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Section F Hormonal Regulation of Reproductive Behavior
Sex Hormones and Other Variables in Human Eroticism
John Money, Ph.D.
Associate Professor Of Medical Psychology And Pediatrics, The Johns Hopkins University, Baltimore
- 1 Sex Hormones and Other Variables in Human Eroticism
- 1.1 I. Introduction
- 1.2 II. Empiric Endocrine Restrictions
- 1.3 III. Morphologic and Behavioral Maturation
- 1.4 V. Cognitional Rehearsals
- 1.5 VII. Concluding Remarks
- 1.6 VIII. References
From the Departments of Psychiatry and Pediatrics, The Johns Hopkins University.
The research program from which many of the data discussed in this chapter were derived has been supported since 1951 bj' a grant from the Josiah Macy, Jr. Foundation. Since 1957 it has been supported also by research grant M-1557 from the National Institute of Mental Health of the National Institutes of Health. The research, formerly under the aegis of Dr. John C. Whitehorn, Professor Emeritus of Psychiatry, is dependent on the unfailing good will of Dr. Lawson Wilkins, Professor of Pediatrics, whose endocrine clinic has provided an indispensable wealth of clinical material.
Investigators of the role of gonadal hormones in human eroticism are more restricted by the mores of our culture than are students in other areas of biologic and medical research. Partly for this reason, our knowledge of the relationship between hormones and reproductive behavior in man is not as advanced as it is for the lower animals generally. There is, however, one compensating factor. Unlike studies of lower mammals which must be on the basis of gestural and nonverbal signs alone, the study of human eroticism is not limited in this way. Eroticism- in man is a complex of signs and signals including physiosomatic signs from the reproductive system, behavioral gestures of premating and mating endeavors, and language messages about sexual sensations, imagery, and expectancy. In the study of human eroticism, therefore, linguistic, verbal signs may be added to the gestural ones, and when interviews are conducted with sufficient skill, objectivity, and regard for the importance of suitable controls, information can be obtained which supplements invaluably that gathered from naturalistic observations and experiments on lower mammals.
The availability of pathologic subjects
" Conceptually, eroticism is a more inclusive term than libido in its traditional sense of sexual desire. In Freudian or Jungian usage, however, libido is vastly more comprehensive a concept than eroticism. To avoid ambiguity the term libido is little used in this chapter.
Sex hormone anomalies and numbers of patients
Unmatured boys aged 16 or older, not dwarfed, with hypogonadisna and sex-hormone deficiency. ... 11 Unmatured girls over 16, dwarfed in stature, with gonadal aplasia and sex-hormone deficiency:
(a) Male sex chromatin 7
(b) Female sex chromatin 2
Woman, sex-hormone deficiency secondary to pituitary deficiency 1
Female hyperadrenocortical hermaphrodites precociously virilized, studied when feminized on
(a) Between ages 8 and 14 years 9
(b) Between ages 15 and 53 years 12
Boys with idiopathic sexual precocity 9
Girls with idiopathic sexual precocity 16
Boys with precocious puberty of hyperadrenocortical onset, not arrested by treatment 8
Various anatomic anomalies affecting eroticism and numbers of patients
Paraplegic men 1
Clitorectomized hermaphrodites 9
Men with penectomy 4
Woman with radical vulvectomy 1
has been important for the furtherance of investigation and development of concepts in human eroticism. It is a well known principle that pathology, by its very exaggerations, helps to sharpen one's acuity for perception of the normal.
On the basis of these principles, this chapter presents material on human eroticism. A part is a review of the many published reports. Much of it, on the other hand, was obtained from people who had some pathologic condition that affected their sexual functioning. INIore than 100 were patients of a variety of ages and types of hermaphroditism and for whom a psychologic study had been completed, the same patients whose histories were the basis of Chapter 23 on the ontogenesis of sexual behavior by the Hampsons. Full details of the endocrine conditions in these patients will be found in Wilkins (1957), the method of psychologic study in Money (1957). Other patients were paraplegics and quadriplegics whose study was made possible through the kindness of Dr. John Neustadt at Baltimore City Hospitals and Dr. Alexander Dowling, Medical Director of Maryland State Chronic Disease Hospitals. The numbers of patients in the different groups are shown in Table 22.1.
Historically, the medical management of hermaphroditism followed the common assumption that masculine eroticism is an attribute of male-hormone functioning and female eroticism of female-hormone functioning. This view receives considerable support in the phenomena of reproductive or sexual behavior in subhuman mammals (see chapter by Young). It was not until 1945 that the significance of hermaphroditism for psychosexual theory was given serious consicleration. In that year Ellis published a review of 84 case reports in the periodical literature. In these reports psychologic information was for the most part anecdotal rather than systematically collected. An important exception was a case study in 1942 by Finesinger, Meigs and Sulkowitch, which still remains the sole example of psychoanalysis applied to an hermaphrodite. The patient was a male hermaphrodite, aged 17, boyish in build and hormonally unfeminized, who had always lived as a girl. Psychoanalytic sessions were conducted daily for 6 months, as a matter of research interest. In a personal communication. Finesinger confirmed that, without exception, the patient's free associations were typically those of a teen-aged girl.
In 1950 Money began a series of hermaphroditic studies^ that have since been expanded in a continuous program in col ^ Money, 1952, 1955; Money, Hampson and Hampson, 1955a, 1955b, 1956, 1957; Hampson, 1955; Hampson, Hampson and Money, 1955; Hampson, Money and Hampson, 1956.
laboration originally with Dr. Joan G. Hampson and latterly with Dr. John L. Hampson and Dr. John W. Shaffer also. Seven variables of sex were examined for incongruities that might exist among them in hermaphroditic patients: (1) nuclear sex, i.e., the sex-chromatin pattern of cell nuclei (Barr, 1957), or the actual chromosome count (Ferguson-Smith, 1960) ; (2) gonadal sex; (3) hormonal sex and pubertal feminization or virilization; (4) the internal accessory reproductive structures ; (5) external genital morphology; (6) sex of assignment and rearing; (7) gender role and sexual orientation established while growing up.
From the evidence these investigators collected, it became quite clear that, among other incongruities in hermaphrodites, the gender role and erotic orientation as man or woman may be independent of hormonal sex. Full elucidation of the relationship between functioning of the sex hormones and of eroticism in men and women is far from complete, however. It is the aim of this chapter to contribute to the elucidation of this problem.
II. Empiric Endocrine Restrictions
Unraveling of erotic-hormonal relationshii)s is subject to three restrictions imposed by the biochemical nature of the sex hormones. The first pertains to study of the effects of hormone administration. It arises from the possible biotransformation of androgens into estrogens, in the male, and of gestagens into androgens in the female. The possibility of such transformations restricts what may be inferred from the results of experimental administration of the sex hormones. It cannot, for instance, be assumed after injection of a specific hormone, say testosterone, that the observed sequelae are direct effects of the testosterone. Some of the hormonal substance may have been converted into an estrogen after its absorption into the blood. Thus, conversion of androgen into estrogen may explain duct proliferation in the breasts (gynecomastia) of a male castrate treated with large doses of testosterone, or in some untreated rapidly maturing, ordinary adolescent males.
The second restriction in the unraveling of erotic-hormonal relationships pertains to determinations of blood and urinary hormone levels. It arises from the fact that the sex hormones, in vivo, are unstable chemical compounds. From a given androgenic steroid, related forms may be derived by molecular rearrangement, addition, or reduction (see chapter by Villee). These changes appear to take place constantly in the biosynthesis and metabolism of androgen in the living body. The same holds true for estrogens and gestagens.
It is impossible at the present stage of scientific knowledge to identify the full variety of related forms of the sex hormones that are functionally active in the body, or to estimate their quantity. Quantitative assessments of androgens or estrogens in routine laboratory practice are not assessments of the hormones actually circulating in the blood and stimulating the cells of the various tissues and organs. Determinations of hormone levels in blood and urine are, in general, determinations of derivative forms and metabolic end-products of the actual compounds active in the body, some of them biologically inert.
Biochemical instability of the sex hormones does not mean that laboratory hormonal measures are useless. What it does mean is that the level of biologically active circulating hormone must be inferred from the laboratory findings. The inference may not be made unless appropriate norms and criteria have been empirically established. In individual cases the further significance of laboratory findings can be pinned down only when they are appraised within the context of clinical and other findings about the patient or subject concerned. To illustrate, an above-average level of urinary ketosteroids in a male may be without specific significance. It would be highly significant, however, if the sample came from a patient who showed other signs such as tumorous enlargement of the adrenal gland or testis, or, in a younger patient, signs of precocious virilism.
The third restriction in unraveling erotic hormonal relationships pertains to bioassays as they are routinely performed. A sex-hormone compound which is judged biologically active, and not inert, is so judged on the basis of its stimulating effect on appropriate morphologic sexual characteristics in test animals. Thus, a biologically active androgen is one that stimulates comb growth in the capon, or prostatic and seminal vesicle secretion in the castrate or immature rat. Similarly, estrogen induces changes in the vaginal epithelium in adult spayed mice or maturation of the uterus in juvenile mice. Gestagens induce the appropriate gestational changes in the endometrium of the uterus. It does not follow that a specified biologically active compound will be equally potent in its stimulating effect on all secondary sexual characteristics. For example, in hypopituitary, eunuchoid youths testosterone induces a fair degree of virilization, except for growth of the beard, although in castrated boys the effect of testosterone on beard growth is excellent. Within a species, some individuals are more sensitive, some more resistant to a prescribed amount of hormone. Within an individual, some target organs are more sensitive or more resistant than others. Examples of such variations are mentioned from time to time in the ensuing text. Because of such variations, there is the possibility that an otherwise biologically active synthetic hormone may fail or partially fail to evoke signs of eroticism, or that it may evoke seemingly incongruous effects.
III. Morphologic and Behavioral Maturation
Androgens are specific for bringing about adult growth of the genital tract in males. Estrogens are similarly specific for females. Further, androgens are antagonistic to feminized genital maturation in females, the classic example being virilizing hyperadrenocorticism in girls.'* These girls, hermaphroditic if born with their adrenal dysfunction, have a precocious and exclusively virilizing puberty. Their ovaries fail to mature and the clitoris becomes hypertrophied to resemble a penis in size. Estrogens are antagonistic to virilized genital maturation in males. They inhibit androgen production in the testes and thus simulate castration. Simultaneously, they stimulate enlargement of the breasts.
Hormonal failure of the gonads, whether through castration or hypofunction of the
- These actions of androgens and estrogens are
described and discussed fully in the chapter by Burns.
gonadal endocrine cells, results in persistence of infantile sexual and body morphology. Patients who reach the middle teenage, or beyond, with sex hormone failure untreated, demonstrate how far-reaching is the morphologic function of the sex hormones.
Teen-agers with morphologically prepubertal bodies are invariably identified by strangers as younger than their age. Old friends, family, and social intimates fall with perilous ease into the custom of treating them as preadolescent juveniles. It is difficult for them to remain persona grata with their age-mates. The boys are worse off than the girls, for it is more feasible for a girl with facial make-up and built-in "falsies" to disguise some of the stigmata of sexual infantilism than it is for a beardless, high-voiced boy. Many girls as well as boys, however, fall by the wayside, physically too immature for their own age group, yet socially too mature for younger children of similar physique. Among the 10 girls and 11 boys of Table 22.1 who were over 16 and still physically unmatured, only 1 girl and 1 boy had a history of a teen-age social and dating life that approached the norm for their group.
When a teen-ager with sex-hormonal failure is responded to as a juvenile from all quarters, the chances are very great that he or she will respond as a juvenile and lag behind in psychologic and behavioral development as he or she gets older.^ The longer this lag persists, the more difficult becomes the problem of social and psychologic adjustment as a maturing and mature adult, after hormonal substitution therapy has been instituted.*' Adolescent psychologic growth cannot be properly achieved in the absence of adolescent physical growth, but it also cannot be properly achieved in the absence of age-mates who are also in the adolescent growth phase. The early teen-years appear
^ This lag is seen even more acutely in dwarfed children whose size belies their age. The converse, an acceleration of psychologic and behavioral development, is possible in children with precocious physical and sexual maturation.
" The same kind of adjustment problem has been seen almost without exception in the post-teen-age women with virilized hyperadrenocortical hermaphroditism, after they have begun to feminize in body morphology following institution of cortisone therapy.
to be the critical period for optimal, allround adolescent development.
The behavioral immaturity that matches sexual immaturity is not necessarily reversible. It may establish itself as a chronic handicap, especially if puberty is too long delayed into and beyond the late teens. Chronic behavioral immaturity then defeats the success of hormonal substitution therapy by militating against success in mating and in adult erotic relationships.
Androgens and estrogens both mature the body, the one in a virilizing, the other in a feminizing way. Androgens, in addition, have a strong positive influence on nitrogen metabolism and so on muscular strength and energy. This muscular effect was particularly noted by two hypogonadal males in the series who worked as manual laborers. Before treatment began they made euphemistic statements about their fatigue threshold. After treatment they were able to make a comparison and so to realize how easily they had tired formerly. The converse effect of androgen suppression producing a diminution of muscular strength and energy is well observed in hyperadrenocortical females when their androgen excesses are first suppressed by cortisone therapy. Two patients have remarked on this phenomenon.
The androgenic effect of stepped-up nitrogen metabolism and increased general muscular size and strength (Kochakian, 1946; Kochakian and Tillotson, 1957) does not seem to have any direct effect on eroticism. The hypogonadal men (Table 22.1) on androgen-substitution therapy became sexually more active and participative, but not because they had formerly been in^'alids too weak to participate. They did not become more violent, assaultive, or aggressive, even though they had more muscle power at their disposal.
Rather than being spread generally over the nervous system and musculature, the specifically erotic action of androgen, insofar as it can be identified, seems to be anatomically localized. In particular, the undeveloped penis or clitoris responds to initial androgen administration with extensive dilation of its vasculature and with growth in size. Thereafter, maintenance of an erection by complete engorgement of the organ with blood is facilitated by androgen. Tumescence of the penis can occur in the absence of androgen, but the erection is generally not complete and long lasting.
The seminal vesicles and prostate remain immature in a male until they are stimulated into growth by androgen. Thereafter, their fluids, which form the medium of sperm conveyance and nourishment, are secreted only so long as there is sufficient androgen in the body.
IV. Eroticism and the Supply of Sex Hormone
A. PREPUBERTAL AND POSTPUBERTAL CASTRATION
Much of the literature bearing on this subject was reviewed by Kinsey, Pomeroy, ]\Iartin and Gebhard (1953). Their summary of the effects of gonadectomy performed prepubertally, as well as the summaries of their predecessors (Lipschiitz, 1924; Commins and Stone, 1932; Beach, 1948), contains abundant evidence for the importance of gonadal hormones in the development of sexual responsiveness in man. The repeated emphasis by Kinsey, Pomeroy, Martin and Gebhard that damage to other endocrine glands such as the pituitary, thyroid, adrenals, and islands of Langerhans, may have similar disastrous effects on the development of the capacity to respond sexually does not weaken the case for the specific role of the gonadal hormones in such development. Debilitating effects on vital functions including reproduction do follow ablation of these other glands, but when the gonads are removed the damaging effects, although general, are nevertheless most sharply focused on reproductive function. The damage is clearly evident in the chronic infantilism that follows prepubertal castration.
The results of postpubertal castration are more variable as Bremer's excellent study (1959) and the reviews by Tauber (1940), Engle (1942), Beach (1948), and Kinsey, Pomeroy, Martin and Gebhard (1953) indicate, some males and females report that the retention of sexual capacities and responsiveness is ostensibly complete following castration, others that decrease or total loss occurs. The most serious obstacle to clarification is the inadequacy of detail in most of the reports available for evaluation, although some reports are exemplary, like those of Foss (1937) and Hamilton (1943). Another part of the difficulty is that the effects of gonadectomy may vary with age, being less marked in older than younger men, with respect to behavior as well as to physical characters. More important, as many writers have emphasized, is the dominance of cognitional and learned factors in the overt expression of eroticism in man. These may include type and extent of prior sexual experience, preference for a given partner, freedom from fear of pregnancy, the degree of respect and affection in the marital relationship, the background of cultural expectancy regarding castration, and so forth.
In efforts to ascertain the effects of castration, as well as in other efforts to determine the role of the gonads in erotic functioning, past experience is always a complicating factor in any type of subject under study. As lower animals and presumably man develop and mature, psychologic or experiential as well as genie factors mold the substrate on which the hormones act in such a way that diversity rather than similarity in response is seen during adulthood (see chapter on hormones by Young) . The range may extend from absence of a response to intense reactions verging on the pathologic in their deviation from what is typical for the species. For the investigator who is expecting a uniformity of response to a given quantity of a hormone, this diversity in response may mask the effect of the hormone and prompt him to minimize the degree of its effectiveness. Or, atypical patterns of behavior such as homosexuality may be seen in individuals in whom there is no evidence of gonadal pathology or of abnormality of secretory function (Kinsey, 1941 ; Sevringhaus and Chornyak, 1945). Cases of this kind do not justify rejection of the concept of hormonal participation in human erotic behavior. They give emphasis instead to the importance of the character of the tissues mediating the behavior, by whatever mechanisms it is brought to expression.
Surgical castration has a functional counterpart in hypogonadism which, like castration, may be of either prepubertal or postpubertal occurrence. Unlike castration, hypogonadism may be partial, with only partial and not complete deficiency of gonadal hormones. Published reports sometimes confuse one type of patient with another, so that findings are somewhat difficult to evaluate. The difficulty is enhanced by the fact that neM^ diagnostic advances in endocrinology and nuclear sexing permit greater precision in the grouping of patients, not possible in the older studies. In general, however, there has been a consensus of opinion that erotic drive in hypogonadal and eunuchoid males of various types is likely to be heightened by androgen administration (McCullagh, IVIcCullagh and Hicken, 1933; Vest and Howard, 1938; Howard and Vest, 1939; Pratt, 1942; Heller and Nelson, 1945; Heller and Maddock, 1947). In cases of partial gonadal failure, control studies with a placebo would have validated the results more convincingly, but in cases of total gonadal failure the results of hormonal substitution therapy are quite dramatic enough to be convincing in themselves. Definitive reports on eroticism in treated hypogonadal females are lacking in the literature.
C. GONADAL FAILURE
Fresh evidence on hypogonadism was obtained from the hypogonadal patients listed in Table 22.1. These patients were completely hypogonadal, that is totally hormone deficient at the age of 16 or older, before treatment began. They were prepubertal, functional castrates.
Juveniles, who are by definition hypogonadal, play erotically together, but it is quite a different matter for a person who has reached the adult years with unmatured genitals to engage in an erotic relationship. Metaphorically speaking, this fact suggests that the first job of the sex hormones is to set the stage properties in order so that the drama of eroticism can be enacted. Support for such a view comes from the circumstance that among the hypogonadal patients entered in Table 22.1, advancing age alone did not change childhood eroticism to adulthood eroticism when sex-hormone failure persisted beyond the age of 16. There were 11 such who were candidates for androgensubstitution treatment and 10 for estrogensubstitution treatment. They evidenced a wide range of variability in their pretreatment erotic development, concomitant with such individual differences as dwarfed stature (among the girls only), inhibitory training in sexual matters, restricted opportunities for social development, and amount of close contact with dating or marriage partners.
Response to treatment may be summed up by saying that these 21 patients went through an erotic development not unlike that of normal boys and girls. It is of interest in addition that they exhibited the same wide range of variability of erotic development and activity after treatment that they had before. It is worth reporting that, among the girls with gonadal aplasia the 2 with a female sex-chromatin pattern (and 1 younger girl like them) did not respond to estrogens in a way conspicuously different from the 7 with a male chromatin pattern.
Eroticism following induced puberty turned out, then, to be not so different from eroticism following ordinary puberty in its character and in its dependence on the presence of sex hormones.
The patients requiring induction of puberty proved more instructive in another way. These individuals, if they omitted their substitution therapy for a period, were in a liosition to give information about adult eroticism in the absence of sex hormones.
D. SUBSTITUTION THERAPY DISCONTINUED
Of the 11 hypogonadal men entered in Table 22.1, there were 5 who discontinued androgen medication for 3 months or longer. In each case, the absence of androgen from the tissues made a decided difference. The most sensitive indicator was the ejaculate.
^ Dwarfed boys who also had gonadal faihiie secondary to h.ypopituitarism were not included in this study. Dwarfed girls were of necessity included, since dwarfism is a frequent accompaniment of gonadal aplasia in girls. Except for one juvenile surgical castrate, there were in the clinic files no nondwarfed girls with sex -hormone failure.
It gradually diminished in volume until no fluid was emitted. In addition, the men reported that they had fewer erections and a lessened initiative to masturbate or to make coital advances. With loss of ejaculation, they also lost erotic ejaculatory dreams. They considered that waking erotic imagery and daydreams diminished in frequency of appearance.
One may generalize and say that these men did not lose completely their erotic imagery, their erotic sensations, or their erotic actions and behavior. What did happen was that eroticism, whether in imagery, sensation, or activity, was not initiated with the same frequency as before. This failure of initiation showed up in the involuntary failure of the penis to erect or to hold an erection, and in the failure of other, more voluntary erotic actions and coordinated endeavors, as well.
The man who was married when he discontinued treatment had a good barometer of the failure of his erotic initiative, namely his wife's comments and complaints. In fact, this man, and the two others who married, found that it paid them not to become lax about their injections. They reported a slackening of erotic initiative, including erectile potency, if they delayed even a week in getting their monthlj'- injection of longacting testosterone.
The conclusion to be drawn from periods of interrupted treatment in men with sexhormone failure is that androgen is necessary, not only to induce morphologic maturity, but also for the maintenance of a well functioning eroticism.
There were four hypogonadal women who discontinued estrogen medication for 3 months or longer. One of them was off treatment for 3 months by request. She married during the first month. Two others were single and celibate. One of the latter discontinued treatment with stilbestrol for 18 months, before resuming on Premarin, because stilbestrol produced unpleasant gastric symptoms. The other single woman had been off treatment for 2 years, having discovered that the only sequel of significance was cessation of the menses. The fourth woman was divorced. She discontinued treatment after a doctor scared her about the carcinogenic dangers of estrogen. She continued to have sexual liaisons during the 5 years off treatment, although eventually she discovered that vaginal tightness and dryness due to lack of estrogen-stimulated secretions was a handicap.
The four women had been on cyclic estrogen therapy so that they menstruated on withdrawal of estrogen for a week each month. Following total withdrawal of estrogen, they ceased to menstruate. Vaginal smears showed that the vaginal mucosa underwent involutional changes as in postmenopausal women. There were no definitive reports of hot flashes or malaise typical of the climacteric, however.
The women reported nothing to indicate any change in their erotic imagery, sensations, or actions. The two who were having intercourse claimed that they reached the climax of orgasm, the same as when taking estrogen.
The evidence from these women fits in with common knowledge concerning postmenopausal disappearance of estrogen in ordinary women. Although there are exceptions, erotic imagery, sensations, and actions are not abolished, often not even lessened, in the usual course of diminished estrogen production at the menopause.
E. IMPOTENCE AND FRIGmiTY
Except in hormone-deficient patients of the hypogonadal and castrate type, impotence in men and frigidity in women are usually unresponsive to treatment with gonadal hormones (Rennie, Vest and Howard, 1939; Cree\y and Rea, 1940; Spence, 1940; Carmichael, Noonan and Kenyon, 1941 ; Kenyon, 1941; Heller and Maddock, 1947; Perloff, 1949) . An exception is that a small proportion of frigid women have responded to androgen therapy (Salmon and Geist, 1943). Impotence not due to hormonal insufficiency may be due to vascular and circulatory impairment of the genital organs. Impotence has also been found associated with diabetes mellitus (Rubin and Babbott, 1958) . Usually, impotence and frigidity not responsive to hormonal treatment are said to be psychogenic in origin, and the argument for psychogenesis is supported when the disorders are not constant but dependent on time, place, and partner.
The possibility that inherited constitutional variations among individuals play a part in impotence and frigidity is suggested by the data of Grunt and Young (1952) and Riss and Young (1954) obtained from studies of the male guinea pig. When highscore and low-score animals were castrated and injected with different quantities of testosterone propionate, the behavior that was exhibited by individual animals was similar to that displayed before gonadectomy and replacement therapy. Furthermore, the administration of large quantities of the hormone did not alter this relationship. Apparently in this species individuals have a characteristic level of responsiveness. Human males may be similar. The tissues that are generally acknowledged to be responsive to sex hormones, such as p?ripheral receptors (see below), probably do not have the same threshold of responsiveness in all persons, and these threshold differences are probably inherited. In such cases as those of impotence and frigidity due to psychic inhibition, where hormone levels are not deficient, additional, exogenous hormone has no effect on a response threshold already well primed with hormone.
F. ANDROGEN, ESTROGEN, AND EROTICISM IN MEN AND WOMEN
The relationship of estrogen to eroticism in the adult female seems, after the pubertal estrogenic function of maturing the reproductive tract and feminizing the body morphology in general, to be restricted to maintaining the lubricant secretions of the vagina preparatory to copulation. The primary estrogenic function would seem to be monitoring endometrial growth in close coordination with the gestagenic function of monitoring nidation and gestation.^
Inasmuch as maintenance of well functioning eroticism in men appears to be dependent on androgen, it would be an odd
^Benedek and Rubenstein (1942) defended the thesis that the content of the dreams and psychoanalytic free-associations of 15 women patients could be used to predict which phase their menstrual cycles had reached, as determined by vaginal smear tests. Their hypothesis has not received general acceptance (see chapter by Hampson and Hampson); the collection of additional data and a reconsideration of the problem would be appropriate.
biologic discontinuity if erotic imagery, sensations, and actions in the female of the human species should be, by contrast with estrous behavior of lower animals, independent of hormonal functioning. A tenable hypothesis is that erotic imagery, sensations, and actions are maintained well functioning in both men and women by androgens. In women, the androgens of eroticism might conceivably be of adrenal origin, or they may be derived from gestagens, or they may be of an origin as yet unknown. The most likely explanation is that these androgens are of adrenal origin. Waxenberg, Drellich and Sutherland (1959) reported an excellent study of eroticism in 29 women who had both ovaries and both adrenals removed in the treatment of breast cancer. Loss of only the ovaries and ovarian hormones had no definite adverse effect on sexual drive, activity, and response, but all three were diminished or abolished in most of the women after their adrenals also had been removed. The adrenals secrete some estrogen, but larger amounts of androgen. The authors concluded that the loss of adrenal androgens was the responsible factor in the women's lessened or abolished eroticism.
Other evidence can be marshaled to support the hypothesis that androgen is the hormone of eroticism in men and women. Many women for whom androgen therapy is prescribed report an increase of sexual desire as a side-effect (Shorr, Papanicolaou and Stimmel, 1938; Salmon, 1941; Greenblatt, Mortara and Torpin, 1942; Greenblatt, 1943; Salmon and Geist, 1943; Foss, 1951; Kupperman and Studdiford, 1953; Dorfman and Shipley, 1956). In the clinical lore of urology, conversely, many men for whom estrogen is prescribed report great diminution or total abolition of sexual desire and activity (Footc, 1944; Paschkis and Rakoff, 1950).
The role of gonadal hormones in stimulating eroticism in males and females is discussed by Carter, Cohen and Shorr (1947) in a review of the use of androgens in women and by Perloff (1949) . It is noted in both articles (1) that the return of normal libido in certain menopausal women under treatment with estrogens may perhaps be related to increased vascularity and epithelial proliferation of the genital tract, and (2) that the increased libido in women who receive testosterone occurs coincidentally with hypertrophy of the clitoris. From these observations, the hypothesis follows that the hormones, especially androgens, although not the exclusive basis of libidinous urge which is multidetermined, may influence the libido by affecting end-organ sensitivity.
Whatever its nature, the androgen-eroticism relationship is far from being a simple linear correlation. There are irregularities that cannot at present be explained. Such irregularities show up in the different reactions in hyperadrenocortical female hermaphrodites before and after their gross excess of adrenal androgens is low^ered by cortisone therapy.
G. Hyperadrenocortical Diversities
There were 21 hyperadrenocortical, hermaphroditic females in the larger series in Table 22.1 who satisfied the triple conditions: (a) they had been brought up and lived as girls and women; (b) they were over the age of 8 and, therefore, precociously virilized before cortisone treatment was instituted; (c) they were interviewed after (in some instances before, also) their virilism had been suppressed and their feminization established by cortisone treatment. Among the 21, 9 were hormonally treated and psychologically studied before the age of 14; 12 were between the ages of 15 and 53.
Erotically, 4 of these 21 patients were noteworthy as follows. One of the younger patients suffered from painful priapism of her enlarged clitoris and, postsurgically, from persistent erections of its stump; the clitoral stump became flaccid and unbothersome after cortisone therapy. Two of the older patients manifested extreme hypereroticism, including compulsive copulatory episodes, before cortisone treatment. Although they reported mild lowering of erotic drive after treatment, they continued to rate higher than all but 1 other of the 21 in posttreatment erotic participation. There was one older patient who showed no evidence of erotic activity, and claimed no erotic sensations or inclinations either before or after the cortisone treatment. Her absence of eroticism is unique in this series of patients and remains a puzzle without explanation.
Between the extremes of hypereroticism and hypo-eroticism, the majority of the patients showed a more moderate eroticism. In the majority group, comprising 17 patients, 8 stated that they had fewer clitoralstump erections or that they masturbated less frequently after cortisone treatment. In the interviews with the other 9, no evidence of change was obtained, but the possibility of change could not be retrospectively excluded with certainty.
None of the 21 patients gave evidence of loss or change of sensitivity of erotic zones, other than the clitoral, while their hormonal feminization was maintained by cortisone treatment, except that sporadically the breasts were reported more sensitive. It is noteworthy that none reported a posttreatment cessation of erotic sensitivity in the clitoral zone — only of erectile autonomy and hypersensitivity of the clitoris, or of its amputated stump. What they lost, therefore, was that autonomous initiatory eroticism of the phallus which seems to be so basic in the eroticism of men. The women were all unequivocally pleased to be relieved of clitoral hypersensitivity; it was the pleasure of being able to feel like a normal woman, several of them explained. In some cases, the clitoris had been amputated in early childhood. Only one of the older patients elected against, and none regretted a decision in favor of this feminizing surgical procedure.
H. Eroticism in Precocious Puberty
An unusual group of children of special interest in the study of childhood eroticism are those with precocious puberty. In children with this diagnosis, the sex hormones function prematurely in relation to the scope and maturity of cognitional functioning as well as to the variety and extent of learning and behavior (Doe-Kulmann and Stone, 1927; Keene and Stone, 1937; Werner, Spector, Vitt, Ross and Anderson, 1942).
In sexual precocity of the idiopathic type, the children are anatomically and physiologically normal in all respects except that
the pubertal alarm clock has, so to speak, sounded too early. With respect to the 25 children with idiopathic sexual precocity (Table 22.1), it can only be said that they showed the same range of variation between sexual expression — childhood sex play and masturbation — and sexual inhibition and self-discipline as one might ordinarily find in a randomly assembled group of sexually immature children matched for age and social background. At least, it would be very easy to find nonprecocious children with matching sexual behavior. The frequency of masturbation seemed increased in some of the boys with idiopathic sexual precocity, as a direct function of their ability to ejaculate. Others who did not masturbate had sleep emissions. In none of the children did premature hormonal function, ipso facto, lead to premature attempts at mating. As in normal children, the latter appears only when it is encouraged by social experiences and opportunities to learn.»
V. Cognitional Rehearsals
Erotic thought and imagery, fantasies and dreams, may all be referred to, for want of a better generic term, as cognitional rehearsals. They may be on-the-spot responses to perceptual images-^sights, sounds, smells, touches, tastes — pertaining to sex. The sexy blonde and the lecherous thoughts of boys on the corner as she l)asses are a stereotyped example. Verbal or graphic reports at second hand may be stimuli that trigger off the derivative sexual imagery and ideas. Erotic thoughts, imagery, and fantasy are not necessarily followed by sexual acts. The rehearsal in thought or in dream may be an entity unto itself, carried no further. Moreover, the rehearsal may also be far removed in time from the day when it will be put into performance.
"A recent report (Gerall, 1958) containing a review of the relevant literature is of interest for the point made in this section. In the guinea pig, but not in the rat, the administration of testosterone propionate to castrated young animals did not advance the appearance of the copulatory pattern or ejaculatory response. In both species injections of te.stosterone propionate produced greater than normal weights of the seminal vesicles, prostate, and coagulating gland. On the other hand, a difference between the species is reflected in the amount of secretory activity. The level of fructose and citric acid in the prostate gland and seminal vesicles of rats injected with testosterone propionate was several times greater than that in normal rats. In the guinea pig testosterone propionate did not produce precocious secretion of these substances.
Erotic rehearsals in dreams and daydreams appear to be spontaneously initiated in many instances. The stimulus may conceivably be a proprioceptive, somesthetic one from the pelvic reproductive system. A memory stimulus may also be the activator of erotic dreams and daydreams, its arousal not manifestly dependent on perceptual signals from the pelvis or elsewhere. The degree of dependence of these rehearsals on gonadal secretions will be considered here.
B. CHILDHOOD REHEARSALS
It is a matter of everyday knowledge that young children, hormonally immature, have thoughts and fantasies, often disclosed in play, that are cognitional rehearsals of eroticism in its broadest sense. Kindergarten children play their way through romantic flirtations, love affairs, and promises to marry when they grow up. Some of these affairs are developed in elaborate makebelieve detail and are viable for months. Explicit genital and copulatory imagery is not a necessary or regular part of these romantic rehearsals. Genital play, however, including copulatory play, is not alien to the play repertory of young children. The frequency of its occurrence seems to depend largely on local cultural and social sanctions. The field studies of cultural anthropologists among ethnic groups geographically as far apart as Africa, Oceana, and the American continents have shown that genital play of all types including copulatory play may be regarded and permitted or encouraged as an expected part of the behavior of normal childhood (Chapter by ]\Iead; Ford and Beach, 1951).
Here is a brief report on the cognitional rehearsals in the 14 of the 25 children in Table 22.1 (12 girls and 2 boys) who had sexually matured physically at or before the age of 8, who were available for psychologic study between the ages of 5 and 12, and who were not too young to be coherent informants.
Among the 14, the occurrence and reporting of sexual dreams and daydreams was variable. Five of the 12 girls said that they had daydreams of boy-friends and romance, and recounted examples — all very stereotyped, Prince-Charming adventures — that excluded genital sex. Romantic sleepdreams were characterized by the same stereotypy in the 2 girls reporting them. None of the girls gave any evidence of orgasm dreams. In fact, there was no evidence that any of them had experienced orgasm, asleep or awake.
Of the 2 older boys, 1 could not recall dreams or daydreams, although he had seen ejaculation stains on his pajamas. He was seen only once, at the age of 7. The other boy (Money and Hampson, 1955) was seen annually between the ages of 5 and 11. He had florid sexy dreams and daydreams of seeing and kissing naked women that were more freely reported before he was 6 and again after he was 9 years old than during the intervening period. He could not produce a specimen of ejaculate at the age of 7. He was 10 before he gave unequivocal accounts of ejaculation. At that time, the erotic imagery of wet dreams and masturbation fantasies was primarily pictorial and represented couples in various copulatory poses and actions.
The data of these too few male cases can be augmented by adding 8 cases (Table 22.1) of precocity in boys that began as hyperadrenocortical and then became testicular precocity after adrenal overactivity was corrected. It is usual for maturational precocity to continue, as in these cases, once a certain level of somatic maturity has been achieved, even after the original stimulus to precocity has been removed.
The 8 boys were all well advanced into puberty by the age of 8. Two of them reported emissions. These 2 had erotic daydreams and, at night, wet dreams, with imagery of erotic play with girls; the older of tlie 2, aged 8, also had imagery of intercourse. Two other boys told of daydreams of kissing and petting with girls, accompanied by erection. A fifth boy reported nonerotic mystery and adventure fantasies accompanied by erection.
Reviewing now the histories of the 22 older children with precocious hormonal puberty, it is noteworthy that like their erotic play, the thematic content of their dreams and daydreams directly reflected not the hormonal age but the social age that each child had achieved (see also Rafferty and Stein, 1958) . The social age agreed with or was in advance or arrears of the birthday age, dependent on the variety and extent of life experiences the child had encountered and, in turn, transacted. Premature puberty made a difference not in the content and imagery of erotic play, dreams and daydreams, but in the frequency of their occurrence.
C. HYPOGONAD.\L ADULTS AND CASTRATES
Orgasm dreams are ordinarily regarded as postpubertal phenomena exclusively. None of the hypogonadal patients of both sexes reported in Table 22.1 reported orgasm dreams before puberty was induced with hormonal substitution therapy. Some of them, however, gave examples of sexy daydreams and fantasies. Hormonal therapy did not unconditionally guarantee the onset or the increased occurrence of erotic dreams or daydreams in these hypogonadal patients, although it had such an effect in most instances.
VI. Sex Hormones, Neural Tissues, and Behavior
In lower animals and presumably in man gonadal hormones act on many target organs or tissues of the body and bring to expression the responses characteristic of those tissues: growth, secretions, motility, sensitization for implantation, alterations in behavior, and others, depending on the tissue. Each tissue in its way, therefore, is with the hormones a coordinate of sexual function. In the case of behavior, neural tissues are in this category. Of these, peripheral receptors have been and are the object of especial attention, for as noted above, the possibility exists that, when sensitized by gonadal hormones, they have a role in the mediation of behavior (see also chapter by Young). For the elucidation of this problem, the verbal reports of human subjects in whom there has been deafferentation of the genital and other erotic areas of the body have provided information which cannot be obtained from lower animals.
In paraplegics, and following penectomy and vulvectomy cognitional eroticism can be studied independently of sensations generated in the genitopelvic area.
A. Paraplegic Cognitional Eroticism
The complete independence of erotic cognition and pelvic signals is demonstrated in the case of paraplegia. The physiologic state in such patients is variable, as is the level of gonadal hormone secretion (Cooper, Rynearson, MacCarty and Power, 1950; Cooper and Hoen, 1952; Bors, Engle, Rosencjuist and Holliger, 1950) . It was found that approximately two-thirds of 500 paraplegic and cjuadriplegic patients were capable of achieving erection, and of these one-third had successful intromission (Talbot, 1955; Zeitlin, Cottrell and Lloyd, 1957), without, however, the gratification obtained before the injury. The dream eroticism of 34 patients was reported by Bors, Engle, Rosenquist and Holliger. Ten patients did not recall any sexual dreams, 14 remembered dreams lacking orgasm, and 10 had complete dry dreams, i.e., including the feelings of orgasm, but without erection or ejaculation.
The independence of erotic cognition and pelvic signals is further demonstrated by an interview with a paraplegic patient (Table 22.1 ) . He was 19 years old and had a paraplegia of 10 months duration. Spinal destruction was at C4 and C5. Loss of feeling and motor function was absolute below the nipples.
In the interview, the man described with exceptional clarity his feelings when his girl-friend kissed him. It seemed as though his penis would be getting into an erection and throbbing, he said. "But when I would look down, nothing was there — it was all in my mind, or something". The phenomenon was similar when he had sexy daydreams, he said. Then he went on, again documenting the discontinuity of cognitional and genitopelvic eroticism. He would think of a time when he had actually had intercourse, he said, as he might formerly have done in a masturbation fantasy. Again he would get a feeling that something was building up pelvically, whereas, in fact, nothing was happening.
The patient introduced the topic of sleeping dreams in which he dreamed of intercourse and orgasm. The dream-orgasm was not accompanied by an erection or ejaculation, but the feeling was the same as it had been in an ordinary nocturnal emission, the man said.
The sexy thoughts and fantasies of a paraplegic patient may serve the literal purpose of rehearsals and lead to attempts at intercourse. Then one has the exceptional phenomenon of cognitional eroticism occurring simultaneously with involuntary, reflex action of the pelvic genitalia, the only possible connection between the two being via the eyes and hands.
So much for paraplegics, the accidental experimental subjects in whom the genital tactile receptors remain intact while their distant connections with the brain are broken. Turn now to the obverse case, patients whose external genitals have been damaged or resected so that, whatever the extent of neural damage, it is local only.
B. Clitorectomy in Hermaphrodites
Genital resection, in the course of surgical reconstruction, is desirable in the management of certain cases of hermaphroditism (Jones and Scott, 1958) . An enlarged clitoris of penis-like proj^ortions is incompatible with complete femininity in the experience of the majority of hermaphrodites living as girls and women. They desire that their masculinized clitoris be amputated.
Such an operation is usually performed on hyperadrenocortical hermaphrodites if they are to be reared as girls, or if they are already psychosexually established as females. For present purposes, the older patients with this adrenogenital type of hermaphroditism (Table 22.1) are of particular interest. Before the introduction of cortisone therapy in 1950, these patients had undergone a precocious but virilizing jniberty in early childhood, under the influence of an excessive supply of adrenal androgen. Under the impact of these androgens, the enlarged, unamputated clitoris was erotically very sensitive. At the same time, the vagina remained unestrogenized and immature. In most instances the vaginal opening was congenitally misplaced in a urogenital sinus and needed surgical reconstruction.
Among 17 older potential informants in this category of hyperadrenocortical hermaphrodites, there were 9 who satisfied the triple condition: (a) they had been brought up as girls and lived as women; (b) they were older than 16 at the time of reporting on eroticism; (c) they had been clitorectomized and reported on their postsurgical eroticism.
The breakdown of findings in these 9 cases was as follows: in 4 the data indicated the patient had not experienced orgasm; in 5 the evidence was that the patient had experienced orgasm, in 2 masturbatory, in 3 coital.
The 9 women were receiving cortisone when they reported on eroticism. There was no evidence, however, to suggest that orgasm might have been lost following clitorectomy only to reappear after hormonal feminization had been established under cortisone therapy, or that orgasm was lost under the influence of cortisone.
There is no ready explanation for the lack of orgasm in the 4 patients where such appeared to be the case. So far as could be ascertained from unstandardized operative notes, lack of orgasm did not correlate with the amount of clitoral tissue removed, or with any other surgical factor.
The point of these data on orgasm and clitorectomy is not, however, that some clitorectomized patients did not experience orgasm. On the contrary, the point is that capacity for orgasm proved compatible with clitorectomy and surgical feminization of the genitalia in some of these patients. Erotically sensitive though it had been, the main body of the clitoris, including the glans of the clitoris, was dispensable with respect to orgasm.
Four patients with an amputated penis (Table 22.1) were available for psychologic study. All reported retention of the capacity for orgasm.
One of the patients lost his penis at the age of 30. Ten years later the prostate was removed and the urethral orifice was relocated near the anus, requiring urination from a sitting position. He was interviewed psychologically at the age of 42. After his injury, he frequently had erection of his penile stump in response to seeing or joking with an attractive woman. In sexual relations, mutual genital friction sufficient to induce mutual orgasm was produced.
Another patient reported loss of the capacity for ejaculation, although he retained the sensation of orgasm in dreams. His was a case of penectomy due to malignancy. There were metastases to both groins requiring radical bilateral groin dissection. The man was 56 when interviewed and had been in the hospital a year. He said he continued to feel sexually aroused, the feelings being the same as they always had been, as for example when his wife came to visit him.
It is roughly accurate to say that the female equivalent of penectomy is radical vulvectomy. An interview was obtained with one patient who had this operation. Fifteen years earlier she had had an epidermoid carcinoma of the vulva which entailed complete resection of the clitoris, labia majora, labia minora, and the mucosa of the introitus. The patient was 45 years old and still premenopausal when she returned, through the courtesy of Dr. Howard W. Jones, Jr., for an interview. She was very frank and spontaneous about her sex life postoperatively. She said that, although she was still self-conscious about her genital appearance, so far as sexual feeling was concerned there was no difference. The feeling or orgasm occurred sometimes in dreams, she said, as well as in coitus.
Summarizing, it is seen that from patients who have undergone extensive surgical resection of the genitals, erotic arousal can be initiated and carried to the completion of orgasm despite the loss of large zones of erotic sensory tissue, including the vulva or the penis itself. From paraplegics one learns that cognitional arousal is possible in subjects in whom the genital tactile receptors are intact while their connections with the brain are broken. From eunuchs and hypogonadal patients (their detailed case illustrations omitted at this point) one learns that erotic arousal and climax can sometimes occur despite hormonal deficiency that in men causes absence of seminal fluid. Thus, among the coordinates of sexual function there are three: local genital surfaces, the brain, the hormones, any one of which can fail in its contribution without total destruction of sexual function. No one of the three can be said to be indispensable more than the others, except insofar as the hormones are indispensable to fertility. Nonetheless, it is evident that loss of any one of the three constituents is an immense handicap to effective sexual functioning.
VII. Concluding Remarks
The sex hormones, it appears, have no direct effect on the direction or content of erotic inclination in the human species. These are assumed to be experientially determined. The importance of experiential factors is nowhere better revealed than in iiermaphrodites.
To refer back to the beginning of the chajiter, the gender role and gender orientation of hermaphrodites became established in accordance with the sex of assignment and rearing. One may say that all through childhood these people accumulated a continuous sequence of cognitional rehearsals in accordance with the encounters and transactions of their sex of assignment. This accumulation exerted its formative influence irrespective of contradictory chromosomal sex, gonadal sex, hormonal sex, or morphologic sex.
Even in those few cases where the gender role and orientation developed ambiguously or contradicted the sex of rearing, the significance of cognitional rehearsals emerged as paramount. In the first place, these were never cases of children who, irrespective of hidden hermaphroditic incongruities, had external genitals that looked perfectly male or perfectly female. The external genitals looked ambiguous, thereby permitting the child from infancy onwards to make comparisons and conjectures about being a boy or a girl. Many of the children had ambiguous external genitals. The few who developed an ambiguous gender role and orientation experienced a reinforcement of ambiguity in the social environment. The parents remained basically unconvinced that they had a son, or a daughter. In some instances there had been also a reassignment of sex. Neighbors remembered and talked about the boy who turned into a girl, and playmates made verbal ammunition of the accusation of being half boy, half girl, or a freak. Even before school age, clear evidence might emerge of a child's personal conviction that everyone was making a mistake and that it was time to begin immediate rehearsals for living as a member of the opposite sex.
It may well be that homosexuality, and other behavioral disorders of sex, are fundamentally disorders of cognitional eroticism, early established and deeply ingrained, their relation to other variables of sex being peripheral at most.
It may be also that disorders of cognitional eroticism should be regarded as imprinting phenomena (chapter by the Hampsons; Fletcher, 1957; Schiller, 1957), imprints that are actually misprints. In accordance with the principles of imprinting, it would be the case that these misprint disorders can be established only at a specified critical period in the life history. The early age, betw^een 18 months and 3 years, after which it becomes psychologically hazardous to make sex reassignments of hermaphrodites suggests that the critical period for the imprinting of gender role and orientation corresponds with the critical period for the establishment of native language. There may conceivably be another critical period for limited modification of gender imprints at the time of puberty.
The phenomenon of the critical imprinting period allows an explanation of the fact that, in the case of homosexuality, a person can engage in a homosexual act when safely past the critical period without becoming a chronic homosexual. Effectively imprinted at the critical period to respond to homosexual stimuli, however, a person becomes a chronic homosexual. Effective stimuli may be extremely specific, and variable from ])erson to person, which may account for the varieties of homosexual preference.
The phenomenon of the critical imprinting period allows an explanation not only of disorders of cognitional eroticism, but also of normal and healthy cognitional eroticism. Just as nonhealthy eroticism may become indelibly imprinted, so also may healthy eroticism, masculine and feminine. Indeed, so fixed is masculinity and femininity of outlook in healthy men and women, respectively, that it has always been assumed that sexual orientation must be determined in some automatic fashion utterly independent of life experience, for example, by genes or hormones. Now it becomes necessary to allow that erotic outlook and orientation is an autonomous psychologic phenomenon independent of genes and hormones and, moreover, a permanent and ineradicable one as well.
The idea of a psychologic phenomenon being autonomous is not new in psychologic theory, but the idea that a psychologic phenomenon may be permanent and ineradicably imprinted is not always hospitably received in the company of present day theories. It is a challenging concept, and one worthy of extensive research within and beyond its application to sex.
The stimuli that bring eroticism to expression remain to be discussed. Are these, too, purely perceptual and learned or do the sex hormones automatically trigger behavior in man as they may do in the lower animals? From the histories of the patients listed in Table 22.1 and the reports herein cited, it is clear that certain changes in behavior are associated with hormone administration: the increased sexual activity of the hypogonadal men who received androgen therapy, the diminution in sexual activity they reported in periods when treatment was discontinued, the increased sexual desire in women following androgen therapy.
Some writers have suggested that psychologic stimuli are sufficient, that in man there has been an emancipation from hormonal control. If it is true, however, as a number of the same writers have postulated, that the sex hormones have a direct effect on the genitalia, maintaining them erotically functional, and on the generation of genitopelvic tactile and somesthetic signals that are relayed to the brain, the role of sex hormones in erotic arousal cannot properly be claimed to have been completely replaced by psychologic stimuli, for indeed the two are not mutually separable.
Tactile and somesthetic signals relayed to the brain from both the genitopelvic area and from other parts of the body may have erotic significance and promote erotic arousal. Erotic arousal may be generated also by signals sent to the brain from the eyes, the ears, and the sense of smell, which signals may be erotically just as potent as genitopelvic tactile signals. An action of sex hormones on certain of these modalities has been claimed (Torda and Wolff. 1944; Beach, 1948; Le Magnen, 1952a, 1952b; Harris, IMichael and Scott, 1958; Schneider, Costiloe, Howard and Wolf, 1958), but the area is largely unexplored, especially in man.
Erotic arousal may be triggered not only by sensory signals but also by memory signals from the brain, witness the erotic dreams of paraplegic patients who are incapable of pelvic sensation and who presumably do not, wdiile sleeping, see, hear, smell or feel anything erotic that induces the dreams. Primarily the function of the brain in eroticism is to coordinate and record signals that arrive by way of the different sensory modalities. Messages so received may then be either inhibited or further processed and transmitted in the service of erotic arousal. The brain can perform its erotic function independently of pelvic participation, witness again the dreams and fantasies of paraplegic patients whose genitalia function only through spinal reflexes and without any neural connection with the brain. The brain may record erotic signals and store them as memories for indefinite periods of time, so that past experiences of critical significance in the life history exert a constant and indelible influence on an individual's erotic inclinations and choices.
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Reference: Young WC. Sex and internal secretions. (1961) 3rd Eda. Williams and Wilkins. Baltimore.
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