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Florence Petit Sorbonne Paris Cité, Université Paris-Diderot, CNRS, INSERM, Biologie Fonctionnelle et Adaptative UMR 8251, Physiologie de l’Axe Gonadotrope U1133, Paris, France

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Victoria Cluzet Sorbonne Paris Cité, Université Paris-Diderot, CNRS, INSERM, Biologie Fonctionnelle et Adaptative UMR 8251, Physiologie de l’Axe Gonadotrope U1133, Paris, France

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Charlotte M François Sorbonne Paris Cité, Université Paris-Diderot, CNRS, INSERM, Biologie Fonctionnelle et Adaptative UMR 8251, Physiologie de l’Axe Gonadotrope U1133, Paris, France

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Frank Giton APHP CIB GHU Sud Henri Mondor, INSERM IMRB U955, Eq.07, Faculté de Médecine, Créteil, France

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Ghislaine Garrel Sorbonne Paris Cité, Université Paris-Diderot, CNRS, INSERM, Biologie Fonctionnelle et Adaptative UMR 8251, Physiologie de l’Axe Gonadotrope U1133, Paris, France

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Joëlle Cohen-Tannoudji Sorbonne Paris Cité, Université Paris-Diderot, CNRS, INSERM, Biologie Fonctionnelle et Adaptative UMR 8251, Physiologie de l’Axe Gonadotrope U1133, Paris, France

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Céline J Guigon Sorbonne Paris Cité, Université Paris-Diderot, CNRS, INSERM, Biologie Fonctionnelle et Adaptative UMR 8251, Physiologie de l’Axe Gonadotrope U1133, Paris, France

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–14 days postnatal (dpn), reaching about five-fold those found in adult cycling females ( François et al. 2017 ). We demonstrated that such FSH concentrations are necessary to induce Cyp19a1 aromatase expression in immature follicles at the preantral

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Rachel A Forsdike Institute of Reproductive and Developmental Biology, Imperial College London, Hammersmith Hospital, Du Cane Road, London W12 0NN, UK
Department of Mathematics, Imperial College London, London SW7 2AZ, UK
Laboratory of Neuroendocrinology, Department of Neurobiology, The Babraham Institute, Babraham, Cambridge CB2 4AT, UK

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Kate Hardy Institute of Reproductive and Developmental Biology, Imperial College London, Hammersmith Hospital, Du Cane Road, London W12 0NN, UK
Department of Mathematics, Imperial College London, London SW7 2AZ, UK
Laboratory of Neuroendocrinology, Department of Neurobiology, The Babraham Institute, Babraham, Cambridge CB2 4AT, UK

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Lauren Bull Institute of Reproductive and Developmental Biology, Imperial College London, Hammersmith Hospital, Du Cane Road, London W12 0NN, UK
Department of Mathematics, Imperial College London, London SW7 2AZ, UK
Laboratory of Neuroendocrinology, Department of Neurobiology, The Babraham Institute, Babraham, Cambridge CB2 4AT, UK

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Jaroslav Stark Institute of Reproductive and Developmental Biology, Imperial College London, Hammersmith Hospital, Du Cane Road, London W12 0NN, UK
Department of Mathematics, Imperial College London, London SW7 2AZ, UK
Laboratory of Neuroendocrinology, Department of Neurobiology, The Babraham Institute, Babraham, Cambridge CB2 4AT, UK

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Lisa J Webber Institute of Reproductive and Developmental Biology, Imperial College London, Hammersmith Hospital, Du Cane Road, London W12 0NN, UK
Department of Mathematics, Imperial College London, London SW7 2AZ, UK
Laboratory of Neuroendocrinology, Department of Neurobiology, The Babraham Institute, Babraham, Cambridge CB2 4AT, UK

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Sharron Stubbs Institute of Reproductive and Developmental Biology, Imperial College London, Hammersmith Hospital, Du Cane Road, London W12 0NN, UK
Department of Mathematics, Imperial College London, London SW7 2AZ, UK
Laboratory of Neuroendocrinology, Department of Neurobiology, The Babraham Institute, Babraham, Cambridge CB2 4AT, UK

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Jane E Robinson Institute of Reproductive and Developmental Biology, Imperial College London, Hammersmith Hospital, Du Cane Road, London W12 0NN, UK
Department of Mathematics, Imperial College London, London SW7 2AZ, UK
Laboratory of Neuroendocrinology, Department of Neurobiology, The Babraham Institute, Babraham, Cambridge CB2 4AT, UK

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Stephen Franks Institute of Reproductive and Developmental Biology, Imperial College London, Hammersmith Hospital, Du Cane Road, London W12 0NN, UK
Department of Mathematics, Imperial College London, London SW7 2AZ, UK
Laboratory of Neuroendocrinology, Department of Neurobiology, The Babraham Institute, Babraham, Cambridge CB2 4AT, UK

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reflects the arrest of follicle development in the late antral stages but recent work from our group ( Webber et al. 2003 ), and that of Erickson and colleagues ( Maciel et al. 2004 ), has highlighted a disorder of early, preantral follicle development

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Mehmet Uzumcu Department of Animal Sciences, School of Environmental and Biological Sciences, Rutgers, The State University of New Jersey, 84 Lipman Drive, New Brunswick, New Jersey 08901-8525, USA

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Peter E Kuhn Department of Animal Sciences, School of Environmental and Biological Sciences, Rutgers, The State University of New Jersey, 84 Lipman Drive, New Brunswick, New Jersey 08901-8525, USA

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Jason E Marano Department of Animal Sciences, School of Environmental and Biological Sciences, Rutgers, The State University of New Jersey, 84 Lipman Drive, New Brunswick, New Jersey 08901-8525, USA

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AnnMarie E Armenti Department of Animal Sciences, School of Environmental and Biological Sciences, Rutgers, The State University of New Jersey, 84 Lipman Drive, New Brunswick, New Jersey 08901-8525, USA

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Lisa Passantino Department of Animal Sciences, School of Environmental and Biological Sciences, Rutgers, The State University of New Jersey, 84 Lipman Drive, New Brunswick, New Jersey 08901-8525, USA

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three layers of cuboidal granulosa cells. Pre-antral: an oocyte surrounded by more than three layers of granulosa cells with no apparent antrum. Antral: an oocyte surrounded by multiple layers of granulosa cells with an antrum. Unknown follicles

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Christopher R Harlow University of Edinburgh Centre for Reproductive Biology, Queen’s Medical Research Institute, 47 Little France Crescent, Edinburgh EH16 4TJ, UK

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Angela C Bradshaw University of Edinburgh Centre for Reproductive Biology, Queen’s Medical Research Institute, 47 Little France Crescent, Edinburgh EH16 4TJ, UK

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Michael T Rae University of Edinburgh Centre for Reproductive Biology, Queen’s Medical Research Institute, 47 Little France Crescent, Edinburgh EH16 4TJ, UK

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Kirsty D Shearer University of Edinburgh Centre for Reproductive Biology, Queen’s Medical Research Institute, 47 Little France Crescent, Edinburgh EH16 4TJ, UK

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Stephen G Hillier University of Edinburgh Centre for Reproductive Biology, Queen’s Medical Research Institute, 47 Little France Crescent, Edinburgh EH16 4TJ, UK

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), chondrocytes ( Eguchi et al. 2001 ), vascular smooth muscle cells ( Fan et al. 2000 ) and renal mesengial cells ( Goppelt-Struebe et al. 2001 ). We previously showed that rat granulosa cells in preantral and early antral follicles abundantly

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M F Machado
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V M Portela Departamento de Reprodução Animal, Centre de recherche en reproduction animale, Departamento de Fisiologia, Departamento de Clínica Veterinária, Faculdade de Medicina Veterinária e Zootecnia, Universidade Estadual Paulista, Botucatu, São Paulo 18618-000, Brazil

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C A Price Departamento de Reprodução Animal, Centre de recherche en reproduction animale, Departamento de Fisiologia, Departamento de Clínica Veterinária, Faculdade de Medicina Veterinária e Zootecnia, Universidade Estadual Paulista, Botucatu, São Paulo 18618-000, Brazil

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I B Costa
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P Ripamonte Departamento de Reprodução Animal, Centre de recherche en reproduction animale, Departamento de Fisiologia, Departamento de Clínica Veterinária, Faculdade de Medicina Veterinária e Zootecnia, Universidade Estadual Paulista, Botucatu, São Paulo 18618-000, Brazil

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R L Amorim Departamento de Reprodução Animal, Centre de recherche en reproduction animale, Departamento de Fisiologia, Departamento de Clínica Veterinária, Faculdade de Medicina Veterinária e Zootecnia, Universidade Estadual Paulista, Botucatu, São Paulo 18618-000, Brazil

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J Buratini Jr Departamento de Reprodução Animal, Centre de recherche en reproduction animale, Departamento de Fisiologia, Departamento de Clínica Veterinária, Faculdade de Medicina Veterinária e Zootecnia, Universidade Estadual Paulista, Botucatu, São Paulo 18618-000, Brazil

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Institute, Cary, NC, USA). Results Immunohistochemistry revealed the presence of FGF17 predominantly in oocytes and granulosa cells of preantral and antral follicles ( Fig. 1 ). Staining was predominant in the nucleus of oocytes in preantral follicles, and

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Karin A Slot Department of Biochemistry & Cell Biology, Faculty of Veterinary Medicine, Utrecht University, Yalelaan 2, 3508 TD Utrecht, The Netherlands
Human and Animal Physiology Group, Department of Animal Sciences, Wageningen University, Haarweg 10, 6709 PJ Wageningen, The Netherlands

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Marsha Voorendt Department of Biochemistry & Cell Biology, Faculty of Veterinary Medicine, Utrecht University, Yalelaan 2, 3508 TD Utrecht, The Netherlands
Human and Animal Physiology Group, Department of Animal Sciences, Wageningen University, Haarweg 10, 6709 PJ Wageningen, The Netherlands

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Mieke de Boer-Brouwer Department of Biochemistry & Cell Biology, Faculty of Veterinary Medicine, Utrecht University, Yalelaan 2, 3508 TD Utrecht, The Netherlands
Human and Animal Physiology Group, Department of Animal Sciences, Wageningen University, Haarweg 10, 6709 PJ Wageningen, The Netherlands

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Harmke H van Vugt Department of Biochemistry & Cell Biology, Faculty of Veterinary Medicine, Utrecht University, Yalelaan 2, 3508 TD Utrecht, The Netherlands
Human and Animal Physiology Group, Department of Animal Sciences, Wageningen University, Haarweg 10, 6709 PJ Wageningen, The Netherlands

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Katja J Teerds Department of Biochemistry & Cell Biology, Faculty of Veterinary Medicine, Utrecht University, Yalelaan 2, 3508 TD Utrecht, The Netherlands
Human and Animal Physiology Group, Department of Animal Sciences, Wageningen University, Haarweg 10, 6709 PJ Wageningen, The Netherlands

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(++). Preantral follicles     Healthy granulosa

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Tatiane da Silva Faria Urogenital Research Unit-UERJ, State University of Rio de Janeiro, Avenue 28 de Setembro, 87, Fundos, FCM, Terreo, 20551-030 Rio de Janeiro, RJ, Brazil

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Flávia de Bittencourt Brasil Urogenital Research Unit-UERJ, State University of Rio de Janeiro, Avenue 28 de Setembro, 87, Fundos, FCM, Terreo, 20551-030 Rio de Janeiro, RJ, Brazil

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Francisco J B Sampaio Urogenital Research Unit-UERJ, State University of Rio de Janeiro, Avenue 28 de Setembro, 87, Fundos, FCM, Terreo, 20551-030 Rio de Janeiro, RJ, Brazil

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Cristiane da Fonte Ramos Urogenital Research Unit-UERJ, State University of Rio de Janeiro, Avenue 28 de Setembro, 87, Fundos, FCM, Terreo, 20551-030 Rio de Janeiro, RJ, Brazil

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final magnification of X400 for primordial and primary follicles, and X100 for preantral, antral, and Graafian follicles and corpus luteum. Photographs of ovarian follicles were analyzed using Image Pro Plus for Windows (version 1.3.2; Media Cybernetics

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H Quesnel
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Binding sites for IGF-I, insulin, and GH were localized by in situ binding of (125)I-labelled hormones to the different compartments of the sow ovary. Binding sites for IGF-I were detected in oocytes, granulosa and thecal cells of healthy and atretic follicles as well as in the antrum and the stroma. Competition of (125)I-labelled IGF-I with IGF-I, insulin and an analogue of IGF-I (Long R(3)IGF-I), which allowed discrimination between binding to binding proteins from binding to type-I receptors, suggested that type-I receptors were present in granulosa cells of healthy follicles, whilst binding in other compartments was mainly due to binding proteins. Binding of insulin was revealed in oocytes, granulosa and theca interna cells of healthy preantral and antral follicles, and, to a lesser extent, in theca externa and stromal cells, and was still observed in granulosa cells of atretic follicles. Labelling with (125)I-labelled bovine GH was demonstrated in oocytes, granulosa cells, theca interna cells, and, although less intense, in theca externa and stromal cells. It disappeared in granulosa cells during atresia. Binding sites for GH were detected at all follicular stages, from preantral to preovulatory stages, but the intensity of labelling in granulosa cells was more intense in preantral than in large follicles. These data support the participation of insulin, GH and IGF-I in oocyte maturation, follicular growth and stromal cell function in swine.

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