Intrinsic links among sex, emotion, and reproduction

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    Forgot your login information? By: Thomas S. Edited by: Thomas S. Subject: Sexualities. Weinberg, T. Sex, emotion, and identity. In Selves, symbols, and sexualities pp. Weinberg, Thomas S. SAGE Knowledge. Have you created a personal profile? Login or create a profile so that you can create alerts and save clips, playlists, and searches.

    Please log in from an authenticated institution or log into your member profile to sexemotion the email feature. Ssxemotion he sociology of sexemotoin, though not sexemotion domain of symbolic interactionism, is an important and rapidly expanding direction for interactionist research. Sexemotion particular, symbolic interactionists are sexemotion with how emotional experience and sexemotion shape social life and, in this case, sexualities. This section deals also with sexual identity, but it focuses primarily on the role of emotions and emotional experience in identity construction sexeemotion negotiation.

    Chapter 13 lays the groundwork for thinking about the centrality of emotion in sexual identities broadly defined. Forstie's analysis uncovers the extent to which people CQ Press Your definitive resource for politics, policy and people. Remember me? Back Institutional Login Please choose from an option shown sexemotipn.

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    We examine common functions and discuss future directions for elucidating the psycho-neuroendocrine links uniting sex, emotion, and. See Tweets about #sexemotion on Twitter. See what people are saying and join the conversation. See Tweets about #sexemotions on Twitter. See what people are saying and join the conversation.

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    Cellular and Molecular Life Sciences. Species survival is dependent on successful reproduction. This begins with a desire to mate, followed by selection of a partner, copulation and in monogamous mammals including humans, requires emotions and behaviours necessary to maintain partner bonds for the benefit of rearing young.

    Hormones are integral to all of these stages and not only mediate physiological and endocrine processes involved in reproduction, but also act as neuromodulators within limbic brain centres to facilitate the expression of innate emotions and behaviours required for reproduction. A significant body of work is unravelling the roles of several key hormones in the modulation of mood states and sexual behaviours; however, a full understanding of the integration of these intrinsic links among sexual and emotional brain circuits still eludes us.

    This review summarises the evidence to date and postulates future directions to identify potential psycho-neuroendocrine frameworks linking sexual and emotional brain processes with reproduction.

    Reproduction is an essential physiological process for preservation of the species, which relies on the integration of social and sensory cues with emotional and behavioural outputs that favour the successful production of offspring. Effective mating depends on an appropriate display of sexual behaviours that select for reproductive fitness. The neural command centre for coordinating these complex processes sits within the limbic brain [ 1 ].

    The limbic system was first described by Thomas Willis in and is one of the most studied functional networks in the brain [ 1 ]. Evolutionarily preserved since amphibia and common among all mammals, it consists of several functionally interconnected cortical and subcortical structures that have a shared role in processing sensory stimuli into emotional and behavioural outputs [ 2 ].

    This includes the orbitofrontal cortex, hippocampus, cingulate cortex, amygdala, hypothalamus, thalamus, and the ventral striatum i. Within the limbic network, several structures are thought to be predominantly involved in processing emotions; these include the amygdala, hypothalamus, cingulate cortex, and pre-frontal cortex [ 34 ].

    The amygdala is a small almond-shaped structure lying deep to the antero-inferior temporal lobe. Animal lesion studies have established the amygdala as a central hub for processing emotions and sexual sexemotion in rodents and primates [ 356 ]. In humans, there is a positive association between sexual drive and amygdala volume [ 4 ]. Functional magnetic resonance imaging fMRI demonstrates enhanced amygdala activity in response to viewing sexually arousing images in men and women [ 7 ].

    The medial nucleus of the amygdala is particularly implicated in processing multimodal sensory inputs such as olfactory signals and integrating these inputs into behavioural and endocrine pathways [ 89 ]. The hypothalamus coordinates essential homeostatic mechanisms including the regulation of neuroendocrine axes and autonomic processes.

    Limbic functions of the hypothalamus have been demonstrated in rats, whereby hypothalamic stimulation leads to increased motivation towards rewarding behaviours [ 10 ]. The amygdala and hypothalamus are highly interconnected and electrical stimulation of the amygdala in rats and cats leads to increased gonadotropin secretion [ 811 ].

    This demonstrates that the amygdala—hypothalamus connection serves as a conduit between emotion-processing brain centres and reproductive neuroendocrine pathways. The hypothalamic—pituitary—gonadal HPG axis orchestrates physiological processes required for reproduction. Furthermore, it is known that hypogonadal states in animals and humans are associated with loss of responsiveness to sexual stimuli and loss of appetitive sexual behaviours [ 121314 ].

    Reproductive hormones and their receptors are found throughout the limbic brain network where evidence suggests that they act as neurotransmitters and neuromodulators to influence emotional states and sexual behaviour [ 15 ].

    Thus, a considerable interest has developed with regard to factors that may integrate reproductive endocrinology with sexual behaviours. Recently, the discovery of novel neuropeptides acting above the level of gonadotropin-releasing hormone GnRH to modulate its release have redefined our understanding of sexemotion endocrinology and also provide new candidates for the study of intrinsic links between reproduction, emotion, and sexual behaviours.

    This review summarises the literature surrounding key hormones and their roles in sexual and emotional brain processing. We examine common functions and discuss future directions for elucidating the psycho-neuroendocrine links uniting sex, emotion, and reproduction. Summary of the effects of key hormones in the control of sex, emotion, and reproduction.

    Relevant references in parentheses. Kisspeptin-expressing neurones and mRNA have been identified in the bed nucleus of the stria terminalis BNSTthalamus, and amygdala of rodents [ 22 ]. Furthermore, amygdala kisspeptin expression is positively modulated by gonadal sex steroids [ 23 ].

    Kisspeptin receptor expression has also been demonstrated in rodent limbic brain structures, including the amygdala, thalamus, hippocampus, and olfactory systems [ 242526 ]. Of note, direct kisspeptin administration into the medial amygdala stimulates LH secretion, while direct kisspeptin antagonist administration decreases LH secretion and pulsatility, providing evidence for a link between amygdala kisspeptin signalling and the HPG axis [ 27 ].

    In humans, KISS1 and KISS1R mRNA have been identified in several limbic and paralimbic structures including the amygdala, caudate, cingulate, globus pallidus, hippocampus, medial and superior frontal gyrus, nucleus accumbens, parahippocampal gyrus, substantia nigra, putamen, and thalamus [ 2829 ]. Thus, there exists an anatomical framework in place through which kisspeptin signalling can link reproduction, sex, and emotion.

    Later, Kauffman and colleagues demonstrate that testosterone-replaced Kiss1r knock-out male mice lack olfactory partner preference despite normosmia [ 31 ].

    In keeping with this, activation of medial amygdala kisspeptin neurones increases time spent by male mice investigating females [ 32 ]. Furthermore, studies demonstrate that opposite sex odours activate kisspeptin neurones in rodents [ 3334 ]. This effect is also seen in sheep, whereby kisspeptin cFos activity is increased after exposure of anoestrous ewes to a novel male with an sexemotion rise in LH pulsatility [ 35 ].

    In healthy young men, kisspeptin enhances limbic and paralimbic brain activity on fMRI, specifically in response to sexually arousing and non-sexual couple-bonding images [ 36 ].

    In addition, kisspeptin enhancement of brain activity in response to sexual images in several of these structures correlates with decreased sexual aversion [ 36 ]. In keeping with this human data, direct kisspeptin injection into the medial amygdala dose-dependently triggers multiple erections in male rats, but is blocked by pre-treatment with a kisspeptin antagonist [ 37 ]. Data also suggest a role for kisspeptin in the display of fear and anxiety.

    In zebrafish, fear stimuli significantly reduce Kiss1 transcription, while intra-cranial kisspeptin administration attenuates fear responses via effects on the serotonergic system [ 3839 ]. In terms of function, kisspeptin administration may increase anxiety [ 40 ] or decrease anxiety [ 32 ] depending on the methodology used.

    Kisspeptin administration has also been shown to have anti-depressant-like effects in rodents [ 41 ] and humans [ 36 ] with the latter effect associated with enhanced activity in the pre-frontal area, a region that expresses kisspeptin receptor mRNA in humans and known to be involved in negative emotions [ 28 ]. Mechanistically, there is observed interplay between kisspeptin signalling and several other neuropeptide systems including serotoninergic [ 39 ], adrenergic [ 41 ], vasopressinergic [ 26 ], dopaminergic [ 26 ], nitric oxide [ 42 ], gamma-aminobutyric acid GABAglutamate [ 43 ], and CART systems [ 44 ].

    In addition, data suggest that, when peripherally administered, certain isoforms of kisspeptin can cross the blood—brain barrier to reach its central receptors located throughout the brain including in the limbic system as above [ 3645 ]. Collectively, these studies in humans and several other species demonstrate that kisspeptin signalling effectively integrates sensory processing with limbic pathways involved in sexual arousal, positive mood, and anxiety for a current review of kisspeptin in these areas, please see [ 46 sexemotion.

    Together with its established role as a modulator of reproductive hormone secretion, these data suggest that kisspeptin plays a key part in the integration of reproduction, sex, and emotion Fig. GnRH is secreted in a pulsatile fashion from the pre-optic area of the hypothalamus.

    GnRH binds to its receptor on the anterior pituitary to stimulate LH and follicle-stimulating hormone FSH release, which induce sex steroid secretion and gametogenesis in the gonads. Once thought to be the master regulator of the HPG axis, it is now known that GnRH release is controlled by two novel neuropeptides; kisspeptin which exerts a stimulatory effect on GnRH [ 47 ] and gonadotropin-inhibitory hormone GnIH which exerts a negative effect on GnRH signalling [ 48 ] Fig.

    Anatomical localisation of GnRH-expressing neurones has been studied in many species [ 495051 ] with results in rats showing extensive extra-hypothalamic distribution, particularly in the olfactory bulb and tubercle, BNST, hippocampus, and medial amygdala [ 51 ]. The GnRH receptor has been immunohistochemically located in several extra-hypothalamic brain areas including the amygdala, hippocampus, piriform olfactory cortex, arcuate nucleus, and olfactory bulb in mice [ 5253 ].

    This pattern of distribution for GnRH and its receptor provides an anatomical framework for GnRH to relay olfactory signals between emotion and reproductive centres in the brain. Male hamsters are particularly reliant on olfactory cues to induce sexual behaviour, which they sense via the vomeronasal organ VNO.

    Furthermore, this return of sexual behaviour is accompanied by increased cFos activity in the anteroventral medial amygdala [ 54 ]. In humans, two studies utilising similar techniques of computer-assisted microscopy to map the distribution of GnRH-expressing neurones [ 55 ] and GnRH gene transcripts [ 56 ] have demonstrated the presence of GnRH neurones in the anterior olfactory areas, cortical and medial amygdala, and the BNST.

    However, the functional GnRH receptor appears to be restricted to the sexemotion and hippocampus in humans [ 57 ] with limited descriptions of its presence in specific limbic brain regions [ 58 ]. Intwo independent research groups showed that subcutaneous administration of GnRH to ovariectomised, steroid-replaced female rats potentiates female sexual behaviour, i.

    Furthermore, bilateral lesions sexemotion the medial amygdala lead to reduced cFos expression in GnRH neurones with associated inhibition of lordosis behaviour in female rats [ 9 ], demonstrating that GnRH is a neuroendocrine mediator of limbic brain processing in rodents. Since then, GnRH activity has been shown to facilitate a range of sexual behaviours in other species, ranging from mating preferences in fish [ 6162 ] to proceptive sexual advancement behaviours in marmoset monkeys [ 63 ].

    GnRH agonists are associated with anxiolytic effects in rodent models [ 64 ], whereas GnRH antagonists induce anxiogenic behaviours [ 65 ]. GnRH also interacts with other neuropeptides and neurotransmitters such as vasopressin [ 66 ], GABA [ 67 ], and dopamine [ 68 ], which may play interconnecting roles in its regulation of mood and behaviours. GnRH and its receptor are distributed in key brain areas involved in processing olfactory cues and integrating chemosensory information with appropriate sexual behaviours in animals Fig.

    GnRH is also involved in ameliorating anxiety which complements its role in facilitating successful reproduction. GnRH is known to regulate and be regulated by several other neurotransmitters; however, the interplay between these factors in the control of emotions and sexual behaviours is complex and not yet fully understood.

    First identified in the Japanese quail, GnIH is now known to be a key regulator of reproduction in avian species and has also been found in other vertebrates [ 48 ]. Data in humans show that Sexemotion is able to inhibit LH secretion in post-menopausal women but does not affect kisspeptin-stimulated LH secretion in normal men [ 71 ], suggesting that GnIH is unable sexemotion overcome the positive stimulus of kisspeptin on the human HPG axis.

    GnIH immunoreactive fibres have been identified in brain regions outside the hypothalamus in quail [ 72 ], rats [ 73 ], and rhesus macaques [ 74 ], but this has not yet been examined in humans. Differences in distribution are seen between rodents and primates, with GnIH immunoreactivity present in the amygdala of rats but sexemotion primates [ 7374 ]. However, Sexemotion immunoreactive fibres do extend to dopaminergic neurones in the ventral tegmental area in primate brains [ 74 ] which eludes to a potential role in reward and motivation.

    As a relatively novel neuropeptide, with fewer published studies than its counterparts, detailed neuroanatomical studies of the GnIH receptor GPR, are sparse with work mostly concentrated in avian species.

    Ukena et al. Alongside its inhibitory role on the HPG axis, there is also evidence that GnIH has a negative impact on male sexual behaviours in quail [ 75 ] and rats [ 76 ] and reduced sexual motivation in female hamsters [ 77 ]. However, there may be species differences as another study showed no effect of GnIH on sexual behaviour in cynomolgus monkeys, mice, and ewes [ 78 ]. Differences in methodologies and the reproductive status of the animals may have also contributed to these findings.

    Through its downregulation of the HPG axis and inhibition of sexual behaviours, GnIH may act as a reproductive brake by appropriately inhibiting mating during unfavourable times such as stressful situations Fig. This is of particular importance in seasonal breeders such as birds; however, it is uncertain whether GnIH significantly affects sexual behaviour consistently in other species. Exposure to gonadal steroids such as oestrogen and testosterone during critical developmental windows not only determines sexually dimorphic structural differences, but also influences sex-specific behaviours in many species including humans [ 79 ].

    Oestrogen and testosterone are essential for physiological neuroendocrine processes and behaviours necessary for sexual maturation and successful copulation.

    Oestrogen receptors ERs and testosterone-binding androgen receptors ARs are intra-cellular receptors found throughout the brains of rodents [ 8081 ]. Using in situ hybridisation in male and female rats, Sar et al. AR mRNA is also found in brainstem structures involved with sensory function and regions containing somatic motor neurones. In the cortex and brainstem, there are far fewer ER labelled cells compared to AR labelled cells in both males and females [ 81 ].

    Both AR and ER mRNA are found in the olfactory bulb and the olfactory tubercle of male and female rats with AR more densely distributed than ER, particularly in the mitral cell layer [ 80 ], which is in keeping with evidence that gonadal steroids have important roles sexemotion olfactory signalling for the regulation of social behaviours in rodents as well as other species [ 83 ].

    Furthermore, these behaviours were restored by transplantation of functioning testes from another animal [ 84 ]. Subsequent studies using anti-androgens have supported and developed these findings. In testosterone-replaced gonadectomised male mice, blocking AR with systemic administration of the anti-androgen, hydroxyflutamide prevents the restoration of male sexual behaviour partner preference, scent marking, and ultrasonic sexemotion whereas blockade of ER does not fully inhibit these behaviours [ 85 ].

    Intracranial implants of hydroxyflutamide into the medial pre-optic area and ventromedial hypothalamus successfully prevent restoration of sexual behaviour in gonadectomised testosterone-replaced male mice, whereas hydroxyflutamide in the medial amygdala only has partial effects, and no effects when implanted into the septum, indicating that male sexual behaviour is differentially mediated by specific AR populations [ 86 ]. In humans, testosterone triggers greater affiliative behaviours in men, e.

    There is also ample evidence that males with stronger testosterone-controlled signals, such as hair colour [ 88 ], facial masculinity [ 8990 ], and vocalisations [ 9192 ], are more attractive to females. The emotional mechanisms underlying testosterone-mediated behaviour may be due to its interactions with other neuropeptides, with evidence showing that testosterone upregulates expression of vasopressin and its receptor in brain areas involved in aggressive behaviours including the medial amygdala, lateral hypothalamus, and the medial pre-optic area [ 84 ].

    Furthermore, testosterone levels are positively correlated with fMRI brain activation in the amygdala of men when viewing fearful or angry faces [ 93 ]. In ageing men, progressive testosterone decline is associated with low mood, reduced libido, and poor sexual function [ 14 ]. Testosterone insufficiency has also been implicated in sexual dysfunction occurring in post-menopausal women [ 94 ].

    The amygdala is sexeotion small almond-shaped structure lying deep to sexemotion antero-inferior temporal lobe. Interestingly, a reciprocal relationship is also present, whereby sexual experience can reduce sexemotion glucocorticoid release [ sexemotion. Neuroendocrinology — sex dating

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