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Emerging concepts in the regulation of female sexual behavior.

Recent advances in the field of steroid hormone action have significantly advanced our understanding of neuroendocrine regulation of sexual behavior in female rodents. The presumed classical steroid receptor-steroid hormone functional relationship has undergone significant modifications to integrate the wider context of cellular signaling mechanisms initiated by sensory and environmental stimuli and their transcriptional regulation of steroid hormone receptors in reproductive behavior. This effort has greatly been aided by recent studies identifying steroid hormone receptors as transcriptional mediators of a variety of ligands, whose functional flexibility is dependent upon their recruitment of coregulators, and the availability of gene knockout animal models. This review provides a framework for current concepts in the field of steroid hormone action in the context of their regulation of female sexual behavior.

Animals↗

Oxytocin and complex social behavior: species comparisons.

The neurohypophyseal peptide hormone oxytocin functions as a neuropeptide in several brain areas in addition to its role as a posterior pituitary hormone. Several studies have determined significant differences in patterns of oxytocin receptor binding in the brains of two closely related species of vole. One of the defining features of these two species is remarkably different reproductive behavior strategies. The prairie vole forms long-term monogamous relationships; the montane vole is polygamous. One potential measure of the formation of a pair bond in prairie voles is the development of intense aggressive behavior directed at male conspecifics following a mating bout. Oxytocin had little effect on aggressive behavior when administered before mating but had profound effects on the aggression of male prairie voles when administered after mating. Oxytocin had relatively modest effects on the behavior of montane voles, and neither the behavior nor the peptide effects were affected by mating experience. The data indicate that differences in peptide binding in these two species of vole may be functionally related to difference in social behavior.

Animals↗

Women's position and family planning in Egypt.

In this report, data from the 1988 Egypt Demographic and Health Survey are used to address some of the most frequently raised questions about the relationship between gender inequality and reproductive behavior. The findings from binomial and multinomial logit models show that while the relationship between women's position and fertility control in Egypt is complex, some clear, broad patterns exist that have important theoretical and policy implications. First, although women's status in Egypt is clearly multidimensional, the reproductive aspect of women's position has a strong connection with the nonreproductive dimensions. Second, the case of the continued use of education and employment as proxies of women's position, especially in relationship to fertility control, is considerably discredited by the results. Finally, the findings indicate that Egyptian culture supports gender equality in the form of interaction and negotiation rather than women's autonomy.

Adolescent↗

An efficient screen for reproductive pathways using mature seeds of monocots and dicots.

Seed samples of 32 species (obligate and facultative sexuals and apomicts of monocots and dicots) were investigated by flow cytometry to reveal the pathway of reproduction. Ten different pathways of seed formation could be reconstructed considering whether the female and/or male gametes were reduced or unreduced, the embryos arose via the zygotic or parthenogenetic route and the endosperm via the pseudogamous or autonomous route. The screen is suited to select sporophytic or gametophytic mutants in sexual species, to identify pure sexual or obligate apomictic genotypes from facultative apomictic species, and to analyze the inheritance of the individual reproductive processes. Corresponding unique results are presented for Arabidopsis, Arabis, Hypericum and Poa. The screen of mature seeds by flow cytometry yielded more information about the reproductive behavior of individual plants than any other available test, and is very useful both in basic research and plant breeding.

Plant Physiological Phenomena↗

Fertility and family planning in Vietnam: evidence from the 1994 Inter-censal Demographic Survey.

Results from the 1994 Vietnam Inter-censal Demographic Survey reveal substantial change over recent years in reproductive behavior and attitudes. Fertility has continued to decline to a level not far above a total fertility rate of three children per woman. Compared with the late 1980s, contraceptive knowledge has broadened and contraceptive prevalence has increased, reaching a level of 65 percent of currently married women of reproductive age. The dominance of the IUD among modern methods has been reduced somewhat. Stated family-size preferences have shifted noticeably downward. Recently married women indicate that they want only 2.3 children, on average, suggesting that fertility will continue to fall in coming years. These findings suggest that Vietnam is in the midst of a transition that will lead to low levels of fertility in the near future.

Abortion, Induced↗

Disruption of sexual behavior in male aromatase-deficient mice lacking exons 1 and 2 of the cyp19 gene.

Aromatase is known to be involved in the control of morphological, neuroendocrine, and behavioral sex differences in the brain. To study the detailed functions of aromatase in mouse brain, we generated aromatase-deficient mice that lack exons 1 and 2 and the proximal promoter region of the cyp19 gene by homologous recombination. The genotypes of offspring from heterozygous matings conformed to the Mendelian rule. The aromatase knockout (ArKO) male and female mice displayed no significant external morphological abnormalities, although the uteri of adult ArKO female mice were remarkably underdeveloped. Male-typical sexual behavior directed toward females was, however, strongly modified in the ArKO male mice. The ArKO male mice showed significantly prolonged latencies to mount and decreased numbers of mounts in response to receptive stimulus females. These results suggest an important participation of the aromatase gene in the reproductive behaviors.

Animals↗

Fos expression in rat pontine tegmental neurons following activation of the medial preoptic area.

Fos immunohistochemistry was used to map the distribution of pontine neurons excited by activation of the medial preoptic area (MPO). Although we have previously shown that Barrington's nucleus receives a very dense focal input from the MPO, electrical stimulation of the preoptic area unexpectedly induced very little Fos expression in Barrington's neurons. These results suggest that the MPO-->Barrington's projection utilizes a transmitter(s) that does not involve transduction of the Fos protein; alternatively, MPO afferents to Barrington's nucleus may be inhibitory in nature. As Barrington's nucleus plays a critical role in micturition, MPO projections to Barrington's nucleus may regulate voiding reflexes during sexual behavior. Interestingly, while the locus coeruleus (LC) proper receives only a sparse projection from the MPO, extensive Fos expression was present in LC. The finding of Fos immunoreactive LC neurons suggests that the excitatory influence of MPO may regulate LC neuronal activity and NE release during reproductive behaviors.

Animals↗

Developmental dissociation of excitability and secretory ability in Aplysia bag cell neurons.

1. Despite the considerable progress made in understanding the role of electrical activity in triggering secretion, the developmental relationships between excitability and secretion are not well understood. The well-characterized bag cell neurons of Aplysia provide an advantageous system in which to investigate developmental interactions of these two key properties of neurons. 2. A prolonged afterdischarge triggers egg laying hormone (ELH) secretion in mature bag cell neurons. To investigate secretion in the developmental framework of excitability, we first examined whether immature neurons, which are incapable of the mature form of excitability (afterdischarge), contain ELH and whether this hormone is packaged in vesicles. We used immunoelectron microscopy to compare vesicular localization of ELH and to compare the size and density of ELH-containing vesicles in neurons from adult and juvenile Aplysia. This comparison revealed that immature neurons contain ELH in vesicles in the size range of secretory vesicles. However, they lack a class of large vesicles (> 250 nm in diameter) that is characteristic of mature neurons. 3. To investigate whether the ELH contained in immature bag cell neurons could be secreted in response to electrical activity, we used the potassium channel blocker tetraethylammonium (TEA) combined with nerve stimulation to depolarize neurons from both juvenile animals (ovotestes do not contain eggs) and from adult Aplysia (ovotestes contain eggs). Using radioimmunoassay, we have found that the duration and amount of ELH secreted from bag cell neurons from juvenile Aplysia in response to TEA does not depend on whether or not the cells can be induced to afterdischarge, and the amount and duration of ELH secreted from bag cell neurons of juvenile Aplysia (whether or not they afterdischarged) differed from those secreted by adult neurons. However, by normalizing for body size, we found that the final estimated hemolymph concentration of ELH would be similar in juvenile and adult animals. 4. We investigated the potential functional significance of secretion of bag cell hormones in juvenile Aplysia by attempting to bypass the bag cell neurons and directly activate downstream elements with extract from adult bag cell neurons (BCE), known to contain ELH and other peptides. We found that juvenile Aplysia exhibit at least one component of egg-laying behavior, cessation of locomotion, in response to BCE during a developmental period (as measured by weight) in which they normally would possess neurons incapable of afterdischarge. Thus developmental regulation of excitability in the bag cell neurons may prevent inappropriate hormone release and subsequent premature expression of reproductive behaviors.

Animals↗

Neurogenesis in juvenile and adult ring doves.

Studies in songbirds suggest that neurogenesis during the first few years of life is related to song learning. In this study, we examined whether postnatal neurogenesis occurs in a nonsongbird, the ring dove (Streptoplia risoria), and whether it persists to old age. Twenty-four hours after a single intramuscular injection of [3H]thymidine, labeled cells were present in the brains, particularly in the lateral wall of the lateral ventricle of juvenile (3-month and 8-month) and adult (1-year to 8-year) doves. Two months after multiple [3H]thymidine injections, there were fewer labeled cells in the ventricular zone (VZ), but many labeled cells with neuronal morphology in the parenchyma of the forebrain; labeled cells were confirmed as neurons by using neuron-specific markers, microtubule-associated protein-2 (MAP-2) and anti-neuronal nucleus (NeuN). In general, new neurons were distributed in the forebrain without clustering in any particular nucleus. During the first year of life, however, neostriatum caudale and hyperstriatum, the regions known to be essential for proper integration of sensory cues and reproductive behavior, contained more new neurons than any other brain regions. These neuronal additions showed an age-related decline; the first reduction coincided with the dove's attainment of adult physical size (about 3 months old) and the second occurred when the dove would normally attain reproductive fitness (about 1 year old). A low level of forebrain neurogenesis persisted up to 8 years of age (the oldest animals studied). These observations suggest that neurogenesis in adulthood is widespread among birds but that the biological significance of adult neurogenesis in the ring dove remains to be determined.

Aging↗

Central administration of chicken gonadotropin-releasing hormone-II enhances courtship behavior in a female sparrow.

Like most vertebrates, birds have two forms of gonadotropin-releasing hormone (GnRH). Chicken GnRH-I (cGnRH-I) is released at the median eminence to elicit gonadotropin release; chicken GnRH-II (cGnRH-II) is thought to be non-hypophysiotropic and its function is unclear. Both forms are hypothesized to act as neurotransmitters in the control of reproductive behavior. In the present study, we implanted chronic cannulae aimed at the third ventricle in female white-crowned sparrows (Zonotrichia leucophrys gambelii) to test the effects of both forms of GnRH on copulation solicitation, a female courtship behavior. This behavior can be elicited in captive, estrogen-primed females by playing a recording of male song. We quantified the behavioral response to recorded song 30, 60, and 90 min after intracerebroventricular infusion of cGnRH-I, -II, or saline. cGnRH-II, but not cGnRH-I, increased solicitation behavior compared with saline 30 min after infusion. Under control conditions, responses to the playback diminish from the 30-min to the 90-min time point. Responses after cGnRH-II infusion followed a similar pattern, whereas after cGnRH-I, there was no significant response decrement. cGnRH-I appears to maintain the level of solicitation seen at 30 min after infusion. Our results suggest a behavioral role for cGnRH-II that may be independent of cGnRH-I.

Animals↗

Phenotypic analysis of mice deficient in the type 2 galanin receptor (GALR2).

Galanin is a neuropeptide implicated in the regulation of feeding, reproduction, cognition, nociception, and seizure susceptibility. There are three known galanin receptor (GALR) subtypes (GALR1, GALR2, and GALR3), which bind to galanin with different affinities and have their own unique distributions, signaling mechanisms, and putative functions in the brain and peripheral nervous system. To gain further insight into the possible physiological significance of GALR2, we created mutant mice that were deficient in GALR2 and compared their phenotype to that of wild-type (WT) littermate or age-matched controls, with respect to basic motor and sensory function, feeding behavior, reproduction, mood, learning and memory, and seizure susceptibility. Phenotypic analysis revealed that animals bearing a deletion of GALR2 did not differ significantly from their WT controls in any of the measured variables. We conclude that either GALR2 plays no role in these physiological functions or through redundancy or compensation these mutant animals can adapt to the congenital absence of GALR2. It is also conceivable that GALR2 plays only a subtle role in some of these functions and that the impact of its loss could not be detected by the analytical procedures used here.

Animals↗

Functional studies of corticosterone receptors in neuronal membranes.

This work reviews evidence that some physiological and behavioral responses to steroid hormones use membrane-associated receptors. The review emphasizes research with an amphibian model, Taricha granulosa, but also cites examples from mammalian research. Many studies document steroid responses that occur within a time frame of a few milliseconds or minutes. In Taricha, corticosterone rapidly inhibits reproductive behaviors and causes site-specific changes in neurotransmitter concentrations and neuronal activity. Ligand binding assays using radiolabeled corticosterone and neuronal membranes from Taricha (and other animals) provide evidence that there are high-affinity steroid receptors in neuronal membranes. Subcellular fractionation, autoradiography, and immunocytochemistry add support to the conclusion that there are steroid receptors in neuronal membranes. Other studies indicate that, in Taricha and other animals, there are two types of membrane-associated steroid receptors--ligand-gated ion channels (specifically, the GABAA receptor) and G-protein coupled receptors.

Animals↗

Saccular potentials of the vocal plainfin midshipman fish, Porichthys notatus.

The plainfin midshipman fish, Porichthys notatus, is a vocal species of teleost fish that generates acoustic signals for intraspecific communication during social and reproductive behaviors. All adult morphs (females and males) produce single short duration grunts important for agonistic encounters, but only nesting males produce trains of grunts and growls in agonistic contexts and long duration multiharmonic advertisement calls to attract gravid females for spawning. The midshipman fish uses the saccule as the main acoustic endorgan for hearing to detect and locate vocalizing conspecifics. Here, I examined the response properties of evoked potentials from the midshipman saccule to determine the frequency response and auditory threshold sensitivity of saccular hair cells to behaviorally-relevant single tone stimuli. Saccular potentials were recorded from the rostral, medial and caudal regions of the saccule while sound was presented by an underwater speaker. Saccular potentials of the midshipman, like other teleosts, were evoked greatest at a frequency that was twice the stimulus frequency. Results indicate that midshipman saccular hair cells of non-reproductive adults had a peak frequency sensitivity that ranged from 75 (lowest frequency tested) to 145 Hz and were best suited to detect the low frequency components (<or=105 Hz) of midshipman vocalizations.

Acoustic Stimulation↗

Insulin-induced repartitioning of metabolic fuels inhibits hamster estrous behavior: role of area postrema.

Excessive diversion of metabolic fuels away from oxidation and into adipose tissue storage depots, such as underfeeding or extraordinary levels of energy expenditure, can induce nutritional infertility. Treatment with pharmacological doses of insulin reduces metabolic fuel availability and suppresses both ovulatory cyclicity and pulsatile luteinizing hormone release in females of several mammalian species, but little attention has been paid to the effects of insulin treatments on reproductive behaviors. Ovariectomized Syrian hamsters were injected with long-acting insulin every 12 h for 72 h and were prevented from overeating by limiting their intake to approximately 110% of pretreatment levels. When given estradiol and progesterone, insulin-treated hamsters exhibited significantly reduced levels of sexual receptivity compared with saline-treated controls. This insulin-induced inhibition of estrous behavior was prevented by lesions of the area postrema. Insulin treatments also caused changes in the number of detectable estrogen receptor immunoreactive cells in the hypothalamus and preoptic area. Therefore, insulin-induced repartitioning of metabolic fuels induces changes in estrous behavior and neural estrogen receptors that are indistinguishable from those caused by food deprivation, cold exposure, or treatment with metabolic inhibitors.

Animals↗

New reproductive anomalies in fruitless-mutant Drosophila males: extreme lengthening of mating durations and infertility correlated with defective serotonergic innervation of reproductive organs.

Several features of male reproductive behavior are under the neural control of fruitless (fru) in Drosophila melanogaster. This gene is known to influence courtship steps prior to mating, due to the absence of attempted copulation in the behavioral repertoire of most types of fru-mutant males. However, certain combinations of fru mutations allow for fertility. By analyzing such matings and their consequences, we uncovered two striking defects: mating times up to four times the normal average duration of copulation; and frequent infertility, regardless of the time of mating by a given transheterozygous fru-mutant male. The lengthened copulation times may be connected with fru-induced defects in the formation of a male-specific abdominal muscle. Production of sperm and certain seminal fluid proteins are normal in these fru mutants. However, analysis of postmating qualities of females that copulated with transheterozygous mutants strongly implied defects in the ability of these males to transfer sperm and seminal fluids. Such abnormalities may be associated with certain serotonergic neurons in the abdominal ganglion in which production of 5HT is regulated by fru. These cells send processes to contractile muscles of the male's internal sex organs; such projection patterns are aberrant in the semifertile fru mutants. Therefore, the reproductive functions regulated by fruitless are expanded in their scope, encompassing not only the earliest stages of courtship behavior along with almost all subsequent steps in the behavioral sequence, but also more than one component of the culminating events.

Animals↗

The brain decade in debate: VIII. Peptide hormones and behavior: cholecystokinin and prolactin.

This article is a transcription of an electronic symposium held on November 28, 2000 in which active researchers were invited by the Brazilian Society of Neuroscience and Behavior (SBNeC) to discuss the advances of the last decade in the peptide field with particular focus on central actions of prolactin and cholecystokinin. The comments in this symposium reflect the diversity of prolactin and cholecystokinin research and demonstrate how the field has matured. Since both peptides play a role in reproductive behaviors, particularly mother-infant interactions, this was the starting point of the discussion. Recent findings on the role of the receptor subtypes as well as interaction with other peptides in this context were also discussed. Another issue discussed was the possible role of these peptides in dopamine-mediated rewarding systems. Both prolactin and cholecystokinin are involved in mechanisms controlling food intake and somatic pain thresholds. The role of peripheral inputs through vagal afferents modulating behavior was stressed. The advent of knockout animals as potential generators of new knowledge in this field was also addressed. Finally, interactions with other neuropeptides and investigation of the role of these peptides in other fields such as immunology were mentioned. Knowledge about the central functions of prolactin and cholecystokinin has shown important advances. The role of these peptides in neurological and psychiatric syndromes such as anorexia, drug abuse and physiological disturbances that lead to a compromised maternal behavior seems relevant.

Brain↗

Central vasotocin-immunoreactive system in a male passerine bird (Junco hyemalis).

Previous investigations have identified regions of the avian brain that contain immunoreactive vasotocinergic (VT-ir) cell bodies and fibers. These studies exclusively used domesticated species, and the relevance of the findings for free-living birds has not been established. The present study used immunocytochemistry to determine the neuroanatomical distribution of the VT-ir system in the brain of a well-studied male passerine bird (dark-eyed junco, Junco hyemalis) obtained from a natural population in interior Alaska (65 degrees N, 147 degrees W). VT-ir cell bodies were observed in several brain regions (paraventricular and supraoptic nuclei, nucleus of the stria terminalis), where they have been described in other oscine species. VT-ir fibers were widespread in many brain regions and were especially abundant in the medial preoptic nucleus, the basal region of the septum, and the hypothalamic-neurohypophyseal tract. Fibers were also present in brain regions that are involved in the control of vocal behavior including the ventromedial capsular region of the nucleus robustus archistriatalis and the dorsomedial portion of the mesencephalic nucleus intercollicularis. The widespread brain distribution of VT-ir cell bodies and fibers in juncos generally resembles that of domestic birds and suggests a role for this neuropeptide in the control of reproductive behavior and physiology.

Animals↗

Molecular phylogeny of the model annelid Ophryotrocha.

Annelids of the genus Ophryotrocha are small opportunistic worms commonly found in polluted and nutrient-rich habitats such as harbors. Within this small group of about 40 described taxa a large variety of reproductive strategies are found, ranging from gonochoristic broadcast spawners to sequential hermaphroditic brooders. Many of the species have a short generation time and are easily maintained as laboratory cultures. Thus they have become a popular system for exploring a variety of biological questions including developmental genetics, ethology, and sexual selection. Despite considerable behavioral, reproductive, and karyological studies, a phylogenetic framework is lacking because most taxa are morphologically similar. In this study we use 16S mitochondrial gene sequence data to infer the phylogeny of Ophryotrocha strains commonly used in the laboratory. The resulting mtDNA topologies are generally well resolved and support a genetic split between hermaphroditic and gonochoristic species. Although the ancestral state could not be unambiguously identified, a change in reproductive strategy (i.e., hermaphroditism and gonochorism) occurred once within Ophryotrocha. Additionally, we show that sequential hermaphroditism evolved from a simultaneous hermaphroditic ancestor, and that characters previously used in phylogenetic reconstruction (i.e., jaw morphology and shape of egg mass) are homoplasic within the group.

Animals↗