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Orgasm
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This study compared the effects of centrally administered oxytocin (OT) and arginine vasopressin (AVP) on partner preference formation and social contact in male and female prairie voles (Microtus ochrogaster). After 1 hr of cohabitation and pretreatment with either AVP or OT, both males and females exhibited increased social contact and significant preference for the familiar partner. After pretreatment with either an OT receptor antagonist (OTA) or an AVP (V1a) receptor antagonist (AVPA), neither OT nor AVP induced a partner preference. In addition, treatment with OT+OTA or AVP+AVPA was associated with low levels of social contact in both sexes. Either AVP or OT is sufficient to facilitate social contact if either the OT or AVP receptor is available. However, the formation of partner preferences may require access to both AVP and OT receptors.
A kinship between cranial and pelvic visceral nerves of vertebrates has been accepted for a century. Accordingly, sacral preganglionic neurons are considered parasympathetic, as are their targets in the pelvic ganglia that prominently control rectal, bladder, and genital functions. Here, we uncover 15 phenotypic and ontogenetic features that distinguish pre- and postganglionic neurons of the cranial parasympathetic outflow from those of the thoracolumbar sympathetic outflow in mice. By every single one, the sacral outflow is indistinguishable from the thoracolumbar outflow. Thus, the parasympathetic nervous system receives input from cranial nerves exclusively and the sympathetic nervous system from spinal nerves, thoracic to sacral inclusively. This simplified, bipartite architecture offers a new framework to understand pelvic neurophysiology as well as development and evolution of the autonomic nervous system.
http://science.sciencemag.org/content/354/6314/893
Background
Although humans experience orgasms with a degree of statistical regularity, they remain among the most enigmatic of sexual responses; difficult to define and even more difficult to study empirically. The question of whether animals experience orgasms is hampered by similar lack of definition and the additional necessity of making inferences from behavioral responses.
Method
Here we define three behavioral criteria, based on dimensions of the subjective experience of human orgasms described by Mah and Binik, to infer orgasm-like responses (OLRs) in other species: 1) physiological criteria that include pelvic floor and anal muscle contractions that stimulate seminal emission and/or ejaculation in the male, or that stimulate uterine and cervical contractions in the female; 2) short-term behavioral changes that reflect immediate awareness of a pleasurable hedonic reward state during copulation; and 3) long-term behavioral changes that depend on the reward state induced by the OLR, including sexual satiety, the strengthening of patterns of sexual arousal and desire in subsequent copulations, and the generation of conditioned place and partner preferences for contextual and partner-related cues associated with the reward state. We then examine whether physiological and behavioral data from observations of male and female rats during copulation, and in sexually-conditioned place- and partner-preference paradigms, are consistent with these criteria.
Results
Both male and female rats display behavioral patterns consistent with OLRs.
Conclusions
The ability to infer OLRs in rats offers new possibilities to study the phenomenon in neurobiological and molecular detail, and to provide both comparative and translational perspectives that would be useful for both basic and clinical research.
We have demonstrated that sexual activity produces transient sympathoadrenal activation and a pronounced, long-lasting increase in prolactin in men and women. However, by analyzing endocrine alterations at 10-min intervals, a precise assignment of these changes to the pre-, peri-and postorgasmic periods was not possible. Thus, the current study aimed to accurately differentiate the endocrine response to sexual arousal and orgasm in men using an automatic blood collection technique with 2-min sampling intervals. Blood was drawn continuously before, during and after orgasm over a total period of 40 min in 10 healthy subjects and were compared with samples obtained under a control condition. Sexual activity induced transient increases of plasma epinephrine and norepinephrine levels during orgasm with a rapid decline thereafter. In contrast, prolactin levels increased immediately after orgasm and remained elevated throughout the experiment. Although oxytocin was acutely increased after orgasm, these changes were not consistent and did not reach statistical significance. Vasopressin, LH, FSH and testosterone plasma concentrations remained unaltered during sexual arousal and orgasm. These data confirm that prolactin is secreted after orgasm and, compared with oxytocin, seems to represent a more reliable and sustained marker for orgasm in man. The results further reinforce a role for prolactin either as a neuroendocrine reproductive reflex or as a feedback mechanism modulating dopaminergic systems in the central nervous system that are responsible for appetitive behavior.
The evolutionary basis of women's orgasm is unknown, but the most favoured theory involves putative physiological processes that accompany orgasm and increase the likelihood of fertilization during intercourse. In a sample of over 8000 female twins and siblings, we show a substantial genetic basis to both orgasm rate and number of offspring. While there was a very weak but significant phenotypic correlation between the two variables, there was no corresponding genetic correlation, suggesting the involvement of environmental confounders. Controlling for relationship length and frequency of sexual intercourse eliminated the significant phenotypic correlation between orgasm rate and number of offspring, which suggests no substantive causal relationship between orgasm and fertility. These results cast doubt on the predominant functional theory of female orgasm.
Article selected to feature in Wiley's News Round-Up (it is a biweekly mailing sent to over 1,800 subscribing journalists). October 06, 2014 Wiley Press Release: http://eu.wiley.com/WileyCDA/PressRelease/pressReleaseId-112610.html
Sexual medicine experts and sexologists must spread certainties on the biological basis of the female orgasm to all women, not hypotheses or personal opinions. Therefore, they must use scientific anatomical terminology. The anatomy of the clitoris and the female orgasm are described in textbooks, but some researchers have proposed a new anatomical terminology for the sexual response in women. The internal/inner clitoris does not exist: the entire clitoris is an external organ. The clitoris is not composed of two arcs but of the glans, body, and crura or roots. “Clitoral bulbs” is an incorrect term from an embryological and anatomical viewpoint: the correct term is “vestibular bulbs.” The bulbocavernosus muscles are implicated in inferior vaginismus, while the pubovaginal muscle is responsible for superior vaginismus. The clitoral or clitoris-urethro-vaginal complex has no embryological, anatomical and physiological support: the vagina has no anatomical relationship with the clitoris, and the clitoris is a perineal organ while the supposed G-spot is in the pelvic urethra. G-spot/vaginal/clitoral orgasm, vaginally activated orgasm, and clitorally activated orgasm, are incorrect terms: like “male orgasm,” “female orgasm” is the correct term. The “vaginal” orgasm that some women report is always caused by the surrounding erectile organs (triggers of female orgasm). The male penis cannot come in contact with the venous plexus of Kobelt or with the clitoris during vaginal intercourse. Also, female ejaculation, premature ejaculation, persistent genital arousal disorder (PGAD), periurethral glans, vaginal–cervical genitosensory component of the vagus nerve, and G-spot amplification, are terms without scientific basis. Female sexual satisfaction is based on orgasm and resolution: in all women, orgasm is always possible if the female erectile organs, i.e. the female penis, are effectively stimulated during masturbation, cunnilingus, partner masturbation, or during vaginal/anal intercourse if the clitoris is simply stimulated with a finger. Clin. Anat., 2014. © 2014 Wiley Periodicals, Inc.
The claims by Emmanuele Jannini, Odile Buisson, Helen O’Connell, Beverly Whipple, Adam Ostrzenski, Stuart Brody, Irwin Goldstein (The Journal of Sexual Medicine), Chiara Simonelli and others, have no scientific basis.
The pituitary gland plays an important role in basic survival mechanisms by releasing fluctuating amounts of hormones into the bloodstream, depending on the circumstances the individual finds itself. However, despite these changes in pituitary hormonal production, neuroimaging studies have never been able to demonstrate changes in the activation level of the pituitary. The most apparent reason is the much higher blood flow rate in the pituitary than in the brain. However, the present PET-scanning study demonstrates for the first time that neuroimaging techniques can identify increased pituitary activity. In a study with 11 healthy women sexual orgasm compared to rest caused an increased blood supply to the pituitary. We assume that this increase signifies elevated pituitary activation in order to produce higher plasma concentrations of oxytocin and prolactin. These hormones induce vaginal and uterus movements, ovulation and enhancement of sperm and egg transport. No increased blood supply was observed comparing clitoral stimulation, orgasm attempt, and faked orgasm with rest. In a study with 11 healthy men comparing ejaculation with rest did not reveal increased pituitary activation, probably because ejaculation causes a much lower increase of oxytocin and prolactin plasma concentration than female orgasm.
Whether women's orgasm is an adaptation is arguably the most contentious question in the study of the evolution of human sexuality. Indeed, this question is a veritable litmus test for adaptationism, separating those profoundly impressed with the pervasive and myriad correspondences between organisms' phenotypes and their conditions of life from those who apply the "onerous concept" of adaptation with more caution, skepticism or suspicion. Yet, the adaptedness of female orgasm is a question whose answer will elucidate mating dynamics in humans and nonhuman primates. There are two broad competing explanations for the evolution of orgasm in women: (1) the mate-choice hypothesis, which states that female orgasm has evolved to function in mate selection and (2) the byproduct hypothesis, which states that female orgasm has no evolutionary function, existing only because women share some early ontogeny with men, in whom orgasm is an adaptation. We review evidence for these hypotheses and identify areas where relevant evidence is lacking. Although additional research is needed before firm conclusions can be drawn, we find that the mate-choice hypothesis receives more support. Specifically, female orgasm appears to have evolved to increase the probability of fertilization from males whose genes would improve offspring fitness.
Sexual dysfunction is a common side effect of antidepressants and can have significant impact on the person’s quality of life, relationships, mental health, and recovery. The reported incidence of sexual dysfunction associated with antidepressant medication varies considerably between studies, making it difficult to estimate the exact incidence or prevalence. The sexual problems reported range from decreased sexual desire, decreased sexual excitement, diminished or delayed orgasm, to erection or delayed ejaculation problems. There are a number of case reports of sexual side effects, such as priapism, painful ejaculation, penile anesthesia, loss of sensation in the vagina and nipples, persistent genital arousal and nonpuerperal lactation in women. The focus of this article is to explore the incidence, pathophysiology, and treatment of antidepressant iatrogenic sexual dysfunction.
In men and women sexual arousal culminates in orgasm, with female orgasm solely from sexual intercourse often regarded as a unique feature of human sexuality. However, orgasm from sexual intercourse occurs more reliably in men than in women, likely reflecting the different types of physical stimulation men and women require for orgasm. In men, orgasms are under strong selective pressure as orgasms are coupled with ejaculation and thus contribute to male reproductive success. By contrast, women's orgasms in intercourse are highly variable and are under little selective pressure as they are not a reproductive necessity. The proximal mechanisms producing variability in women's orgasms are little understood. In 1924 Marie Bonaparte proposed that a shorter distance between a woman's clitoris and her urethral meatus (CUMD) increased her likelihood of experiencing orgasm in intercourse. She based this on her published data that were never statistically analyzed. In 1940 Landis and colleagues published similar data suggesting the same relationship, but these data too were never fully analyzed. We analyzed raw data from these two studies and found that both demonstrate a strong inverse relationship between CUMD and orgasm during intercourse. Unresolved is whether this increased likelihood of orgasm with shorter CUMD reflects increased penile-clitoral contact during sexual intercourse or increased penile stimulation of internal aspects of the clitoris. CUMD likely reflects prenatal androgen exposure, with higher androgen levels producing larger distances. Thus these results suggest that women exposed to lower levels of prenatal androgens are more likely to experience orgasm during sexual intercourse.
The purpose of this study was to determine whether plasma oxytocin (OT) levels change during human sexual responses and, if so, to demonstrate the temporal pattern of change. Plasma OT levels were measured by RIA before, during, and after private self-stimulation to orgasm in normal men (n = 9) and women (n = 13). Blood samples were collected continuously through indwelling venous catheters. The subjects pressed a signal to indicate the start and finish of orgasm/ejaculation. Objective assessment of sexual arousal and orgasm was obtained by measuring blood-pulse amplitude and electromyographic activity, recorded continuously throughout testing from an anal device containing a photoplethysmograph and electromyograph electrodes connected to a polygraph located in an adjacent room. These measures allowed collection of data from men and women of changes in blood flow and muscle activity in the lower pelvic/pubic area. Plasma OT levels increased during sexual arousal in both women and men and were significantly higher during orgasm/ejaculation than during prior baseline testing. We suggest that the temporal pattern of secretion could be related to smooth muscle contractions of the reproductive system during orgasm.
The authors analyzed the incidence of sexual dysfunction (SD) with different selective serotonin reuptake inhibitors (SSRIs; fluoxetine, fluvoxamine, paroxetine, and sertraline) and hence the qualitative and quantitative changes in SD throughout time in a prospective and multicenter study. Outpatients (192 women and 152 men; age = 39.6 +/- 11.4 years) under treatment with SSRIs were interviewed with an SD questionnaire designed for this purpose by the authors and that included questions about the following: decreased libido, delayed orgasm or anorgasmia, delayed ejaculation, inability to ejaculate, impotence, and general sexual satisfaction. Patients with the following criteria were included: normal sexual function before SSRI intake, exclusive treatment with SSRIs or treatment associated with benzodiazepines, previous heterosexual or self-erotic current sexual practices. Excluded were patients with previous sexual dysfunction, association of SSRIs with neuroleptics, recent hormone intake, and significant medical illnesses. There was a significant increase in the incidence of SD when physicians asked the patients direct questions (58%) versus when SD was spontaneously reported (14%). There were some significant differences among different SSRIs: paroxetine provoked more delay of orgasm or ejaculation and more impotence than fluvoxamine, fluoxetine and sertraline (chi 2, p < .05). Only 24.5% of the patients had a good tolerance of their sexual dysfunction. Twelve male patients who suffered from premature ejaculation before the treatment preferred to maintain delayed ejaculation, and their sexual satisfaction, and that of their partners, clearly improved. Sexual dysfunction was positively correlated with dose. Patients experienced substantial improvement in sexual function when the dose was diminished or the drug was withdrawn. Men showed more incidence of sexual dysfunction than women, but women's sexual dysfunction was more intense than men's. In only 5.8% of patients, the dysfunction disappeared completely within 6 months, but 81.4% showed no improvement at all by the end of this period. Twelve of 15 patients experienced total improvement when the treatment was changed to moclobemide (450-600 mg/day), and 3 of 5 patients improved when treatment was changed to amineptine (200 mg/day).
Recent studies from our laboratory have investigated the hormonal response to various forms of sexual stimulation, including film, masturbation, and coitus in both men and women. This series of studies clearly demonstrated that plasma prolactin (PRL) concentrations are substantially increased for over 1h following orgasm (masturbation and coitus conditions) in both men and women, but unchanged following sexual arousal without orgasm. Here we discuss evidence suggesting that the PRL response to orgasm may play an important role in the control of acute sexual arousal following orgasm. Supporting this position, chronic elevations of PRL (hyperprolactinemia) produce pronounced reductions in animal sexual activity, and significant reduction of libido and gonadal function in both men and women. These data suggest that PRL may represent a peripheral regulatory factor for reproductive function, and/or a feedback mechanism that signals CNS centres controlling sexual arousal and behaviour. Thus, we propose a theoretical model of the role of PRL as a neuroendocrine reproductive reflex.
The initial observations, made in our laboratory with Knut Larsson, of the ability of vaginocervical stimulation (VCS) to block withdrawal responses to foot pinch in rats has led to findings of multiple behavioral, autonomic, and neuroendocrine effects of this potent stimulus in rats and also in women. It has led to an understanding of: (1) the neuroanatomical and neurochemical basis of a novel and potent pain-blocking mechanism; (2) likely neuroanatomical pathways mediating both the Ferguson reflex and a specific autonomic response - the pupil-dilating effect of VCS; (3) a role for oxytocin as a putative central nervous system neurotransmitter that stimulates autonomic sympathetic preganglionic neurons within the spinal cord; and (4) a novel pathway that can convey sensory activity from the cervix, adequate to induce orgasm, via the vagus nerves. This latter pathway bypasses the spinal cord and projects directly to the medulla oblongata, and thus can convey genital afferent activity despite complete spinal cord injury at any level.
Women diagnosed with complete spinal cord injury (SCI) at T10 or above report vaginal-cervical perceptual awareness. To test whether the Vagus nerves, which bypass the spinal cord, provide the afferent pathway for this response, we hypothesized that the Nucleus Tractus Solitarii (NTS) region of the medulla oblongata, to which the Vagus nerves project, is activated by vaginal-cervical self-stimulation (CSS) in such women, as visualized by functional magnetic resonance imaging (fMRI). Regional blood oxygen level-dependent (BOLD) signal intensity was imaged during CSS and other motor and sensory procedures, using statistical parametric mapping (SPM) analysis with head motion artifact correction. Physiatric examination and MRI established the location and extent of spinal cord injury. In order to demarcate the NTS, a gustatory stimulus and hand movement were used to activate the superior region of the NTS and the Nucleus Cuneatus adjacent to the inferior region of the NTS, respectively. Each of four women with interruption, or "complete" injury, of the spinal cord (ASIA criteria), and one woman with significant, but "incomplete" SCI, all at or above T10, showed activation of the inferior region of the NTS during CSS. Each woman showed analgesia, measured at the fingers, during CSS, confirming previous findings. Three women experienced orgasm during the CSS. The brain regions that showed activation during the orgasms included hypothalamic paraventricular nucleus, medial amygdala, anterior cingulate, frontal, parietal, and insular cortices, and cerebellum. We conclude that the Vagus nerves provide a spinal cord-bypass pathway for vaginal-cervical sensibility in women with complete spinal cord injury above the level of entry into spinal cord of the known genitospinal nerves.
We present a comprehensive account of clitoral anatomy, including its component structures, neurovascular supply, relationship to adjacent structures (the urethra, vagina and vestibular glands, and connective tissue supports), histology and immunohistochemistry. We related recent anatomical findings to the historical literature to determine when data on accurate anatomy became available.
An extensive review of the current and historical literature was done. The studies reviewed included dissection and microdissection, magnetic resonance imaging (MRI), 3-dimensional sectional anatomy reconstruction, histology and immunohistochemical studies.
The clitoris is a multiplanar structure with a broad attachment to the pubic arch and via extensive supporting tissue to the mons pubis and labia. Centrally it is attached to the urethra and vagina. Its components include the erectile bodies (paired bulbs and paired corpora, which are continuous with the crura) and the glans clitoris. The glans is a midline, densely neural, non-erectile structure that is the only external manifestation of the clitoris. All other components are composed of erectile tissue with the composition of the bulbar erectile tissue differing from that of the corpora. The clitoral and perineal neurovascular bundles are large, paired terminations of the pudendal neurovascular bundles. The clitoral neurovascular bundles ascend along the ischiopubic rami to meet each other and pass along the superior surface of the clitoral body supplying the clitoris. The neural trunks pass largely intact into the glans. These nerves are at least 2 mm in diameter even in infancy. The cavernous or autonomic neural anatomy is microscopic and difficult to define consistently. MRI complements dissection studies and clarifies the anatomy. Clitoral pharmacology and histology appears to parallel those of penile tissue, although the clinical impact is vastly different.
Typical textbook descriptions of the clitoris lack detail and include inaccuracies. It is impossible to convey clitoral anatomy in a single diagram showing only 1 plane, as is typically provided in textbooks, which reveal it as a flat structure. MRI provides a multiplanar representation of clitoral anatomy in the live state, which is a major advantage, and complements dissection materials. The work of Kobelt in the early 19th century provides a most comprehensive and accurate description of clitoral anatomy, and modern study provides objective images and few novel findings. The bulbs appear to be part of the clitoris. They are spongy in character and in continuity with the other parts of the clitoris. The distal urethra and vagina are intimately related structures, although they are not erectile in character. They form a tissue cluster with the clitoris. This cluster appears to be the locus of female sexual function and orgasm.
The evolutionary explanation of female orgasm has been difficult to come by. The orgasm in women does not obviously contribute to the reproductive success, and surprisingly unreliably accompanies heterosexual intercourse. Two types of explanations have been proposed: one insisting on extant adaptive roles in reproduction, another explaining female orgasm as a byproduct of selection on male orgasm, which is crucial for sperm transfer. We emphasize that these explanations tend to focus on evidence from human biology and thus address the modification of a trait rather than its evolutionary origin. To trace the trait through evolution requires identifying its homologue in other species, which may have limited similarity with the human trait. Human female orgasm is associated with an endocrine surge similar to the copulatory surges in species with induced ovulation. We suggest that the homolog of human orgasm is the reflex that, ancestrally, induced ovulation. This reflex became superfluous with the evolution of spontaneous ovulation, potentially freeing female orgasm for other roles. This is supported by phylogenetic evidence showing that induced ovulation is ancestral, while spontaneous ovulation is derived within eutherians. In addition, the comparative anatomy of female reproductive tract shows that evolution of spontaneous ovulation is correlated with increasing distance of clitoris from the copulatory canal. In summary, we suggest that the female orgasm-like trait may have been adaptive, however for a different role, namely for inducing ovulation. With the evolution of spontaneous ovulation, orgasm was freed to gain secondary roles, which may explain its maintenance, but not its origin.
The review critically examines two functional claims involving the human female orgasm. The first is that it is involved in the transport of spermatozoa by its release of oxytocin to create uterine contractions that suck up more semen at a faster rate. There is no physiological evidence for this scenario and the published experimental studies with oxytocin do not mimic the conditions of natural coitus. There is no evidence that orgasm has a role to play in reproductive fitness. The second claim is that as penile-vaginal intercourse (PVI) and its induced orgasm is the only sexual arousal that involves possible reproduction, evolution has rewarded it, and not clitorally-induced arousal (which it punishes), with highly specific health rewards. Apparently these cannot be generated by clitoral stimulation, moreover, its employment previous to and even at the same time as PVI negates these health advantages. The studies rely not only on women's retrospective self-reports of the genital structures creating their orgasms, which are known to have an “ambiguity problems”, but also on correlations from which causality is claimed or inferred. They have not been confirmed by independent researchers. Despite all women having the same genital structures from which sexual arousal is generated and that orgasm is not involved in reproduction via sperm transport, it is more than surprising that evolution has apparently created a situation where the majority of women do not or cannot generate orgasms from PVI alone and are thus imbued with poorer physical and mental health. This is even more unexpected when it is known that PVI actually stimulates the clitoris through thrusting traction on its attached ligaments via the anterior vaginal wall.
Previous data have indicated that orgasm produces marked alterations in plasma prolactin concentrations in men and women. Thus, the current study aimed to extend these data by examining prolactin response to coitus in healthy males and females. Ten pairs of healthy heterosexual couples participated in the study. Blood was drawn continuously for 20 min before, during, and until 60 min following sexual intercourse and orgasm. Plasma was subsequently analysed for prolactin concentrations. Coitus-induced orgasm produced a marked elevation of plasma prolactin in both males and females. Plasma prolactin concentrations remained elevated 1 h following orgasm. These data, together with previous evidence that masturbation-induced orgasm produces pronounced, long-lasting increases in plasma prolactin concentrations in both males and females, suggest a role for acute prolactin alterations in modifying human sexual desire following orgasm.
Spermatozoal uptake, facilitated by uterine contractions induced by oxytocin at orgasm during coitus, has been a long term concept. Studies attempting its support, however, have been poorly examined especially in the context of the changes in the female genital tract activated by sexual arousal.
To examine experimental support for the concept.
Using a variety of search engines, mainly peer reviewed articles and un-reviewed books were examined relating to sperm transport and function in the human female genital tract in the absence and presence of arousal to orgasm.
Identifying evidence-based data to support authority-based opinion.
All the experimental observations of sperm or model substitute's transport have been undertaken in women who were not sexually aroused. They fail to take into account that arousal creates vaginal tenting lifting the cervico-uterine complex into the false pelvis away from the ejaculated semen. This delays sperm uptake and transport making conclusions from these observations invalid in relation to transport during coitus. Studies injecting oxytocin have not used women in their sexually aroused state and used supraphysiological doses unlikely to be comparable with coitus and orgasm. The proposal that the transport of extra sperm by oxytocin-induced uterine contractions at orgasm is needed to facilitate fertility ignores possible harm from increased sperm numbers creating polyspermy and sperm enzyme release causing ovum degeneration, leading to decreased fertility. The role of sperm motility in their uptake from the vagina into the cervix as opposed to en bloc transfer through uterine archimyometrial-mediated transport in the absence of orgasm is at present unresolvable because of conflicting studies.
The bulk of the reported evidence favors the conclusion that the female orgasm, with its concomitant central release of oxytocin, has little or no effective role in the transport of spermatozoa in natural human coitus.
Investigación sobre la sexualidad femenina hecha en Estados Unidos con base en una encuesta aplicada a 3 mil mujeres. Aborda los siguientes temas: masturbación, orgasmo, coito, estimulación clitórica, lesbianismo, esclavitud sexual, la revolución sexual, mujeres mayores y hacia una nueva sexualidad femenina.
Serum prolactin has been measured in blood samples collected daily during 51 menstrual cycles using an homologous human radioimmunoassay for 17 cycles and an homologous ovine radioimmunoassay for 34 cycles. There was a progressive and significant increase in serum prolactin during the late follicular phase, with a maximal value concomitant to the LH peak. Serum prolactin levels were also significantly higher during the luteal phase than during early follicular phase. In some 70% of the individual cycles, the highest serum prolactin level was found at mid-cycle. Similar patterns were obtained with both radioimmunoassays. However, when using the same laboratory serum standard, the average serum prolactin level calculated for the entire cycle was 1.6 times higher with the homologous ovine assay than with the homologous human assay. The overall pattern of serum prolactin during the menstrual cycle resembles that reported for circulating 17beta-estradiol.
Immunocytochemical methods were used to localize the protein product of the immediate-early gene, c-fos, in male rats after exposure to, or direct physical interaction with, oestrous females. Increasing amounts of physical contact with a female, with resultant olfactory-vomeronasal and/or genital-somatosensory inputs, caused corresponding increments in c-fos expression in the medial preoptic area, the caudal part of the bed nucleus of the stria terminalis, the medial amygdala, and the midbrain central tegmental field. Males bearing unilateral electrothermal lesions of the olfactory peduncle showed a significant reduction in c-fos expression in the ipsilateral medial amygdala, but not in other structures, provided their coital interaction with oestrous females was restricted to mount-thrust and occasional intromissive patterns due to repeated application of lidocaine anaesthetic to the penis. No such lateralization of c-fos expression occurred in other males with unilateral olfactory lesions which were allowed to intromit and ejaculate with a female. These results suggest that olfactory inputs, possibly of vomeronasal origin, contribute to the activation of c-fos in the medial amygdala. However, lesion-induced deficits in this type of afferent input to the nervous system appear to be readily compensated for by the genital somatosensory input derived from repeated intromissions. Unilateral excitotoxic lesions of the medial preoptic area, made by infusing quinolinic acid, failed to reduce c-fos expression in the ipsilateral or contralateral medial amygdala or central tegmental field following ejaculation. By contrast, combined, unilateral excitotoxic lesions of the medial amygdala and the central tegmental field significantly reduced c-fos expression in the ipsilateral bed nucleus of the stria terminalis and medial preoptic area after mating; no such asymmetry in c-fos expression occurred when lesions were restricted to either the medial amygdala or central tegmental field. This suggests that afferent inputs from the central tegmental field (probably of genital-somatosensory origin) and from the medial amygdala (probably of olfactory-vomeronasal origin) interact to promote cellular activity, and the resultant induction of c-fos, in the ipsilateral bed nucleus of the stria terminalis and medial preoptic area. The monitoring of neuronal c-fos expression provides an effective means of studying the role of sensory factors in governing the activity of integrated neural structures which control the expression of a complex social behaviour.
Most studies investigating the behavioral effects of centrally administered oxytocin (OT) have been confined to single acute injections followed by brief behavioral observations lasting up to 90 min. The present study examines the behavioral effects of chronic, centrally administered OT in male rats observed continuously for prolonged periods of time. Either artificial cerebrospinal fluid or OT was centrally infused (via osmotic minipump) to gonadally intact male rats. Behavioral observations were made on males paired with either ovariectomized or estrous females during a 6-h time period. Most striking was the observation that durations of physical contact were doubled in pairs containing OT-infused males, even in the absence of sexual interactions. Also, OT-infused males showed significantly higher levels of anogenital sniffing of females and autogrooming; however, sexual interactions were unaffected by chronic OT. Chronic OT had no effect on body temperature, analgesia, or exploratory behavior in an open field. These findings suggest that chronic OT in male rats has behavioral effects that may significantly enhance adult social (nonsexual) interactions, possibly through alterations in olfactory and somatosensory information processing.
The neurohypophyseal hormone oxytocin has been implicated in many aspects of reproduction including sexual behavior. This review considers the hypotheses that oxytocin and/or the neural events surrounding the release of oxytocin may have behavioral effects during sexual arousal, orgasm, sexual satiety and other aspects of sociosexual interactions.
Studies of normal luteal phase women have shown that increases in serum LH and PRL are commonly synchronous. This study was designed to investigate the possible neuroendocrine mechanism(s) underlying this phenomenon. Six normal women were studied during the midluteal phase of 2 cycles. In the first cycle, they had blood samples collected at 15-min intervals for 6 h on 3 occasions during which time they received an infusion of normal saline or naloxone (1 mg/h) or a bolus of metoclopramide (10 mg, iv). In a second cycle, they received GnRH in increasing iv doses of 1, 10, and 50 micrograms at 2-h intervals. During the saline infusion, 11 of the 16 serum LH pulses (69%) were accompanied by an increase in serum PRL, and in 5 of the subjects, the first pulse of LH was synchronous with that of PRL (P = 0.0015). Naloxone increased the number of LH pulses from 16 to 20 and the number of PRL pulses from 12 to 16, all of which were synchronous with LH pulses. Administration of metoclopramide caused a substantial increase in PRL and a loss of further PRL pulsatility; however, LH pulsatility remained unaffected. Even after the smallest dose of GnRH (1 microgram), there was an increase in serum PRL [basal level, 11.8 +/- 2.1 (+/- SE) micrograms/liter; peak level, 16.5 +/- 3.3 micrograms/liter] as well as LH and FSH. The increase in serum PRL was, unlike the gonadotropin response, maximal after the 10-microgram dose of GnRH (peak level, 23.2 +/- 6 micrograms/liter) and did not increase further after the 50-micrograms dose (peak level, 18.5 +/- 2.4 micrograms/liter). These studies demonstrate that there is a PRL response to GnRH in the luteal phase and suggest that the observed synchrony in LH and PRL secretion at this time results from a physiological response of both the gonadotrope and the lactotrope to endogenous GnRH.
Thirty-eight spinal cord injured (SCI) males (median age = 26) completed an 80-item multiple choice questionnaire (median 37 months postinjury) which assessed sexual functioning pre- and post-spinal cord injury in four areas: (i) sexual activities and preferences, (ii) sexual abilities, (iii) sexual desire, arousal, and satisfaction, and, (iv) sexual adjustment. Frequency of sexual activity decreased following SCI with a reduction in intercourse and increased interest in alternative sexual activities. Of complete quadriplegic subjects 38% reported the ability to have an orgasm accompanied by ejaculation underscoring the need for physiological studies. Partner's desire for sex as perceived by the SCI individual was correlated with frequency of sex and numbers of sexual partners postinjury. Subject's perceptions of their own and partner's sexual desire decreased following SCI. Sexual satisfaction decreased postinjury and was positively correlated with both the patients' and their partners' interest in penile-vaginal intercourse. Of the subjects, 27% reported sexual adjustment difficulties and 74% relationship difficulties but only 22% received counseling. Results indicate the importance of the availability and desire of a sexual partner in the sexual activities and satisfaction of the SCI individual. SCI patient and staff sexual education and counseling continue to be strong needs.
The present study investigated the cardiovascular, genital, and endocrine changes in women after masturbation-induced orgasm because the neuroendocrine response to sexual arousal in humans is equivocal.
Healthy women (N = 10) completed an experimental session, in which a documentary film was observed for 20 minutes, followed by a pornographic film for 20 minutes, and another documentary for an additional 20 minutes. Subjects also participated in a control session, in which participants watched a documentary film for 60 minutes. After subjects had watched the pornographic film for 10 minutes in the experimental session, they were asked to masturbate until orgasm. Cardiovascular (heart rate and blood pressure) and genital (vaginal pulse amplitude) parameters were monitored continuously throughout testing. Furthermore, blood was drawn continuously for analysis of plasma concentrations of adrenaline, noradrenaline, cortisol, prolactin, luteinizing hormone (LH), beta-endorphin, follicle-stimulating hormone (FSH), testosterone, progesterone, and estradiol.
Orgasm induced elevations in cardiovascular parameters and levels of plasma adrenaline and noradrenaline. Plasma prolactin substantially increased after orgasm, remained elevated over the remainder of the session, and was still raised 60 minutes after sexual arousal. In addition, sexual arousal also produced small increases in plasma LH and testosterone concentrations. In contrast, plasma concentrations of cortisol, FSH, beta-endorphin, progesterone, and estradiol were unaffected by orgasm.
Sexual arousal and orgasm produce a distinct pattern of neuroendocrine alterations in women, primarily inducing a long-lasting elevation in plasma prolactin concentrations. These results concur with those observed in men, suggesting that prolactin is an endocrine marker of sexual arousal and orgasm.
The c-fos polyclonal anti-c-fos antibody was used to examine the effects of mating on Fos expression in brain neurons of 11 male macaques. Behavior tests were for 30 min, five males were unmated, four were mated, and two were social controls. Mated males were killed 60 min after ejaculation. Social controls were paired with females, but mating did not occur. Fos immunoreactive (Fos-ir) neuronal nuclei were counted in nine brain regions extending from the medial preoptic to the mammillary body area of all males. In contrast to previous reports on nonprimate laboratory species, overall there was as much Fos-ir in unmated as in mated males. Moreover, there was significantly less Fos expression in four brain regions (known to contain steroid receptors), namely, ventromedial hypothalamus, arcuate nucleus, lateral mammillary area, and bed nucleus of stria terminalis, of mated than of unmated males. There were no significant differences between mated and unmated males in the 5 other brain regions studied. These findings may reflect taxonomic differences between primates and nonprimates, or result from greater neural activation in feral animals maintained in a laboratory than in domesticated, inbred laboratory species. The simplest interpretation would be that neural activity in the male primate is turned off by mating in some brain sites but not in others.
This critical review presents a synthesis of the available theoretical and empirical literatures on human orgasm. Findings from both normal and clinical human populations are included. Two major trends in the literature, the dichotomization of biological and psychological perspectives and the assumption of gender differences, are highlighted. A new multidimensional model of the psychological experience of orgasm is described with a view to futhering a biopsychological approach applicable to both sexes. Clinical applications of this new model are discussed.
Vaginocervical stimulation (VS) significantly elevated the concentration of oxytocin (OT) in spinal cord superfusates of 8 intact urethane-anesthetized rats measured 10-15 min after VS (median [interquartile range]: 1.7 [1.00-3.37] pg/ml) compared to that measured 10-15 min before VS (1.1 [1.01-1.40] pg/ml). When VS was administered once (n = 8), it produced a 55% increase over baseline values; when administered a second time 45 min later (n = 6), it produced only a 22% increase over pre-VS values. The effects of estrogen on the VS-induced release of OT were then investigated using ovariectomized rats that were treated either with estradiol benzoate (EB; 10 microg/100 g bw) (n = 6) or with an oil vehicle (n = 6) subcutaneously for 3 days. The EB treatment significantly elevated the basal levels of OT released into spinal cord superfusates above vehicle control levels. Within 5-10 min after the onset of VS, OT concentrations in the superfusates were significantly higher in EB-treated than in vehicle-treated rats. The vehicle-treated rats did not show a significant elevation in OT concentration following VS. To rule out the possibility that the posterior pituitary gland was the source of this OT, the effect of hypophysectomy (HYPOX) was assessed on the VS-induced release of OT into spinal cord superfusates and plasma. The concentration of OT in spinal cord superfusates of both the HYPOX (n = 5) and intact rats (n = 6) increased significantly from 5.8 [4.4-6.5] pg/ml pre-VS to 7.9 [6.7-10.3] pg/ml immediately after VS, and from 4.4 [3.8-5] pg/ml pre-VS to 5.1 [4.6-5.7] pg/ml immediately after VS, respectively. There was no significant difference in baseline levels of OT in cerebrospinal fluid between the two groups. By contrast, plasma OT levels, while significantly elevated in response to VS from 3.42 [2.9-5.34] pg/ml baseline to 7.25 [5.33-15.77] pg/ml in the intact group, failed to respond significantly to VS in the HYPOX group (n = 5). The present findings provide evidence of a direct estrogen-dependent release of OT within the spinal cord in response to VS, presumably via descending oxytocinergic neurons.
Ovarian steroids and oxytocin (OT) have been implicated in the regulation of social behaviors. The purpose of the present study was to examine hormonal substrates of aggression and affiliation in the female Mongolian gerbil (Meriones unguiculatus), a highly social, monogamous rodent. Sexually naive adult females were paired with sexually experienced males for 48 h and their interactions videotaped. Females were gonadally intact and tested during vaginal estrus (INT) or ovariectomized and observed after the following treatments, administered by means of sc injections: EBEB (7 days of estradiol-benzoate); EBP (2 days of EB followed by progesterone), SALEB (saline, days 1-5 then 2 days of EB), OTEB (OT for days 1-5 then 2 days of EB); OTOIL (OT for days 1-5 then 2 days of OIL); or SALOIL (saline days 1-5 then 2 days of OIL). During the first hour of pairing INT females displayed higher levels of affiliation and lower levels of sniffing and agonistic behavior than SALOIL females. All hormonal treatments reduced agonistic behaviors when compared to SALOIL, although none of the hormonal treatments restored affiliation to INT levels. During the 48-h test overt aggression varied by treatment with INT, EBEB, EBP, and OTEB females displaying lower levels than SALOIL, while all groups displayed similar levels of affiliation. The results indicate that OT and E play a significant role in regulating male-directed aggressive behavior in females and that the presence of ovarian hormones as well as OT can increase affiliation during initial contact. Over a sustained period of cohabitation social cues appear to be more important in regulating affiliation than gonadal hormones.
In a prior [O]-H2O positron emission tomographic study we reported brain regions involved in human male ejaculation. Here, we used another, more recently acquired data set to evaluate the methodological approach of this previous study, and discovered that part of the reported activation pattern was not related to ejaculation. With a new analysis of these ejaculation data, we now demonstrate ejaculation-related activations in the deep cerebellar nuclei (dentate nucleus), anterior vermis, pons, and ventrolateral thalamus, and, most importantly, ejaculation-related deactivations throughout the prefrontal cortex. This revision offers a new and more accurate insight into the brain regions involved in human male ejaculation.
The way women experience orgasm is of interest to scientists, clinicians, and laypeople. Whereas the origin and the function of a woman's orgasm remains controversial, the current models of sexual function acknowledge a combined role of central (spinal and cerebral) and peripheral processes during orgasm experience. At the central level, although it is accepted that the spinal cord drives orgasm, the cerebral involvement and cognitive representation of a woman's orgasm has not been extensively investigated. Important gaps in our knowledge remain. Recently, the astonishing advances of neuroimaging techniques applied in parallel with a neuropsychological approach allowed the unravelling of specific functional neuroanatomy of a woman's orgasm. Here, clinical and experimental findings on the cortico-subcortical pathway of a woman's orgasm are reviewed and compared with the neural basis of a man's orgasm. By defining the specific brain areas that sustain the assumed higher-order representation of a woman's orgasm, this review provides a foundation for future studies. The next challenge of functional imaging and neuropsychological studies is to understand the hierarchical interactions between these multiple cortical areas, not only with a correlation analysis but also with high spatio-temporal resolution techniques demonstrating the causal necessity, the temporal time course and the direction of the causality. Further studies using a multi-disciplinary approach are needed to identify the spatio-temporal dynamic of a woman's orgasm, its dysfunctions and possible new treatments.
The pleasure bond: A new look at sexuality and commitment
- W H Masters
- V E Johnson
- R J Levin
- WH Masters
Textbook of sexual medicine
- R C Kolodny
- W H Masters
- V E Johnson
- RC Kolodny
The lagoon. How Aristotle invented science
- A M Leroi
- AM Leroi