Neuroanatomy and Neurochemistry of Sexual Desire, Pleasure, Love and Orgasm
H. Ümit Sayin and Carlos H. Schenck
Pleasure is a mental and emotional state that humans and other animals experience as positive, enjoyable, satisfying, giving joy and happiness or worth seeking. It may include other mind states such as happiness, entertainment, enjoyment, ecstasy, and euphoria, with peak experiences. According to the “incentive salience model” pleasure has three components: “wanting and motivation”, “learning” and “liking-pleasure”. Different brain regions cooperatively work together for each of these constituents. Since most of the data to establish this model came from animal studies, for human beings a more profound and complex model needs to be established. During the onset of pleasure or sensuality, dopaminergic neurons in ventral tegmental area (VTA) fire and their projections to the nucleus accumbens (NA) release dopamine which participates in the complex psychological processes of learning and liking. Similar episodes occur during the onset of sexual pleasure and human orgasm. Some complex human phenomena, such as passionate love or expanded orgasms, can be regarded as peak experiences, which the animal kingdom does not seem to experience. During passionate love, for instance, the release of some neurotransmitters, such as dopamine, oxytocin, and norepinephrine, is enhanced, while serotonin levels drop; in some fMRI studies, it is shown that some brain regions and “hedonic hot spots” are activated. During the female orgasm, VTA, NA, anterior cingulate, insula, amygdala, hippocampus, and hypothalamus are activated. In this article neuroanatomical, neurophysiological, neurochemical components of pleasure, love, motivation and orgasm, which lead to impressive behavioral changes, are discussed. Also, the phenomenon of experiencing sexual pleasure during the sleep found with the parasomnia “Sexsomnia” and also “Epileptic Sexsomnia”, both for the affected person and the bed partner will be discussed, with further research encouraged on this intriguing new clinical and clinical research area.
KEY WORDS: pleasure, hedonic hot spot, reward, orgasm, sexual pleasure, pleasure center, cingulate, insula, amygdala, hippocampus, hypothalamus, n. accumbens, ventral tegmental area, addiction, dopamine, oxytocin, liking, motivation, sexsomnia, epileptic sexsomnia
SexuS Journal ● Winter 2019 ● 4 (11): 907-946
Acevedo BP, Aron A, Fisher HE & Brown LL. Neural 45 The neuroimaging of love and desire Clinical Neuropsychiatry Correlates of long-term intense romantic love. Social Cognitive and Affective Neuroscience 2012; published online January 5 2011 doi:10.1093/scan/nsq092, 2012.
Acevedo-Rodriguez A, Mani SK, Handa RJ. Oxytocin and Estrogen Receptor β in the Brain: An Overview. Frontiers in Endocrinology 2015; 6:160.
Adams S, Heckard D, Hassell J, Uphouse L. Factors influencing fluoxetine-induced sexual dysfunction in female rats. Behav Brain Res 2012; 235: 73-81.
Ahlenius S, Larsson K. Physiological and pharmacologi cal implications of speciﬁc effects by 5HT1A agonists on rat sexual behavior. In 5HT1A Agonists, 5HT3 Antagonists and Benzodiazepines: Their Comparative Behavioral Pharmacol- ogy, ed. R. J. Rodgers & S. J. Cooper. New York: John Wiley, 1991.
Aizenberg D, Zemishlany Z, Dorfman-Etrog P, Weizman A. Sexual dysfunction in male schizophrenic patients. J Clin Psychiatry. 1995; 56:137- 143.
American Academy of Sleep Medicine. International classification of sleep disorders, 3rd ed. Darien, Illinois, 2014.
Anderson-Hunt M, Dennerstein L. Increased female sexual response after oxytocin [editorial].BMJ. 1994; 309:929.
Anderson-Hunt M, Dennerstein L. Oxytocin and female sexuality. Gynecol Obstet Invest. 1995; 40:217-221.
Argiolas A. Neuropeptide and sexual behaviour. Neurosci & Biobehav Rev 1999; 23: 1127–1142.
Arias M. Neurology of ecstatic religious and similr experiences: ecstatic, orgasmic and musicogenic seizures. Stendhal syndrome and autoscopic phenomena. Neurologia 2019; 34 (1): 55-61.
Armagan N, Sayin HÜ, Kocatürk A. Can Sexual Response be Enhanced and Expanded in the Human Female: Preliminary Findings and a Proposed Psychometric Scale for Expanded Sexual Response (ESR). 38th Annual Meeting of IASR (International Academy of Sex Research), Lisbon-Portual, July 8-11, 2012; Abtract Book, 38: 9, 2012.
Aron A, Fisher H, Mashek DJ, Strong G, Li H & Brown LL. Reward motivation and emotion systems associated with early-stage intense romantic love. Journal of Neurophysiology 2005; 94: 327-337.
Atzil S, Hendler T, Zagoory-Sharon O, Winetraub Y, Feldman R. Synchrony and specificity in the maternal and the paternal brain: relations to oxytocin and vasopressin. J Am Acad Child Adolesc Psychiatry 2012; 51(8):798-811. doi: 10.1016/j.jaac.2012.06.008.
Bancroft J. Human sexuality and its problems. New York: Churchill Livingstone, 1989.
Bancroft J. The endocrinology of sexual arousal. Journal of Endocrinology 2005; 186:411–427.
Bartels A, Zeki S. The neural basis of romantic love. Neuroreport 2000; 27;11:3829-34.
Bartels A, Zeki S. The neural correlates of maternal and romantic love. Neuroimage 2004; 21(3):1155-66.
Bernard J. Crespi. Oxytocin, testosterone, and human social cognition Biol. Rev. 2015; 91: 390-408.
Berridge KC, Valenstein ES What psychological process mediates feeding evoked by electrical stimulation of the lateral hypothalamus? Behav Neurosci 1991; 105:3–14.
Berridge KC, Robinson TE. What is the role of dopamine in reward: hedonic impact, reward learning, or incentive salience? Brain Res Rev 1998; 28:309–369.
Berridge KC, Robinson TE. Parsing Reward. Trends in Neurosciences 2003-a; 26(9):507–13.
Berridge KC. Pleasures of the brain. Brain Cogn 2003b; 52:106–28.
Berridge KC, Kringelbach ML. Affective Neuroscience of Pleasure: Reward in Humans and Animals. Psychopharmacology 2008. 199:457–80.
Berridge KC. The Debate over Dopamine’s Role in Reward: The Case for Incentive Salience. Psychopharmacology 2007-a; 191(3):391–431.
Berridge KC & Morten L. Kringelbach, Affective neuroscience of pleasure: reward in humans and animals, Psychopharmacology 2008-a;199:457–480.
Berridge KC, Aldridge JW Decision utility, incentive salience, and cue-triggered ‘wanting’. In: Bargh J, Morsella E (eds) The psychology of action. Oxford University Press, New York, 2008-b.
Berridge KC, Robinson TE, Aldridge JW. Dissecting components of reward: ‘liking’, ‘wanting’, and learning, Curr Opin Pharmacol 2009; 9(1): 65–73.
Berridge KC, Kringelbach ML "Pleasure systems in the brain". Neuron 2015. 86 (3): 646–664.
Bethlehem RAI, van Honk J, Auyeung B, Baron-Cohen S. Oxytocin, brain physiology, and functional connectivity: A review of intranasal oxytocin fMRI studies. Psychoneuroendocrinology 2013; 38: 962-974.
Blaicher W, Gruber D, Bieglmayer C, Blaicher AM, Knogler W, Huber JC. The role of oxytocin in relation to female sexual arousal. Gynecol Obstet Invest 1999; 47: 125–126.
Blumer D. Hypersexual episodes in temporal lobe epilepsy. Am J Psychiatry 1970, 126 (8): 1099-1106
Bowers MB, Woert MV, Davis L. Sexual behavior during L-dopa treatment for parkinsonism. Am J Psychiatry 1971;127:1691-1693.
Bownds M D , Biology of Mind, Maryland: Fitzgerald Science Press, pp: 46-47,1999.
Brody S, T.H. Krüger. The post-orgasmic prolactin increase following intercourse is greater than following masturbation and suggests greater satiety, Biol Psychiatry 2006; 71(3):312-315.
Brody S, Wiess P. Vaginal orgasm is Associated with Vaginal (not clitoral) sex education, focusing on mental attention on vaginal sensations, intercourse duration and a preference for a longer penis. J. Sex Med 2010; 7:2774-2781.
Brunton L, Chabner B, Bjorn Knollman B. Goodman and Gilman's The Pharmacological Basis of Therapeutics, Twelfth Edition, New York: Mc Graw Hill, 2011.
Burkett JP, Young LJ. The behavioral, anatomical and pharmacological parallels between social attachment, love and addiction. Psychopharmacology (Berl) 2012; 224(1):1-26.
Cantor JM, Binik YM & Pfaus JG. 1999 Chronic fluoxetine inhibits sexual behavior in the male rat: reversal with oxytocin. Psychopharmacology 1999; 144: 355–362.
Carellas B. Urban Tantra. New York: Celestial Arts, 2007.
Carmichael MS, Humbert R, Dixen J, Palmisano G, Greenleaf W Davidson JM. Plasma oxytocin increases in the human sexual response. J Clin Endoc Met 1987; 64: 27–31.
Carmichael MS, Warburton VL, Dixen J, DavidsonJM. Relationships among cardiovascular, muscular, and oxytocin responses during human sexual activity. Arch Sex Behav. 1994; 23:59-77.
Carter CS. Oxytocin and sexual behavior. Neurosci Biobehav Rev 1992; 16: 131-144.
Casey KL, Minoshima S, Berger KL, et al. Positron emission tomographic analysis of cerebral structures activated specifically by repetitive noxious heat stimuli. J Neurophysiol. 1994;71:802–807.
Casey KL, Morrow TJ, Lorenz J, Minoshima S. Temporal and spatial dynamics of human forebrain activity during heat pain: analysis by positron emission tomography. J Neurophysiol. 2001; 85:951–959.
Castro DC, Berridge KC. "Opioid and orexin hedonic hotspots in rat orbitofrontal cortex and insula". Proceedings of the National Academy of Sciences of the United States of America 2017; 114 (43): E9125–E9134.
Chua Chee Ann. A proposal for a radical new sex therapy technique for the management of vasocongestion and orgasmic dysfunction in women: the AFE zone stimulation technique. Sex Marital Ther 1997;12: 357–70.
Chuang YC, Lin TK, Lui CC, Chen SD, Chang CS. Tooth-brushing epilepsy with ictal orgasms. Seizure 2014; 13: 179-182.
Courty E, Durif F, Zenut M, Courty P, Lavarenne J. Psychiatric and sexual disorders induced by apomorphine in parkinson’s disease. Clin Neuropharmacol. 1997; 20:140-147.
Demerdash A, Shaalan M, Midani A, Kamel F, Bahri M. Sexual behavior of sample of females with epilepsy. Epilepsia 1991; 32 (1): 82-85
Danjou P, Alexandre L, Warot D, Lacomblez L, Puech AJ. Assessment of erectogenic proper ties apomorphine and yohimbine in man. Br J Clin Pharmacol 1988; 26:733-739.
de Jong TR, Veening JG, Olivier B, Waldinger MD. Oxytocin involvement in SSRI-induced delayed ejaculation: a review of animal studies. J Sex Med 2007; 4: 14-28.
Delgado PL, Brannan SK, Mallinckrodt CH, Tran PV, McNamara RK, Wang F, et al. Sexual functioning assessed in 4 double-blind placebo- and paroxetinecontrolled trials of duloxetine for major depressive disorder. J Clin Psychiatry 2005; 66: 686-692.
Diamond LM & Dickenson JA. The Neuroimaging of Love and Desire: Review and Future Directions. Clinical Psychiatry, 9 (1): 39-46, 2012.
Emanuele E, Politi P, Bianchi M, Minoretti P, Bertona M, Geroldi D. Raised plasma nerve growth factor levels associated with early-stage romantic love. Psychoneuroendocrinology 2006; 31: 288-294.
Escapa R, Bizzare Sex, London: Grafton Books, 1989.
Esch T and Stefano G.B. The neurobiology of love. Neuro Endocrinol Lett 2005; 26: 175-192.
Exton MS, Bindert A, Kruger T, Scheller F, Hartmann U, Schedlowski M. Cardiovascular and endocrine alterations after masturbation-induced orgasm in women. Psychosom Med. 1999; 61: 280-289.
Febo M, Shields J, Ferris C, King J. Oxytocin modulates unconditioned fear response in lactating dams: An fMRI study. Brain Res. 2009; 1302:183–193.
Feiger A, Kiev A, Shrivastava RK, Wisselink PG, Wilcox CS. Nefazodone versus sertraline in outpatients with major depression: focus on effi cacy,tolerability, and effects on sexual function and satisfaction. J Clin Psychiatry. 1996; 57 (suppl 2): 53-62.
Fischetti Mark. Passionate Love in the Brain as Revealed by MRI Scans, Scientific American; Feb. 14, 2011, pp: 18-21.
Fisher HE, Aron A, Brown LL. Romantic love: a mammalian brain system for mate choice. Philos Trans R Soc Lond B Biol Sci 2006; 361: 2173-2186.
Fisher HE, Brown LL, Aron A, Strong G, Mashek D. Reward, addiction, and emotion regulation systems associated with rejection in love. J Neurophysiol 2010; 104: 51-60.
Fisher S. Understanding the Female Orgasm, Lon-don: Penguin Books, 1973-1977.
Flynn MA, Weaver-Osterholtz D, Sharpe-Timms KL, Allen S, Krause G. Dehydroepiandrosterone replacement in aging humans. J Clin Endocrinol Metab 1999; 84:1527-1533.
Fonteille V & Stoleru S. The cerebral correlates of sexual desire: Functional neuroimaging approach. Sexologies 10.1016/j.sexol.2010.03.011. 2010.
Freud S. Beyond the pleasure principle. New York: Bantam, 1950.
Futuyma D. Evolutionary Biology. NY: Sinaur Associates, 1997.
FXu X, Aron A, Brown L, Cao G, Feng T, Weng X. Reward and motivation systems: a brain mapping study of early-stage intense romantic love in Chinese participants. Hum Brain Mapp 2011;32(2): 249-57.
Georgiadis RJ, R Kortkaas, R Kuipers, A Neiuwenburg, J Pruim, AAT Simone Reinders, G Holstege. Regional cerebral blood flow changes associated with clitorally induced orgasm in healthy women. Eur J Neurosci , 2006 ; 24:3305-16.
Georgiadis RJ. Men versus women on Sexual Brain Function: Prominent differences during tactile genital stimulation, but not orgasm. Hum Brain Map , 2009; 30(10):3089-101.
Gillman M & Lichtigfield F. The effects of nitrous oxide and naloxone on orgasm in human females: a preliminary report. J Sex Res 1983; 19 49–57.
Gouin JP, Carter CS, Pournajafi-Nazarloo H, Glaser R, Malarkey WB, Loving TJ, Stowell J, Kiecolt-Glaser JK. Marital behavior, oxytocin, vasopressin, and wound healing. Psychoneuroendocrinology 2010; 35 (7): 1082–90.
Guzmán YF, Tronson NC, Jovasevic V, Sato K, Guedea AL, Mizukami H, Nishimori K, Radulovic J. ear-enhancing effects of septal oxytocin receptors. Nature Neurosci 2013; 16 (9): 1185–7.
Halpern CR, Udry JR, Suchindran C. Monthly measures of salivary testosterone predict sexual activity in adolescent males. Arch Sex Behav 1998; 27:445-465.
Halpern CT, Udry JR, Campbell B, Suchindran C, Mason GA. Testosterone and religiosity as predictors of sexual attitudes and activity among adolescent males: a biosocial model. J Bioso Sci. 1994; 26: 217-234.
Halpern CT, Udry JR, Suchindran C. Testosterone predicts initiation of coitus in adolescent females. Psychosom Med. 1997; 59: 161-171.
Heinrichs M, Baumgartner T, Kirschbaum C, Ehlert U. Social support and oxytocin interact to suppress cortisol and subjective responses to psychosocial stress. Biological Psychiatry 2003; 54 (12): 1389–98.
Hertlein K, Gerald R Weeks, Nancy Gambescia. Systemic Sex Therapy, New York: Routledge, 2009.
Huber D, Veinante P, Stoop R. Vasopressin and oxytocin excite distinct neuronal populations in the central amygdala. Science 2005; 308 (5719): 245–8.
Hull EM. Sex, drugs and gluttony: how the brain controls motivated behaviors. Physiol Behav 2011; 104: 173-177.
Hyyppa M, Rinne OK, Sonnine V. The activating effect of L-dopa treatment on sexual functions and its experimental background. Acta Neurol Scand Suppl. 1970; 43(suppl 46):223-224.
Jannini E.A, Wise N. Frangos E. and Komisaruk BR. Peripheral and Central Neural Bases of Orgasm. In Texbook of Sexual Function and Dysfunction: Diagnosis and Treatment, First Edition, Edited by Sue W. Goldstein, Noel N. Kim, Anita H. Clayton. New York: John Wiley & Sons Ltd, 2018. Chapter 13, pp:179-195; 2018.
Kaplan Helen Singer. The New Sex Therapy, New York: Penguin Books, 1981.
Karama S, Lecours AR, Leroux J-M, Bourgouin P, Beaudoin G, Joubert S et al. Areas of brain activation in males and females during viewing of erotic film excerpts. Human Brain Mapping 2002; 16: 1-13.
Katzung Bertram G, Masters Susan, B. Tevor Antony J. Basic and Clinical Pharmacology, Lange Publications, 12th edition, Chicago: Mc Graw Hill, 2012.
Kawabata H, Zeki S. Neural correlates of beauty. J Neurophysiol 2004; 91(4):1699-705.
Kendrick K.M. The neurobiology of social bonds. J Neuroendocrinol 2004; 16: 1007-1008.
Kim WK, Schenck CH, Grant JE, Yoon G, Dosa PI, Odlaug BL, Shreiber, LRN, Hurwitz TD, Pfaus JG. Neurobiology of Sexual Desire. SexuS Journal 2015; 1 (1): 044-076.
Kirsch P, Esslinger C, Chen Q, Mier D, Lis S, Siddhanti S, Gruppe H, Mattay VS, Gallhofer B, Meyer-Lindenberg A. Oxytocin modulates neural circuitry for social cognition and fear in humans. J. Neurosci 2005; 25 (49): 11489–93.
Komisaruk BR, Beverly W. The suppression of pain by genital stimulation in females. Ann Rev Sex Re 1995; 6: 151 –186.
Komisaruk BR, Beverly W. Love as a sensory stimulation: Physiological Consequences of its deprivation and expression. Psychoneuroendocrinology 1998; 8:927-944.
Komisaruk BR, Sansone G. Neural pathways mediating vaginal function: The vagus nerves and spinal cord oxytocin. Scandinavian J Psychiatry 2003; 44:241-250.
Komisaruk BR, Whipple B, Crawford A, Grimes S, Liu WC, Kalnin A, Mosier K. Brain activation during vaginocervical self-stimulation and orgasm in women with complete spinal cord injury: fMRI evidence of mediation by vagus nerves. Brain Research 2004; 1024:77-88.
Komisaruk BR, Beverly W. Functional MRI of the brain during orgasm in women. Ann Rev Sex Res 2005; 16: 62-86.
Komisaruk BR, Beyer-Flores C, Bevely W. The Science of Orgasm, Baltimore: John Hopkins University Press, 2006.
Komisaruk BR, N Wise, EF, Liu WC, Allen K, Brody S. Women's clitoris, vagina, and cervix mapped on the sensory cortex: fMRI evidence, J Sex Med 2011; 8(10):2822-30.
Komisaruk BR and Whipple B. Non-genital Orgasms. Sex Rel Ther 2012; 1-17.
Komisaruk BR, Allen K, Wise N, et al. Men’s genital structures mapped on the sensory cortex: fMRI evidence. Soc for Neurosc Ann Conference. 2013.
Koob GF, Le Moal M. Drug abuse: hedonic homeostatic dysregulation. Science 1997; 278: 52–58.
Koob GF, Volkow ND. Neurocircuitry of Addiction. Neuropsychopharmacology 2010; 35:217-238.
Koob GF, Simon EJ. The Neurobiology of Addiction: Where we have been and where are we going? J Drug Issues 2009; 39 (1) 115-132.
Kosfeld M, Heinrichs M, Zak P, Fischbacher U, Fehr E. Oxytocin increases trust in humans. Nature 2005; 435:673–676.
Kringelbach ML, Berridge KC. The Joyful Mind. Scientific American 2012; 307 (2): 44–45.
Krüger TH, Haake P, Chereath D, Knapp W, Janssen OE, Exton MS, Schedlowski M, Hartmann U. Specificity of the neuroendocrine response to orgasm during sexual arousal in men. J Endocrinol 2003; 177 (1): 57–64.
Kuchinska S. The Chemistry of Connection: How the Oxytocin Response Can Help You Find Trust, Intimacy and Love, New York: New Harbinger Publications, 2009.
Kumsta R, Heinrichs M. Oxytocin, stress and social behavior: neurogenetics of the human oxytocin system. Curr Opin Neurobiol 2012; 4388 (12) 146-148.
Ladas AK, Beverly W, Perry JD. The G-Spot and other Discoveries About Human Sexuality, New York: Henry Holtand Company, (first publication 1982), 2005.
Lal S, Ackman D, Thavundayil JX, Kiely ME, Eti Enne . Effect of apomorphine, a dopamine re Captor agonist, on penile tumescence in normal subjects. Prog Neuropsychopharmacol Biol Psychiatry 1984; 8:695-699.
Landén M, Högberg P, Thase ME. Incidence of sexual side effects in refractory depression during treatment with citalopram or paroxetine. J Clin Psychiatry 2005; 66: 100-106.
Langeslag SJ, Muris P, Franken IH. Measuring Romantic Love: Psychometric Properties of the Infatuation and Attachment Scales. J Sex Res 2012 Oct 25.
Lee HJ, AH Macbeth, JH Pagani, WS Young WS. Oxytocin: The Great Facilitator of Life, Progress in Neurobiology; 88 (2): 127–51, June 2009.
Levin RJ. The breast/nipple/areola complex and human sexuality. Sex Relat Ther 2006; 21:237–49.
Levin RJ. The pharmacology of the human female orgasm — Its biological and physiological backgrounds. Pharmacol Biochem Behav 2014; 121: 62-70.
Lischke A, Gamer M, Berger C, Grossmann A, Hauenstein K, Heinrichs M, Herpertz SC, Domes G. Oxytocin increases amygdala reactivity to threatening scenes in females. Psychoneuroendocrinology 2012; 37 (9): 1431–1438.
Lopez AM, Weintraub D, Claassen DO. Impulse Control Disorders and Related Complications of Parkinson's Disease Therapy. Semin Neurol 2017;37(2):186-192.
Luef GJ. Epilepsy and Sexuality. Seizure 2008; 17: 127-130.
Lundeberg T, Meister B, Bjo¨ rkstrand E, Uvna¨ s-Moberg K. Oxytocin modulates the effects of galanin in carrageenan-induced hyperalgesia in rats. Brain Res 1993; 608:181–185.
Magon N, Kalra S. The orgasmic history of oxytocin: Love, lust, and labor. Indian J Endoc & Metabol 2011; Suppl 3:S156-61.
Maravilla KR & Yang CC. Magnetic resonance imaging and the female sexual response: Overview of techniques results and future directions. J Sexual Med 2008; 5: 1559-1571.
Maravilla KR & Yang CC. Sex and the brain: The role of fMRI for assessment of sexual function and response. Int J Impotence Res 2007; 19: 25-29.
Marazziti D and Canale D. Hormonal changes when falling in love. Psychoneuroendocrinology 2004; 29: 931-936. Marazziti D and Cassano G.B. The neurobiology of attraction. J Endocrinol Invest 2003; 26:58-60.
Marazziti D, Akiskal HS, Rossi A et al., Alteration of the platelet serotonin transporter in romantic love. Psychol Med 1999; 29:741–745.
Marazziti D, Canale D. Hormonal changes when falling in love. Psychoneuroendocrinology 2004; 29 (7): 931–936.
Marazziti D, Consoli G, Albanese F, Laquidara E, Baroni S, Catena Dell'osso M. Romantic attachment and subtypes/dimensions of jealousy. Clin Pract Epidemiol Ment Health 2010; 8(6):53-8.
Marazziti D, Dell'Osso B, Baroni S, Mungai F, Catena M, Rucci P, Albanese F, Giannaccini G, Betti L, Fabbrini L, Italiani P, Del Debbio A, Lucacchini A, Dell'Osso L. A relationship between oxytocin and anxiety of romantic attachment. Clinical Practice and Epidemiology in Mental Health 2006; 2 (1): 28.
Marder SR, Meibach RC. Risperidone in the treatment of schizophrenia. Am J Psychiatry. 1994; 151:825-835.
Martin-Du Pan RC. Oxytocin: the hormone of love, trust and social bond. Clinical use in autism and social phobia. Rev Med Suisse 2012; 21(8): 627-30.
Maslow A. Toward a Psychology of Being, New York: Von Nostrand Reinhold Company, 1968.
Master SL, Eisenberger NI, Taylor SE, Naliboff BD, Shirinyan D, Lieberman MD. A Picture’s Worth: Partner Photographs Reduce Experimentally Induced Pain. Psychological Science 2009; 20: 1316-1318.
Masters W, Johnson V. Human Sexual Response. NY: Little Brown Co. 1966.
Matsuzaki M, Matsushita H, Tomizawa K, Matsui H. Oxytocin: a therapeutic target for mental disorders. J Physiol Sci 2012; 62 (6): 441–4.
McQuaid RJ, McInnis OA, Abizaid A, Anisman H. Making room for oxytocin in understanding depression. Neurosci & Biobehav Rev 2014; 45: 305–22.
Melis MR, Argiolas A. Dopamine and sexual behavior. Neuro science and Biobehav Rev 1995; 19:19–38.
Meston C, Frohlich P. The Neurobiology of Sexual Function. Arch Gen Psychiatry 2000; 57: 1012-1030.
Meston CM, Gorzalka BB, Wright JM. Inhibition of subjective and physiological sexual arousal in women by clonidine. J Psychosom Med 1997; 59: 399-407.
Meston CM, Gorzalka BB. Psychoactive drugs and human sexual behavior: the role of serotoner gic activity. J Psychoactive Drugs 1992; 24: 1-40.
Meston CM, Gorzalka BB. The effects of imme diate,delayed, and residual sympathetic activa tion on sexual arousal in women. Behav Res Ther 1996; 34:143-148.
Meston CM, Gorzalka BB. The effects of sympathetic activation on physiological and subjec Tive sexual arousal in women. Behav Res Ther 1995; 33:651-664.
Meston CM, Heiman JR. Ephedrine-activated sexual arousal in women. Arch Gen Psychiatry. 1998; 55:652-656.
Montejo-Gonzalez AL, et al. SSRI Induced sexual dysfunction: fluoxetine, parox etine,sertraline, and fluvoxamine in a prospective, multicenter, and descriptive clinical study of 344 patients. J Sex Marital Ther 1997; 23: 176-193.
Moore TJ, Glenmullen J, Mattison DR. Reports of pathological gambling, hypersexuality, and compulsive shopping associated with dopamine receptor agonist drugs. JAMA Intern Med 2014; 174(12):1930-3.
Morris D. The Naked Woman: A Study of the Female Body. London: St. Martin's Griffin, 2004.
Moulier V, Mouras H, Pelegrini-Isaac M, Glutron D, Rouxel R & Grandjean B. Neuroana-tomical correlates of penile erection evoked by photographic stimuli in human males. Neuroimage 2006; 33.
Murphy MR, Seckl JR, Burton S, Checkley SA, Lightman SL. Changes in oxytocin and vasopressin secretion during sexual activity in men. Jf Clin Endoc & Metab 1987; 65 (4): 738–41.
Nakajima M, Görlich A, Heintz N. Oxytocin modulates female sociosexual behavior through a specific class of prefrontal cortical interneurons. Cell 2014; 159 (2): 295–305.
Odendaal JS, Meintjes RA. Neurophysiological correlates of affiliative behaviour between humans and dogs. Veterinary J 2003; 165 (3): 296–301.
Ortigue S & Bianchi-Demicheli F. Interactions between human sexual arousal and exual desire: a challenge for social neuroscience. Revue Medicale Suisse 2007; 3, 809-813.
Ortigue S, Bianchi-Demicheli F de C, Hamilton AF & Grafton ST. The neural basis of love as a subliminal prime: An event-related functional magnetic resonance imaging study. J Cog Neurosci 2007; 19: 1218-1230.
Ortigue S, Bianchi-Demicheli F, Patel N, Frum C, Lewis JW. Neuroimaging of love: fMRI meta-analysis evidence toward new perspectives in sexual medicine. J Sex Med 2010.
Özkara C, Özdemir S, Yilmaz A, Uzan M, Yeni N, Özmen M. Orgasm-induced seizures: A study of six patients. Epilepsia 2006; 47 (12) 2193-2197
Patterson WM. Fluoxetine-induced sexual dysfunction [letter]. J Clin Psychiatry 1993; 54: 71.
Pfaus JG & Gorzalka BB. Opioids and sexual behavior. Neurosci Biobehav Rev 1987; 11: 1–34.
Pfaus JG, Kippin TE, Coria-Avila GA, Gelez H, Alfonso VM, Ismail N, et al. Who, what, where, when (and maybe even why)? How the experience of sexual reward connects sexual desire, preference, and performance. Arch Sex Behav 2012; 41: 31-62.
Pfaus JG, Tse TL, Werk CM, Chanda ML, Leblonde A, Harbour VL, et al. Enhanced synaptic responses in the piriform cortex associated with sexual stimulation in the male rat. Neuroscience 2009; 164: 1422-1430.
Phelps E, LeDoux J. Contributions of the amygdala to emotion processing: from animal models to human behavior. Neuron 2005; 48:175–187.
Poryazova R; Khatami R; Werth E; Bassetti CL. Weak with Sex: Sexual Intercourse as a Trigger for Cataplexy. J Sex Med 2009; 6(8):2271-7.
Pukall CF, Strigo IA, Binik YM, et al. Neural correlates of painful genital touch in women with vulvar vestibulitis syndrome. Pain 2005; 115:118–127.
Reich W. The function of the orgasm: sex – economic problems of biological energy. New York: Farrar, Straus and Giroux (V.R. Carfagno, Translation; original work published 1942), 1973.
Reinert AE, Simon JA. "Did You Climax or Are You Just Laughing at Me?" Rare Phenomena Associated With Orgasm. Sex Med Rev 2017; 5:275-281.
Richard JM, Castro DC, Difeliceantonio AG, Robinson MJ, Berridge KC. Mapping brain circuits of reward and motivation: in the footsteps of Ann Kelley. Neurosci Biobehav Rev 2013; 37 (9 Pt A): 1919–1931.
Robinson TE & Berridge KC The psychology and neurobiology of addiction: an incentive-sensitization view. Addiction 2000; 95 (Suppl 2) S91–S117.
Sansone GR, Gerdes CA, Steinman JL, et al. Vaginocervical stimulation releases oxytocin within the spinal cord in rats. Neuroendocrinology 2002; 75:306–315.
Sansone GR, Komisaruk BR. Evidence that oxytocin is an endogenous stimulator of autonomic sympathetic preganglionics: the pupillary dilatation response to vaginocervical stimulation in the rat. Brain Res. 2001; 898:265–271.
Sayin HÜ. Artırılmış Cinsel Doyum: ESR; Kadınlarda Ultra Orgazm (Expanded Sexual Response: ESR; Ultra Orgasm in Women), İstanbul: Tantra Akademi/Onur Publications, 2012-a. (336 pages) ISBN: 978-605-87091-0-2.
Sayin HÜ. Doors of female orgasmic consciousness: New theories on the peak experience and mechanisms of female orgasm and expanded sexual response. (Research and Review) NeuroQuantology 2012-b; 10 (4): 692-714.
Sayin HÜ. Cinsellikte Farklı Boyutlar (Derin Sex) (Different Dimensions of Sexuality: Deep Sex), second expanded edition, two volumes) Second expanded-extended updated edition, 2 volumes. İstanbul: Tantra Akademi/Onur Publications, 2014. (880 pages) ISBN: 978-60587091-9-5
Sayin HÜ. Kadınlarda Orgazmın Psikolojisi (Psychology of Orgasm in Women). İstanbul: Tantra Akademi/Onur Publications, 2015-a. (304 pages) ISBN: 978-605-85249-3-4
Sayin HÜ. Altered States of Consciousness Occurring During Expanded Sexual Response (ESR) in the Human Female: Preliminary Definitions (Review). SexuS Journal 2015-b; 1 (1): 077-088. www.sexusjournal.com
Sayin HÜ. Psikoaktif Bitkilerin Tarihi ve Farklı Bilinç Hallerinin Psikofarmakolojisi (The History of Psychoactive Plants and the Psychopharmacology of Altered States of Consciousness), Ders kitabı-Textbook, İstanbul: Tantra Akademi/Onur Publications, (324 pages), 2016. ISBN: 978-605-85249-6-5
Sayin HÜ. Kadın ve Orgazm: Orgazm Yöntemleri ve Yeni Cinsel Terapi Teorileri (Women and Orgasm: Orgasm Methods and Novel Sex Therapy Theories), İstanbul: Tantra Akademi/Onur Publications (480 pages). 2017-a. ISBN: 978-605-2141-02-1
Sayin HÜ. Tantra, ESR and the Limits of Female Potentials (Review). SexuS Journal 2017-b; 2 (3): 055-074. www.sexusjournal.com
Sayin HÜ. Female Orgasmic Consciousness: New Horizons (Review). SexuS Journal 2017-c; 2 (4): 117-145. www.sexusjournal.com
Sayin HÜ. Psychoactive Plants Consumed in Religious Rituals: Common Archetypal Symbols & Figures in Myths & Religions (Review). SexuS Journal 2017-d; 2 (5): 201-236. www.sexusjournal.com
Sayin HÜ. Neurons’ Secret Archaic Optic Language: Entoptic Images, Pshophenes and Archetypes (Review). SexuS Journal 2017-e ; 2 (6): 308-348. www.sexusjournal.com
Sayin HÜ. Cinsel Beyin (Sexual Brain-Sexual Mind). İstanbul: Tantra Akademi/Onur Publications (384 pages). 2018-a. ISBN: 978-605-2141-19-9
Sayin HÜ. A Comparative Review of Psychopharmacology of Hallucinogen-Induced Altered States of Consciousness: Relation to Sexuality (Review). SexuS Journal 2018-b; 3 (7): 413-450. www.sexusjournal.com
Sayin HÜ & Kocatürk A. Expanded Sexual Response in the Human Female: The Mechanisms of Expanded Orgasms in Women. SexuS Journal 2018-c; 3 (8): 533-548. www.sexusjournal.com
Sayin HÜ. The Neural Correlates of the “ID”. E-Cronicon Neurology 2019; 11(4): 256-273.
Schenck CH, Arnulf, I, Mahowald MW. Sleep And Sex: What Can Go Wrong? A Review Of The Literature On Sleep Related Disorders And Abnormal Sexual Behaviors And Experiences. Sleep 2007; 30: 683-702.
Schreiner-Engel P, Schiavi RC, Smith H, White D. Sexology: sexual biology, behavior and sexual function: a critical review. Clin Psychopharmacol 1999; 19: 67-85.
Schultz W. Neuronal reward and decision signals: from theories to data. Physiological Reviews 2015; 95 (3): 853–951.
Sherwin BB. The free testosterone index as a predictor of sexual behavior in postmenopausal therapy. In: Hoch Z and Lief HI, eds. Selected Papers Of The Fifth World Congress Of Sexology. Amsterdam, the Netherlands: Exerpta Medica; 1982: 88-92.
Steinman JL, Komisaruk BR, Yaksh TL, Tyce GM. Spinal cord monoamines modulate the antinociceptive effects of vaginal stimulation in rats. Pain 1983; 16:155–166.
Succu S, Sanna F, Argiolas A, Melis MR. Oxytocin injected into the hippocampal ventral subiculum induces penile erection in male rats by increasing glutamatergic neurotransmission in the ventral tegmental area. Neuropharmacology 2011; 61: 181-188.
Succu S, Sanna F, Melis T, Boi A, Argiolas A, Melis MR. Stimulation of dopamine receptors in the para-ventricular nucleus of the hypothalamus of male rats induces penile erection and increases extra-cellular dopamine in the nucleus accumbens: Involvement of central oxytocin. Neuropharmacology 2007; 52: 1034-1043.
Surbeck W, Bouthillier A, Nguyen DK. Bilateral cortical representation of orgasmic ecstasy localized by depth electrodes. Epilepsy & Behav Case Rep 2013; 1: 62-65.
Tarlaci S. Brain Chemistry of love and sex. SexuS Journal 2017; 2 (5): 189-200, 2017. www.sexusjournal.com
Taylor P. PhD Thesis: An Observational and Comparative Study of Practitioners of Expanded Orgasm: An Investigation of an Effective and Accessible Path to Transcendent States of Consciousness, Submitted in Partial Fulfillment of the Requirements for the Degree of Philosophy in Transpersonal Psychology, International University of Professional Studies, Maui, Hawaii, 2000.
Taylor P. Expanded Orgasm, Soar to Ecstasy at your Lover's Every Touch, New York: Sourcebooks, 2002.
Udry JR, Billy JO, Morris NM, Groff TR, Raj MH. Serum androgenic hormones motivate sexual behavior in adolescent boys. Fertil Steril 1985; 43: 90-94.
Uitti RJ, Tanner CM, Rajput AH, Goetz CG, Klawans HL, Thiessen B. Hypersexuality with antiparkinsonian therapy. Clin Neuropharmacol 1989; 12: 375-383.
van Leengoed E, Kerker E, Swanson HH. Inhibition of post-partum maternal behaviour in the rat by injecting an oxytocin antagonist into the cerebral ventricles. J Endocrinol 1987; 112 (2): 275–82.
Veronneau‐Longueville F, Rampin O, Freund‐Mercier MJ, et al. Oxytocinergic innervation of autonomic nuclei controlling penile erection in the rat. Neuroscience 1999; 93:1437–1447.
Viero C, Shibuya I, Kitamura N, Verkhratsky A, Fujihara H, Katoh A, Ueta Y, Zingg HH, Chvatal A, Sykova E, Dayanithi G. Oxytocin: Crossing the Bridge between Basic Science and Pharmacotherapy. CNS Neurosci & Therap 2010; 16 : e138–e156.
Vitalo A, Fricchione J, Casali M, et al. Nest making and oxytocin comparably promote wound healing in isolation reared rats. PLoS One 2009; 4:e5523.
Viviani D, Charlet A, van den Burg E, Robinet C, Hurni N, Abatis M, Magara F, Stoop R. Oxytocin selectively gates fear responses through distinct outputs from the central amygdala. Science 2011. 333 (6038): 104–7.
Volkow, N.D., Wang, G.J., Telang, F., Fowler, J.S., Logan, J., Jayne, M., Ma, Y., Pradhan, K., Wong, C. Profound decreases in dopamine release in striatum in detoxified alcoholics: possible orbitofrontal involvement. J. Neurosci.; 27: 12700–12706, 2007.
Walter M, Bermpohl F, Mouras H, Schiltz K, Tempelmann C, Rotte M et al. Distinguishing specific sexual and general emotional effects in fMRI-subcortical and cortical arousal during erotic picture viewing. Neuroimage 2008; 40: 1482-1494.
Wei D, Lee D, Cox CD, Karsten CA, Peñagarikano O, Geschwind DH, Gall CM, Piomelli D. Endocannabinoid signaling mediates oxytocin-driven social reward. Proc Nat Acad Sci USA 2015; 112 (45): 14084–9.
Whipple B, Komisaruk BR. Analgesia produced in women by genital self-stimulation. J Sex Res 1988; 24: 130-140.
Whipple B, Komisaruk BR. Elevation of pain threshold by vaginal stimulation in women. Pain 1985; 21:357–367.
Wise NJ, Frangos E, Komisaruk BR. Brain activity Unique to Orgasm in Women: An fMRI Analysis. J Sexual Med 2017; 14 (11): 1380-1391.
Xu X, Aron A, Brown L, Cao G, Feng T & Weng X. Reward and motivation systems: A brain mapping study of early-stage intense romantic love in Chinese participants. Hum Brain Mapp 2010; 32, 249-257.
Xu X, Wiesenfeld-Hallin Z. Intrathecal oxytocin facilitates the spinal nociceptive flexor reflex in the rat. Neuroreport 1994; 5:750–752.
Yang J, Yang Y, Chen J, Liu W, Wang C, Lin B. Central oxytocin enhances antinociception in the rat. Peptides 2007; 28:1113–1119.
Yang J. Intrathecal administration of oxytocin induces analgesia in low back pain involving the endogenous opiate peptide system. Spine (Phila Pa 1976) 1994; 19:867–871.
Young, L, HH Z. Oxytocin. In: Pfaff ED AM, editor. Molecular mechanisms of hormones actions on behavior, San Diego, CA: Academic Press, pp: 783–802, 2009.
Younger J, Aron A, Parke S, Chatterjee N, Mackey S. Viewing Pictures of a Romantic Partner Reduces Experimental Pain: Involvement of Neural Reward Systems. 2010; 5(10): e13309. doi:10.1371/journal.pone.0013309.
Zdrok V. Anatomy of Pleasure. Philadelphia: Infinity Publishing Co., 2004.
Zeki S, Romaya JP. The brain reaction to viewing faces of opposite- and same-sex romantic partners. PLoS One. 2010; 5(12):e15802.
Zeki S. Neurobiology of love. FEBS Letters 2007; 581: 2575-2579