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CONTENTS ACKNOWLEDGEMENTS PREFACE 1. BACKGROUND/INTRODUCTION 1.1. The Concept of Sexual Dimorphisms 1.1.1. McEwen BS and Goy R (1980) Sexually Dimorphic Behavior: Definition and the Organizational Hypothesis. In: Sexual Differentiation of the Brain: Based on a Work Session of the Neurosciences Research Program. MIT Press, Cambridge, pp. 1-12. 1.1.2. McEwen BS and Goy R (1980) Sex Differences in Behavior: Rodents, Birds, and Primates. In: Sexual Differentiation of the Brain: Based on a Work Session of the Neurosciences Research Program. MIT Press, Cambridge, pp. 13-58. 1.1.3. Beach FA (1941) Female mating behavior shown by male rats after administration of testosterone propionate. Endocrinol. 29: 409- 412. 1.1.4. Beach FA and Rasquin P (1942) Masculine copulatory behavior in intact and castrated female rats. Endocrinol. 31: 393-409. 1.2. The Hypothalamic-Pituitary Axis 1.2.1. Raisman G (1997) An urge to explain the incomprehensible: Geoffrey Harris and the discovery of the neural control of the pituitary gland. Ann Rev Neurosci. 20: 533-566 1.2.2. Harris G (1937) The induction of ovulation in the rabbit, by electrical stimulation of the hypothalmo-hypophysial mechanism. Proc Roy Soc Lon B. 612:374-394. 1.2.3. Harris GW and Jacopsohn D (1950) Proliferative capacity of the hypophysial portal vessels. Nature. 165: 854. 1.2.4. Campbell HJ, Feuer G, Harris GW (1964) The effect of intrapituitary infusion of median eminence and other brain extracts on anterior pituitary gonadotrophic secretion. J Physiol. 170:474-486. 1.3. Sexual Differentiation 1.3.1. McClusky N and Naftolin F (1981) Sexual differentiation of the central nervous system. Science 211:1294-1302. 1.3.2. Sinclair AH, Berta P, Palmer MS, Hawkins JR, Griffiths BL, Smith MJ, Foster JW, Frischauf A-M, Lovell-Badge R, Goodfellow PN (1990) A gene from the human sex-determining region encodes a protein with homology to a conserved DNA-binding motif. Nature 346:240- 244. 1.3.3. Haqq CM, King C-Y, Ukiyama E, Falsafi S, Haqq TN, Donahoe PK, Weiss (1995) Molecular basis of mammalian sexual determination: Activation of Mullerian inhibiting substance gene expression by SRY. Science 266:1494-1500. 1.4. The Alignment of Chromosomes, Phenotype and Gender 1.4.1. Page, D.C., Brown, L.G., and De la Chapelle, A. (1987). Exchange of terminal portions of X- and Y- chromosomal short arms in human XX males. Nature 328, 437-440. 1.4.2. Imperato-McGinley J, Guerrero L, Gautier T, Peterson RE (1974) Steroid 5-alpha-reductase deficiency in man: An inherited form of male pseudohermaphroditism. Science 186:1213-1215. 1.4.3. Ahmed SF, Cheng A, Dovey L, Hawkins JR, Martin H, Rowland J, Shimura N, Tait AD, Hughes, IA (2000) Phenotypic features, adnrogen receptor binding, and mutational analysis in 278 clinical cases reported as androgen insensitivity syndrom. J Clin Endo Metab. 85:658-665. 1.4.4. Saavedra-Castillo E, Cortes-Gutierrez El, Davila-Rodriguez MI, Reyes-Martinez ME, Oliveros-Rodriguez A. (2005) 47,XXY female with testicular feminization and positive SRY: a case report. J Repro Med. 50:138-140. 1.4.5. Fausto-Sterling A (1993) The five sexes. The Sciences, March/April, 20-24. 1.5 The Biochemistry and Actions of Steroid Hormones 1.5.1. MacLusky NJ, Clark AS, Naftolin F, Goldman-Rakic PS (1987) Estrogen formation in the mammalian brain: Possible role of aromatase in sexual differentiation of the hippocampus and neocortex. Steroids 50:459-474. 1.5.2. Tsai M, O¿Malley BW (1994) Molecular mechanisms of action of steroid/thyroid receptor superfamily. Ann Rev Biochem. 63:451- 486. 1.5.3. Mani SK, Blaustein JD, Allen JMC, Law SW, O¿Malley BW, Clark JH (1994) Inhibition of rat sexual behavior by antisense oligonucleotides to the progesterone receptor. Endocrinol. 135:1409-1414. 1.5.4. Law SW, Apostolakis EM, Samora PJ, O¿Malley BW, Clark JH (1994) Hormonal regulation of hypothalamic gene expression: Identification of multiple novel estrogen induced genes. J Steroid Biochem. Molec. Biol. 51:131-136. 1.5.5. Smith EP, Boyd J, Frank, GR, Hioyuki T, Cohen RM, Specker B, Williams TC, Lubahn DB, Korach KS (1994) Estrogen resistance caused by a mutation in the estrogen-receptor gene in a man. New England Journal of Medicine 331:1056- 1061. 1.6. Organization and Activation 1.6.1. Phoenix CH, Goy RW, Gerall AA, Young WC (1959) Organizing action of prenatally administered testosterone propionate on the tissues mediating mating behavior in the female guinea pig. Endocrinol. 65:369-382. 1.6.2. Young WC, Goy RW, Phoenix CH (1961) Hormones and sexual behavior. Science 143: 212- 218. 1.6.3. Meany MJ and Steward J (1981) Neonatal androgens influence the social play of prepubescent rats. Hormones and Behavior 15:197-213. 2. CENTRAL NERVOUS SYSTEM DIMORPHISMS 2.1. The Song Bird 2.1.1. Nottebohm F and Arnold A (1976) Sexual dimorphism in vocal control areas of the songbird brain. Science 194:211-213. 2.1.2. Nottebohm F (1980) Testosterone triggers growth of brain vocal control nuclei in adult female canaries. Brain Research 189:429-436. 2.1.3. Brenowitz EA (1991) Altered perception of species-specific song by female birds after lesions of a forebrain nucleus. Science 251: 303-305. 2.2. The Mammalian Spinal Cord 2.2.1. Breedlove M and Arnold AP (1980) Hormone accumulation in a sexually dimorphic motor nucleus of the rat spinal cord. Science 210:564-566. 2.2.2. Breedlove M and Arnold AP (1983) I. Complete demasculinization of the male rat spinal nucleus of the bulnocavernosus using the anti-androgen flutamide. J Neurosci. 3:417-423. 2.2.3. Forger NG and Breedlove M (1986) Sexual dimorphism in human and canine spinal cord: Role of early androgen. Proc. Natl. Acad. Sci. 83:7527-7531. 2.3. The Mammalian Brain 2.3.1. The Anatomy of Cycling 2.3.1.1. Raisman G and Field PM (1973) Sexual dimorphism in the neuropil of the preoptic area of the rat and its dependence on neonatal androgen. Brain Res. 54:1-29. 2.3.1.2. Allen LS and Gorski RA (1990) Sex difference in the bed nucleus of the stria terminalis of the human brain. J Comp Neurol. 302:697-706. 2.3.2. The Anatomy of Difference 2.3.2.1. Gorski RA, Gordon JH, Shryne JE, Southam AM (1978) Evidence for a morphological sex difference within the medial preoptic area of the rat brain. Brain Res.148:333-346. 2.3.2.2. Swaab DF and Fliers E (1985) A sexually dimorphic nucleus in the human brain. Science 228:112-115. 2.3.2.3. Allen LS, Hines M, Shryne JE, Gorski RA (1989) Two sexually dimorphic cell groups in the human brain. J Neurosci. 9:497-506. 2.3.3. Physiological Correlates 2.3.3.1. Arendash GW and Gorski RA (1983) Effects of discrete lesions of the sexually dimorphic nucleus of the proptic area or other medial preoptic regions on the sexual behavior of male rats. Brain Res Bull. 10:147-154. 2.3.3.2. Perachio AA, Marr LD, Alexander M (1979) Sexual behavior in male rhesus monkeys elicited by electrical stimulation of preoptic and hypothalamic areas. Brain Res. 177:127-144. 2.3.3.3. Oomura Y, Yoshimatsu H, Aou S (1983) Medial preoptic and hypothalamic neuronal activity during sexual behavior of the male monkey. Brain Res. 266:340-343. 2.3.4. Neurotransmitter Systems 2.3.4.1. Simerly RB, Swanson LW, Gorski RA. (1984) Demonstration of a sexual dimorphism in the distribution of serotonin- immunoreactive fibers in the medial preoptic nucleus of the rat. J Comp Neurol. 225:151-166. 2.3.4.2. Simerly (1985) Reversal of the sexually dimorphic distribution of serotonin-immunoreactive fibers in the medial preoptic nucleus by treatment with perinatal androgen. Brain Res. 340:91-98. 2.3.4.3. De Vries GJ, Buijs RM, van Leeuwen FW (1984) Sex differences in vasopressin and other neurotransmitter systems in the brain. Prog Brain Res. 61:185-197. 3. MECHANISMS FOR CREATING DIMOPHISMS 3.1. Receptor Mediated Estrogenic Effects 3.1.1. Sar M and Stumpf WE (1977) Distribution of androgen target cells in rat forebrain and pituitary after [3H]-dihydrotestosterone administration. J Ster Biochem. 1131-1135. 3.1.2. Shughrue PJ, Stumpf WE, MacLusky NJ, Zielinski JE, Hochberg RB (1990) Developmental changes in estrogen receptors in mouse cerebral cortex between birth and postweaning: Studied by autoradiography with 11b-methoxy-16a- [125I]iodoestradiol. Endocrinol. 126:1112-1124. 3.1.3. Kudwa A, Bodo EC, Gustafsson J-A, Rissman EF (2005) A previously uncharacterized role for estrogen receptor beta: defeminization of male brain and behavior. PNAS. 102:4608- 4612. 3.1.4. Toran-Allerand CD, Guan X, MacLusky NJ, Horvath TL, Diano S, Singh M, Connolly Jr ES, Nethrapalli IS, Tinnikov A. (2002) ¿ER-X¿: A novel, plasma-membrane-associated, putative estrogen receptor that is regulated during development and following ischemic brain Injury. J. Neurosci., 22 8391-8401. 3.2. Estrogens and Growth Factors 3.2.1. Toran-Allerand D CD (1976) Sex steroids and the development of the newborn mouse hypothalamus and preoptic area in vitro: Implications for sexual differentiation. Brain Res. 106:407-412. 3.2.2. Riesert I, Han V, Lieth E, Toran- Allerand DL, Pilgrim C, Lauder J (1987) Sex steroids promote neurite growth in mesencephalic tyrosine hydroxylase immunoreactive neurons in vitro. Int J Dev Neurosci. 5:91-98. 3.2.3. Toran-Allerand CD, Miranda RC, Bentham WDL, Sohrabji F, Brown TJ, Hochberg RB, MacLusky NJ (1992) Estrogen receptors co- localize with low-affinity nerve growth factor receptors in cholinergic neurons of the basal forebrain. Proc Nat Acad of Science, 89:4668- 4672. 3.3. Estrogens and Plasticity 3.3.1. Jones KJ, Pfaff DW, McEwen BS (1985) Early estrogen-induced nuclear changes in rat hypothalamic ventromedial neurons: An ultrastructural and morphometric analysis. J Comp Neurol. 239:255-266. 3.3.2. Frankfurt M, McEwen BS (1991) 5,7- dihydroxytryptamine and gonadal steroid manipulation alter spine density in ventromedial hypothalamic neurons. Neuroendocrinol. 54:653-657 3.3.3. Woolley CS and McEwen BS (1992) Estradiol mediates fluctuation in hippocampal synapse density during the estrous cycle in the adult rat. J Neurosci. 12:2549-2554. 3.4. Genes 3.4.1. Beyer C, Kolbinger W, Forehlich U, Pilgrim C, Reisert I (1992) Sex differences of hypothalamic prolactin cells develop independently of the presence of sex steroids. Brain Res. 593:253-256. 3.4.2. De Vries GJ, Rissman EF, Simerly RB, Yang LY, Scordalakes EM, Auger CJ, Swain A, Lovell-Badge R, Burgoyne PS, Arnold AP. (2002) A model system for study of sex chromosome effects on sexually dimorphic neural and behavioral traits. J Neurosci.15: 9005-14. 3.5. Experience 3.5.1. McEwen BS (1988) Steroid hormones and the brain: Linking ¿nature¿ and ¿nurture.¿ Neurochem Res. 13:663-669. 3.5.2. Modney BK and Hatton GI (1990) Motherhood modifies magnocellular neuronal interrelationships in functionally meaningful ways. In Krasnegor NA and Bridges RS (Eds) Mammalian Parenting: Biochemical, Neurobiological, and Behavioral Determinants. New York, Oxford University Press. 3.5.3. Xerri C, Stern JM, Merzenich MM (1994) Alterations of the cortical representation of the rat ventrum induced by nursing behavior. J Neurosci. 14:1710-1721. 4. DIMORPHISMS AND COGNITION 4.1 Intellectual and Spatial Abilities 4.1.1. Goldman P, Crawford HT, Stokes LP, Galkin TW, Rosvold HE (1974) Sex-dependent behavioral effects of cerebral cortical lesions in the developing Rhesus monkey. Science 186:540-542. 4.1.2. Witelson SF (1976) Sex and the single hemisphere: Specialization of the right hemisphere for spatial processing. Science 193:425-427. 4.1.3. Inglis J and Lawson JS (1981) Sex differences in the effects of unilateral brain damage on intelligence. Science 212:693-695. 4.1.4. Geschwind N and Behan P (1982) Left- handedness: Association with immune disease, migraine, and developmental learning disorder. Proc Nat Acad Science 79:5097-5100. 4.1.5. Imperato-McGinley M, Pichardo, M, Gautier T, Voyer D, Bryden P (1991) Cognitive abilities in androgen-insensitive subjects: comparison with control males and females from the same kindred. Clin Endocrin. 34:341-347. 4.1.6. Gur RC, Mozley LH, Mozley PD, Resnick SM, Karp JS, Alavi A, Arnold ST, Gur RE (1995) Sex differences in regional cerebral glucose metabolism during a resting state. Science 267:528-531. 4.2. Language Abilities 4.2.1. Kimura D (1983) Sex differences in cerebral organization for speech and praxic functions. Can J Psych. 37:19-35. 4.2.2. Witelson SF, Glezer II, Kigar DL (1995) Women have greater density of neurons in posterior temporal cortex. J Neurosci.15:3418- 3428. 4.2.3. Shaywitz BA, Shaywitz SE, Pugh KR, Constable RT, Skudlarski P, Fulright RK, Bronen RA, Fletcher JM, Shankweiler DP, Katz L, Gore JC (1995) Sex differences in the functional organization of the brain for language. Nature 373:607-609. 4.2.4. Rossell SL, Bullmore ET, Williams CR, David AS (2002) Sex differences in functional brain activation during a lexical visual field task. Brain and Lang 80:97-105. 4.3. Hemispheric Specialization 4.3.1. DeLacoste-Utamsing C and Halloway RL (1982) Sexual dimorphism in the human corpus callosum. Science 216:1431-1432. 4.3.2. Oppenheim JS, Benjamin AB, Lee CP, Nass R, Gazzaniga MS (1987) No sex-related differences in human corpus callosum based on magnetic resonance imagery. Ann Neurol. 21:604- 606. 4.3.3. Halloway RL, Anderson PJ, Defendini R, Harper C (1993) Sexual dimorphism of the human corpus callosum from three independent samples: Relative size of the corpus callosum. Am J Phys Anthropol. 92:481-498. 4.3.4. Allen LS and Gorski RA (1991) Sexual dimorphism of the anterior commissure and massa intermedia of the human brain. J Comp Neurol. 312:97-104. 4.3.5. Allen LS, Richey MF, Chai YM, Gorski RA (1991) Sex differences in the corpus callosum of the living human being. J Neurosci. 11:933- 942. 4.3.6. Rissman EF, Heck AL, Leonard JE, Shupnik MA, Gustafsson JA. (2002) Disruption of estrogen receptor beta gene impairs spatial learning in female mice. PNAS. 99:3996-4001. 4.3.7. Jones BA, Watson NV (2005) Spatial memory performance in androgen insensitive male rats. Physiol Behav 85:135-141. 5. DIMORPHISMS AND IDENTITY 5.1. Female and Male 5.1.1. Slijper FME (1984) Androgens and gender role behaviour in girls with congenital adrenal hyperplasia. Prog Brain Res. 61:417-422. 5.1.2. Mazur T (2005) Gender dysphoria and gender change in androgen insensitivity or micropenis. Arch Sex Behav 34:411-421. 5.1.3. Deeb A, Mason C, Lee YS, Hughes IA (2005) Correlation between genotype, phenotype, and sex of rearing in 111 patients with partial androgen insensitivity syndrome. Clin Endo 63:56-62. 5.2. Gay and Straight 5.2.1. Slimp JC, Hart BL, Goy RW (1978) Heterosexual, autosexual and social behavior of adult male Rhesus monkeys with medial preoptic- anterior hypothalamic lesions. Brain Res. 142:105-122. 5.2.2. Ehrhardt AA, Meyer-Bahlburg HFL, Rosen LR, Feldman JF, Veridiano NP, Zimmerman I (1985) Sexual orientation after prenatal exposure to estrogens. Arch Sex Behav. 14:57. 5.2.3. Swaab DF and Hofman MA (1990) An enlarged suprachiasmatic nucleus in homosexual men. Brain Res. 537:141-148. 5.2.4. LeVay S (1991) A difference in hypothalamic structure between heterosexual and homosexual men. Science 253:1034-1037. 5.2.5. Allen LS and Gorski RA (1992) Sexual orientation and the size of the anterior commissure in the human brain. Proc Nat Acad Sci. 89:7199-7202. 5.2.6. Savic I, Berglund H, Lindstr÷m P (2005) Brain response to putative pheromones in homosexual men. PNAS. 102:7356-7361. 5.3. Transgendered and Gendered 5.3.1. Zucker KJ, Green RJ (1992) Psychosexual disorders in children and adolescents. Child Psychol Psych. 33:107-51. 5.3.2. Gooren LJG, Rao R, van Kessel H, Harmsen-Louman W (1984) Estrogen positive feedback on LH secretion in transsexuality. Psychoneuroendochrinol. 9:249-259. 5.3.3. Zhou J-N, Hofman MA, Gooren LJG, Swaab DF (1995) A sex difference in the human brain and its relation to transsexuality. Nature 378:68-70. 5.3.4. Kruijver F, Zhou J-N, Pool C, Hofman M, Gooren L, Swaab D (2002) Male to female transsexual individuals have female neuron numbers in the central subdivision of the bed nucleus of the stria terminalis. J Clin Endo Metab 85:2034-2041. 6.1. EPILOGUE 6.1.1. Beach FA (1981) Review: Historical origins of modern research on hormones and behavior. Horm Behav. 15: 325-376.
Library of Congress Subject Headings for this publication:
Sex differences.
Sex (Biology).
Sexual dimorphism (Animals).
Brain -- physiology -- Collected Works.
Sex Characteristics -- Collected Works.
Behavior, Animal -- physiology -- Collected Works.
Brain -- anatomy & histology -- Collected Works.
Gonadal Steroid Hormones -- Collected Works.
Sexual Behavior -- physiology -- Collected Works.