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Review| Volume 154, ISSUE 1, P3-8, January 2011

Embryology and anatomy of the vulva: the female orgasm and women's sexual health

      Abstract

      Sexual health is vital to overall well-being. Orgasm is a normal psycho-physiological function of human beings and every woman has the right to feel sexual pleasure. The anatomy of the vulva and of the female erectile organs (trigger of orgasm) is described in human anatomy textbooks. Female sexual physiology was first described in Dickinson's textbook in 1949 and subsequently by Masters and Johnson in 1966. During women's sexual response, changes occur in the congestive structures that are essential to the understanding of women's sexual response and specifically of their orgasm. Female and male external genital organs arise from the same embryologic structures, i.e. phallus, urogenital folds, urogenital sinus and labioscrotal swellings. The vulva is formed by the labia majora and vestibule, with its erectile apparatus: clitoris (glans, body, crura), labia minora, vestibular bulbs and corpus spongiosum. Grafenberg, in 1950, discovered no “G-spot” and did not report an orgasm of the intraurethral glands. The hypothetical area named “G-spot” should not be defined with Grafenberg's name. The female orgasm should be a normal phase of the sexual response cycle, which is possible to achieve by all healthy women with effective sexual stimulation. Knowledge of the embryology, anatomy and physiology of the female erectile organs are important in the field of women's sexual health.

      Keywords

      1. Introduction

      Orgasm is a normal psycho-physiological function of human beings [
      • Masters W.H.
      • Johnson V.E.
      Human sexual response.
      ]. Women have the right to feel sexual pleasure: “sexual health is a global issue that is vital to overall well-being” [
      • Abdool Z.
      • Thakar R.
      • Sultan A.H.
      Postpartum female sexual function.
      ]. The anatomy of the vulva and of the female erectile organs (trigger of orgasm) is described in human anatomy textbooks with the clitoris considered as homologous to the male penis [
      • Masters W.H.
      • Johnson V.E.
      Human sexual response.
      ,
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ]. Female sexual physiology was first described in Dickinson's textbook in 1949 and subsequently by Masters and Johnson in 1966 [
      • Masters W.H.
      • Johnson V.E.
      Human sexual response.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ].
      For women, clitoral stimulation is important for achieving orgasm. Distinguishing between clitoral and vaginal orgasm is not correct from a physiological point of view [
      • Masters W.H.
      • Johnson V.E.
      Human sexual response.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ,
      • Mah K.
      • Binik Y.M.
      The nature of human orgasm: a critical review of major trends.
      ,
      • Laqueur T.
      Making sex: body and gender from the Greeks to Freud.
      ]. The vagina is mainly a reproductive organ with little sensitivity [
      • Masters W.H.
      • Johnson V.E.
      Human sexual response.
      ,
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ,
      • Mah K.
      • Binik Y.M.
      The nature of human orgasm: a critical review of major trends.
      ], the vaginal orgasm is not corroborated by anatomical evidence and the term was invented by Freud in 1905. For Laqueur the theory of Freud is an instrument for making women accept their social role [
      • Laqueur T.
      Making sex: body and gender from the Greeks to Freud.
      ]. During women's sexual response, changes occur in the congestive structures that are essential to the understanding of their sexual response and specifically of their orgasm.

      2. The embryological development of the vulva

      In anatomy textbooks there is a separation between the embryological development of the internal and external genital organs in the male and in the female. It is important to know this because it is related to the function of these organs, i.e. the internal genitals have a reproductive function, while the external ones have the function of giving pleasure [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ].
      There is an important difference between the two genders: in the male, ejaculation and the release of sperm happen at the same time as orgasm. In the female, however, orgasm is not connected to reproductive function, and ovulation is not accompanied by orgasm. The ovum is fertilized when it is in the uterine tubes, which are internal organs, but the clitoris and the other female erectile structures are located externally [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ].
      In male and female embryos the external and the internal genital organs are “indifferent” for the first few weeks. At 8 weeks of gestation the external genitalia of males and females begin to develop from the urogenital sinus (which is divided into pelvic and phallic parts), from the genital tubercle or phallus, from the urogenital folds and from the labioscrotal swellings [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ]. The corpora cavernosa of the clitoris and the glans with the prepuce are formed from the phallus (which bends forward or caudally) [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ]. In the female these structures are not so large as in the male, although in the early phases the clitoris exceeds the length of the homologous male counterpart [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ].
      The vestibule of the vagina, the labia minora, the vestibular bulbs and the female corpus spongiosum are formed by the pelvic and phallic part of the urogenital sinus and from the urogenital folds, which do not fuse together. The labioscrotal swellings do not meld and they form the labia majora [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Van Turnhout A.A.
      • Hage J.J.
      • Van Diest P.J.
      The female corpus spongiosum revisited.
      ].
      In the female, the pelvic part of the urogenital sinus moves down, is incorporated by the phallic part and opens externally as the vaginal and urethral orifices. In the male, the pelvic part of the urogenital sinus corresponds to the internal portion of urethra located under the seminal collicle (veru montanum), a prominence in the dorsal surface of the prostatic urethra where the ejaculatory ducts open. Between the two ejaculatory ducts is the prostatic utricle (i.e. the “male vagina”: it is the homologous to the female vagina) [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ]. Sometimes the ejaculatory ducts open into the prostatic utricle instead of the prostatic urethra. The small segment of the prostatic urethra which reaches the seminal collicle from the bladder corresponds to the whole female urethra [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ].
      The main difference between male and female genitalia is the lack of the development in the female of the external urethra, as consequence of the non-fusion of the urogenital folds. However, the structures responsible for forming the external urethra in the male are also present in the female and correspond to the vestibule of the vagina and to the internal surface of the labia minora [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ].
      The vulva, in fetal life and after birth, appears with the labia majora closed: to visualize the vaginal opening it is necessary to open the labia, and therefore the labia minora are in contact with each other. The picture of the normal vulva should be shown with the labia majora and the labia minora closed.

      3. The vulva

      The vulva is formed by the labia majora and vestibule, with its erectile apparatus: labia minora, clitoris (glans, body, crura), and vestibular bulbs with the corpus spongiosum. These structures are localized under the urogenital diaphragm, behind the pubic symphysis in the anterior perineal region [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ].

      3.1 The vestibule

      The vaginal vestibule is triangular in shape. It corresponds to the dorsal wall of the male cavernosa urethra. Here the external orifice of the urethra with the paraurethral (Skene's) ducts opening on both sides are localized [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ]. Their length is 0.5–3 cm and they are found, in women, with the intraurethral (Skene's) gland which has been named the female prostate [
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ,
      • Zaviacic M.
      • Jakubovská V.
      • Belošovi R.M.
      • Breza J.
      Ultrastructure of the normal adult human female prostate gland (Skene's gland).
      ]. This structure can be affected by the same diseases as the male counterpart, including carcinoma and prostatitis; the secretion of these glands, expelled during the female orgasm (female ejaculation) into the vaginal vestibule through the urethral meatus or through the orifices of the paraurethral ducts, contains prostate-specific antigen [
      • Zaviacic M.
      • Jakubovská V.
      • Belošovi R.M.
      • Breza J.
      Ultrastructure of the normal adult human female prostate gland (Skene's gland).
      ].
      From a physiological point of view the term “female emission” is more accurate than female ejaculation (in a few women there is a powerful expulsion of this fluid); in the male it corresponds to the phase of emission of seminal liquid which is ejected out in the prostatic urethra [
      • Giuliano F.
      • Clément P.
      Physiology of ejaculation: emphasis on serotonergic control.
      ]. The lack of the ejaculation phase in the female could explain why women do not have a refractory period and are able to have multiple orgasms.
      In the mid-line of the vestibule is located a longitudinal formation 4–5 mm wide, with defined margins and more pale than the nearby tissue, thin and well wedged in the dermis of the vestibule, which goes up from the urethral orifice to the sulcus of the inferior surface of the corpora cavernosa of the clitoris up to the glans, described first by Pozzi in 1884 and named “masculine bride” (habenulae uretrales). It corresponds to the dorsal part of the male corpus spongiosum of urethra: it is the female corpus spongiosum [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ,
      • Van Turnhout A.A.
      • Hage J.J.
      • Van Diest P.J.
      The female corpus spongiosum revisited.
      ].
      The vagina is an internal genital organ and the vaginal opening (introitus) is usually a median fissure, under the urethral meatus. The anterior vaginal wall is separated from the posterior urethral wall by the urethrovaginal septum and there are no secretory glands in the vagina [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ].
      The Bartholin's glands end up on the sides of the vaginal opening, and are homologous to the male bulbourethral glands or Cowper's gland [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ]. According to some studies, pre-ejaculatory fluid secreted by Cowper's gland in the distal urethra, during sexual stimulation, lacks in sperm so that it cannot cause a pregnancy after coitus interruptus [
      • Zukerman Z.
      • Weiss D.B.
      • Orvieto R.
      Does preejaculatory penile secretion originating from Cowper's gland contain sperm?.
      ]. Bartholin's glands secrete a sticky liquid similar to the mucus preceding orgasm and, as in the male, this pre-orgasmic secretion is light and more frequent after prolonged sexual stimulation [
      • Masters W.H.
      • Johnson V.E.
      Human sexual response.
      ]. Endocrine cells producing serotonin, calcitonin, bombesin, katacalcin and alpha-hCG have been found in Bartholin's glands [
      • Fetissof F.
      • Arbeille B.
      • Bellet D.
      • Barre I.
      • Lansac J.
      Endocrine cells in human Bartholin's glands. An immunohistochemical and ultrastructural analysis.
      ]. Furthermore in the vestibule many mucous glands open, called minor vestibular glands [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ].

      3.2 The labia minora

      The labia minora, or nymphs, are two small cutaneous folds, 3–4 cm long. They are normally approximated together. They correspond to the ventral wall of both the cavernosa urethra and the corpus spongiosum of the urethra in the male [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ]. They vary in size. They may be almost unrecognisable or may protrude from the labia majora (“hypertrophic” labia minora should not be considered a malformation). In addition they can be asymmetrical or double on one or both sides [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ,
      • Sakamoto H.
      • Ichikawa G.
      • Shimizu Y.
      • Kikuchi A.
      • Yamamoto T.
      Extreme hypertrophy of the labia minora.
      ,
      • Pardo J.
      • Solà V.
      • Ricci P.
      • Guilloff E.
      Laser labioplasty of labia minora.
      ]. In some races the labia minora can be very large because of the practice of stretching them: in some African populations they can be as large as 20 cm and are known as “Hottentot apron” [
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ]; today this is classified in the type IV female genital mutilations [
      • Catania L.
      • Abdulcadir O.
      • Puppo V.
      • Baldaro Verde J.
      • Abdulcadir J.
      • Abdulcadir D.
      Pleasure and orgasm in women with female genital mutilation/cutting (FGM/C).
      ].
      The upper points converge to form the prepuce and frenulum of the clitoris; posteriorly they form the frenulum of the labia minora but they can be also separated [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ]. The labia minora contain erectile tissue constituted by a thick connective tissue rich in small blood vessels. With sexual arousal the labia minora, because of engorgement with blood, become turgid, doubling or tripling in thickness. They have a great sensitivity, due to the considerable number of nerve endings and sensory receptors. Among these, as in the male, the most important for erogenous sensitivity are the genital corpuscles (typical receptors of the external genitals); the Krause–Finger corpuscles (corpuscles of voluptuousness) are predominant. Pacini and Meissner corpuscles (cutaneous mechanoreceptors distributed on various areas of the skin, but concentrated in areas especially sensitive to light touch) are also present [
      • Masters W.H.
      • Johnson V.E.
      Human sexual response.
      ,
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ,
      • Shafik A.
      • Shafik A.A.
      • Ahmed I.
      Response of the labia majora and minora to clitoral stimulation.
      ,
      • Yang C.C.
      • Cold C.J.
      • Yilmaz U.
      • Maravilla K.R.
      Sexually responsive vascular tissue of the vulva.
      ].

      3.3 The clitoris

      The clitoris is usually considered the homologue of the male penis. It is formed by two corpora cavernosa and the glans, covered by the prepuce [
      • Masters W.H.
      • Johnson V.E.
      Human sexual response.
      ,
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ,
      • Laqueur T.
      Making sex: body and gender from the Greeks to Freud.
      ,
      • Yang C.C.
      • Cold C.J.
      • Yilmaz U.
      • Maravilla K.R.
      Sexually responsive vascular tissue of the vulva.
      ]. The corpora cavernosa are two cylindrical organs, made of cavernous erectile tissue that becomes turgid and erect with sexual arousal; the crura represent the hidden part of the clitoris located in contact with the ischiopubic ramus, covered by the ischiocavernosus muscle, a superficial perineal muscle [
      • Masters W.H.
      • Johnson V.E.
      Human sexual response.
      ,
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ,
      • Yang C.C.
      • Cold C.J.
      • Yilmaz U.
      • Maravilla K.R.
      Sexually responsive vascular tissue of the vulva.
      ].
      The perineal muscles are innervated by branches of the pudendal nerve coming Onuf's nucleus; “pudendal nerve integrity may play a role in female sexual dysfunction” [
      • Connell K.
      • Guess M.K.
      • La Combe J.
      • et al.
      Evaluation of the role of pudendal nerve integrity in female sexual function using noninvasive techniques.
      ]. Onuf's nucleus is located at the sacral level of the spinal cord and is formed by motoneurons innervating the perineal muscles; the number and the size of these neurons are sexually dimorphic: this dimorphism is mediated by androgens (in absence of these hormones the motoneurons die by apoptosis) with ciliary neurotrophic factor (CNTF) and other trophic factors [
      • Catala M.
      Control of the development of Onuf's spinal nucleus.
      ]. Because of this, in females the ischiocavernosus muscles are much thinner than the male counterparts. Their contraction during female arousal determines a surge of blood in the crura towards the corpora cavernosa of the clitoris and a compression of the deep dorsal vein, contributing therefore to erection of the clitoris [
      • Masters W.H.
      • Johnson V.E.
      Human sexual response.
      ,
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ].
      The ischiocavernosus muscles, like the bulbocavernosus, are type mixed muscles (even if histologically they are striated). During erection, they are implicated in producing a continuous involuntary reflex hypertonic contraction, important not only for the rigidity of the penis but also for the maintenance of the erection [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Shafik A.
      Response of the urethral and intracorporeal pressures to cavernosus muscle stimulation: role of the muscle in erection and ejaculation.
      ].
      The crura are joined beyond and in front of the pubic symphysis, forming the body of the clitoris (which in the flaccid state is 1–3 cm long), which goes back and up to the glans. The suspensory ligament links the clitoral body to the mons pubis and pubic symphysis [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ,
      • Yang C.C.
      • Cold C.J.
      • Yilmaz U.
      • Maravilla K.R.
      Sexually responsive vascular tissue of the vulva.
      ]. The erectile tissue of the clitoris is made up of caverns covered by the tunica albuginea: the two corpora cavernosa have the same structure as the male penis; in some studies a subalbugineal layer in the corpora cavernosa is not present and the subalbugineal venous plexus is not located internally [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ,
      • Toesca A.
      • Stolfi V.M.
      • Cocchia D.
      Immunohistochemical study of the corpora cavernosa of the human clitoris.
      ]. The glans is 4–7 mm long and covers the distal part of the corpora cavernosa, from which it is independent [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ].
      The prepuce covers all or part of the glans, its size varies considerably, and it is just like the foreskin of the male penis; it is a specialized, specific erogenous tissue in both males and females. The fetal development of the prepuce and the glans in the male and female is similar: they are fused together during the first year after the birth. If they are not divided it can cause adhesions and phimosis, as in the male. An accumulation of smegma in infants and preadolescent girls under the prepuce is responsible for adhesions between the glans and prepuce and for phimosis of the clitoris [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ,
      • Goldstein A.T.
      • Burrows L.J.
      Surgical treatment of clitoral phimosis caused by lichen sclerosus.
      ].
      The glans contains cavernous tissue, as in the male, in direct contact with the skin due to the absence of the albuginea. It is rich in nerve endings and in genital and Krause–Finger corpuscles, and corpuscles of Pacini and Meissner [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ,
      • Yang C.C.
      • Cold C.J.
      • Yilmaz U.
      • Maravilla K.R.
      Sexually responsive vascular tissue of the vulva.
      ]. These receptors, because of the smaller dimensions of the clitoris, are more concentrated in the female than in the male glans and they probably do not require to surface to be excited by mechanical stimuli. Some studies have suggested that penile mechanoreceptors are more responsive when the penis is erect or near body temperature [
      • Johnson R.D.
      • Kitchell R.L.
      Mechanoreceptor response to mechanical and thermal stimuli in the glans penis of the dog.
      ]. In fact the glans has a protopathic sensitivity and it can be maintained, much more easily than in the male, in a state of arousal without an evident increase in its dimensions, with an easier receptiveness of the Krause–Finger corpuscles [
      • Masters W.H.
      • Johnson V.E.
      Human sexual response.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ,
      • Halata Z.
      • Munger B.L.
      The neuroanatomical basis for the protopathic sensibility of the human glans penis.
      ].
      Verkauf et al. with prospective measurement of clitoral dimensions in 200 normal women at routine gynecologic examination in an office setting, suggested that the clitoris is not influenced by age, height, weight, or current use of oral contraceptives, but in pregnant women it was significantly larger [
      • Verkauf B.S.
      • Von Thron J.
      • O’Brien W.F.
      Clitoral size in normal women.
      ]; the clitoral sexual response is not affected by aging [
      • Masters W.H.
      • Johnson V.E.
      Human sexual response.
      ].
      As in males, in females it is possible to have priapism of the clitoris, a rare condition associated with prolonged erection of the corpora cavernosa, an erection lasting for more than six hours and unassociated with sexual arousal, causing engorgement, swelling, and pain of the clitoris and immediate adjacent area. The cause of priapism, in male and female, is impaired outflow of blood from the corpora cavernosa because of venous obstruction or because of failure of the alpha-adrenergic relaxation system [
      • Medina C.A.
      Clitoral priapism: a rare condition presenting as a cause of vulvar pain.
      ,
      • Arntzen B.W.
      • de Boer C.N.
      Priapism of the clitoris.
      ]. “Most reported cases of female priapism describe the association with the use of antidepressant and other psychotropic drugs, all with alpha-adrenergic blocking potential, such as trazodone, buproprion citalopram and nefazodone. … Treatment consisted of discontinuing the offending medication or providing symptomatic pain relief. Serious permanent damage where treatment has been delayed has been reported in men but not in women. Furthermore, the association between congenital clitoromegaly and priapism has also not been reported previously. With this concern in mind, we felt justified to resort to management options described for male priapism but hitherto not for female priapism, i.e. the direct injection with epinephrine and heparin, followed by aspiration to provide immediate decompression” [
      • Arntzen B.W.
      • de Boer C.N.
      Priapism of the clitoris.
      ].

      3.4 The vestibular bulbs and the female corpus spongiosum

      The vestibular bulbs are two erectile organs and correspond to the urethral bulb in the male. The bulbs are situated in the anterior region of the perineum, the bulboclitoral region, which is homologous to penile region in the male. Their dimensions in the flaccid state are 3–4 cm in length and they become engorged or “erected” during arousal, when in the lower third of the vagina the “orgasmic platform” of Masters and Johnson is reached [
      • Masters W.H.
      • Johnson V.E.
      Human sexual response.
      ,
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ,
      • Van Turnhout A.A.
      • Hage J.J.
      • Van Diest P.J.
      The female corpus spongiosum revisited.
      ,
      • Yang C.C.
      • Cold C.J.
      • Yilmaz U.
      • Maravilla K.R.
      Sexually responsive vascular tissue of the vulva.
      ].
      The two bulbs are joined together, under the vestibule of the vagina, by the commissure of the bulbs and, through the corpus spongiosum or “pars intermedia”, they extend to the base of the glans. The corpus spongiosum is constituted by cavernous tissue which thins towards the glans and is situated underneath the vestibular epithelium [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ,
      • Van Turnhout A.A.
      • Hage J.J.
      • Van Diest P.J.
      The female corpus spongiosum revisited.
      ].
      In the angle of the clitoris, in the space between the corpora cavernosa and the female corpus spongiosum, there is the venous plexus of Kobelt, responsible for communication between the venous circulation of the bulbs and of the corpora cavernosa of the clitoris; it corresponds to the inferior veins of the male corpora cavernosa, which open in the inferior median sulcus between corpora cavernosa and the male urethra, and receives the veins coming from the upper part of the male corpus spongiosum [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ].
      The bulbs are covered by the bulbocavernosus muscles, which are implicated in inferior vaginismus, while the pubovaginal muscle is responsible for superior vaginismus. Some studies have suggested that the components of the levator ani muscle are the puborectal, iliococcygeal, pubovisceral muscles, further subdivided into pubovaginal, puboperineal and puboanal. This terminology was accepted in 1998 by the Federative Committee on Anatomical Terminology [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ,
      • Kearney R.
      • Sawhney R.
      • DeLancey J.O.
      Levator ani muscle anatomy evaluated by origin insertion pairs.
      ]. The rhythmic contractions of lower third of the vagina during orgasm are mainly due to the contractions of the bulbocavernosus muscle [
      • Masters W.H.
      • Johnson V.E.
      Human sexual response.
      ,
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ].

      3.5 The corpus spongiosum of the female urethra

      The corpus spongiosum of the urethra is present in all women, as in the male. It is a cavernous tissue rich in veins, situated at submucosal level and among the muscular bundles of the smooth muscular tunica of the urethral wall. It becomes engorged, or “erect”, with sexual arousal and Grafenberg in 1950 wrote “In the course of sexual stimulation the female urethra begins to enlarge” [
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Yang C.C.
      • Cold C.J.
      • Yilmaz U.
      • Maravilla K.R.
      Sexually responsive vascular tissue of the vulva.
      ,
      • Grafenberg E.
      The role of urethra in female orgasm.
      ]. Female urethral sensibility has been little investigated until now, though Dickinson wrote in 1949 “Indeed, the meatus is largely endowed with special sensibility” [
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ], and Grafenberg in 1950 described some cases of female and male urethral masturbation [
      • Grafenberg E.
      The role of urethra in female orgasm.
      ].
      Therefore the erectile structures are the same in female and in male. The clitoris is the homologue of the two corpora cavernosa, glans and prepuce of the male penis, and the male erection is equivalent to the erection of all female erectile organs (vaginal lubrication is a consequence of congestion) [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ,
      • Yang C.C.
      • Cold C.J.
      • Yilmaz U.
      • Maravilla K.R.
      Sexually responsive vascular tissue of the vulva.
      ]. The female external genitals are joined together even though separated by the presence of the vaginal opening, and they represent the penis and the scrotum of the male (Fig. 1, Fig. 2) [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ,
      • Van Turnhout A.A.
      • Hage J.J.
      • Van Diest P.J.
      The female corpus spongiosum revisited.
      ]. This has been recognized for many years. In 1949 Dickinson wrote: “Vulvo-vaginal changes in excitement. Protrusion of erectile bulbs of vestibule. Clitoris erection. Labial erection.” [
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ]. Laqueur wrote that Realdus Columbus, in 1559, named “the love or sweetness of Venus” (“amoris dulcedo”) what he found in the nature, a female penis [
      • Laqueur T.
      Making sex: body and gender from the Greeks to Freud.
      ,
      • Stringer M.D.
      • Becker I.
      Colombo and the clitoris.
      ].
      Figure thumbnail gr1
      Fig. 1I, The male penis. II – III, The female erectile organs (without vestibular bulb). Modified from: Puppo V. La sessualità umana e l’educazione a fare l’amore. Firenze: Loggia de’ Lanzi, 2005. I - 1, Glans. 2, Correspondent part to the labia minora. 3, Scrotum. 4, Anus. II - 1, Glans. 2, Labia minora. 3, Labia majora. 4, Anus. 5, Frenulum. 6, Sulcus nympholabialis. III - 1, Corpora cavernosa. 2, Septum. 3, Corpus spongiosum. 4, Kobelt's venous plexus.
      Figure thumbnail gr2
      Fig. 2The female erectile organs (The “female penis” without labia minora) Modified from: Puppo V. La sessualità umana e l’educazione a fare l’amore. Firenze: Loggia de’ Lanzi, 2005. 1, Body. 2, Glans. 3, Suspensory ligament. 4, Angle of clitoris. 5, Vestibular bulb. 6, Crus. 7, Corpus spongiosum.

      4. Discussion

      Addiego et al. in 1981 wrote: “At the April, 1979, testing session, the subject identified an erotically sensitive spot, palpable through the anterior wall of her vagina. We subsequently named this area the Grafenberg spot, in recognition of the person who wrote of its existence and relationship to female ejaculation (Grafenberg, 1950)” [
      • Addiego F.
      • Belzer E.G.
      • Comolli J.
      • Moger W.
      • Perry J.D.
      • Whipple B.
      Female ejaculation: a case study.
      ]. Subsequently the “Grafenberg spot” (i.e. intraurethral glands or female prostate) was seized on by the popular media, which called it the “G-spot”. The G-spot is currently an hypothesis: Grafenberg, in 1950, discovered no G-spot [
      • Grafenberg E.
      The role of urethra in female orgasm.
      ,
      • Hines T.M.
      The G-spot: a modern gynecologic myth.
      ]. In his article he did not describe a vaginal spot, but “the role of the urethra in female orgasm” (not the role of the vagina): the urethra, on the anterior wall of the vagina, is a “distinct erotogenic zone” in women [
      • Grafenberg E.
      The role of urethra in female orgasm.
      ].
      Grafenberg described some cases of female urethral masturbation and the corpus spongiosum of the female urethra, stating that, “analogous to the male urethra, the female urethra also seems to be surrounded by erectile tissues like the corpora cavernosa”. He did not report an orgasm of intraurethral glands, but wrote: “Involuntary expulsion of urine is reported in sex literature. In the cases observed by us, the fluid was examined and it had no urinary character. I am inclined to believe that urine reported to be expelled during female orgasm is not urine, but only secretions of the intraurethral glands” [
      • Grafenberg E.
      The role of urethra in female orgasm.
      ].
      In 1950 Grafenberg wrote about the corpus spongiosum of the female urethra: “In the course of sexual stimulation the female urethra begins to enlarge and can be felt easily. It swells out greatly at the end of orgasm. The most stimulating part is located at the posterior urethra, where it arises from the neck of the bladder” [
      • Grafenberg E.
      The role of urethra in female orgasm.
      ]. The so-called “G-spot” of the anterior vaginal wall is located in Pawlick's triangle (which corresponds to Lieutaud's triangle in bladder) that has a smooth vaginal mucosa and it is only a space with minor resistance [
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ].
      Hines in 2001 wrote: “Grafenberg discusses no evidence for a G-spot. …[Some female patients], he says, derived sexual pleasure from inserting objects, such as hat pins, into their urethras. Just how later writers (i.e. 2) transformed these reports into evidence for a G-spot is unclear. … Women who fail to find their G-spot, because they fail to respond to stimulation as the G-spot myth suggests that they should, may end up feeling inadequate or abnormal.” [
      • Hines T.M.
      The G-spot: a modern gynecologic myth.
      ].
      Burri et al. in 2010 wrote: “The existence of the G-spot seems to be widely accepted among women, despite the failure of numerous behavioral, anatomical, and biochemical studies to prove its existence. …there is no physiological or physical basis for the G-spot.” [
      • Burri A.V.
      • Cherkas L.
      • Spector T.D.
      Genetic and environmental influences on self-reported G-spots in women: a twin study.
      ]. Besides, there are no ultrasonographic images or anatomical pictures of the G-spot, and the female prostate has no anatomical structure that can cause an orgasm [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ]. The hypothetical area named G-spot should not be defined with Grafenberg's name.
      “G-spot amplification” is a procedure that is not medically indicated. The American College of Obstetricians and Gynecologists states: “So-called vaginal rejuvenation, designer vaginoplasty, revirgination, and G-spot amplification are vaginal surgical procedures being offered by some practitioners. Women should be informed about the lack of data supporting the efficacy of these procedures and their potential complications, including infection, altered sensation, dyspareunia, adhesions, and scarring” [
      • American College of Obstetricians and Gynecologists (ACOG)
      Committee opinion No. 378: vaginal “rejuvenation” and cosmetic vaginal procedures.
      ].
      The importance of orgasm has been questioned: “Orgasm and resolution are not essential in Basson's model of the sexual response cycle” [
      • Rosen R.C.
      • Barsky J.L.
      Normal sexual response in women.
      ]. Basson et al. in 2005 wrote, “Orgasm may or may not be necessary for sexual satisfaction” [
      • Basson R.
      • Brotto L.A.
      • Laan E.
      • Redmond G.
      • Utian W.H.
      Assessment and management of women's sexual dysfunctions: problematic desire and arousal.
      ]. According to Masters and Johnson, however: “Physiologically sexual response could be described as a cycle with four phases: excitement, plateau, orgasm, and resolution” (i.e. “the period of return to the unaroused state”) … If there has been considerable excitement but orgasm has not occurred, resolution takes a longer time … there is sometimes a lingering sensation of pelvic heaviness or aching that is due to continued vasocongestion, …this may create a condition of some discomfort, particularly if high levels of arousal were prolonged” [
      • Masters W.H.
      • Johnson V.E.
      Human sexual response.
      ,
      • Masters W.H.
      • Johnson V.E.
      • Kolodny R.C.
      Masters and Johnson on sex and human loving.
      ].
      In 1949 Dickinson wrote about long-term sequelae of female sexual excitement without orgasm: “Vulvo-vaginal changes in excitement. Chronic or persisting alterations due to strong, long continued and close-set repetition of excitation are the following: (1) Inflammation at mouth of duct of vulvo-vaginal gland. (2) Prominent veins in certain locations about vulva. (3) Varicosities of bulb, and of pars intermedia toward clitoris. (4) Varicosities in base of broad ligament. (5) Varicosities in upper part of broad ligament. (6) Enlargement of clitoris. (7) Labial enlargement. (8) Levator thickening and irritability, followed by relaxation. (9) Hymen gaping, worn, ironed out or disappearing. (10) Chronic bladder base congestion, (11) Hemorrhoids.” [
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ]. The evidence for these statements is unclear but physiologically, female sexual satisfaction is based on orgasm and resolution [
      • Masters W.H.
      • Johnson V.E.
      Human sexual response.
      ,
      • Masters W.H.
      • Johnson V.E.
      • Kolodny R.C.
      Masters and Johnson on sex and human loving.
      ,
      • Puppo V.
      • Mannucci A.
      • Abdulcadir J.
      • Puppo G.
      Clarifications about some theories in sexology and about a correct sexual terminology.
      ].

      5. Conclusion

      Dickinson wrote: “Exalting vaginal orgasm while decrying clitoris satisfaction is found to beget much frustration. Orgasm is orgasm, however achieved” [
      • Dickinson R.L.
      Atlas of human sex anatomy.
      ]. Masters and Johnson agreed. “Physiologically all female orgasms follow the same reflex response patterns, no matter what the source of sexual stimulation. An orgasm that comes from rubbing the clitoris cannot be distinguished physiologically from one that comes from intercourse or breast stimulation alone” [
      • Masters W.H.
      • Johnson V.E.
      Human sexual response.
      ,
      • Masters W.H.
      • Johnson V.E.
      • Kolodny R.C.
      Masters and Johnson on sex and human loving.
      ]. Clitoral/vaginal/uterine orgasm, “G/A/C/U spot” orgasm, are terms that should not be used by sexologists, women and mass-media [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Puppo V.
      • Mannucci A.
      • Abdulcadir J.
      • Puppo G.
      Clarifications about some theories in sexology and about a correct sexual terminology.
      ].
      The female orgasm (and resolution) should be a normal phase of the sexual response cycle, which is possible to achieve by all healthy women with effective sexual stimulation, and “aging alone does not diminish female sexual interest or the potential of the woman to be sexually responsive if her general health is good” [
      • Masters W.H.
      • Johnson V.E.
      Human sexual response.
      ,
      • Masters W.H.
      • Johnson V.E.
      • Kolodny R.C.
      Masters and Johnson on sex and human loving.
      ].
      Clitoral bulbs, clitoral or clitoris–urethrovaginal complex, urethrovaginal space, periurethral glans, genitosensory component of the vagus nerve, and G-spot, are terms used by some sexologists but they are not accepted or shared by experts in human anatomy [
      • Williams P.L.
      • Bannister L.H.
      • Berry M.M.
      • et al.
      Gray's anatomy.
      ,
      • Testut L.
      • Latarjet A.
      Traité d’Anatomie Humaine, neuvième édition.
      ,
      • Puppo V.
      • Mannucci A.
      • Abdulcadir J.
      • Puppo G.
      Clarifications about some theories in sexology and about a correct sexual terminology.
      ]. Findings from the disciplines of embryology, anatomy, and physiology about the congestive structures of women's urogenital areas, which are homologous to men's erectile organs, should form the basis of the discourse about the biological basis women's orgasm.

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