Neuroendocrine regulation of GnRH release in induced ovulators. (original) (raw)
Article (Scientific journals)
Neuroendocrine regulation of GnRH release in induced ovulators.
; Baum, M. J.
2000 • In Frontiers in Neuroendocrinology, 21 (3), p. 220-62
Peer Reviewed verified by ORBi
Files (1)Send toDetailsStatisticsBibliographySimilar publications
Files
Full Text
All documents in ORBi are protected by a user license.
Send to
Details
Keywords :
Animals; Female; Follicular Phase/metabolism; Gonadotropin-Releasing Hormone/metabolism; Hormones/metabolism/physiology; Humans; Luteinizing Hormone/metabolism; Neuropeptides/physiology; Neurosecretory Systems/physiology; Neurotransmitter Agents/physiology; Ovary/metabolism; Ovulation/physiology; Receptors, LHRH/physiology
Abstract :
[en] GnRH is the key neuropeptide controlling reproductive function in all vertebrate species. Two different neuroendocrine mechanisms have evolved among female mammals to regulate the mediobasal hypothalamic (MBH) release of GnRH leading to the preovulatory secretion of LH by the anterior pituitary gland. In females of spontaneously ovulating species, including rats, mice, guinea pigs, sheep, monkeys, and women, ovarian steroids secreted by maturing ovarian follicles induce a pulsatile pattern of GnRH release in the median eminence that, in turn, stimulates a preovulatory LH surge. In females of induced ovulating species, including rabbits, ferrets, cats, and camels, the preovulatory release of GnRH, and the resultant preovulatory LH surge, is induced by the receipt of genital somatosensory stimuli during mating. Induced ovulators generally do not show "spontaneous" steroid-induced LH surges during their reproductive cycles, suggesting that the positive feedback actions of steroid hormones on GnRH release are reduced or absent in these species. By contrast, mating-induced preovulatory surges occasionally occur in some spontaneously ovulating species. Most research in the field of GnRH neurobiology has been performed using spontaneous ovulators including rat, guinea pig, sheep, and rhesus monkey. This review summarizes the literature concerning the neuroendocrine mechanisms controlling GnRH biosynthesis and release in females of several induced ovulating species, and whenever possible it contrasts the results with those obtained for spontaneously ovulating species. It also considers the adaptive, evolutionary benefits and disadvantages of each type of ovulatory control mechanism. In females of induced ovulating species estradiol acts in the brain to induce aspects of proceptive and receptive sexual behavior. The primary mechanism involved in the preovulatory release of GnRH among induced ovulators involves the activation of midbrain and brainstem noradrenergic neurons in response to genital-somatosensory signals generated by receipt of an intromission from a male during mating. These noradrenergic neurons project to the MBH and, when activated, promote the release of GnRH from nerve terminals in the median eminence. In contrast to spontaneous ovulators, there is little evidence that endogenous opioid peptides normally inhibit MBH GnRH release among induced ovulators. Instead, the neural signals that induce a preovulatory LH surge in these species seem to be primarily excitatory. A complete understanding of the neuroendocrine control of ovulation will only be achieved in the future by comparative studies of several animal model systems in which mating-induced as well as spontaneous, hormonally stimulated activation of GnRH neurons drives the preovulatory LH surge.
Research Center/Unit :
Boston University
Disciplines :
Neurosciences & behavior
Endocrinology, metabolism & nutrition
Author, co-author :
Bakker, Julie ; Université de Liège - ULiège > Département des sciences biomédicales et précliniques > Biologie de la différenciation sexuelle du cerveau
Baum, M. J.
Title :
Neuroendocrine regulation of GnRH release in induced ovulators.
Journal title :
Frontiers in Neuroendocrinology
Publisher :
Academic Press, Orlando, United States - Florida
Peer reviewed :
Peer Reviewed verified by ORBi
Funders :
NIH - National Institutes of Health
Commentary :
Copyright 2000 Academic Press.
Available on ORBi :
since 18 May 2011
Statistics
Number of views
1058 (6 by ULiège)
Number of downloads
1759 (7 by ULiège)
Scopus citations®
without self-citations
175
Bibliography
- Adams C E. Observations on the induction of ovulation and expulsion of uterine eggs in the mink, Mustela vision. J Reprod Fertil. 63:1981;241-248.
- Adelman J P, Mason A J, Hayflick J S, Seeburg P H. Isolation of the gene and hypothalamic cDNA for the common precursor of gonadotropin-releasing hormone and prolactin release-inhibiting factor in human and rat. Proc Natl Acad Sci USA. 83:1986;179-183.
- Aiyer M S, Chiappa S A, Fink G. A priming effect of luteinising hormone releasing factor on the anterior pituitary gland in the female rat. J Endocrinol. 62:1974;573-588.
- Anselmo-Franci J A, Franci C R, Krulich L, Antunes-Rodrigues J, McCann S M. Locus coeruleus lesions decrease norepinephrine input into the medial preoptic area and medial basal hypothalamus and block the LH, FSH, and prolactin preovulatory surge. Brain Res. 767:1997;289-296.
- Antunes-Rodrigues J, McCann S M. Effect of suprachiasmatic lesions on the regulation of luteinizing hormone secretion in the female rat. Endocrinology. 81:1967;666-670.
- Asdell S A. Patterns of Mammalian Reproduction. 1964;Comstock, Ithaca.
- Bakker J, Baum M J. Gonadal steroids modulate pituitary LH secretion, but not mediobasal hypothalamic GnRH mRNA levels in the ferret. J Reprod Fertil. July 2000.
- Bakker J, Rubin B S, Baum M J. Changes in mediobasal hypothalamic gonadotropin-releasing hormone messenger ribonucleic acid levels induced by mating or ovariectomy in a reflex ovulator, the ferret. Endocrinology. 140:1999;595-602.
- Barraclough C A, Wise P M. The role of catecholamines in the regulation of pituitary luteinizing hormone and follicle-stimulating hormone secretion. Endocr Rev. 3:1982;91-119.
- Barry J. Characterization and topography of LHRH neurons in the rabbit. Neurosci Lett. 2:1976;201-205.
- Baum M J, Bressler S C, Daum M C, Vega C A, McNamee C S. Ferret mothers provide more anogenital licking to male offspring: Possible contribution to psychosexual differentiation. Physiol Behav. 60:1996;353-359.
- Baum M J, Carroll R S, Cherry J A, Tobet S A. Steroidal control of behavioural, neuroendocrine and brain sexual differentiation: Studies in a carnivore, the ferret. J Neuroendocrinol. 2:1990;401-418.
- Baum M J, Gerlach J L, Krey L C, McEwen B S. Biochemical and radioautographic analysis of estrogen-inducible progestin receptors in female ferret brain and pituitary: Correlations with effects of progesterone on sexual behavior and gonadotropin-releasing hormone-stimulated secretion of luteinizing hormone. Brain Res. 368:1986;296-309.
- Baum M J, Keverne E B, Everitt B J, Herbert J, de Greef W J. Effects of progesterone and estradiol on sexual attractivity of female rhesus monkeys. Physiol Behav. 18:1977;659-670.
- Baum M J, Schretlen P J. Oestrogenic induction of sexual behaviour in ovariectomized ferrets housed under short or long photoperiods. J Endocrinol. 78:1978;295-296.
- Berria M, Pau K-Y F, Spies H G. Evidence for alpha 1-adrenergic involvement in neuropeptide Y-stimulated GnRH release in female rabbits. Neuroendocrinology. 53:1991;480-486.
- Berriman S J, Wade G N, Blaustein J D. Expression of Fos-like proteins in gonadotropin-releasing hormone neurons of Syrian hamsters: Effects of estrous cycles and metabolic fuels. Endocrinology. 131:1992;2222-2228.
- Beyer C, McDonald P. Hormonal control of sexual behaviour in the female rabbit. Adv Reprod Physiol. 6:1973;185-219.
- Bibeau C E, Tobet S A, Anthony E LP, Carroll R S, Baum M J, King J C. Vaginocervical stimulation of ferrets induces release of luteinizing hormone-releasing hormone. J Neuroendocrinol. 3:1991;29-36.
- Billings H J, Katz L S. Progesterone facilitation and inhibition of estradiol-induced sexual behavior in the female goat. Horm Behav. 31:1997;47-53.
- Birkhead T R. Sperm competition: Evolution and mechanisms. Curr Top Dev Biol. 33:1996;103-158.
- Bissonette T H. Modification of mammalian sexual cycles: Reactions of ferrets of both sexes to electric light added after dark in November and December. Proc R Soc London Biol. 110:1932;322.
- Blaustein J D, Fleder H H. Cytoplasmic progesterone receptors in guinea pig brain: Characterization and relationship to the induction of sexual behavior. Brain Res. 169:1979;481-497.
- Bohler H C, Tracer H, Merriam G R, Petersen S L. Changes in proopiomelanocortin messenger ribonucleic acid levels in the rostral periarcuate region of the female rat during the estrous cycle. Endocrinol. 128:1991;1265-1269.
- Brann D W, Mahesh V B. Excitatory amino acids: Function and significance in reproduction and neuroendocrine regulation. Front Neuroendocrinol. 15:1994;3-49.
- Bronson F H. Rodent pheromones. Biol Reprod. 4:1971;344-357.
- Brown-Grant K, Raisman G. Abnormalities in reproductive function associated with the destruction of the suprachiasmatic nuclei in female rats. Proc R Soc London B Biol Sci. 198:1977;279-296.
- Caba M, Beyer C, González-Mariscal G, Silver R. Sexual difference in the effect of mating on gonadotropin releasing hormone (GnRH) neurons expressing Fos protein in the rabbit forebrain. Society for Neuroscience meeting. 1998.
- Caraty A, Locatelli A. Effect of time after castration on secretion of GnRH and LH in the ram. J Reprod Fert. 82:1988;263-269.
- Carmel P W, Araki S, Ferin M. Pituitary stalk portal blood collection in rhesus monkeys: Evidence for pulsatile release of gonadotropin-releasing hormone (GnRH). Endocrinology. 99:1976;243-248.
- Carroll R S, Baum M J. Evidence that oestrogen exerts an equivalent negative feedback action on LH secretion in male and female ferrets. J Reprod Fertil. 86:1989;235-245.
- Carroll R S, Erskine M S, Baum M J. Sex difference in the effect of mating on the pulsatile secretion of luteinizing hormone in a reflex ovulator, the ferret. Endocrinology. 121:1987;1349-1359.
- Carroll R S, Erskine M S, Doherty P C, Lundell L A, Baum M J. Coital stimuli controlling luteinizing hormone secretion and ovulation in the female ferret. Biol Reprod. 32:1985;925-933.
- Carter C S, Getz L L. Social and hormonal determinants of reproductive patterns in the prairie vole. Gilles R, Balthazart J. Neurobiology. 1985;18-36 Springer-Verlag, Berlin.
- Carter C S, Getz L L, Gavish L, McDermott J L, Arnold P. Male-related pheromones and the activation of female reproduction in the prairie vole (Microtus ochrogaster). Biol Reprod. 23:1980;1038-1045.
- Carter C S, Witt D M, Manock S R, Adams K A, Bahr J M, Carlstead K. Hormonal correlates of sexual behavior and ovulation in male-induced and postpartum estrus in female prairie voles. Physiol Behav. 46:1989;941-948.
- Cattanach H M, Iddon C A, Charlton H M, Chiappa S, Fink G. Gonadotropin-releasing hormone deficiency in a mutant mouse with hypogonadism. Nature. 269:1977;338-340.
- Charlton H M, Naftolin F, Sood M C, Worth R W. The effect of mating upon LH release in male and female voles of the species Microtus agrestis. J Reprod Fertil. 42:1975;167-170.
- Chemineau P. Possibilities for using bucks to stimulate ovarian and oestrus cycles in anovulatory goats. Livest Prod Sci. 17:1987;135-147.
- Chen W-P, Witkin J W, Silverman A J. Beta-endorphin and gonadotropin-releasing hormone synaptic input to gonadotropin-releasing hormone neurosecretory cells in the male rat. J Comp Neurol. 286:1989;85-95.
- Chen W P, Witkin J W, Silverman A J. Sexual dimorphism in the synaptic input to gonadotropin releasing hormone neurons. Endocrinology. 126:1990;695-702.
- Ching M. Morphine suppresses the proestrous surge of GnRH in pituitary portal plasma of rats. Endocrinology. 112:1983;2209-2211.
- Clark I J, Thomas G B, Yao B, Cummins J T. GnRH secretion throughout the ovine estrous cycle. Neuroendocrinology. 46:1987;82-88.
- Cohen-Parsons M, Carter C S. Males increase serum estrogen and estrogen receptor binding in brain of female voles. Physiol Behav. 39:1987;309-314.
- Cohen-Parsons M, Carter C S. Males increase progestin receptor binding in brain of female voles. Physiol Behav. 42:1988;191-197.
- Conaway C H. Ecological adaptation and mammalian reproduction. Biol Reprod. 4:1971;239-241.
- Concannon P, Hodgson B, Lein D. Reflex LH release in estrous cats following single and multiple copulations. Biol Reprod. 23:1980;111-117.
- Crews D, Grassman M, Lindzey J. Behavioral facilitation of reproduction in sexual and unisexual whiptail lizards. Proc Natl Acad Sci USA. 83:1986;9547-9550.
- Crews D P, Silver R. Reproductive physiology and behavior interactions in nonmammalian vertebrates. Adler N, Pfaff D, Goy R W. Handbook of Behavioral Neurobiology, Vol 7. 1985;101-182 Plenum, New York.
- Crighton D B, Foster J P. Luteinising hormone release after two injections of synthetic luteinising hormone-releasing hormone in the ewe. J Endocrinol. 72:1977;59-67.
- Crowder M E, Herring R D, Nett T M. Rapid recovery from gonadotrope function after down-regulation of receptors for GnRH in ewes. J Reprod Fertil. 78:1986;577-585.
- Cunningham E TJ, Sawchenko P E. Anatomical specificity of noradrenergic inputs to the paraventricular and supraoptic nuclei of the rat hypothalamus. J Comp Neurol. 274:1988;60-76.
- DeBold J F, Martin J V, Whalen R E. The excitation and inhibition of sexual receptivity in female hamsters by progesterone: Time and dose relationships, neural colocalization and mechanisms of action. Endocrinology. 99:1976;1519-1527.
- Dellovade T L, King J A, Millar R P, Rissman E F. Presence and differential distribution of distinct forms of immunoreactive gonadotropin-releasing hormone in the musk shrew brain. Neuroendocrinology. 58:1993;166-177.
- Dellovade T L, Ottinger M A, Rissman E F. Mating alters gonadotropin-releasing hormone cell number and content. Endocrinology. 136:1995;1648-1657.
- Dempsey E W, Searles H F. Environmental modification of certain endocrine phenomena. Endocrinology. 32:1943;119-128.
- Dewsbury D A. Copulatory behavior of montane voles (Microtus montanus). Behaviour. 44:1973;186-202.
- Dhillon H, Dunn A M, Esquival E, Hamernik D L, Wise M E. The estradiol-induced luteinizing hormone surge in the ewe is not associated with increased gonadotropin-releasing hormone messenger ribonucleic acid levels. Biol Reprod. 57:1997;107-111.
- Dluzen D E, Carter C S. Ovarian hormones regulating sexual and social behaviors in female prairie voles, Microtus Ochrogaster. Physiol Behav. 23:1979;597-600.
- Doan A, Urbanski H F. Diurnal expression of Fos in luteinizing hormone releasing hormone neurons of Syrian hamsters. Biol Reprod. 50:1994;301-308.
- Donovan B T, ter Haar M B. Effects of LH-releasing hormone on plasma FSH and LH levels in the ferret. J Endocrinol. 73:1977;37-52.
- Donovan B T, Matson C, Kilpatrick M J. Effect of exposure to long days on the secretion of oestradiol, oestrone, progesterone, testosterone, androstenedione, cortisol and follicle-stimulating hormone in intact and spayed ferrets. J. Endocrinol. 99:1983;361-368.
- Dryden G L. Reproduction in Suncus murinus. J Reprod Fert Suppl. 6:1969;377-396.
- Dufy-Barbe L, Franchimont P, Faure J M. Time courses of LH and FSH release after mating in the female rabbit. Endocrinology. 92:1973;1318-1321.
- Edwards R G. Test-tube babies, 1981. Nature. 293:1981;253-256.
- Emanuele N V, Jurgens J, La Paglia N, Williams D W, Kelley M R. The effect of castration on steady state levels of luteinizing hormone-releasing hormone (LHRH) mRNA and proLHRH processing: Time course study utilizing semi-quantitative reverse transcription/polymerase chain reaction. J Endocrinol. 148:1996;509-515.
- Enders R K. Reproduction in the mink (Mustela vision). Proc Am Philos Soc. 96:1952;691-755.
- Eskay R L, Mical R S, Porter J C. Relationship between luteinizing hormone releasing hormone concentration in hypophysical blood and luteinizing hormone release in intact, castrated, and electrochemically-stimulated rats. Endocrinology. 100:1977;263-270.
- Everett J W. Spontaneous persistent estrus in a strain of albino rats. Endocrinology. 25:1939;123-127.
- Everett J W. The mammalian reproductive cycle and its controlling mechanisms. Young W. Sex and Internal Secretions. 1961;497-555 Williams & Wilkins, Baltimore.
- Fadem B H. The effects of pheromonal stimuli on estrus and peripheral plasma estradiol in female gray short-tailed opossums (Monodelphis domestica). Biol Reprod. 41:1989;231-217.
- Ferin M, Van Vugt D, Wardlaw S. The hypothalamic control of the menstrual cycle and the role of endogenous opioid peptides. Recent Prog Horm Res. 40:1984;441-485.
- Fernandez-Baca S, Madden D HL, Novoa C. Effect of different mating stimuli on induction and ovulation in the alpaca. J Reprod Fertil. 22:1970;261-267.
- Finn P D, Steiner R A, Clifton D K. Temporal patterns of gonadotropin-releasing hormone (GnRH), c-fos, and galanin gene expression in GnRH neurons relative to the luteinizing hormone surge in the rat. J Neurosci. 18:1996;713-719.
- Fortune J E, Eppig J J, Rissman E F. Mating stimulates estradiol production by ovaries of the musk shrew (Suncus murinus). Biol Reprod. 46:1992;885-891.
- Gautron J P, Leblanc P, Bluet-Pajot M T, Pattou E, L'Héritier A, Mounier F, Ponce G, Audinot V, Rasolonjanahary R, Kordon C. A second endogenous molecular form of mammalian hypothalamic luteinizing hormone-releasing hormone (LHRH), (hydroxyproline9) LHRH, releases luteinizing hormone and follicle-stimulating hormone in vitro and in vivo. Moll Cell Endocrinol. 85:1992;99-107.
- Gibson M J, Wu T J, Miller G M, Silverman A J. What nature's knockout teaches us about GnRH activity: Hypogonadal mice and neuronal grafts. Horm Behav. 31:1997;212-220.
- Goldsmith P C, Ganong W F. Ultrastructural localization of luteinizing hormone-releasing hormone in the median eminence of the rat. Brain Res. 97:1975;181-193.
- Gomendio M, Harcourt A H, Roldan E RS. Sperm competition in mammals. Birkhead T R, Møller A P. Sperm Competition and Sexual Selection. 1998;667-756 Academic Press, London.
- Goodman A L, Neill J D. Ovarian regulation of postcoital gonadotropin release in the rabbit: Reexamination of a functional role for 20 αdihydroprogesterone. Endocrinology. 99:1976;852-860.
- Goodrowe K L, Chakraborty P K, Wildt D E. Pituitary and gonadal response to exogenous LH-releasing hormone in the male domestic cat. J. Endocrinol. 105:1985;175-181.
- Gore A C, Roberts J L. Regulation of gonadotropin-releasing hormone gene expression in the rat during the luteinizing hormone surge. Endocrinology. 136:1995;889-896.
- Gore A C, Roberts J L. Regulation of gonadotropin-releasing hormone gene expression in vivo and in vitro. Front Neuroendocrinol. 18:1997;209-245.
- Gray G D, Davis H N, Kenney Amc M, Dewsbury D A. The effect of mating on plasma levels of LH and progesterone in montane voles (Microtus montanus). J Reprod Fertil. 47:1976;89-91.
- Gudermuth D F, Newton L, Daels P, Concannon P. Incidence of spontaneous ovulation in young, group-housed cats based on serum and faecal concentrations of progesterone. J Reprod Fertil Suppl. 51:1997;177-184.
- Guillemin R. Purification, isolation, and primary structure of the hypothalamic luteinizing hormone-releasing factor of ovine origin. A historical account. Am J Obstet Gynecol. 129:1977;214-218.
- Hamada T, Nakajima M, Takeuchi Y, Mori Y. Pheromone-induced stimulation of hypothalamic gonadotropin-releasing hormone pulse generator in ovariectomized, estrogen-primed goats. Neuroendocrinology. 64:1996;313-319.
- Hammond J, Marshall F HA. Oestrus and pseudopregnancy in the ferret. Proc R Soc. B105:1930;607-630.
- Harris G W, Jacobsohn D. Functional grafts of the anterior pituitary gland. Proc Soc London Biol. 139:1952;263-270.
- Harris G W, Naftolin F. The hypothalamus and control of ovulation. Br Med Bull. 26:1970;3-9.
- Harris T G, Robinson J E, Evans N P, Skinner D C, Herbison A E. Gonadotropin-releasing hormone messenger ribonucleic acid expression changes before the onset of the estradiol-induced luteinizing hormone surge in the ewe. Endocrinology. 139:1998;57-64.
- Hasler J F, Banks E M. The behavioural and somatic effects of ovariectomy and replacement therapy in female collared lemmings (Dicrostonyx groenlandicus). Biol Reprod. 12:1976;647-656.
- Herbison A E. Noradrenergic regulation of cyclic GnRH secretion. Rev Reprod. 2:1997;1-6.
- Herbison A E. Multimodal influence of estrogen upon gonadotropin-releasing hormone neurons. Endocr Rev. 19:1998;302-330.
- Herbison A E, Robinson J E, Skinner D C. Distribution of estrogen receptor-immunoreactive cells in the preoptic area of the ewe: Co-localization with glutamic acid decarboxylase but not luteinizing hormone-releasing hormone. Neuroendocrinology. 57:1993;751-759.
- Hilliard J, Eaton L WM. Estradiol-17β, progesterone and 20α-hydroxypregn-4-en-3-one in rabbit venous plasma. II. From mating through implantation. Endocrinology. 89:1971;522-527.
- Hilliard J, Penardi R, Sawyer C H. A functional role for 20-alpha-hydroxypregn-4-en-3-one in the rabbit. Endocrinology. 80:1967;901-909.
- Hind R A. Interaction of internal and external factors in integration of canary reproduction. Beach F A. Sex and Behavior. 1965;381-415 Wiley, New York.
- Hoffman G E, Smith M S, Verbalis J G. C-fos and related immediate early gene products as markers of activity in neuroendocrine systems. Front Neuroendocrinol. 14:1993;173-213.
- Hokfelt T, Martenson R, Bjorklund A, Kleinau S, Goldstein M. Distributional maps of tyrosine-hydroxylase-immunoreactive neurons in the rat brain. Bjorklund A, Hokfelt T. Handbook of Chemical Neuroanatomy, Vol 2. Classical Neurotransmitters in the CNS, Part I. 1984;277-379 Elsevier Science, Amsterdam.
- Hubbart J I, Hyland B I, Sirett N E. Stimulation of the LH release by naloxone in anaesthetized cats after ovariectomy. Neurosci Lett. 108:1990;295-302.
- Huck U W, Carter C S, Banks E M. Estrogen and progesterone interactions influencing sexual and social behaviour in the brown lemming (Lemmus trimucronatus). Horm Behav. 12:1979;40-49.
- Jöchle W. Coitus induced ovulation. Contraception. 7:1973;523-564.
- Jöchle W. Current research in coitus-induced ovulation: A review. J Reprod Fertil Suppl. 22:1975;165-207.
- Jones E F, Bain J B, Odell W D. Postcoital luteinizing hormone release in male and female rabbits as determined by radioimmunoassay. Fertil Steril. 27:1976;848-852.
- Johnson L M, Gay V L. Luteinizing hormone in the cat. I. Tonic secretion. Endocrinology. 109:1981;240-246.
- Johnson L M, Gay V L. Luteinizing hormone in the cat. II. Mating-induced secretion. Endocrinology. 109:1981;247-252.
- Kalra S P. Mandatory neuropeptide-steroid signaling for the preovulatory luteinizing hormone-releasing hormone discharge. Endocr Rev. 14:1993;507-538.
- Kalra S P, Kalra P S. Neural regulation of luteinizing hormone secretion in the rat. Endocr Rev. 4:1983;311-351.
- Kalra S P, Kalra P S. Do testosterone and estradiol-17β enforce inhibition or stimulation of luteinizing hormone-releasing hormone secretion? Biol Reprod. 41:1989;559-570.
- Karsch F J, Cummins J T, Thomas G B, Clarke I J. Steroid feedback inhibition of pulsatile secretion of gonadotropin-releasing hormone in the ewe. Biol Reprod. 36:1987;1207-1218.
- Karsch F J, Weick R F, Butler W R, Dierschke D J, Krey L C, Weiss G, Hotchkiss J, Yamaji T, Knobil E. Induced LH surges in the rhesus monkey: Strength-duration characteristics of the estrogen stimulus. Endocrinology. 92:1973;1740-1747.
- Kasten T L, White S A, Norton T T, Bond C T, Adelman J P, Fernald R D. Characterization of two new preproGnRH mRNAs in the tree shrew: First direct evidence for mesencephalic GnRH gene expression in a placental mammal. Gen Comp Endocrinol. 104:1996;7-19.
- Kaynard A H, Pau K-Y F, Hess D L, Spies H G. Gonadotropin-releasing hormone and norepinephrine release from the rabbit mediobasal and anterior hypothalamus during the mating-induced luteinizing hormone surge. Endocrinology. 127:1990;1176-1185.
- Kelly M J, Garrett J, Bosch M A, Roselli C E, Douglass J, Adelman J P, Ronnekleiv O K. Effects of ovariectomy on GnRH mRNA, proGnRH and GnRH levels in the preoptic hypothalamus of the female rat. Neuroendocrinology. 49:1989;88-97.
- Kenney Amc M, Dewsbury D A. Effect of limited mating on the corpora lutea in montane voles, Microtus montanus. J Reprod Fertil. 49:1977;363-364.
- Khorram O, Pau K-Y F, Spies H G. Release of hypothalamic neuropeptide Y and effects of exogenous NPY on the release of hypothalamic GnRH and pituitary gonadotrophs in intact and ovariectomized does in vitro. Peptides. 9:1988;411-417.
- King J A, Steneveld A A, Curlewis J D, Rissman E F, Millar R P. Identification of chicken GnRH II in brains of metatherian and early-evolved eutherian species of mammals. Regul Pep. 54:1994;467-477.
- Krsmanovic L Z, Stojilkovic S S, Mertz L M, Tomic G, Catt K J. Expression of gonadotropin-releasing hormone receptors and autocrine regulation of neuropeptide release in immortalized hypothalamic neurons. Proc Natl Acad Sci USA. 90:1993;3908-3912.
- Lambert G M, Erskine M S, Baum M J. Effect of naloxone on the pulsatile secretion of luteinizing hormone in gonadectomized male and female ferrets before and after oestradiol replacement. J Neuroendocrinol. 2:1990;701-705.
- Lambert G M, Rubin B S, Baum M J. Sexual dimorphism in the effects of mating on the in vitro release of LHRH from the ferret mediobasal hypothalamus. Physiol Behav. 52:1992a;809-813.
- Lambert G M, Rubin B S, Baum M J. Sex difference in the effect of mating on c-fos expression in luteinizing hormone-releasing hormone neurons of the ferret forebrain. Endocrinology. 131:1992b;1473-1480.
- Lawler D F, Johnston S D, Hegstad R L, Keltner D G, Owens S F. Ovulation without cervical stimulation in domestic cats. J Reprod Fertil Suppl. 47:1993;57-61.
- Leavitt W W, Chen T J, Allen T C, Johnston T ON. Regulation of progesterone receptor formation by estrogen action. Ann NY Acad Sci. 286:1977;210-225.
- Lee W S, Smith S, Hoffman G E. Luteinizing hormone-releasing hormone neurons express fos protein during the proestrous surge of luteinizing hormone. Proc Natl Acad Sci USA. 87:1990;5163-5167.
- Lehman M N, Karsch F J. Do gonadotropin-releasing hormone, tyrosine hydroxylase- and β-endorphin-immunoreactive neurons contain estrogen receptors? A double-label immunocytochemical study in the suffolk ewe. Endocrinology. 133:1993;887-895.
- Lehrman D S. Interaction between internal and external environments in the regulation of the reproductive cycle of the ring dove. Beach F A. Sex and Behavior. 1965;355-380 Wilkins, New York.
- Léránth C, Seguraum L MG, Palkovits M, MacLusky J, Shanabrough M, Naftolin F. The LH-RH-containing neuronal network in the preoptic area of the rat: Demonstration of LH-RH-containing nerve terminals in synaptic contact with LH-RH neurons. Brain Res. 345:1985;332-336.
- Lescheid D W, Terasawa E, Abler L A, Urbanski H F, Warby C M, Millar R P, Sherwood N M. A second form of gonadotropin-releasing hormone (GnRH) with characteristics of chicken GnRH-II is present in the primate brain. Endocrinology. 138:1997;5618-5629.
- Levine J E, Pau K-Y F, Ramirez V D, Jackson G L. Simultaneous measurement of luteinizing hormone-releasing hormone and luteinizing hormone release in unanesthetized, ovariectomized sheep. Endocrinology. 111:1982;1449-1455.
- Levine J E, Ramirez V D. Luteinizing hormone-releasing hormone release during the rat estrous cycle and after ovariectomy, as estimated by push-pull cannulae. Endocrinology. 111:1982;1439-1448.
- Levine J E, Ramirez V D. Measurements of neuropeptide release: In vitro and in vivo procedures. Methods Enzymol. 124:1986;466-494.
- Li W, Pelletier G. Opioid regulation of gonadotropin-releasing hormone gene expression in the male rat brain as studied by in situ hybridization. Neuroreport. 4:1993;331-333.
- Lin W W, Ramirez V D. Effect of pulsatile infusion of progesterone on the in vivo activity of the luteinizing hormone-releasing hormone neural apparatus of awake unrestrained female and male rabbits. Endocrinology. 122:1988;868-876.
- Lin W W, Ramirez V D. Infusions of progestins into the hypothalamus of female New Zealand white rabbits: Effect on in vivo luteinizing hormone-releasing hormone release as determined with push-pull perfusion. Endocrinology. 126:1990;261-272.
- Lin W W, Ramirez V D. Effect of mating behavior on luteinizing-hormone releasing hormone release in female rabbits as monitored with push-pull cannulae. Neuroendocrinology. 53:1991;229-235.
- MacLusky N J, Lieberburg I, Krey L C, McEwen B S. Progestin receptors in the brain and pituitary of the bonnet monkey: Differences between the monkey and the rat in the distribution of progestin receptors. Endocrinology. 106:1980;185-191.
- MacLusky N J, McEwen B S. Progestin receptor in the rat brain: Distribution and properties of cytoplasmic binding sites. Endocrinology. 106:1980;192-202.
- Malik K F, Silverman A J, Morrell J I. Gonadotropin-releasing hormone mRNA in the rat: Distribution and neuronal content over the estrous cycle and after castration of males. Anat Rec. 231:1991;457-466.
- Marie M, Anouassi A. Mating-induced luteinizing hormone surge and ovulation in the female camel (Camelus dromedarius). Biol Reprod. 35:1986;792-798.
- Marion G D. The effect of sterile copulation on time of ovulation in dairy heifers. J Dairy Sci. 33:1950;885-889.
- Marsden H M, Bronson F H. Estrous synchrony in mice: Alteration by exposure to male urine. Science. 144:1964;3625.
- Marshall F HA. The oestrous cycle of the common ferret. Q J Microsc Sci. 48:1904;323-345.
- McCann S M, Taleisnik S, Friedman H M. LH-releasing activity in hypothalamic extracts. Proc Soc Exp Biol. 104:1960;432-434.
- Mendonca M T, Crews D. Effect of fall mating on ovarian development in the red-sided garter snake. Am J Physiol. 257:1989;R1548-1550.
- Merchenthaler I, Görcs T, Sétáló G, Petrusz P, Flerkó B. Gonadotropin-releasing hormone (GnRH) neurons and pathways in the rat brain. Cell Tissue Res. 237:1984;15-29.
- Millar R P, King J A. Plasticity and conversation in gonadotropin-releasing hormone structure. Davey K G, Peter R E, Tobe S S. Perspectives in Comparative Endocrinology. 1994;129-136 National Research Council of Canada, Ottawa.
- Milligan S R. Mating, ovulation, and corpus luteum function in the vole, Microtus agrestis. J Reprod Fertil. 42:1975;35-44.
- Milligan S R. The feedback of exogeneous steroids on LH release and ovulation in the intact female vole (Microtus agrestis). J Reprod Fertil. 54:1978;309-311.
- Milligan S R. The effect of bromocriptine treatment on the ovulatory response to oestradiol benzoate in the reflex ovulator, Microtus agrestis. J Endocrinol. 84:1980;315-316.
- Milligan S R. Analysis of the LH surges induced by mating and LH-RH in the vole, Microtus agrestis. J Reprod Fertil. 63:1981;39-45.
- Milligan S R. Induced ovulation in mammals. Finn C A. Oxford Reviews of Reproductive Biology. 1982;1-46 Clarendon Press, London.
- Mitsushima D, Hei D L, Terasawa E. Gamma-aminobutyric acid is an inhibitory neurotransmitter restricting the release of luteinizing hormone-releasing hormone before the onset of puberty. Proc Natl Acad Sci USA. 91:1994;395-399.
- Mock O M, Conaway C H. Reproduction of the least shrew. (Cryptitus parva) in captivity. Antikatzides Th, Spiegel A. The Laboratory Animal in the Study of Reproduction, 6th symposium of the International Committee on Laboratory Animals, Thessalonika. 1976;59-74 Fisher, Stuttgart.
- Moenter S M, Karsch F J, Lehman M N. Fos expression during the estradiol-induced gonadotropin-releasing hormone (GnRH) surge of the ewe: Induction in GnRH and other neurons. Endocrinology. 133:1993;896-903.
- Moffatt C A, Gerber J M, Blom J M, Kriegsfeld L J, Nelson R J. Photoperiodic effects on steroid negative feedback in female prairie voles. Gen Comp Endocrinol. 100:1995;92-95.
- Moguilewskey M, Raynaud J-P. Estrogen-sensitive progestin-binding sites in the emale brain and pituitary. Brain Res. 164:1979;165-175.
- Moricard R. Wolstenholme G EW. Ciba Foundation Symposium: Mammalian Germ Cells. 1953;187-197 Churchill, London.
- Muske L E. Evolution of gonadotropin-releasing hormone (GnRH) neuronal systems. Brain Behav Evol. 1993;215-230.
- Nadler R D. A biphasic influence of progesterone on sexual receptivity of spayed female rats. Physiol Behav. 5:1970;95-97.
- Neill J D. Sexual differences in the hypothalamic regulation of prolactin secretion. Endocrinology. 90:1972;1154-1159.
- Nett T M, Crowder M E, Moss G E, Duello T M. GnRH-receptor interaction. V. Down-regulation of pituitary receptors for GnRH in overiectomized ewes by infusion of homologous hormone. Biol Reprod. 24:1981;1145-1155.
- Orstead K M, Spies H G. Inhibition of hypothalamic gonadotropin-releasing hormone release by endogenous opioid peptides in the female rabbit. Neuroendocrinology. 46:1987;14-23.
- Park O-K, Gugneja S, Mayo K E. Gonadotropin-releasing hormone gene expression during the rat estrous cycle: Effects of pentobarbital and ovarian steroids. Endocrinology. 127:1990;365-372.
- Parker G A. Sperm competition and its evolutionary consequences in the insects. Biol Rev. 45:1970;525-567.
- Pau K-Y F, Khorram O, Kaynard A H, Spies H G. Simultaneous induction of neuropeptide Y and gonadotropin-releasing hormone in the rabbit hypothalamus. Neuroendocrinology. 49:1989;197-201.
- Pau K-Y F, Lee C J, Cowles A, Yang S P, Hess D L, Spies H G. Possible involvement of norepinephrine transporter activity in the pulsatility of hypothalamic gonadotropin-releasing hormone release: Influence of the gonad. J Neuroendocrinol. 10:1998;21-29.
- Pau K-Y F, Ma Y J, Yu J, Yang S P, Airhart N, Spies H G. Topographic comparison of the expression of norepinephrine transporter, tyrosine hydroxylase and neuropeptide Y mRNA in association with dopamine β-hydroxylase neurons in the rabbit brainstem. Mol Brain Res. 48:1997;367-381.
- Pau K-Y F, Orstead K M, Hess D L, Spies H G. Feedback effects of ovarian steroids on the hypothalamic-hypophyseal axis in the rabbit. Biol Reprod. 35:1986;1009-1023.
- Pau K-Y F, Spies H G. Estrogen-dependent effects of norepinephrine on hypothalamic gonadotropin-releasing hormone release in the rabbit. Brain Res. 399:1986;15-23.
- Petersen S L, Keller M L, Carder S A, McCrone S. Differential effects of estrogen and progesterone on levels of POMC mRNA levels in the arcuate nucleus: Relationship to the timing of LH surge release. J Neuroendocrinol. 5:1993;643-648.
- Pfaus J G, Jakok A, Kleopoulos S P, Gibbs R B, Pfaff D W. Sexual stimulation induces Fos immunoreactivity within GnRH neurons of the female rat preoptic area: Interaction with steroid hormones. Neuroendocrinol. 60:1994;283-290.
- Pfeiffer C A. Sexual differences of the hypophysis and their determination by the gonads. Am J Anat. 58:1936;195-225.
- Porkka-Heiskanen T, Urban J H, Turek F W, Levine J E. Gene expression in a subpopulation of luteinizing hormone-releasing hormone (LHRH) neurons prior to the preovulatory gonadotropin surge. J Neurosci. 14:1994;5548-5558.
- Price C S, Dyer K A, Coyne J A. Sperm competition between Drosophila males involves both displacement and incapacitation. Nature. 400:1999;449-452.
- Rajendren G, Dudley C A, Moss R L. Influence of male rats on the luteinizing hormone releasing hormone neuronal system in female rats: Role of the vomeronasal organ. Neuroendocrinol. 57:1993;898-906.
- Ramirez V D, Chen J C, Nduka E, Lin W, Ramirez A D. Push-pull perfusion (PPP) of the hypothalamus and the caudate nucleus (CN) in conscious, unrestrained animals. Ann NY Acad Sci. 473:1986a;434-448.
- Ramirez V D, Ramirez A D, Slamet W, Nduka E. Functional characteristics of the luteinizing hormone-releasing hormone pulse generator in conscious unrestrained female rabbits: Activation by norepinephrine. Endocrinology. 118:1986;2331-2339.
- Ramirez V D, Soufi W L. The neuroendocrine control of the rabbit ovarian cycle. Knobil E, Neill J D. The Physiology of Reproduction. 1994;585-612 Raven Press, New York.
- Richmond M, Stehn R. Olfaction and reproductive behavior in microtine rodents. Doty R L. Mammalian Olfaction, Reproductive Processes, and Behavior. 1976;197-217 Academic Press, New York.
- Rissman E F. Behavioral regulation of gonadotropin-releasing hormone. Biol Reprod. 54:1996;413-419.
- Rissman E F, Clendenon A L, Krohmer R W. The role of androgens in the regulation of sexual behavior in the female musk shrew. Neuroendocrinology. 51:1990;468-473.
- Rissman E F, Crews D. Hormonal correlates of sexual behavior in the female musk shrew: The role of estradiol. Physiol Behav. 44:1988;1-7.
- Rissman E F, Li X. Sex differences in mammalian and chicken-II gonadotropin-releasing hormone immunoreactivity in musk shrew brain. Gen Comp Endocrinol. 112:1998;346-355.
- Rissman E F, Silveira J, Bronson F H. Patterns of sexual receptivity in the female musk shrew (Suncus murinus). Horm Behav. 22:1988;186-193.
- Robinson A. XIII. The formation, rupture, and closure of ovarian follicles in ferrets and ferret-polecat hybrids, and some related phenomena. Trans R Soc Edinburgh. 52:1918;303-362.
- Roselli C E, Snipes C A. Cytoplasmic progestin receptors in the hypothalamus-preoptic area of the mouse. Effect of estradiol priming. J Steroid Biochem. 19:1983;1571-1575.
- Ryan K D. Hormonal correlates of photoperiod-induced puberty in a reflex ovulator, the female ferret. Biol Reprod. 31:1984;925-935.
- Sadlier R MFS. The Ecology of Reproduction in Wild and Domestic Animals. 1969;Methuen, London.
- Sagrillo C A, Grattan D R, McCarthy M M, Selmanoff M. Hormonal and neurotransmitter regulation of GnRH gene expression and related reproductive behaviors. Behav Gen. 26:1996;241-277.
- Sarkar D K, Chaippa S A, Fink G, Sherwood N M. Gonadotrophin-releasing hormone surge in pro-oestrous rats. Nature. 264:1976;461-463.
- Sarkar D K, Yen S S. Changes in beta-endorphin-like immunoreactivity in pituitary portal blood during the estrous cycle and after ovariectomy in rats. Endocrinology. 116:1985;2075-2079.
- Sawchenko P E, Swanson L W. The organization of noradrenergic pathways from the brainstem to the paraventricular and supraoptic nuclei in the rat. Brain Res Rev. 257:1982;275-325.
- Sawchenko P E, Swanson L W, Grzanna R, Howe P RC, Bloom S R, Polak J M. Colocalization of neuropeptide Y immunoreactivity in brainstem catecholaminergic neurons that project to the paraventricular nucleus of the hypothalamus. J Comp Neurol. 241:1985;138-153.
- Sawyer C H. Stimulation of ovulation in the rabbit by intraventricular injection of epinephrine or norepinephrine. Anat Rec. 112:1952.
- Sawyer C H. Control of secretion of gonadotropins. Cole H H. Gonadotropins. 1964;113-159 Freeman, San Francisco.
- Sawyer C H, Markee J E. Estrogen facilitation of release of pituitary ovulating hormone in the rabbit in response to vaginal stimulation. Endocrinology. 65:1959;614-621.
- Sawyer C H, Markee J E, Hollinshead W H. Inhibition of ovulation in the rabbit by the adrenergic-blocking agent dibenamine. Endocrinology. 41:1947;395-402.
- Schally A V. Aspects of hypothalamic regulation of the pituitary gland. Science. 202:1978;18-28.
- Schiml P A, Rissman E F. Effects of gonadotropin-releasing, corticotropin-releasing hormone, and vasopressin on female sexual behavior. Horm Behav. 37:2000;212-220.
- Schiml P A, Wersinger S R, Rissman E F. Behavioral activation of the female neuroendocrine axis. Wallen K, Schneider J E. Reproduction in Context: Social and Environmental Influences on Reproduction. 1999;MIT Press.
- Schwagmeyer P L, Parker G A. Male mate choice as predicted by sperm competition in 13-lined ground squirrels. Nature. 348:1990;62-64.
- Shivers B D, Harlan R E, Morrell J I, Pfaff D W. Absence of estradiol concentration in cell nuclei of LHRH-immunoreactive neurons. Nature. 304:1984;345-347.
- Signoret J P, du Mesnil du Buisson F, Mauléon P. Effect of mating on the onset and duration of ovulation in the sow. J Reprod Fertil. 31:1972;327-330.
- Silverman A J, Desnoyers P. Ultrastructural immunocytochemical localization of LHRH in the median eminence of the guinea pig. Cell Tissue Res. 169:1976;157-166.
- Skynner M J, Sim J A, Herbison A E. Detection of estrogen receptor α and β Messenger ribonucleic acids in adult gonadotropin-releasing hormone neurons. Endocrinology. 140:1999;5195-5201.
- Smith R L. Sperm Competition and the Evolution of Animal Mating Systems. 1984;Academic Press, Orlando.
- Spies H G, Pau K-Y F, Yang S P. Coital and estrogen signals: A contrast in the preovulatory neuroendocrine networks of rabbits and rhesus monkeys. Biol Reprod. 56:1997;310-319.
- Stockman E R, Albers H E, Baum M J. Activity in the ferret: Oestradiol effects and circadian rhythms. Anim Behav. 33:1985;150-154.
- Sullivan K A, Witkin J W, Ferin M, Silverman A J. Gonadotropin-releasing hormone neurons in the rhesus macaque are not immunoreactive for the estrogen receptor. Brain Res. 685:1995;198-200.
- Tang Y P, Kashon M L, Sisk C L. Brain region-specific regulation of luteinizing hormone-releasing hormone messsenger ribonucleic acid in the male ferret: Interactions between pubertal maturation and testosterone. Endocrinology. 138:1997;4740-4747.
- Terkel J. Neuroendocrine processes in the establishment of pregnancy and pseudopregnancy in rats. Psychoneuroendocrinology. 13:1988;5-28.
- Thind K K, Goldsmith P C. Infundibular gonadotropin-releasing hormone neurons are inhibited by direct opioid and autoregulatory synapses in juvenile monkeys. Neuroendocrinology. 47:1988;203-216.
- Traczyk W Z, Pau K-Y F, Kaynard A H, Spies H G. Modulatory role of substance P on gonadotropin and prolactin secretion in the rabbit. J Physiol Pharmacol. 43:1992;279-297.
- Tsuruo Y, Hitoshi K, Hisano S, Kagotani Y, Daikoku S, Zang T, Yanaihara N. Substance P-containing neurons innervating LHRH-containing neurons in the septo-preoptic area of rats. Neuroendocrinology. 53:1991;236-245.
- Turzillo A M, Di Gregoria G B, Nett T M. Messenger ribonucleic acid for gonadotropin-releasing hormone receptor and numbers of gonadotropin-releasing hormone receptors in ovariectomized ewes after hypothalamic-pituitary disconnection and treatment with estradiol. J Anim Sci. 73:1995;1784-1788.
- Van der Beek E M, Horvath T L, Wiegant V M, Van den Hurk R, Buijs R M. Evidence for a direct neuronal pathway from the suprachiasmatic nucleus to the gonadotropin-releasing hormone system: Combined tracing and light and electron microscopic immunocytochemical studies. J Comp Neurol. 384:1997;569-579.
- Versi E, Chiappa S A, Fink G, Charlton H M. Effect of copulation on the hypothalamic content of gonadotrophic hormone-releasing hormone in the vole, Microtus agrestis. J Reprod Fertil. 64:1982;491-494.
- Villars T A, Erskine M S, Lambert G M, Jacobson D, Weaver D E, Baum M J. Endocrine correlates of mating-induced reductions in estrous behavior in an induced ovulator. Horm Behav. 24:1990;198-214.
- Vreeburg J T, van der Vaart P D, van der Schoot P. Prevention of central defeminization but not masculinization in male rats by inhibition neonatally by oestrogen biosynthesis. J Endocrinol. 74:1977;375-382.
- Wang C F, Lasley B L, Lein A, Yen S SG. A functional change of the pituitary gonadotrophs during the menstrual cycle. J Clin Endocrinol Metab. 42:1976;718-728.
- Warembourg M, Leroy D, Peytevin J, Martinet L. Estrogen receptor and progesterone receptor-immunoreactive cells are not co-localized with gonadotropin-releasing hormone in the brain of the female mink (Mustela vison). Cell Tissue Res. 291:1998;33-41.
- Waring D W, Turgeon J L. Luteinizing hormone-releasing hormone-induced luteinizing hormone secretion in vitro: Cyclic changes in responsiveness and self-priming. Endocrinology. 106:1980;1430-1436.
- Watson R E, Langub M C, Landis J W. Further evidence that most luteinizing hormone-releasing hormone neurons are not directly estrogen-responsive: Simultaneous localization of luteinizing hormone-releasing hormone and estrogen receptor immunoreactivity in the guinea pig brain. J Neuroendocrinol. 4:1992;311-317.
- Watts A G, Fink G. Constant light blocks diurnal but not pulsatile release of luteinizing hormone in the ovariectomized rat. J Endocrinol. 89:1981;141-146.
- Weiland N G, Barraclough C A. Neonatal castration of male and female rats affects luteinizing hormone responses to estrogen during adulthood. Biol Reprod. 31:1984;942-949.
- Wersinger S R, Baum M J. The temporal pattern of mating-induced immediate early-gene product immunoreactivity in LHRH and non-LHRH neurons of the estrous ferret forebrain. J Neuroendocrinol. 8:1996;345-359.
- Wersinger S R, Baum M J. Sexually dimorphic processing of somatosensory and chemosensory inputs to forebrain luteinizing hormone-releasing hormone neurons in mated ferrets. Endocrinology. 138:1997;1121-1129.
- Wersinger S R, Baum M J. Sexually dimorphic activation of midbrain tyrosine hydroxylase neurons after mating or exposure to chemosensory cues in the ferret. Biol Reprod. 56:1997;1407-1414.
- Whalen R E. Sexual behavior of cats. Behaviour. 20:1963;321-342.
- Whalen R E, Hardy D F. Induction of receptivity in female rats with oestrogen and testosterone. Physiol Behav. 5:1970;529-533.
- White R B, Eisen J A, Kasten J L, Fernald R D. Second gene for gonadotropin releasing hormone in humans. Proc Natl Acad Sci USA. 95:1998;305-309.
- Whitney L F, Underwood A B. The Racoon. 1952;Practical Science, Orange.
- Whitten W K. Pheromones and mammalian reproduction. McLaren A. Advances in Reproductive Physiology. 1966;155-177 Academic Press, New York.
- Wiemann J N, Clifton D K, Steiner R A. Gonadotropin-releasing hormone messenger ribonucleic acid levels are unaltered with changes in the gonadal hormone milieu of the adult male rat. Endocrinology. 127:1990;523-532.
- Wildt D E, Chan S YW, Seager S WJ, Chakrabory P K. Ovarian activity, circulating hormones, and sexual behavior in the cat. I. Relationships during the coitus-induced luteal phase and the estrous period without mating. Biol Reprod. 25:1981;15-28.
- Wildt D E, Guthrie S C, Seager S WJ. Ovarian and behavioral cyclicity of the laboratory maintained cat. Horm Behav. 10:1978;251-257.
- Wildt D E, Seager S WJ, Chakraborty P K. Effect of copulatory stimuli on incidence of ovulation and on serum luteinizing hormone in the cat. Endorinology. 107:1980;1212-1217.
- Wise M E, Nieman D, Stewart J, Nett T M. Effect of number of receptors for gonadotropin-releasing hormone on the release of luteinizing hormone. Biol Reprod. 31:1984;1007-1013.
- Witkin J W, Silverman A J. Synaptology of LHRH neurons in the rat preoptic area. Peptides. 6:1985;263-271.
- Wood R I, Newman S W, Lehman M N, Foster D L. GnRH neurons in the fetal lamb hypothalamus are similar in males and females. Neuroendocrinology. 55:1992;427-433.
- Wray S, Gainer H. Effect of neonatal gonadectomy on the postnatal development of LHRH cell subtypes in male and female rats. Neuroendocrinology. 45:1987;413-419.
- Wray S, Hoffman G. A developmental study of the quantitative distribution of LHRH neurons within the central nervous system of postnatal male and female rats. J Comp Neurol. 252:1986;522-531.
- Wright D E, Jennes L. Origin of noradrenergic projections to GnRH perikarya-containing areas in the medial septum-diagonal band and preoptic area. Brain Res. 621:1993;272-278.
- Wu T J, Segal A Z, Miller G M, Gibson M J. FOS expression in gonadotropin-releasing hormone neurons: Enhancement by steroid treatment and mating. Endocrinology. 131:1992;2045-2050.
- Yang S P, Pau K-Y F, Airhart N, Spies H G. Attenuation of gonadotropin-releasing hormone reflex to coitus by alpha1-adrenergic receptor blockade in the rabbit. Proc Soc Exp Biol Med. 218:1998;204-209.
- Yang S P, Pau K-Y F, Hess D L, Spies H G. Sexual dimorphism in secretion of hypothalamic gonadotropin-releasing hormone and norepinephrine after coitus in rabbits. Endocrinology. 137:1996;2683-2693.
- Yang S P, Pau K-Y F, Spies H G. Tyrosine hydroxylase and norepinephrine transporter mRNA levels increase in locus coeruleus after coitus in rabbits. J Mol Endocrinol. 19:1997;311-319.
- Yen S S, Quigley M E, Reid R L, Ropert J F, Cetel N S. Neuroendocrinology of opioid peptides and their role in the control of gonadotropin and prolactin secretion. Am J Obstet Gynecol. 152:1985;485-493.
- Younglai E V, Wilkinson M, Thompson N. Effects of ovariectomy and estradiol replacement on naloxone induced LH secretion in the female rabbit. J Steroid Biochem. 29:1988;347-351.
- Zarrow M X, Clark J H. Ovulation following vaginal stimulation in a spontaneous ovulator and its implications. J Endocrinol. 69:1968;183-191.
- Zoeller R T, Seeburg P H, Young W S III. In situ hybridization histochemistry for messenger ribonculeic acid (mRNA) encoding gonadotropin-releasing hormone (GnRH): Effect of estrogen on cellular levels of GnRH mRNA in female rat brain. Endocrinology. 122:1988;2570-2577.
- Zohar Y, Elizur A, Sherwood N M, Powell J FF, Rivier J E, Zmora N. Gonadotropin-releasing activities of the three native forms of gonadotropin-releasing hormones present in the brain of gilthead seabream, Sparus aurata. Gen Comp Endocrinol. 97:1995;289-299.
- Zucker I, Goy R W. Sexual receptivity in the guinea pig: Inhibitory and faciltatory actions of progesterone and related compounds. J Comp Physiol Psychol. 64:1967;378-383.