Reproduction

The subject of this book is reproduction-specifically, the interplay between reproductive physiology (especially neural and endocrine events) and behavior. In presenting this topic, there are two expository goals. The first is to study repro­ duction at all of the major levels of biological organization-from the molecular (e. g. , hormone receptors in the brain), through the cellular (e. g. , ovarian morphogene­ sis), systemic (e. g. , operation of the hypothalamo-pituitary-ovarian axis), and the organismic levels of organization. Analogously, behavior is treated from the most molecular, elementary, and fundamental components (e. g. , copulatory reflexes), through behavior in the reproductive dyad (e. g. , analysis of female sexual behav­ ior), to complex social behavior (e. g. , the interaction of social context and behav­ ioral sex differences). To the extent that these levels of biological and behavioral organization rep­ resent a "vertical axis" in behavioral neurobiology, a second goal is to treat the "horizontal axis" of biological organization, viz. , time. There are, therefore, treat­ ments of evolutionary origins (e. g. , a phylogenetic survey of psychosexual differ­ entiation), genetic origins in the individual (e. g. , sexual organogenesis), ontoge­ netic development (e. g. , behavioral sexual differentiation), and the immediate physiological precursors of behavior (e. g. , hormonal and nonhormonal initiation of maternal behavior). In addition to tracing the origins of reproduction and reproductive behavior, one extends the time-line from the behavior to its physio­ logical consequences (e. g. , neuroendocrine consequences of sexual behavior).

I Differentiaton and Organization of Reproductive Processes.- 1 Sexual Organogenesis.- Body Sex.- Role of the Fetal Testis.- Dual Nature of the Testicular Chemical Signals.- Testicular Androgens.- Müllerian Inhibitor (or Anti-Müllerian Hormone).- Estrogen Production by the Fetal Ovary.- Gonadal Sex.- Testicular Organogenesis.- Ovarian Morphogenesis.- Control of Gonadal Differentiation.- References.- 2 Gonadal Hormones during Sexual Differentiation in Vertebrates.- Steroid Hormone Production in Developing Birds and Amphibia.- Morphology of the Embryonic Chick Testes.- Biosynthetic Capacity of Chick Gonads for Steroid Production.- Effects of Sex Steroids on Amphibian Development.- Steroid Production of Endocrine Tissues of Developing Amphibia.- Steroid Hormones in Developing Rodents.- Morphological Considerations.- Steroid Hormone Production by Fetal and Neonatal Gonads of Short-Gestation Rodents.- Steroid Hormone Production by Fetal Gonads in Long-Gestation Rodents.- Trophic Control of the Fetal Gonads in Rodents.- Functional Considerations of Fetal Rabbit Gonads.- Fetal Hormones of Domestic Animals.- Production of Steroid Hormones in Primate Fetuses.- Differentiation of the Fetal Testes.- Capacity of the Fetal Testes for Steroid Biosynthesis.- Androgens in the Systemic Circulation of Primate Fetuses.- Other Hormones and Sexual Differentiation.- Summary.- References.- 3 Nonmammalian Psychosexual Differentiation.- Psychosexual Differentiation in the Domestic Chicken (Gallus gallus).- Sex Differences and the Reproductive Behavior of Chickens.- Effects of Hormonal Manipulations before Hatching on the Behavior of Chickens.- Neurobiological Mechanisms of Psychosexual Differentiation in Domestic Fowl.- Psychosexual Differentiation in Japanese Quail (Coturnix coturnix japonica).- Sex Differences and the Reproductive Behavior of Quail.- Effects of Hormone Manipulations before Hatching on Reproductive Behavior.- Psychosexual Differentiation in Zebra Finches (Poephila guttata).- Sex Differences and the Reproductive Behavior of Zebra Finches.- Effect of Early Hormone Manipulations on Zebra Finches.- Psychosexual Differentiation in Other Avian Species.- Psychosexual Differentiation in Reptiles.- Psychosexual Differentiation in Amphibians.- Psychosexual Differentiation in Teleost Fish.- Hermaphroditism in Teleost Fish.- Hormonal Mechanisms of Spontaneous Sex Reversal.- Questions and Theoretical Issues.- Are Paradoxical Effects of Early Hormone Treatments due to Conversion of Androgen to Estrogen?.- Are Early Hormones Responsible for the Differentiation of Ovulation Control Mechanisms?.- Does Any Behavior Not Directly Related to Reproduction Also Differentiate under the Influence of Early Hormones in Nonmammalian Species?.- How Useful Is the Organization/Activation Model for Understanding Nonmammalian Psychosexual Differentiation?.- Is the Empirically Determined Pattern of Hormone Secretion of Embryos or Young Consistent with Conclusions Derived from the Behavioral Data?.- Do Sex Chromosomes or Genes Have a Direct (Nonhormonally Mediated) Role in Psychosexual Differentiation?.- Are Patterns of Psychosexual Differentiation Correlated with Morphological Differentiation or with Sex Chromosome Type in the Different Vertebrate Classes?.- Overview.- Conclusions.- References.- 4 Sexual Behavior Differentiation: Effects of Prenatal Manipulations in Rats.- Sexual Differentiation Theory.- Aromatization Hypothesis.- Critical Period: Postnatal or Pre-Plus Postnatal.- Postnatal Effects.- Strategies for Manipulating Androgenic Hormone Levels in Fetuses.- Additive Effects of Pre-Plus Postnatal Manipulations.- Exchange of Androgen among Fetuses.- Effects of Prenatal Stress on Sexual Differentiation.- Behavioral Syndrome.- Alterations in Plasma Testosterone Levels and Testicular Enzyme Activity.- No Persisting Modification of Morphology.- Therapy for Prenatally Stressed Males.- Prenatal Testosterone Synthesis Inhibition by Opiates.- Prenatal Opioid Syndrome.- Effects on Fetal Plasma Testosterone and Testicular Enzyme Activity.- Possible Role of Endogenous Opiate Release in Mediating the Prenatal Stress Syndrome.- Conclusions.- References.- II Hormonal Activation of Behavior.- 5 Reproductive Physiology and Behavior Interactions in Nonmammalian Vertebrates.- Teleost Fishes.- The Hypothalamo-Pituitary-Gonadal Axis.- Hormone—Behavior Interactions in Teleost Fish.- Amphibians.- The Hypothalamo-Pituitary-Gonadal Axis.- Physiological Factors Controlling Amphibian Reproductive Behavior.- Reptiles.- The Hypothalamo-Pituitary-Gonadal Axis.- Endocrine and Neural Control of Reptilian Reproductive Behavior.- Birds.- The Hypothalamo-Pituitary-Gonadal Axis.- Endocrine and Neural Control of Avian Reproductive Behavior.- Concluding Remarks.- References.- 6 The Role of Gonadal Hormones in the Activation of Feminine Sexual Behavior.- Historical Concepts of Feminine Sexual Behavior.- Structure and Biosynthesis of Gonadal Steroids.- Structure of Gonadal Steroids.- Biosynthesis of Gonadal Steroids.- Steroid Structure and Estrus Induction.- Estrous Cyclicity and Hormone Levels in Intact Females.- Spontaneous Ovulation.- Induced Ovulation (Reflex Ovulation).- Analysis of Feminine Sexual Behavior.- Attractivity.- Proceptivity.- Receptivity.- Conclusions.- References.- 7 Maternal Behavior among the Nonprimate Mammals.- Evolution of Modes of Reproduction among the Mammals.- Patterns of Maternal Care.- Onset of Maternal Behavior during Pregnancy and Near Term.- Behavioral Changes during Pregnancy and Near Term.- Hormonal Basis of the Onset of Maternal Behavior.- Endocrine Control of Pregnancy and Parturition.- Endocrine Stimulation of Maternal Behavior.- Analytical Studies of the Hormonal Basis of Maternal Behavior.- Postpartum Regulation of Maternal Behavior: Behavioral Synchrony between Mother and Young.- Mother’s Dependence on the Young to Maintain Her Maternal Responsiveness.- Nonhormonal Basis of Postpartum Maternal Behavior.- Mother—Young Interaction.- Parturition: The Transition Phase.- Lactation and Maternal Behavior.- Weaning.- Summary.- References.- 8 Peripheral Plasma Levels of Gonadal Steroids in Adult Male and Adult, Nonpregnant Female Mammals.- Androgens in the Systemic Circulation of Adult Male Mammals.- Insectivora and Chiroptera.- Nonhuman Primates.- Humans.- Lagomorpha.- Rodentia.- Cetacea.- Carnivora.- Proboscidea and Hyracoidea.- Perissodactyla.- Artiodactyla.- Estrogens and Progestins in Adult, Nonpregnant Female Mammals.- Insectivora and Chiroptera.- Nonhuman Primates.- Humans.- Lagomorpha.- Rodentia.- Cetacea.- Carnivora.- Proboscidea.- Perissodactyla.- Artiodactyla.- Plasma Steroid Data: Uses and Limitations.- Biochemical Factors That Complicate Interpretation of Plasma Steroid Data.- Physiological Factors That Complicate Interpretation of Plasma Steroid Data.- Environmental and Psychological Factors That Complicate Interpretation of Plasma Steroid Data.- Concluding Remarks.- References.- III Neurological Bases of Mammalian Reproductive Behavior.- 9 Neurological Bases of Male Sexual Behavior: A Comparative Analysis.- Sensory Mechanisms.- Main and Accessory Olfactory System.- Genital Sensory Mechanisms.- Genital Effector Mechanisms.- Penile Erection.- Ejaculation.- Spinal Reflexes.- General Comments.- Sexual Reflexes of Male Dogs.- Sexual Reflexes of Male Rats.- Brain Mechanisms.- Approaches and Problems.- Cerebral Cortex.- Limbic System.- Medial Preoptic-Anterior Hypothalamic Continuum.- Pathways Associated with the Output of the Medial Preoptic-Anterior Hypothalamic System.- Rostral Midbrain Area.- Neural Involvement in Heterotypical Behavior.- Concluding Comment.- References.- 10 Neural Mechanisms of Female Reproductive Behavior.- Approaches to the Neural Mechanisms of Behavior.- Hormones and Reproductive Behavior.- The New Reflexology.- Organization of the Chapter.- Mechanisms of Hormonal Control.- Hormones Involved.- Binding of Sex Hormones in the Brain.- Behavioral and Physiological Effects of Estrogen in Brain Regions Where It Is Bound.- Result of Decreased Estrogen Binding.- Summary.- Sensory Control of the Lordosis Reflex.- Definition of Stimuli That Evoke Lordosis in Female Rats.- Peripheral and Central Sensory Pathways for Lordosis.- Motor Components of Lordosis.- Description of the Response.- Muscles.- Motoneurons.- Descending Pathways Controlling Lordosis.- Hypothalamic Mechanisms.- Brainstem Modules.- Pathways from Brainstem to Spinal Cord.- Working Models of Female Reproductive Behavior Mechanisms.- Reflex Nature of Lordosis Behavior.- Neural Circuitry and Dynamics Underlying Hormone Effects and Lordosis Mechanisms.- References.- 11 Neuropharmacology, Neurotransmitters, and Sexual Behavior in Mammals.- Neuronal Interaction.- Chemical Transmission.- Neuropsychopharmacology.- Neuropsychopharmacology of Sexual Behavior in the Female.- Methods.- Monoamines and Copulatory Behavior in the Female.- Acetylcholine and Copulatory Behavior.- Monoamines and Noncopulatory Sexual Responses.- Neuropsychopharmacology of Sexual Behavior in the Male.- Methods.- Monoamines and Copulatory Behavior in the Male.- Acetylcholine and Copulatory Behavior in the Male.- Monoamines and Noncopulatory Sexual Responses.- Neuropsychopharmacology of Heterotypical Copulatory Behavior.- Neuropeptides, Monoamines, and Sexual Responses.- Neuroanatomical Approaches to the Study of Monoamines in Sexual Behavior.- Effects of Lesions to 5-HT Neurons Using 5, 7-Dihydroxytryptamine.- Effects of Lesions to NA Neurons Using 6-Hydroxydopamine.- Effects of Lesions to DA Neurons Using 6-Hydroxydopamine.- Summary and Conclusions.- References.- 12 Brain Mechanisms and Parental Behavior.- Sensory Mechanisms.- The Development of the Concept of Multisensory Control.- Olfactory Input and the Role of the Olfactory Bulb.- The Role of Afferent Feedback from the Mammary Gland and Nipples.- Additional Afferent Input.- Comments and Conclusions.- Neocortical Mechanisms.- Experimental Findings.- Comments.- Limbic Mechanisms.- Cingulate Cortex.- Hippocampus.- Amygdala.- Septum.- Conclusions.- Hypothalamic Mechanisms.- The Lateral Hypothalamus.- The Medial Preoptic Area and Maternal Behavior in the Rat.- Comparisons between the Neurohormonal Mechanisms Underlying Female Maternal Behavior with Those Underlying Male Sexual Behavior and Female Sexual Behavior in the Rat.- Neurochemical Correlates of Maternal Behavior.- A Neurobehavioral Model.- References.- IV Biochemical Models of Hormonal Action.- 13 The Role of Metabolism in Hormonal Control of Sexual Behavior.- The Prohormone Concept.- Male Sexual Behavior.- Circulating Androgens and Estrogens in Males.- Hypothesis: T Mediates Male Sexual Behavior via Metabolism to A.- Hypothesis: T Mediates Male Sexual Behavior via Metabolism to DHT.- Hypothesis: T Mediates Male Sexual Behavior via Metabolism to Other Androgens.- Hypothesis: T Mediates Male Sexual Behavior via Metabolism to Estrogen.- Hypothesis: T Mediates Male Sexual Behavior via Metabolism to Estrogen plus DHT.- Female Sexual Behavior.- Secretion Products of the Ovaries: Estrogens.- Secretion Products of the Ovaries: Progestins.- Progestin Metabolism.- Inhibitory Effects of Progestins.- Sexual Differentiation.- The Role of 5?-Reduction.- The Role of Other Androgens.- The Role of Aromatization.- Conclusions.- References.- 14 Steroid Hormone Receptors in Brain and Pituitary: Topography and Possible Functions.- Cellular Mechanism of Steroid Hormone Action.- Visualization of Genomic Effects of a Steroid Hormone.- Cellular Uptake and Retention of Radioactive Steroid Hormones.- Cell Nuclear Retention of Radioactive Steroid Hormones.- The Search for Cell Nuclear Acceptor Sites.- Effects of Steroid Hormones on Gene Transcription.- Critera for “Receptors”.- Properties and Topography of Putative Steroid Receptors in Brain.- Estradiol.- Testosterone.- Progestins.- Glucocorticoids.- Mineralocorticoids.- Neurochemical Consequences of Steroid Interactions with Putative Neural Receptors as Illustrated for Estradiol.- Temporal Relationships of Receptor Occupation to Physiological Effects.- Alterations of RNA Formation.- Changes in Protein Formation.- Estrogen Effects on Pituitary Sensitivity to LHRH.- Hypothalamic Peptidases.- Oxidative Metabolism.- Catecholamines and Serotonin.- Acetylcholine.- Enzymes as Neurochemical Markers for Estrogen Receptor Function.- Summary and Prospectus: A Model for Steroid Hormone Action on Neuronal Functioning.- References.- V Social and Experiential Influences on Reproduction.- 15 Neuroendocrine Consequences of Sexual Behavior.- Hormone—Behavior Interactions.- Patterns of Neuroendocrine Control: Ovulation and Luteal Function.- Behavioral Effects on Ovulation.- Reflexive Ovulation.- Spontaneous Ovulation.- Behavioral Induction of Progestational Hormone Secretion.- Components of Mating.- Receptors and Afferent Pathways.- Pituitary Involvement.- Storage of the Copulatory Stimulus.- CNS Mediation.- Summary.- References.- 16 On Measuring Behavioral Sex Differences in Social Contexts.- Objections to Behavioral-Sex-Differences Research in Animals.- The Generalization Problem.- The “Assumed Biological Mediation” Problem.- Behavioral Endocrinology and Sex Differences Research.- Measuring Behavioral Sex Differences.- Definition of Behavioral Sex Differences.- Dimorphic Behavior Assessed Individually.- Dimorphic Behavior Assessed in Dyads.- Dimorphic Behavior Assessed in Groups.- Conclusion.- References.