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I have decided to take early retirement in September 2020. During the many years online I have received wonderful feedback from many readers, researchers and students interested in human embryology. I especially thank my research collaborators and contributors to the site. The good news is Embryology will remain online and I will continue my association with UNSW Australia. I look forward to updating and including the many exciting new discoveries in Embryology!
Young WC. Sex and internal secretions. (1961) 3rd Eda. Williams and Wilkins. Baltimore.
Section A Biologic Basis of Sex Cytologic and Genetic Basis of Sex | Role of Hormones in the Differentiation of Sex
Section B The Hypophysis and the Gonadotrophic Hormones in Relation to Reproduction Morphology of the Hypophysis Related to Its Function | Physiology of the Anterior Hypophysis in Relation to Reproduction
The Mammalian Testis | The Accessory Reproductive Glands of Mammals | The Mammalian Ovary | The Mammalian Female Reproductive Cycle and Its Controlling Mechanisms | Action of Estrogen and Progesterone on the Reproductive Tract of Lower Primates | The Mammary Gland and Lactation | Some Problems of the Metabolism and Mechanism of Action of Steroid Sex Hormones | Nutritional Effects on Endocrine Secretions
Section D Biology of Sperm and Ova, Fertilization, Implantation, the Placenta, and Pregnancy Biology of Spermatozoa | Biology of Eggs and Implantation | Histochemistry and Electron Microscopy of the Placenta | Gestation
Section E Physiology of Reproduction in Submammalian Vertebrates Endocrinology of Reproduction in Cold-blooded Vertebrates | Endocrinology of Reproduction in Birds
Section F Hormonal Regulation of Reproductive Behavior The Hormones and Mating Behavior | Gonadal Hormones and Social Behavior in Infrahuman Vertebrates | Gonadal Hormones and Parental Behavior in Birds and Infrahuman Mammals | Sex Hormones and Other Variables in Human Eroticism | The Ontogenesis of Sexual Behavior in Man | Cultural Determinants of Sexual Behavior
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Section F Hormonal Regulation of Reproductive Behavior

The Ontogenesis of Sexual Behavior in Man

THE ONTOGENESIS OF SEXUAL BEHAVIOR IN MAN

John L. H amp son, M.D.

ASSOCIATE PROFESSOR OF PSYCHIATRY AND PEDIATRICS, THE JOHNS HOPKINS UNIVERSITY, BALTIMORE, MARYLAND

and Joan G. Hampson, M.D.

ASSISTANT PROFESSOR OF PSYCHIATRY AND PEDIATRICS, THE JOHNS HOPKINS UNIVERSITY, BALTIMORE, MARYLAND


I. Introduction 1401 I. Introduction

II. Heredity, Environment and the

Instinct Controversy 1401 Adam and Eve — the ancient account of

A. Animals Studies and Human Sex their creation is firmly anchored in our cul R T^u^'rf T' V P ■ Au\^. ■ !im ^^^^^ traditions as symbolic of the primeval

a. 1 he Critical renod Hypothesis. .. .1404 cr^^^,.^^-^^,^^r. r™ i i r i -i-^T , •, •

C. Imprinting 1404 separateness of male and female. Plato, it is

III. The Establishment of Psychologic ^^^^> re-endorsed an ancient legend of the

Sex 1400 original hermaphroditism of Man, and

A. The Evidence of Human Her- Freud rehabilitated for Science a bisexual

niaphroditism 1406 concept of the human psyche. Yet the di 1. Chromosomal sex 408 ^^^^^ ^^ ^^^j^ ^^^ ^^^^^^^ .^ ^^ entrenched

z. ijonaclai sex l4Uo . ,, ■ i i m i ,

3. Hormonal sex 1409 ^^ colloquial philosophy that even in rigor 4. Internal accessory organs 1410 o^s scientific thinking it is difficult to tran 5. External genital appearance 1411 scend.

6. Assigned sex and rearing 1413 Embryologists have long known of the

7. Psychologic sex: gender role. .1413 original developmental hermaphroditism of

B. The Influence of the Physical Sex the human fetus. It is well established by

Variables on the Establishment of biochemists that the sex hormones are

lender Kole: Body Image 1414 , , i ■ • j^i • i • ■, ,

C. Social Learning and Gender Role . . . 1415 f^^'^^^f f,^^" ^^ ^^^'l chemical structure. Al 1. Social environment and the es- though the overlap between male and female

tablishment of gender role .... 1417 ^as been conceded, psychosexual orienta 2. Gender role rehearsal 1419 tion in the two sexes has, in the final analy 3. Gender role identification and sis, been attributed to two separate instincts.

gender role preference 1420 These instincts have been vaguely ascribed

IV. Parental Behavior IN Humans 1421 to innate and constitutional sources. The v. 1 HE Sexual Cycle IN Women : Pyscho- ;j„„ f ^ , , i x tx

SEXUAL Concomitants 1423 '^'t °^, ^ complete psychosexual neutrality

VI. Disorders of Psychologic Sex; ^" ^^e human infant seemed too farfetched

Psychopathology 1425 ^o be entertained seriously.

A. Gender Role Distortions 1425 it u i'» r • . i ^i

r. TT u J , _ J Tj 1 **• Heredity, fcinvironment and the

B. Hermaphroditism and Psvchopa- ^ V ^

thology 1428 Instinct Controversy

VII. Concluding Remarks 1429 Since the beginning of history man has

VIII. References 1430 assumed himself to be separate and apart

1401


1402


HORMONAL REGULATION OF BEHAVIOR


from other creatures in a myriad of ways. The theologians of our time, no less than the Greek philosophers of the fourth century B.C. have held to a view that infrahuman creatures function instinctively, reserving intellect, reasoning, and will power for Man alone. But as Beach (1955) has written: "prescientific concepts of instinct were not deduced from the facts of nature ; they were necessitated by the demands of philosophic systems based on supernatural conceptions of nature."

It would be foolish to quibble here about the place of Man in Nature; it is assumed, a priori, the Man belongs in the system of biologic continuities. It is also assumed that a student of behavior, even as a student of biochemistry or anatomy, can expect to observe differences both between species and within a species. These differences do not in themselves contradict the assumption of a system of continuities.

The concept of instinct, quite apart from having served to set the other animals apart from man, has been used also as a means of explaining and understanding behavioral phenomena which emerged so spontaneously and with such predictability that they seemed to have arisen preformed from some inner source. At the same time the terms innate, constitutional, and genetically determined attained explanatory significance in some quarters. Although the instinctive explanation of behavior still enjoys currency, the concept has not gone unchallenged (Beach and Jaynes, 1954, 1955). Experimental and clinical psychology embrace a theory of instincts ; psychoanalytic theory of personality relies heavily on instinctive explanations and commonly regards psychologic disorder as due to some disorganization of instinctual life. There is, however, an accumulating body of evidence which strongly suggests that the usual instinctual explanation of behavior is a gross oversimplification; as an adequate foundation for a science of behavior the traditional form of the concept of instinct has proven less than adequate.

One factor in this inadequacy is the regrettable alliance of the term innate wuth the concept of instinct. Innate and instinctive have come to be equated in common usage in a way that excludes the influence of


experience and learning. The more rigorous scientific viewpoint holds that a distinction between learned and unlearned behavior is not only impossible to dichotomize in any meaningful way but is, in any case, not a very useful distinction. For as Beach (1955) has pointed out: "the final form of any response is affected by a multiplicity of variables only two of which are genetical and experiential factors." Thus in considering any behavior pattern one is obliged to take cognizance of the interplay between two broad categories of variables: (1) those variables which are intrinsic to the organism, and (2) those variables which are extrinsic. Much of the present volume deals in detail with the intrinsic variables which one must consider in studying sexual behavior. In his chapter on hormones. Young takes into account one of these intrinsic variables, gonadal hormones, and the influence on reproductive behavior in infrahuman species. Cultural influence is an important extrinsic variable dealt with in the chapter by Mead.

A. ANIMAL STUDIES AND HUMAN SEX BEHAVIOR

Social behavior in man, including sexual behavior, is undeniably complex and frequently baffling. It is not a new observation that many of the social attitudes and behavior patterns in man are acquired through one or another process of learning during an individual's lifetime. It is relatively recent, however, that scientifically sound evidence has been collected which spotlights the earliest months and years of life as a highly critical learning period of inexorable importance to later psychologic functioning. As yet the bulk of the experimental evidence for this has come from animal experimentation; experimentation in humans has of necessity been limited to chance occurrences and the "experiments of Nature."

In the area of social perception and social responses some of the most important work in recent years has been done by the European zoologists who have elected to be called ethologists. These experimentalists, notably Lorenz and Tinbergen, have devoted much attention to the observation that, in many animal species, virtually all social behavior is based on, or is an elaboration of, spe


ONTOGENESIS OF SEXUAL BEHAVIOR


1403


cific stereotyped behavior that can be elicited by specific sign stimuli. These stereotyped behavior patterns they consider to be innate in the traditional sense of "instinctive." It has been postulated that special neurophysiologic brain mechanisms, referred to as Innate Releasing ^Mechanisms (IRM), operate to release the impounded "innate reaction." Operation of an IRM is a response to specific sign stimuli (the terms cue stimuli and social releasers have also been applied) in the environment. In the herring gull, for example, the red spot on the beak of the adult has been found to serve as a trigger stimulus which elicits food-begging behavior in the fledgling.

Systematic study of similar phenomena in humans has not yet been undertaken. A well known human example is the smiling response in infants (Spitz and Wolf, 1946). The smiling response becomes active somewhere between the fourth and tenth week of life and is elicited by the sign stimulus of a slowly moving human face or, alternatively, the essential elements comprising the gestalt of the human face. Thus a drawing of two circles for eyes and a mark for a nose and mouth moved slowdy in the infant's visual field will suffice.

A word about terminology is in order at this point. Some psychiatrists, knowing the imjiortance of the theory of instinct in psychoanalytic doctrine, have been eager to construe ethologic findings as an experimental validation of this theory. Doubtless the overlap in terminology in the two fields has encouraged this not entirely justifiable practice and the result has been a semantic entente rather than an identity of operational definitions. In writing this chapter, we have been in no position to be laboratory purists in the matter of operational definitions; on the other hand, in pointing up the similarity, where it occurs, between concepts derived from ethology and from our own work we have endeavored to avoid the worst sins of argument by analogy. In the context of human psychololgy, we have deliberately avoided some of the ethologic vocabulary as being arbitrarily mechanistic and too likely to breed anachronistic misunderstanding among those chiefly acquainted with psychodynamic concepts.

The premise that behavior is based pri


marily on instincts is gradually disai^pearing from scientific writing and the traditional concept of instinct is undergoing revision and modification.^ In its place has emerged the view that early experience importantly structures subsequent behavior. This is not to say, lest misunderstanding arise, that the animal organism, human or .subhuman, is merely a blank slate to be written upon by the capricious finger of life experiences. Quite the contrary, for there are now many studies in the literature dealing with genetic constitution and the inheritance of basic capacities affecting later learning, temperament and personality (Medawar, 1947; Scott, 1953; Scott and Charles, 1953, 1954; Palowski and Scott, 1956; Goy and Young, 1957).

With increasing sophistication in these matters behavioral scientists have begun to abandon the fruitless effort to determine what proportion environmental or hereditary factors contribute to a given behavior pattern. Instead, greater attention is being given to the question, "hoiv do these factors operate in structuring behavior?"

Anastasi (1958) has recently appraised the heredity-environment issue and the multifocal research approaches required to investigate the question "how?" That author rightly points out that the influence of hereditary or environmental factors is always indirect; the more indirect the connection the wider the range of variation of possible outcomes.

With respect to early experience Beach and Jaynes (1954) see three possible answers to the question of how its influence is mediated: (1 ) habits formed in early life may persist in adult behavior; (2) the individual's perceptual capacities may be so structured by early experience as to affect adult behavior; and (3) during specific

^ Two exten8i\e ievie\v.s dealing with instinctive behavior have recently been published. In the one, Fletcher (1957) has surveyed the work in ethology, comparative psychology, and social and educational theory. Fletcher endeavors to integrate these new insights with psychoanalytic instinct theory; regrettably he omits mention of such important American and Canadian work as that of Scott, Nissen, Skinner, Hebb, and Young, to mention only a few. The other review (Schiller, 1957) is more narrowly restricted to a presentation of a few important, perhaps classic, studies in animal behavior.


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HORMONAL REGULATION OF BEHAVIOR


stages in ontogeny ("critical periods") certain types of behavior are indelibly shaped and molded for the life of the animal.

B. THE CRITICAL PERIOD HYPOTHESIS

Research advances in psychology during the past decade have added so much information relevant to the establishment of the modalities of social behavior that it may be anachronistic to discuss the "critical period" for learning as merely an hypothesis. Stated briefly, it is now well established that there are limited and often highly specific periods during the early life of an animal during which important social learning takes place, more or less permanently affecting the subsequent social behavior of the individual. There is known to be considerable species variation in the occurrence and duration of these critical learning periods. In the graylag goose, for example, one critical period can be identified as beginning immediately after hatching (Lorenz, 1935, 1950, 1952) . Scott and his co-workers (Scott, Fredericson and Fuller, 1951 ; Scott and Marston, 1950; Scott, 1958) in a meticulously detailed series of observations have identified five distinct periods in the life history of the dog. In this species, the period from 3 to 7 weeks (Period III, Primary Socialization I is the time when primary social modalities become established. Physiologically this period is characterized by advancing but incomplete myelinization and development of the central nervous system and associated sense organs ; for the first time conditioning becomes possible. Critical periods have also been identified and studied in the mouse (Williams and Scott, 1953), sheep (Scott, 1945), howling monkey (Carpenter, 1934), and red deer (Darling, 1937).

The finding that a critical period for primary socialization is such a widespread phenomenon in animals gives us reason to expect a continuity of this situation in the human species. Scott (1958) speculates, on the basis of his findings in dogs, that one might expect the earliest critical period in humans to begin between the 6th to 24th week of life and to last two years or possibly longer. Needless to say, much has already been written in the psychiatric and psychoanalytic literature to support a contention


that these earliest years of life are crucial ones in personality development. The findings in regard to the establishment of psychologic sex reported in detail on the pages to follow are in substantial agreement with the view that critical learning periods in the human are indeed an established reality.

C. IMPRINTING

There is a particular phenomenon well described by the ethologists which deserves special mention, namely, the phenomenon of imprinting. Our work (Ilampson and Money, 1955; Money, 1955; Money and Hampson, 1955; Money, Hampson and Hampson, 1955a, b, 1957; Hampson, 1955; Hampson, Hampson and Money, 1955; Hampson, Money and Hampson, 1956) with hermaphroditic children has led us to see an analogy between establishment of gender role in early childhood and the phenomenology of imprinting as described in lower animals. The following are two examples which illustrate the basic similarities involved.

L In >e;us past the sight of a string of day-old goslings following their parent would have been cited as an example of innate, unlearned in,stinctive behavior. Heinroth (1910), Lorenz (1935), and others have shown how, in birds, species recognition as well as appropriate behavior in response to another member of its species, is not pre-established but is subject to modification through early life experience. A greylag gosling, for example, that has lived a few days with its parents will never respond to a human as if the human were its parent. On the other hand, to quote Lorenz (in Schiller, 1953), "if a greylag gosling is taken into human care immediately after hatching, all the behavior patterns which are slanted to the parents respond at once to the human being. In fact, only very careful treatment can induce incubator-hatched

"Lorenz (in Schiller, 1957) specified two features characteristic of the phenomenon of imprinting which differentiate it from other types of learning. (1) Imprinting is limited to a very definite and often extremely short phase of ontogeny, the "critical period." (2) The result of this process of determination is irreversible. In this context the term learning is used in the broad sense referring to a family of processes inferred from the observation that animals learn. To quote Verplanrk (1957): "When we say that an animal learns, we are stating that, other things being equal, some behavior now occurs in a situation in which it had not occurred previously, or that the behavior nowoccurring in a given situation is different from the behavior that occurred on the last occasion the animal was in that situation."


ONTOGENESIS OF SEXUAL BEHAVIOR


1405


TABLE 23.1 Varieties of ambisexual incongruities

1. Conyenital hyperadrenocortical females: externally hermaphroditic; normal internal reproductive

organs ; female sex chromatin pattern (36

2. Hermaphrodites with ambiguous or masculinized external genitals: normal functional female inter nal reproductive structures and ovaries; female sex chromatin pattern 6

3. Classical true hermaphroditism: testicular and ovarian tissue both present; enlarged phallus; vari able development of the genital ducts; male or female sex chromatin pattern 1

4. Cryptorchid hermaphrodites with relatively complete MUllerian differentiation: penis hypospadic or

normal; possibly virilizing at puberty; male sex chromatin pattern 7

5. Cryptorchid hrrniaphrodilcs with relatively incomplete Mullerian differentiation: hypospadic oi- cli toral phallus; possibly virilizing at pubertj^; male sex chromatin pattern

a. With urogenital sinus 18

b. With blind vaginal pouch 2

G. Simulant feniales with fendnizing inguinal testes and vestigial Miillerian differentiation: blind vaginal pouch; male sex chromatin pattern 13


greylag goslings to follow a grey mother goose. They must not be allowed to see a human being from the moment they break their shell to the time they are placed under the mother goose. If they do, they follow the human being at once." It is also remarkable that almost any object between the size of a small chicken and a human being which moves and makes a noise can imprint the following-response in a newborn greylag.

In the experimental goslings the imprint of filial response to a decoy parent, with a human-being as parental model, was fixed and irreversible.

2. There is a relatively rare variety of hermaphroditism (Variety 2, Table 23.1) in which a baby is born in every respect a regular, normal female except that the labia are fused to look like a scrotum and the clitoris is enlarged to look like an unfinished penis; there is a gutter in the place of a penile urethra so that urination is accomplished in a sitting position (Fig. 23.1). So confusing is the ambiguity of such genitalia that the baby may be considered either a boy or a girl. Studies have been made, one of which has been reported by Money, Hampson, and Hampson (1956), of two such people reared as boys who have now reached adulthood. W^e have also studied three similar people, still juvenile, being reared as girls. It is not surprising that the latter are growing up as ordinary girls psychologically undifferentiable from their normal sisters and schoolgirl friends. What is quite remarkable, however, is that two individuals reared as boys matured with a regular, masculine psychology, and sexual orientation. At the time of the first signs of pubertal feminization they had recoiled from any suggestion that they change to live as a girl and subsequently they had been pleased and benefited by surgical and hormonal masculinization. Erotically they have been beset by the handicap of an undersized and malformed penis and scrotum. Nonetheless they were living happy, successful lives as men, one of them old enough to be a liusband and an adoptive father. The imprint of


VV


^




i


k


Fig. 23.1. Six-day-old female infant with masculinized external genitals, female sex chromatin pattern. (Courtesy of Dr. Lawson Wilkins.)

psychologic masculinity, from multiple male models and examples, was fixed and irreversible.

Psychiatrists have long realized that some modalities and disorders of psychologic fimctioning in humans resist all efforts at modification. The traditional explanation of this has rested heavily on constitutional and


1406


HORMONAL REGULATION OF BEHAVIOR


genetic considerations. In recent years animal studies such as these cited, together with new findings in the fields of psychiatry and clinical psychology, suggest possible alternative explanations.^ The investigations into the nature of psychologic sex provide an example in point, for a person's psychosexual orientation, once established, becomes an ineradicable part of personality functioning.

III. The Establishment of Psychologic Sex

In the human psychologic sexuality is not differentiated when the child is born. Rather, psychologic sex becomes differentiated during the course of the many experiences of growing up, including those experiences dictated by his or her own bodily equipment. Thus, in the place of the theory of an innate, constitutional psychologic bisexuality such as that proposed by Freud — a concept already questioned on theoretical grounds by Rado (1940), among others — we must substitute a concept of psychologic sexual neutrality in humans at birth. Such psychosexual neutrality permits the development and perpetuation of divers patterns of psychosexual orientation and functioning in accordance with the life experiences each individual may encounter and transact.

A. THE EVIDENCE OF HUMAN HERMAPHRODITISM

The evidence for the foregoing statement has, in part, emerged from the study of human hermaphroditism.^ The sexual in '^ Others have also pointed out analogies between the ethologic findings in lower animals and certain psychologic and behavioral phenomena in humans. Lorenz, Bowlby and Walter, to name but a few (see Tanner and Inhelder, 1953, 1954, 1955), have discussed possible relationships implicit in animal findings to the ontogeny and phylogeny of psychologic development in children. Russell and Russell (1957) have made the suggestion that certain aspects of human behavior might best be approached and studied from an ethologic point of view.

  • The term hermaphrodite is used here to describe not only those individuals with completely

ambiguous external genital development but also to include all instances in which a contradiction exists between the predominant external genital appearance on the one hand and the sex chromatin pattern, gonads, hormones, or internal accessory structures, singly or severally, on the other. In this


congruities which occur in hermaphroditism involve contradictions, singly or in combination, between six variables of sex. These variables, the first five of which are dealt with in specific detail by other writers in this volume, are: (a) chromosomal sex: (b) gonadal sex; (c) hormonal sex; (d) internal accessory reproductive structures; (e) external genital morphology; (f) the sex of assignment and rearing. Hermaphroditic patients, showing various combinations of these six sexual variables, may be appraised with respect to a seventh variable, (g) Gender role^ or psychologic sex. In this way one can ascertain something about the relative importance of each of the six variables in relation to the seventh.

A nineteenth century classification of hermaphrodites was based on the assumption that the microscopic structure of the gonads was the ultimate criterion for purposes of

sense the older terms pseudohermaphrodite and inlersexuality are superfluous and unnecessarily confusing and have not been retained. Undeniably no classificatory scheme is perfect or sacrosanct; other classifications of the hermaphroditic anomalies have been published (c/. Wilkins 1957, Jones and Scott, 1958).

^ By the term, gender role, is meant all those things that a person says or does to disclose himself or herself as having the status of boy or man, girl or woman, respectively. It includes, but is not restricted to sexuality in the sense of eroticism. Gender role is appraised in relation to the following: general mannerisms, deportment and demeanor; play preferences and recreational interests; spontaneous topics of talk in unprompted conversation and casual comment; content of dreams, daydreams, and fantasies; replies to oblique inquiries and projective tests; evidence of erotic practice and finally, the person's own replies to direct inquiry. Lest there be misunderstanding, the term gender role is not identical and synonymous with the term sex of assignment and rearing. Sex status can be assigned to a child by parental, medical, or legal decision. The psj^chologic phenomenon which we have termed gender role, or psychosexual orientation, evolves gradually in the course of growing up and cannot be assigned or discarded at will. The components of gender role are neither static nor imiversal. They change with the times and are an integral part of each culture and subculture. Thus one may expect important differences in what is to be considered typical and appropriate masculine or feminine gender role as displayed by a native of Thailand and a native of Maryland, or as displayed by the pioneer contemporaries of Peter Stuyvesant and by their descendants in Westchester County suburbia of the 1960's.


ONTOGENESIS OF SEXUAL BEHAVIOR


1407


differentiation (Klebs, 1876). In his classification Klebs recognized true hermaphrodites, who possessed both ovarian and testicular tissue, male pseudohermaphrodites with only testicular tissue and female pseudohermaphrodites with only ovarian tissue. In studying the determinants of the psychologic |)henomena of sexual outlook and orientation — gender role — Kleb's classification has proven too anachronistic to be useful, for modern discoveries in endocrinology, including recent work in sex chromatin determination (Moore and Barr, 1953; Moore, (iraham, and Barr, 1953; Grumbach and Barr, 1958) make it clear that multiple variables are involved. The following list of the varieties of ambisexual incongruities was drawn up with these considerations in mind and is based in part on Dr. .John ^loney's unpublished doctoral thesis in which he reviewed the medical literature of over 300 cases of hermaphroditism.

It would be inappropriate here to expound at length on the differential diagnosis of the various clinical types of ambisexual incongruities but it may be relevant to comment briefly on the six varieties listed in Table 23.1.^

Variety 1. Congenital hyperadrenocortical females. In this group of patients the external genitals may appear almost normally female with slight to medium enlargement of the clitoris; or there may be a single urogenital orifice with enlargement of the clitoris. Rarely, a penile urethra and a fused empty scrotal sac are found. Without l)enefit of suppressive cortisone therapy physical growth and development is precocious and virilizing. Diagnosis is now possible during llic neonatal [leriod by means of urinary IT-koto-lrionl assessment, without laparotomy. Plural inciileiiee m a family is common, but is usually restricted to a single generation and a single marriage (Childs, Grumbach and Van Wyk, 1956). The Miillerian ?y.stem and ovaries are normal; the sex chromatin l^attern is female.

Variety 2. Hermaphrodites with ambiguous or masculinized external genitals. Unlike females with liyperadrenocorticism, the hermaphrodites in this siroup, although born with varying degrees of aml)iguous or masculinized genitals, do not show progressive virilization. On the contrary, since the ovaries and accessory internal reproductive structures are normal, secondary feminization at pul)erty is the rule and reproduction is iio.-sil)le. The enlarged phallus usually re.scnililes a liypospadic penis with chordee. Labial fusion may be pronounced or even complete. The vagina usually opens into a urogenital sinus but occasionally opens independently. The sex chromatin pattern is fe


male. Wilkins, Jones, Holman and Stempfel (1958) have published a rei)ort on 21 such hermaphrodites; in all but 3 the mother had received progestinic medication (usually 17-ethinyltestosterone) beginning in the first trimester of pregnancy.

Variety 3. Classical true hermaphroditism. Ovarian and testicular tissue is present, in one ovotestis, the other gonad being an ovary or a testis, in two ovotestes, or in one ovary and one testis. The internal genital structures as well as the external genitalia show various degrees of ambiguity. Pubertal development of the secondary sexual characteristics may be masculine, feminine, or ambiguous. Diagnosis requires laparotomy and biopsy of the gonads. In an unpublished study of 20 patients with classical true hermaphroditism, M. L. Barr found that 15 had a female sex chromatin pattern.

Variety 4- Cryptorchid hermaphrodites with relatively complete Miillerian differentiation. The penis may be fully formed with a penile urethra or it may be hypospadic. One or both testes may by cryptorchid; one testis may be atrophic. Miillerian structures not infrequently herniate into the groin or scrotum in the company of one testis. At puberty the secondary sexual development is nearly always masculine although a eunuchoid habitus is sometimes observed, especially in those instances of bilateral cryptorchidism and hypospadias. Diagnosis requires surgical exploration; the sex chromatin pattern is probably always male although it is conceivable that a female pattern may be possible.

Variety 5. Cryptorchid hermaphrodites with relatively incomplete Miillerian differentiation. Historically, hypospadiacs have not been called hermaphrodites imless the hypospadias is severe, the scrotum bifid, and the testes undescended. Careful examination, however, often reveals that such hypospadiacs have a blind vaginal pouch hidden beyond the single external urogenital orifice ; or there may be a separate external orifice for a blind vaginal pouch. In some instances the phallus may be only sHghtly larger than a clitoris. The Miillerian and Wolffian systems are malformed or vestigial. A virilizing puberty cannot be predicted; in some patients pubertal development may be weakly feminine resulting in breast enlargement. Plural incidence within a family may occur. Diagnosis requires testicular biopsy ; the sex chromatin pattern is probably always male although future findings may require a revision of this assumption.

Variety 6. Simulant females with feminizing inguinal testes and vestigial Miillerian differentiation. In this group of patients the external genital appearance completely simulates the normal female. The vagina is a blind pouch and, with few exceptions, the Miillerian system is a cord-like vestige; the Wolffian system is malformed or vestigial. The testes may remain intra-abdominal or herniate into the groin; even in adults the microscopic appearance of the testes is best described as immature and poorly differentiated. At puberty the development of the female secondary sexual characteristics is nearly always complete except that menstruation does not occur. In about one


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HORMONAL REGULATION OF BEHAVIOR


TABLE 23.2 Sex chromatin pattern and rearing contradictory: 20 cases


Variety of Ambisexual Incongruities


Sex Chromatin Pattern


Assigned Sex and Rearing


Gender Role


Hyperadrenocortical females (variety 1): total, 3

Hermaphrodites with ambiguous or masculinized external genitals (variety 2): total, 1

Classical true hermaphroditism (variety 3) : total, 1

Cryptorchid hermaphrcjdites (varieties 4, 5): total, 8

Simulant females with feminizing testes (variety 6): total, (i


3 9

1 9

1 9 8 d" 6 d"


3 d"

1 d"

1 cf 8 9 6 9


3 cf

1 d"

1 d^

8 9 6 9


Additional Data


Gonads


Endogenous Hormonal Sex


Internal Accessory Organs


External Genital Morphology


3 9


3 cf


3 9


3 ?


1 9


1 9


1 9


1 ^


9 left


1 ^


1 ^


1 ^


cf right





8 &


2 c^, 1 9


4 c^, 3 9


8 ^



5 juvenile


1 ^



6 cf


2 9

4 juvenile


G vestigial


6 9


third of the cases sexual hair fails to appear despite normal adult estrogen levels. Diagnosis requires gonadal biopsy and the demonstration of a male sex chromatin pattern. Familial incidence in several generations is common.

The following findings and conclusions emerged from the study^ of a series of over 110 hermaphroditic individuals. The tables do not include an entry for every single patient. Some of the patients were too young at the time of study for meaningful conclusions to be drawn; others failed in one or another detail to fulfill the criteria necessary for inclusion. On the other hand, some patients, because of their multiple hermaphroditic manifestations, ciualified for inclusion in more than one of the following tables.

1. Chromosomal Sex

There were 20 patients in this series of hermaphroditic individuals who had been assigned to and reared in a sex contrary to

"Under the aegis of John C. Whitehorn, Professor of Pediatrics, The Johns Hopkins University. The research was supported by grants from the Josiah Macy, Jr., Foundation, and the Public Health Service (USPHS Grant M-1557).


their sex chromatin pattern as established by skin biopsy or the buccal smear technique (Grumbach and Barr, 1958)." Without a single exception, it was found that the gender role and orientation as man or woman, boy or girl was in accordance with the assigned sex and rearing rather than in accord with the chromosomal sex (Table 23.2 ) . It seems convincingly clear, therefore,, that gender role, orientation as male or female, does not correspond automatically with the chromosomal sex; rather, it is in some way related to assigned sex and rearing.

'3. Gonadal Sex

There were 30 hermaphroditic patients in whom a contradiction was found between

• Tables 23.2 through 23.7 are revisions of tabular material published earlier (Money, Hampson and Hampson, 1955b) and include patients studied since the original report. The authors are indebted to John Money, Ph.D., for his part in collecting psychologic data on which the tables are based. The responsibility, however, for the additions toand modifications of the earlier tables was assumed by the authors of this chapter.


ONTOGENESIS OF SEXUAL BEHAVIOR


1409


TABLE 23.3

Gonads and rearing contradictory: 30 cases


Variety of Ambisexual Incongruity


Gonads


Assigned Sex and Rearing


Gender Role


Hyperadrenocortical females (variety 1): total, 5

Hermaphrodites with ambiguous or masculinized external genitals (variety 2): total, 2

Crvptorchid hvpospadic hermaphrodites (varieties 4, 5): total. 14

iSimulant females with feminizing testes (variety 6): total, 9


5 9

2 9

14 d"

9 c^


5 d"

2 c^

13 9

1 9 ^ cf

9 9


5 rf 2 cf 11 9

3 ^ 9 9



Addition


al Data



Sex Chromatin Pattern


Endogenous Hormonal Sex


Internal Accessory Organs


External Genital Morphology


5 9


5 d"


5 9


5 ?


1 9

1 9 (?)


2 9


2 9


2 ^


9 d" 5 o^(?)


6 d" 1 9

7 juvenile


8 vestigial 1 ^ 5 9


14 ^


7 cf 2 c^(?)


2 9

7 juvenile


9 vestigial


9 9


the sexual status of the gonads and the sex of assignment and rearing^ (Table 23.3). In all but 3, psychologic studies revealed a gender role fully concordant with the sex of rearing. As a prognosticator of a person's gender role and orientation, gonadal structure, per se, thus proved to be unreliable; again gender role was in greatest accord with the assigned sex and rearing.

3. Hormonal Sex

It is of course necessary to consider hormonal sex separately from gonadal histology, for ovaries do not always secrete effective estrogens, nor testes effective androgens. The ovaries of females with hyperadrenocorticism are suppressed in function and, although their adrenals produce an excess of estrogens as well as of androgens, the androgenic activity dominates, inducing excessive virilization of the body. The gonads of simulant females with ijiguinal testes produce, in most instances, estrogens which feminize the body.

The data on 21 patients who went into or beyond puberty or, in the cases of patients


wuth hyperadrenocorticism, a precocious puberty-equivalent in which hormonal influences induced secondary sexual changes contradictory of the sex in which the individual had been living, are summarized in Table 23.4. In all patients the contradiction was subsequently corrected with hormonal therapy and with plastic surgery when indicated.

Of the 31 patients whose sex hormones and secondary sexual body development contradicted their assigned sex and rearing, only 5 became ambivalent with respect to their gender role. Four of the 5 had been reared as girls. One of these, a man in his thirties when studied by Dr. John Money, had acted on his own initiative and changed to live as a man from the age of 16 onward. The other four, although living as women, displayed disordered gender role as evidenced by some degree of bisexual erotic inclination. The impracticability of publishing here the extensive psychiatric and psychologic records of these 5 patients requires the authors to submit, instead, their considered opinion that these 5 patients do not.


1410


HORMONAL REGULATION OF BEHAVIOR


TABLE 23.4

EndogeiKnis honnonal sex and rearing contradictory: 31 cases


Variety of Ambisexual Incongruity


Endogenous Hormonal Sex


Assigned Sex and Rearing


Gender Role



Hyperadrenocortical females (variety 1): total, 21


21 d^


21 9


19 2


9



Hermaphrodites with ambiguous or masculinized external genitals (variety 2): total, 2


2 9


2 cf


2


d^



Classical true hermaphroditism (variet>- 3): total, 1


1 ^


1 cf


1


d'



Cryptorchid hermaphrodites with relatively complete Miillerian differentiation (variety 4): total, 2


2 cf


2 9


2


9



Crvptorchid hermaphrodite, feminizing (variety 5): total, 1


1 9


1 d^


1


d'



Cryptorchid hermaphrodite, virilizing {variety 5): total, 4


4 ^


4 9


1 3


9



Additional Data


External Genital Morphology


Sex Chromatin Pattern


Gonads


Internal Accessory Organs


21 ^


21 9(?)


21 9


21 9


2 ^


1 9

1 9 (?)


2 9


2 9


1 ?


1 9


1 ^


1 ^


2 ^


1 d

1 di?)


2 d


2 9


1 ^


1 c^(?)


1 d^


1 vestigial


4 ?


1 d 3 c?(?)


4 d


4 vestigial


in themselves, seem to offer any convincing evidence that sex hormones act as a single causal agent in the establishment of an individual's gender role and psychosexual orientation. Restoration of normal female estrogen balance did not alter the psychosexual functioning of the 4 patients who had lived as women; the patient who had changed to live as a man declined androgen treatment after surgical castration for malignancy. The remaining 26 patients in the group established a gender role consistent with their assigned sex and rearing, despite the embarrassment and difficulties of living with contradictory secondary sexual development.


4- Internal Accessory Organs

The fourth sex variable embraces the internal derivatives of the Miillerian and Wolffian duct systems. The uterus, as the organ of menstruation, and the prostate and seminal vesicles as organs concerned with the secretion of seminal fluid, are of particular theoretic importance psychologically.

There were 25 instances in the group of patients studied in whom the assigned sex and rearing was not in accord with the predominant male or female internal accessory structures (Table 23.5). In 22 of the 25 the individual's gender role agreed with the assigned sex and rearing. The 3 remaining


ONTOGENESIS OF SEXUAL BEHAVIOR


1411


were the same three who deviated in Table 23.3; they appeared again as deviants in Table 23.4.

It is rare in hermaphroditism for either the uterus or the prostate to reach functional maturity without medical intervention. There were 3 patients among the 25 in Table 23.5 for whom this statement did not hold; despite the fact that the 3 had grown up with a normal uterus, they had been reared as boys and had a thoroughly masculine gender role. In view of these findings there seems no reason to suspect any correlation between gender role and the internal accessory organs.

5. External Genital Appearance

The a]iiiearance at birth of the external genitals usually dictates the sex to which a baby is assigned and in which it is reared.


As a child grows older the appearance of the external genitals is a bodily feature of importance to the child's private assuredness of being a boy or a girl.

When a hermaphroditic baby is born the medical attendants are likely to be less casual in declaring its sex from external appearance alone. Nevertheless in reviewing the histories of a large number of babies born with anomalous external genitalia it was our impression that in the majority of cases the external genital appearance was a primary consideration in the initial assignment of sex. It is perhaps remarkable that some children, in fact, are reared in a sex contradicting their predominant external genital appearance. Moreover, it is even possible for such hermaphrodites to establish a gender role entirely in agreement with assigned sex and rearing, despite the para


TABLE 23.5 Internal accessory organs and rearing contradictory: 25 cases


Variety of Ambisexual Incongruity


Internal Accessory Organs


Assigned Sex and Rearing


Gender Role


Hj'peradrenocortical females (variety 1): total, 5

Hermaphrodites with ambiguous or masculinized external genitals (variety 2): total, 2

Classical true hermaphroditism (variety 3): total, 1


5 9

2 9 > cf 1 ^


5 cf 2 d"

1 cf


5 cf^ 2 cf

1 d^


Cryptorchid hermaphrodites with incomplete Mlillerian differentiation (varietv 5): total, 8


Simulant females with feminizing testes (varietv 6): total, 9


8 cT > 9 (vestigial)


9cf > 9


6 9

1 cf

1 9

9 9


5 9 3 ^


9 9


Additional Data


Sex Chromatin Pattern


Gonads


Endogenous Hormonal Sex


External Genital Morphology


5 9(?)


5 9


5 d"


4 ^ 1 c^


1 9


2 9


2 9


2 ^


1 9(?)





1 9


1 ^


1 ^


1 9'


4 cT


8 cf


4 d"


8 ^


4 d^(?)



4 juvenile



7 d"


9 cT


2 9


9 9


2 cT



7 juvenile



1412


HORMONAL REGULATION OF BEHAVIOR


doxical appearance of their external genitalia.

In our series of hermaphrodites there were 25 with a marked degree of contradiction between their external genital appearance and their assigned sex and rearing (Table 23.6) . Of the 16 older patients, all had lived at least to teen-age (in one case as long as 47 years!) with ambiguous or contradictory genital appearance. Of the 9 under 12 years of age all had lived for at least 8 years of their life surgically uncorrected. All but 2 of the 25 individuals had been able to come to terms with his or her anomalous appearance and had established a gender role wholly consistent with assigned sex and rearing. The exceptions, one a young adult, the other a middle-aged man, had profound psychologic problems, some of which were clearly concerned with an ambivalent gender identification. It is of interest that both


of these patients had had a reassignment of sex, the one by medical decision at the age of 5 from female to male; the other, at the age of 16, of his own initiative changed to live as a male.

Lest the foregoing be misunderstood, the table documents the possibility of a person's establishing a gender role consistent with the sex of assignment and rearing despite ambiguous looking or even contradictory-looking genitals. The Table does not, however, document the enormity of the problem these people had to surmount in coming to terms psychologically with their paradoxical appearance. It has been our experience that more than anything else, the visible anatomic genital or bodily contradictions occasion the grestest psychologic distress. Although none of this group of patients had ever had a psychotic illness, many displayed a moderate degree of psy


TABLE 23.G External genital appearance and rearing contradictory: 25 cases


Variety of Ambisexual Incongruity


Predominant

External Genital

Appearance


Assigned Sex and Rearing


Gender Role


Hyperadrenocortical females (variety 1): total, 16

Cryptorchid hermaphrodites with relatively complete Miillerian differentiation (variety 4): total, 3

Cryptorchid hermaphrodites with relatively incomplete Miillerian differentiation (variety 5): total, 3

Cryptorchid hermaphrodites with relatively incomplete Miillerian development (variety 5): total, 2

Cryptorchid hermaphrodites, etc., feminizing (breasts) variety 5): total, 1


<r > 9 & > 9

9 > &

cf > 9

9 > &


16 9 3 9

1 d^

2 9 ^ d^

2 9

1 0^


16 9 3 9

1 d^

2 ^

2 9 1 d^


Additional Data


Sex Chromatin Pattern


Gonads


Endogenous Hormonal Sex


Internal Accessory Organs


16 9(?)


16 9


16 cf


16 9


1 &


3 d^


1 d^


3 9


2 cf (?)



2 juvenile



1 c^


3 c^


1 c^


3 vestigial


2 cf(?)



2 juvenile



2 c^


2 d^


1 cf

1 juvenile


2 vestigial


1 cf (?)


1 &


1 9


1 vestigial


ONTOGENESIS OF SEXUAL BEHAVIOR


1413


chologic nonhealthiness. The importance of body appearance in the establishment of gender role is dealt with more fully in a later section.

6. Assigned Sex and Rearing

In the foregoing tables, chromosomal sex, gonadal sex, hormonal sex, internal reproductive organs, and external genital appearance have been considered in turn with respect to (1) the assigned sex and rearing, and (2) the gender role established by each individual. In only 7 of the cases represented in these tables was there any inconsistency between the sex of rearing and gender role despite other incongruities between these and the other variables of sex. Parenthetically, it is to be noted that 3 of the 7 appear in more than one of the tables.

Thus it appears legitimate, once again, to conclude that there is a very close relationship between the sex of assignment and rearing and the establishment of a masculine or feminine gender role and psychosexual orientation.

7. Psychologic Sex: Gender Role

The evidence of human hermaphroditism indicates that psychologic maleness or femaleness in the human is not to be attributed to any single one of the physical variables of sex, i.e., the gonads, sex hormones, sex chromatin pattern, or the morphology of the external genitals and internal rejiroductive structures. It is, of course, conceivable that some other intrinsic body factor could have an important bearing on psychosexual development. Anthropometrists in Europe (Vague, 1953) and in this country (Sheldon and Stevens, 1942) have suggested a relationship between body build and psychosexual orientation. Their data, however, are not beyond alternative interpretation insofar as the psychologic importance of body structure is concerned.

In the late nineteenth century von KrafftEbing (1890) suggested special bisexual brain centers as an explanation of the psychologic differences between men and women. There have never been established any anatomic or neurophysiologic data to support such a conjecture. From 1890 to the present day, however, von Krafft-Ebing's views on the l)isexual na


TABLE 23.7

Gender rule in patients with same diagnosis reared male or female: 65 cases


Variety of Ambisexual Incongruity


Assigned Sex and Rearing


Gender Role


Hyperadrenocortical

females (variety 1)

Cryptorchid hermaphrodites (varieties 4, 5)


39 9 15 9


37 9 29^ 5c^

12939^ 6d^


ture of man have been taken up in turn by such theorists as H. Ellis, Hirschfeld, Freud and others in a way that has shaped psychiatric theory (see also Rado, 1940). The presence of chromosomal, hormonal, gonadal, or genital incongruities in an individual does not automatically confer incongruous or disordered masculine or feminine psychologic development. It would seem that the theory of bisexuality must be laid to rest when one considers the evidence of hermaphroditic individuals with the same diagnosis some of whom have been reared as boys and some as girls (Table 23.7).

There are 65 cases represented in Table 23.7 including 5 of the 7 in the entire series studied in whom ambivalence of gender role was found. These 5 had all been reared as girls. Of the 60 remaining, the 49 patients reared as girls had established an entirely feminine gender role, whereas the 11 reared as boys had established a masculine gender role and orientation.

One can conclude that an individual's gender role and orientation as boy or girl, man or woman does not have an innate, preformed instinctive basis as some have maintained. Instead the evidence supports the view that psychologic sex is undifferentiated at birth, a sexual neutrality in the place of the Freudian bisexuality, and that the individual becomes differentiated as masculine or feminine, psychologically, in the course of the many experiences of growing up.

The comjK'lling quality of the genital erotic component of sex role has led to the utilization of a concept of drive as an explanation of sex role differences. In that event sex drive should be considered genderless at birth and can be assumed to have no


1414


HORMONAL REGULATION OF BEHAVIOR


somatic basis other than the highly innervated and erotically sensitive areas of the body. As an alternative to a drive concept some will find it preferable to say simply that the erotically sensitive parts of the human body can be used and stimulated by oneself or another person and that during the process of psychologic growth and development erotic sensations become firmly associated with and inextricably a part of adult gender role.

B. THE INFLUENCE OF THE PHYSICAL SEX

VARIABLES ON THE ESTABLISHMENT OF

GENDER role: BODY IMAGE

To say that a person's gender role is a correlation of the kind of learning experiences encountered and transacted is not to endorse an oversimplified version of social and environmental determinism. It is readily seen that in a very real sense the somatic variables of sex constitute an important part of a person's environment.

For example, although hormonal functioning does not directly or automatically determine maleness or femaleness of gender role an important relationship is involved, namely, the effect of hormones on body morphology and their sensitizing influence on genital sensory structures. Before birth hormonal functioning has a major role to play in embryonic differentiation of both the internal and external genital structures. When a child is born, needless to say, it is the morphology of the external genitalia which dictates or guides the assignment of sexual status. Although we have found that most children can overcome the quandary of anomalous or ambiguous genital appearance and grow up with a sexual orientation appropriate to the sex in which they were reared, it is by no means true that they do so without difficulty or with complete success. Broadly speaking, the more pronounced and obvious the anomaly the greater the concomitant feelings of bashfulness, shame, and differentness with which the person must contend. Some children, in the face of ambiguous genital appearance, may privately construe that an error has been made; they are then able to adapt to a psychosexual orientation appropriate to their assigned sex only by paying the penalty of one or another kind of psychologic


symptomatology. Parents, too, react to their child's ambiguous genital development with concerns and worries about the correctness of the child's sexual identity thus importantly structuring the family environment in which gender learning has its experiential origins.

The sex hormones also play an important role at puberty in establishing the secondary sexual characteristics of the body. Ordinarily the transition from prepuberty to adolescence is a gradual one that occurs in the setting of pubertal physical changes and social interaction. A child's anticipation of being grown up and of being considered grown up by others directs attention to body appearance. More than any other indicator the physical changes at puberty indicate to other people that this person is now ready for the social transactions of adolescence. Thus prepubertal children usually anticipate eagerly the first signs of breast development, of beard growth or voice change, as evidence that they are growing up, and they compare their progress with that of their age-mates. The appearance of menses in girls and nocturnal emissions in boys are important private signs which enhance a child's sense of confidence in his or her masculinity or femininity. In the absence of visible bodily signs of maturity the youngster suffers not only in lowered self-esteem but equally important is excluded by his contemporaries from a host of adolescent activities which are indispensable to psychologic and social maturation. Secondary sexual development may, however, be incongruous (as in hyperadrenocortical female hermaphrodites), precocious (as in cases of idiopathic sexual precocity), or delayed (as in hypogonadal males and girls with gonadal dysgenesis). Incongruity, precocity, and delay in secondary sexual development inevitably have important effects psychologically. The more publicly evident the incongruity the more distressing the person finds it. Thus hirsute girls and women with hyperadrenocorticism have almost invariably reported to us that they would, more than anything else like to be rid of their unfeminine-looking body hair; their hyperplastic clitoris and deep voice assumed a secondary importance beside the hirsutism. Precocious appearance of the hormonally


ONTOGENESIS OF SEXUAL BEHAVIOR


1415


induced signs of puberty may, on the one hand, open the doors to adolescent social encounters before a child is psychologically ready and able to cope with such learning experiences (see Hampson, 1955; Money, 1955). On the other hand pubertal delay permits a lag in social and psychologic development by making the child so out of step with his contemporaries that the job of catching up may be slow or even impossible. Without belaboring the point, body appearance does have an important, indirect bearing on the development of psychologic functioning, including that which we term gender role or psychosexual orientation.

There is yet another link between hormonal functioning and gender role which is specifically relevant to the genital erotic component of psychologic sex. To be brief, for this topic is taken up more fully in jMoney's chapter on sex hormones and eroticism, both androgens and estrogens can act on external genital structures so as to intensify genital erotic sensation. In the absence of one or the other sex hormone genital erotic sensation is relatively weak. We would stress that genital eroticism, however important, is but one facet of the constellation of elements comprising psychosexual functioning. One may, therefore, expect to find individual differences, both pathologic and nonpathologic, in this component alone, or in combination with the other components of gender role behavior.

C. SOCIAL LEARNING AND GENDER ROLE

It is not a new idea that humans learn a great deal during their earliest years. From the beginning of psychodynamic research, much information has been collected about the importance of learning experiences in man and the influence of such experience in shaping those psychologic modalities broadly spoken of as personality.*^

Freud (1910) was among the first to pre " It is beyond the scope of this chapter to discuss the many theories of personality extant in psychology and psychiatry. The interested reader will find an excellent review of the more important of these theories in Theories of Personality by Hall and Lindsey (1957). Murphy (1947), with considerable ai>propriateness, defined personality as "the integration of all the roles that a particular person has to enact."


sent a comprehensive theory of psychosexuality. Freud apparently based his views on a postulate von Krafft-Ebing had also embraced some years before, namely, that every person is inherently bisexual, or, in another sense, homosexual as well as heterosexual tendencies are originally present in everyone. The Freudian theory of psychosexual development embodied a succession of oral, anal, and genital phases and held that the psychologic differences between the sexes emerged during the first 5 years of life.'-' Freud saw the third to the fifth year

"Freud (1949) attached great importance to a biologic hypothesis that "man is descended from a mammal which reached maturity at the age of five, but that some great external influence was brought to bear upon the species and interrupted the straight line of de^'elopment of sexuality. This may also have been related to some other transformations in the sexual life of man as compared with that of animals, such as the suppression of the periodicity of the libido and the exploitation of the part played by menstruation in the relation between the sexes." Freud (1927) gave the following account of the maturation and development of infantile sexuality : "The sexual function ... is in existence from the very beginning of the individual's life, though at first it is assimilated to the other vital functions and does not become independent of them until later ; it has to pass through a long and complicated process of development before it becomes what we are familiar with as the normal sexual life of the adult. It begins by manifesting itself in the activity of a whole number of component instincts. These are dependent upon erotogenic zones in the body; some of them make their appearance in pairs of opposite impulses (such as sadism and masochism or the impulse to look and to be looked at) they operate independently of one another in their search for pleasure, and they find their object for the most part in the subject's own body. Thus to begin with they are noncentralized and predominantly auto-erotic. Later they begin to be co-ordinated ; a first step of organization is reached under the dominance of the oral components, an anal-sadistic stage follows and it is only after the third stage has at last been reached that the primacy of the genitals is established and that the sexual function begins to serve the ends of reproduction."

In Freud's view it was the phallus which dictated the organization of infantile sexuality: "The difference between the two — the infantile genital organization and the final genital organization of the adult — constitutes at the same time the main characteristic of the infantile form, namely that for both sexes in childhood only one kind of genital organ comes into account, the male. The primacy reached is, therefore, not a primacy of genital but of the phallus" (Freud, 1924). Freud considered there was no distinction between maleness and femaleness in the anal-sadistic stage; in the in


1416


HORMONAL REGULATION OF BEHAVIOR


of life as largely occupied by OedipaP" operations culminating in the resolution of the innate bisexual conflict.

Despite the fact that the concept of innate bisexual instincts has been repeatedly challenged on theoretical grounds (Rado, 1940) during the past two decades, the notion continues to enjoy doctrinaire status. That this is so is rather remarkable, for the bulk of modern research, both in animals and man, points unequivocally to the importance of learning in the establishment of those behavioral and psychologic characteristics which in humans we speak of as personality. Thus an individual learns ways of behaving through the experiences he encounters and transacts and not by virtue of an endowment of inbuilt imperatives for one or another mode of functioning and behaving. The evidence of human hermaphroditism, as cited earlier, is substantially in agreement with this point of view provided it be allowed that learning includes an individual's experiences, as mediated by his own uniquely individual perceptual-cognitional capacities, with his own body morphology and physiologic functioning.

The accrued evidence strongly suggests that the beginnings of gender-specific psychologic characteristics are, to an important extent, acquired, and so become manifest, very early in life, probably during the first 2V2 or 3 years. This early period for gender role learning may quite legitimately, it is thought, be designated a critical -period, the effects of which persist throughout the life of the individual. One item of evidence for this comes from the psychologic studies (Hampson, 1955) of children and adults who had undergone a reassignment of sex status at some time in their lives. Some, but not all of these hermaphroditic individuals

fantile genital stage maleness had emerged but no femaleness. Not until puberty was the polarity of male and female established.

'" Stated briefly, the Freudian Oedipus complex consists of a sexual attraction for the parent of the opposite sex and a hostile dislike for the parent of the same sex. The child's sexual identification is then thought to occur as a defense against his projected hostile impulses and a fear of retaliation. Although it cannot be denied that this sequence of events may sometimes occur it is contrary to the facts to consider seriously that this is the only, or even the most important means by which a child establishes his gender identity.


were hyperadrenocortical hermaphrodites. Data on 12 of these patients are shown in Figure 23.2. Only patients who had been studied at least 6V2 years and up to a maximum of 20 years after the reassignment of sex were included. In each instance an appraisal was made of the pervasive thoroughness of sexual orientation in the sex assigned at the change using the criteria for gender role discussed earlier in this chapter. An appraisal was also made in each case of the successfulness and stability of day-to-day adjustment with deliberate intention, if any doubt existed, of erring on the side of overscrupulousness rather than allowing any benefit of the doubt.

Of the 5 who were reassigned before their first birthday all but 1 had no subsequent disturbance psychologically. This one child, living as a girl, growing uj) in circumstances of family turmoil and chaos, and subjected to repeated thoughtless comments about the time "when you were a boy" had begun to disi)lay truant behavior at school and some degree of uncertainty about her gender status. With only 1 exception the 7 patients reassigned later than the first birthday were rated as inadequately adjusted. The exception was a boy who, in all fairness, had to be rated as adequately adaptive, inasmuch as the problems he had were typical for many boys of his religious and moral upbringing, and likely to be transient. The three patients reassigned at the age of 4 or after had varying degrees of confusion and ambivalence with regard to their gender status.

The importance of the earliest years of life for gender role learning is illustrated in Figure 23.2. However, one should not infer from such data any hard and fast date-line for either the beginning or the ending of the early critical period for gender orientation but merely the broad limits. Such a view is in agreement with the findings of other investigators who have seen gender-role differentiation beginning about the second year of life and becoming firmly established by the third year, by which age roughly 75 per cent of children can distinguish between the sexes and identify themselves as boys or girls (Gesell, Ilg and Ames, 1940; Seward, 1946). Although the development of gender role is a continuing process throughout the


ONTOGENESIS OF SEXUAL BEHAVIOR


1417


I6 +


16


7


Age in Years at 5 Time of

Change 4


AGE AT TIME OF REASSIGNMENT OF SEX

AND SUBSEQUENT ADEQUACY

OF PSYCHOLOGIC ADJUSTMENT

A Healthy Adjustment

A Mildly Non-Healthy Adjustment

A Moderately Non-Healthy Adjustment


t



Syr lOyr 7yr 8yr lOyr 9yr 20yr I4yr llyr I4yr 22yr37yr Assigned Sex and Age of 12 Patients at Time of Study Fig. 23.2


growing up years and into adulthood, the first few years of a child's life are significantly characterized by mutually reciprocating expectancies on the part of parents and child alike regarding gender-appropriate behavior which are particularly influential in structuring personality functioning. Speaking metaphorically, the learning experiences during the first 2 to 3 years ap])ear to be critical in that they set the stage for the dramatization to follow.

1. Social Environment and the Establishment of Gender Role

In working with hermaphroditic children and their parents, it has Ijecome clear that the establishment of a child's psychosexual orientation begins not so much with the child as with his parents. Faced with the dilemma of a sexually ambiguous-looking baby it is next to impossible for parents to


avoid assigning sex status to the child. If medical indecision about the child's sex status is delayed too long such parents, understandably, find it difficult to refer to their baby as "it" indefinitely. Under ordinary circumstances, there is no such quandary about the sex status of a newborn and appropriate pronouns are used from the outset. From cigars, w^ hich announce a new son, to a mother's insistence on sex-appropriate pink or blue cradle accessories, our cultural folkways provide ample evidence that parental attitudes and expectancies are set in operation virtually the moment a child is born. Thus the assigned sex-status, as well as such body signs as the external genitals,"

" It is only partly facetious to say that even the hair-cut is important. In a clear and unmistakable way the short hair-cut of a boy and the long tresses of the girl provide a clear sign to others which in effect announces "I am a boy (girl) — treat me as such." Many preschool youngsters when


1418


HORMONAL REGULATION OF BEHAVIOR


provides an important stimulus to parents which helps determine the gender-species role training they will provide.

Rabban (1950) studied the influence of social class status on sex-role identification in children and reported that boys are more clearly aware of sex-appropriate behavior than girls in both middle- and working-class groups. On the other hand, he found that boys and girls of the working-class group were not only earlier, but more clearly, aware of gender-appropriate behavior than were either boys or girls of the middle-class group. Clearly class level provides the social frame of reference within which such diverse influences as parental models, playmates, social mores, attitudes, and opportunities can operate to structure gender role and personality structure.

In their excellent study of how 379 American mothers brought up their children from birth to kindergarten. Sears, Maccoby and Levin (1957) correctly point out that, although sex role in modern American culture is constantly changing and shifting in terms of what the culture regards as appropriate, there is no culture that does not make some distinction between masculine or feminine role behavior (see also the Mead's chapter, in this volume). Although it may not be a logical absurdity to regard masculine and feminine behavior as biologically innate phenomena, the study of sex role in other cultures makes it clear that all too often we have wrongly identified that which is current and prevalent in our own culture as the natural and normal state of affairs. It seems more in accord with the findings in both humans and other animal forms to regard humans, male and female alike, as capable of acquiring, during early life, an almost infinite variety of behavior patterns some of which will be approved of as serving the individual and society well, whereas others will be considered unfortunate, if not actually detrimental, both biologically and culturally (cf. Mead's comments on the folkways of the Mundugumors) .

Sears, Maccoby and Levin (1957) distinguished three kinds of learning occurring in association with the development of social

asked about the difference between boys and girls will give an answer in terms of the hair-cut rather than the genitals.


behavior appropriate to the child's own sex: (1) trial and error, (2) direct tuition by the parents, and (3) role practice or rehearsals. From their experience these authors were inclined to believe that very little learning of social behavior and social values ever occurs by simple trial and error. They point out an important difference between learning by direct tuition and guidance and learning through role rehearsal, namely that, in the latter, the child selects the actions to perform from his own observations of what the role requires rather than from what the instructions of his parents may be. This being so one must immediately consider individual differences in cognitional and perceptual inheritance as important variables affecting the final outcome of such learning.

In studying the intentional teaching of gender-appropriate behavior by the 379 parents in their study. Sears, Maccoby and Levin found important differences in the kind of rearing meted out to the two sexes. Among the items studied it was with respect to aggressive behavior that the greatest distinctions were made by parents; boys were allowed significantly more freedom than girls to fight back in encounters with nonsiblings. Many mothers actively encouraged their sons to be aggressive and provided ample opportunity for learning aggressive behavior and, as one parent put it, to be a "real boy." Parental attitude toward such diverse things as household tasks and chores and school achievement differed, too, with respect to sons and daughters. Of the 91 mothers of boys and the 83 mothers of girls who were most strongly inclined to differentiate sex-role in their rearing practices, it was found that definitely higher expectations were put on girls for table manners, neatness and orderliness, and instant obedience. Techniques of child training differed too: even with the nonaggressive group of children, boys were more commonly administered physical punishment than girls. Girls were disciplined more often by so-called love-oriented techniques of praise for good behavior and the display of disappointment at unacceptable behavior. Interestingly, many of the mothers did not recognize any efforts they were making to evoke appropriate sex-role behavior in their child and those


ONTOGENESIS OF SEXUAL BEHAVIOR


U19


who did were inclined to interpret the difference as their natural reaction to the "innate" sex-determined temperament of the child.

2. Gender Role Rehearsal

Role rehearsal stands apart as being of unique importance to the acquisition of role behavior. Needless to say this mode of learning operates hand in hand with parental expectancies and experiential opportunities.

Gender role rehearsals in childhood occur as pretend operations. Much of the time these rehearsals occur in fantasy only, although it is but a short step for a child from fantasy and daydreams to active play. Sears, Maccoby and Levin (1957) point out that such role practice involves more than simple imitation of single aspects of the model's behavior in that the child "takes on the role itself, at least momentarily, with all the feelings, attitudes, values and actions that he attributes to the person who actually occupies the role."*- In this way a child "tries out" many roles, some of which he will keep and others discard. Thus, typically, a child does not lose his sexual identity by switching from pretending to be a nurse or a mother one time and a fireman sometime later. The available evidence strongly suggests, moreover, that it is social endorsement and approbation of any given pattern of identification which governs the final constitution of gender role. A boy, for instance, will therefore come to share many of his mother's attitudes and values while abhorring for himself the use of feminine cosmetics.

In the absence of the possibility for controlled experimental studies in this area with humans, questions as to the determinants of gender-specific adolescent phenomena have remained matters of conjecture. The temporal coincidence of bodily maturational changes with the emergence of psychologic signs of heterosexual interest

'- This mode of learning is usually referred to as identification in the psychiatric literature. Kagan (1958) has attempted an analysis of the concept of identification in an effort to place the concept within the framework of learning theory. In this he defined identification as an accjuired cognitive resjionse within a person which can lead to similarities in behavior between a subject and a model.


has been taken by some as evidence of a common origin in sex-specific instincts or gonadal hormones. As pointed out earlier, there are undeniably physiologic correlates of gender role in that body image and body functioning are an inevitable ingredient of experience. The development of breasts and a more feminine figure in adolescence, for example, increases the likelihood of a girl's being included in teen-age social activities such as dating. Nonetheless, role enactment is preceded by role rehearsal in games of pretend, daydreams, and fantasies during the childhood and pre-adolescent years in the course of which gender role becomes initially structured and, usually, indelibly so.

The girls and women born with the condition known as gonadal dysgenesis provide an unplanned experiment-of-Nature which sheds some light on the origins of gender role fantasies and sexual enactment. This group of patients, reported on earlier by Hampson, Hampson and Money (1955), and now found to have an XO chromatin pattern rather than a typical female XX pattern, is of relevance in that despite normal female external genital appearance their gonads are dysplastic and do not secrete female sex hormones. A measure of control over the timing of puberty is possible, for at the socially appropriate age, treatment with estrogens is begun so that secondary sexual physical development proceeds along typically feminine lines.

Table 23.8 summarizes the psychosexual data on 13 such individuals; in four instances psychiatric and psychologic studies were obtained both before and after the administration of estrogens to induce somatic puberty. Without exception all 13 had established an unequivocally feminine gender role and psychosexual orientation. Also without exception daydreams and fantasies of heterosexual courtship, romance, and sometimes of marriage, motherhood and erotic play had not only occurred but in most cases were a conspicuous feature of their psychosexual expression. The daydreams and fantasies of these individuals were recognizably feminine in quality and content whether or not treatment with estrogen had been instituted, although, to be sure, increasing age and experience seemed


1420


HORMONAL REGULATION OF BEHAVIOR


TABLE 23.8


Psychosexual data for 13 patients wi


th gonadal dysgenesis and XO sex


"hromatin pattern




Daydreams




Hetero





and Fanta







Patient and Age at Time of Study


sies of Heterosexual Courtship, Motherhood and Erotic


Auto-erotic Genital Practices


Heterosexual Social Dating


Erotic

Activity:

Kissing,

Petting,

etc.


Heterosexual Intercourse


Homosexual

Eroticism

of Any Sort




Play






Before estrogen


1, age 91^


+


+



_


_


_


substitution


2, age 12


+







therapy


3, age 121^


+


+





+



4, age 13


+


Inconclusive data







5, age 13


+


+


+






6, age 14


+








7, age 151^


+


+


+






8, age IG


+



+


+





9, age 17


+


Inconclusive data


+


'


"


"


After estrogen


3, age 15


+


+






substitution


4, age 17


+


+


+





therapy


6, age 17


+


+


+


+





8, age 17


+


+


+


+





10, age 19


+



+


+


Inconclusive data




11, age 23


+


+


+


+





12, age 26


+


+


+


+





13, age 27


+



+


+


+



to result in more sophisticated fantasy material. The data also show that, although auto-erotic genital activity, usually masturbatory, occurred more often among girls who had been supplied with estrogen, such activity was by no means precluded by the total absence of gonadal hormones. The one patient noted in the column for homo-erotic activity of any sort had engaged in transient homo-erotic play during her preteen years; this girl has been described in greater detail elsewhere (Hampson, Hampson and Money, 1955). In a general way, it appeared that with increasing age, the likelihood of at least some heterosexual social and erotic encounters was increased even in the absence of estrogen therapy. In this regard, however, both qualitative and quantitative differences were observed before and after the secondary sexual changes of puberty had been induced with estrogen treatment. It seemed likely that this difference could be accounted for by the increased social opportunities opened up by a maturing body appearance.


3. Gender Role Identification and Gender Role Preference

It is a common observation, substantiated by clinical and experimental findings (brown, 1956; Rabban, 1950), that beginning as early as the third year of life some children express a preference to have been born, or even to be, of the sex opposite that in which they are being reared. This unexpected role preference, in our western civilization at least, is encountered far more frequently in girls than in boys. In Brown's ( 1958) study of middle-class children between the ages of 3V2 to IP/2 years, the boys expressed a considerably stronger preference for the masculine role than did the girls for the feminine role. A number of studies have shown that even among adults, up to 12 times as many women as men report sometimes wishing that they were of the opposite sex (Terman, 1938; Gallup, 1955) . Unless one is aware of this phenomenon and the reasons for its occurrence it may be a major source of error in the


ONTOGENESIS OF SEXUAL BEHAVIOR


1421


evaluation of psychosexual functioning and its origins and development.

One explanation (and one which has had little or no scientific evidence to support it) was Freud's proposal that the young girl regularly felt envious of the male phallus ("penis envy") in the absence of a similar organ herself. The resolution of "penis envy" was held by Freud to be an essential part of psychosexual development during childhood. The psychoanalytic position on this matter has changed considerably over the years, however, and at present seems more in accord with the sociologic findings (Thompson, 1943).

Two factors seem to have an imjiortant bearing on expressed role preference. For one, our western culture is more apt to permit a girl to express a contrary gender role preference than it is a boy. Parents may permit, even encourage, tomboyishness in a daughter; a sissy son would likely evoke concern or even embarrassment. Secondly, and more important, is the child's growing awareness of the favored position of the male in the culture, a position carrying with it greater prestige and privilege than does being a female. Children may very early in life see that this is so and, unless wise parental guidance intervenes, the young girl becomes understandably envious of the masculine status which she does not have.

Although in typical, healthy psychosexual development gender role identification and gender role preference do not come into serious or permanent conflict, it is important to recognize that this may not invariably be so. Brown (1958a), writing of nonhermaphroditic girls and women, cites three major gender role patterns: (a) identification with and preference for the gender role of one's own sex, (b) identification with the gender role of one's own sex but preference for the gender role of the opposite sex, and (c) identification with the gender role of the opposite sex but preference for the gender role of one's own sex. One could find clinical justification for yet a fourth pattern, namely (d) identification with the gender role of the opposite sex combined with a solidly pervasive preference for the gender role of the opposite sex.


The relevance of this diversity of possible gender role patterns for such disorders of psychologic sex as homosexuality and transvestism is obvious and is dealt with in another section.

Although reliable well documented evidence has only in recent years begun to appear in the scientific literature, it is probably safe to say that gender role development during childhood is continuously and directly related to adult sexual l)ehavior in humans.

IV. Parental Behavior in Humans

Aliead of the human infant at birth lies the prospect of a more protracted period of immature dependency than for any other neonatal animal. That the human race has survived at all is testimony to the fact that in some way humans are capable of nurturing their young through this period of dependent immaturity. As with other human activities, parental behavior has been variously ascribed to innate, automatic instincts or drives, or, alternatively, considered by some to be largely determined by experience and social learning.

In other animal species maternal care has been considered the paradigm of instinctive behavior. Without belaboring the point, it may safely be said that both experimental biologists and comparative psychologists have come to see the problem as vastly more complex than the traditional assumptions have previously allowed. Whenever a species has been systematically and properly observed and studied, that behavior previously ascribed, by default, to "instinct" has been revealed as being the product of a great many variables. In this respect maternal behavior in infrahuman species has proven no exception and the concept of a "maternal instinct" operating without prior learning or experience now lacks scientific endorsement.

Insofar as maternal behavior in the human species is concerned, much the same predicament has prevailed as for other species; inadequate or improper study and experimentation had left a gap between common observation and scientifically established fact which the concept of a


1422


HORMONAL REGULATION OF BEHAVIOR


maternal instinct seemed to bridge. One of the chief difficulties in studying humans has been the practical one of observing individuals systematically and carefully over a long period of years. Moreover, many psychoanalytic and psychiatric studies have dealt with maternal attitudes rather than with mothering behavior in its broadest sense. Brody (1956) made a scholarly and comprehensive review of the literature pertaining to maternal behavior as of that date, dividing her survey into three main categories: (1) experimental studies in subhuman species, (2) anthropologic reports, particularly as they pertain to the feeding and handling of young children, and (3) clinical data culled from the literature of pediatrics, clinical psychology, psychiatry, and psychoanalysis. She noted with considerable relevance that the bulk of the material in the literature on human maternal behavior pointed to the interest of investigators in finding etiologic factors for the character disturbances and other psychopathologies,^^ and that there was a dearth of systematic investigation of the mothers themselves.

In an effort to devise a systematic method for the investigation of some significant aspects of maternal behavior, Brody made a content analysis of the behavioral records of 32 mothers whose behavior with their infants was observed for an approximately 4-hour period under standard conditions. Her research was also intended to test the hypothesis, namely, that the mother-infant interaction during feeding is the salient index of maternal behavior, "serving better than any other activity as a model of a mother's over-all behavior toward a given infant." Making use of explicitly defined quantitative methods, the observational rec " The empiric data bearing on those theories alleging that certain features of parental care of infants determine adult personality was reviewed, as of 1949, by Orlansky (1949). "Orlansky found much to doubt and criticize in the widely popular psychoanalytic opinions and emphasized instead the importance of constitutional and cultural factors as well as post-infantile experience in personality formation. In recent years, however, so much has been published concerning the inter-relatedness of parental and child behavior that cognizance must be taken of this variable in personality development. The question is no longer whether but how such influences act to structure personality.


ords were subjected to systematic classification and comparison. The 32 maternal records lent themselves to classification into four groups, each group having a statistically distinctive pattern of general maternal behavior. From this general typology four types of maternal feeding behavior were described. Although the hypothesis regarding feeding as an index of general maternal behavior was not proven for mothers of infants as young as 4 weeks of age, it was convincingly proven for mothers of older infants. Brody's work demonstrates the feasibility of quantitative examination and evaluation of parental behavior.

Sears, Maccoby, and Levin's (1957) study of the child rearing practices of 379 American mothers is another important recent contribution toward filling the hiatus in our knowledge of parental behavior in humans. The relevance of such studies to a better understanding of psychosexual development was referred to in an earlier section.

Present day psychiatric thinking about ]iarental behavior has been importantly structured by psychoanalytic theory. Freud regarded motherhood as the culmination of psychosexual development, the course of which he considered to be impelled by instinctual drives. Subsequent workers operating in the psychoanalytic frame of reference using, a 'priori, a concept of a maternal instinctual drive have further developed these ideas {e.g., Bonaparte, 1953; Deutsch, 1945).

On the other hand, there have been workers who have sought to prove that the psychologic manifestations of the maternal state are to be understood only in terms of the influence of social institutions and ideologies. Briffault (1931), representing an extreme in this point of view, maintained that all social feelings are derived from the mother-child relationship and considered the complexity of human social organization to be a direct correlate of the prolonged period of dependency of the human infant on his mother.

Until recently, the purely biologic factors in human maternal behavior have been subjected to very little direct scrutiny. Benedek (1949) postulated that the motherchild symbiosis begins at conception and continues through and beyond parturition


ONTOGENESIS OF SEXUAL BEHAVIOR


1423


'to lactation. Progestin and/or prolactin are seen by Benedek as inducing not only somatic changes but also the "registering of these changes in the psychic apparatus"; she viewed motherliness as stemming from the biologic effects and specific concomitants of these hormones. Such a view cannot be substantiated in the present state of knowledge concerning the specificities of psychologic-hormonal relationships.

Levy (1942) attempted to tease out some of the psychosomatic correlates of maternal behavior. Based on interviews involving much self -appraisal, his finding that there is "a rather high and significant positive correlation between maternal behavior and duration of menstrual flow" (r = 0.579) would have to be challenged on several grounds, the most significant of which is that the study on which the findings were based did not include any direct observations of maternal behavior. A decade later (Levy and Hess, 1952) Levy himself established the lack of positive correlation between interview and self-appraisal data on the one hand and direct observational ratings on the other.

The ethologists, having identified seemingly ready-made neurojjhysiologic pathways or releasing mechanisms for certain social responses in lower animals, have suggested that parental behavior can be viewed in much the same light. Lorenz is quoted by Tinbergen (1951) as considering that "parental behavior, a subinstinct of the major reproductive instinct, is responsive to sign-stimuli provided by the human baby." Although no controlled experimentation has been done in this area, Lorenz suggested that the human parental instinct, or IRM, responds to the gestalt of (1) a short face in relation to a large forehead, (2) ])rotrudind cheeks, and (3) maladjusted limb movements. Engaging though such a concept may ])e, no one accjuainted with the complexity of human psychologic development and functioning would seriously embrace such a view as adequately explaining the totality of parental or maternal behavior.

Without intending to oversimplify an obviously complex feature of human behavior the authors submit that parental behavior may be considered to be a facet of and an extension of gender role. Like the other as


pects of gender role that have been considered, parental role behavior does not require and is not dependent on innate sex-specific mechanisms. That it is teleologically purposive is fortuitous rather than directly causal. Like other aspects of gender role, parental behavior is accjuired largely through a process of learning during childhood. Direct tuition, identification with single or multiple parental models, rehearsals through play and fantasy— all come to be involved in the learning process. Like other aspects of gender role, the acciuisition of parental role behavior may go smoothly and normally, or become disordered.^'*

V. The Sexual Cycle in Women: Psychosexual Concoinitants

Cyclic variations in the sexual behavior of lower mammals and infrahuman primates as they relate to the hormonal events of the reproductive cycle have received much attention. Clearcut rhythms of heightened sexual activity have been described in the females not only of the lower mammalian species but of monkeys and apes as well. The occurrence of heightened sexual responsiveness and activity coinciding with the follicular (high estrogen) phase of the cycle in these subhuman species has been viewed as biologically purposeful in that it serves in the perpetuation of the species. The quest for evidences of similar estral correlates in the human female has been an inevitability.

It is probably safe to say that in terms of the complexity of motivation of behavior Man presents a considerably wider range of variables than any infrahuman species. It has become apparent, even in infrahuman species, however, that a single index, such as coital frequency, is unsatisfactory in studying over-all sexual behavior.

In studying Man one difficulty has been in getting agreement as to the components encompassed by the totality of sexual behavior. A second difficulty has been the es " A discussion of disordered parental beliavior is beyond the scope of this chapter. Brief reference here to "maternal overprotectiveness," "maternal underprotectiveness" (see Levy, 1943), and to allegedly "schizophrenogenic" parental behavior (Tietze, 1949: Reichard and Tillman, 1950) will serve to illustrate the point.


1424


HORMONAL REGULATION OF BEHAVIOR


tablishment of generally acceptable indices for the measurement of these components. Benedek and Rubinstein ( 1942) sought to circumvent these difficulties in the study of cyclic sexual behavior in the human female. They argued that: "Sexual desire, urge, and tenderness alternate with affective behavior of other sorts or may find substitution in activities and fantasies which are far different from overt sexual behavior, although all may be consequences of sexual stimulation." They made the assumption that the sexual rhythm of woman can be detected only "by psychoanalytic interpretation of the various preconscious and unconscious representations of the sexual drive" and therefore regarded the analysis of dreams, fantasies, and free-associative material as the primary data to be correlated with the cyclic physiologic changes. In brief, their study of 15 neurotic women by this approach revealed: (1) the estrogenic phase of the cycle corresponds to active heterosexual tendencies, the object of which is the sexual partner and coitus, fusing with (2) a passive receptive tendency correlated with progesterone activity, (3) a sudden decrease of active libido following ovulation which is assumed to correlate with the increase in progesterone levels, with interest for the sexual partner diminishing and with "emotional preparation for the function of motherhood" becoming the object of the sexual drive, and (4) disappearance of the emotional concentration on motherhood if pregnancy did not occur (decreased production of progesterone). Benedek and Rubinstein allowed that basic capacities for love, motherliness, constructive activity or their lack are present before the maturation of sexual functioning and that they do not cease to exist after the decline of hormone regulation. They acknowledged further that the processes which they described are "like faint ripples on a large body of water as compared with the constitutional basis of personality." For those who were able to accept the basic premises of this approach, Benedek and Rubinstein's work can be considered to verify the biologic assumptions implicit in Freud's concept of sexual drive. On the other hand the many other workers (e.g., Davis, 1926; Hamilton, 1929; Dickinson and Bean, 1931) who have given


more consideration to the statements of women regarding their conscious awareness of erotic desire and readiness for sexual activity as well as the reports of these women regarding overt sexual activity, do not agree with the findings of Benedek and Rubinstein. Although women show great individual diversity, their testimony in general is that erotic feeling and desire increase immediately before and again immediately after menstruation. Aware of the possible pitfalls in questionnaire type of data-gathering, where failures and inaccuracies in both memory and reporting may seriously impair the validity of the conclusions, McCance, Luff and Widdowson (1937) analyzed the daily entries in the special diary-type records of 780 complete menstrual cycles kept for them by 167 women. They found that nearly half of the subjects reported some degree of variation in sexual feeling during a menstrual cycle. The main peak of heightened eroticism occurred about the 8th day following the onset of menstrual flow (average duration of flow, 4.5 days) with a second smaller peak just preceding the onset of the next menstrual flow. These authors also systematically surveyed other menstrual molimina, recording the periodicity of each subjective phenomenon.

Kinsey, Pomeroy, Martin and Gebhard (1953) seeking a still more objective measure of female erotic arousal and responsiveness than that provided by a woman's subjective self-appraisal, collected data concerning variations in the quantity of mucous vaginal secretions (both Bartholin's and cervical glands) during the menstrual cycle. Their data indicated that "there is considerable variation in the quantity of the vaginal secretions among different females. There may also be variation in the quantity of secretion at different times in the same individual." Further, their data indicated that "the time of maximal mucous secretion and the time of maximal erotic responsiveness are almost always the same." About 59 per cent of their sample of women with coital experience recognized a monthly fluctuation in their vaginal secretions during erotic arousal. In general, these women reported increases to occur premenstrually, and, somewhat less generally, postmenstruallv. The data are in accord with the


ONTOGENESIS OF SEXUAL BEHAVIOR


1425


consensus of earlier workers pointing to evidence of pre- and postmenstrual enhancement of sexual responsiveness.

These findings have been puzzling in that they seemed contrary to biologic ends and to the observed increase in coital frequency just before ovulation in lower primates. Several explanations for the differences between these findings in humans and those in lower mammals have been suggested, important among which are the social attitudes bearing on sexual activity. In the majority of human societies sexual intercourse during active menstrual bleeding is contravened by prohibitions ranging from esthetic distaste to rigidly ritualized taboo. Moreover, the human female is probably alone among menstruating creatures in being able to anticipate and plan in terms of menstruation. The significance of the intermenstrual period and its association with ovulation and pregnancy is, in more sophisticated societies, so widely known as to constitute yet another variable that must be taken into account. In this sense it is even possible that the observed "peaks" of heightened eroticism are artifacts dependent on many factors other than variations in the supply of female sex hormones. It is in any case apparent that the increasing importance of nonhormonal control of sexual functioning through agencies of social learning and cognitional functioning to be observed even in monkeys and apes has reached its highest development in the human species.

VI. Disorders of Psychologic Sex: Psychopathology

A. GENDER ROLE DISTORTIONS

Of all the disorders of psychologic sex, homosexuality and transvestism have seemed to be the most baffling and difficult to explain. As is common when ignorance prevails, causal theories have grown up, some outrageous, some fanciful, some provocative, but none that has received the universal endorsement of medical science. Terms such as intersexuality, psychologic hermaphroditism, and others have found their way into the literature, often being used as if the words themselves constituted a scientific explanation.

One of the common eiTors made in con


sidering the problem of psychologic sex disorders, particularly homosexuality, has been to conceive of men and women as divided each into two camps: those who are heterosexual, and therefore spoken of as normal, and those who are homosexual. Such a notion is both conceptually and statistically incorrect and unw^arranted. The comprehensive studies on sex behavior in Americal men and women (Kinsey, Pomeroy and Martin, 1948; Kinsey, Pomeroy, Martin and Gebhard, 1953) have produced some of the best evidence relative to the incidence of homosexual behavior. According to those authors, 37 per cent of the white male population and 13 per cent of the white female population have had at least some overt homosexual experience to the point of orgasm between adolescence and old age. On the other hand, only 4 per cent of the white males and 1 to 3 per cent of the white females were found to be exclusively homosexual in their erotic activities throughout their lives. Between these extremes these authors found a wide range of sexual behavior involving varying proportions of heterosexual and homosexual orientation and activity which could, with good justification, be arranged on, and described by, a 7-point (0 to 6) rating scale. Such findings make it clear that one is never justified in speaking of homosexuality as if it w^ere a single descriptive or clinical entity as some authors have done. The point is well taken (Kinsey, Pomeroy and Martin, 1948, p. 617) that: "It would encourage clearer thinking on these matters if persons were not characterized as heterosexual or homosexual, but as individuals who have had certain amounts of heterosexual experience and certain amounts of homosexual experience. Instead of using these terms as substantives wdiich stand for persons, or even as adjectives to describe persons, they may better be used to describe the nature of the overt sexual relations, oi\ of the stimuli to which an individual erotically responds^^ (italics ours). For the purposes of the study, Kinsey, Pomeroy, Martin and Gebhard deliberately restricted themselves to an investigation of those sexual activities which culminate in orgasm. They doubtless would have allowed that, although overt homosexual stimulation to the Doint of orgasm, is one index of


1426


HORMONAL REGULATION OF BEHAVIOR


homosexuality it is not the only index. Psychologic sex is not restricted to sexuality in the sense of genital eroticism, but includes, for example, wishes and fantasies, demeanor and interests.

By way of illustrating this point as it pertains to homosexuality, we cite an excerpt from the letter to the authors from a 24-year-old woman. A Latin- American university student, the woman was not fluent in idiomatic English, but nonetheless, clearly expressed her dilemma.

"...I want to add that my attitudes, my way of thinking and feehng, my viewpoints regarding life, love, and everything else that make up and define the personality toward a specific sex have been masculine since I can remember. Later on, when a person reaches that complicated and difficult age when he discovers the difference between men and women, I suddenly realized that I liked girls very much, but not precisely as friends. I have never practiced homosexuali.sm, which I consider dishonest and abnormal. I have struggled to adapt myself to the behavior of girl friends, in an endeavor to be like them, but I have failed completely. I know it is u.seless to pretend to adjust myself to a feminine life because the problems, behavior, and attitudes of such sex are so strange and far away from me that I could not make them a part of my everyday life I have used lipstick; I have tried to dress differently — more like other girls, but I have not known how to select a more attractive dress; and to go to a dress-maker fills me with an anger that I do not understand. I have tried to widen the circle of my acquaintances by joining a girls' club, but I have dropped out because I cannot bear their small talk, which, even if I respect, I cannot share. I have gone out with some good male friends who have shown themselves to be interested in talking with me about the current things, but I have not been able to dance once, and one night I made myself ridiculous by leaving without any explanations because of my deep feelings of inadequacy "

This patient, like many others seen clinically, displayed a host of signs characteristic of a pervasive identification with a male model. Such a person may be said to have acquired what Brown (1958b) regards as an inverted gender identification. Almost invariably'^ the person with inverted gender identification desires sexual activity with a person of the same anatomic sex and in that sense can be regarded as homosexual in his

^^One exception would seem to be the case of transvestism. Transvestites often eschew homosexual sex activity and may have established a heterosexual adjustment of sorts (Hamburger, Stiirup and Dahl-Iversen, 1953; Benjaniin, 1954).


or her preference of sex partner whether or not homosexual activity has actually takeji place. In this light homosexuals as a group have as their common denominator only an erotic preference for sexual partners of the same anatomic or biologic sex. Brown has proposed the terms inverted homosexual and noninverted homosexual as more accurately describing the psychologic features in the gender role identification of these individuals. Such terminology does have the undoubted advantage of avoiding the imprecision inherent in the older and more commonly used adjectives "active" and "passive," which refer only to erotic behavior as such.

With the identification of the sex hormones, it was doubtless unavoidable that homosexuality, transvestism and other deviant patterns of sexual behavior in humans came to be ascribed, popularly, to an imbalance of androgen or estrogen. Such a belief was fostered and kept alive by reports of studies in lower animals revealing that injection of sex hormones of the opposite sex could evoke inverted sex behavior; that is to say, such animals failed to conform to the usual pattern of sexual aggressiveness or receptiveness and, further, assumed coital positions typical of the ojiposite sex (Young and Rundlett, 1939; Ball, 1940; Beach, 1941, 1942a, 1942b, 1945). But anthropomorphic interpretations are always risky and in this instance have proven deceptive and unwarranted insofar as the psychologic sex disorders in humans is concerned. Studies purporting to demonstrate an excess of estrogen in the urinary androgen-estrogen levels of homosexual men (Glass, Deuel and Wright, 1940) have not been verified (Perloff, 1949) . It is common clinical experience that the treatment of homosexual persons with sex hormones, although often intensifying erotic genital sensation, does not bring about any change in sex-role or psychosexual orientation. This is not particularly surprising because, as previously stated, androgens and estrogens have an indirect, rather than a direct influence on gender role and erotic orientation. There does not seem to be a valid basis for the endorsement of any theory of a simple and direct hormonal determination of human sexual behavior, either typical or atypical.


ONTOGENESIS OF SEXUAL BEHAVIOR


1427


Several investigators have studied homosexuality from the hereditary point of view. Lang (1940) reported findings which suggested to him that male homosexuals were genotypically female; he considered such individuals to be "male sex intergrades which are genetically female but have lost all morphologic sex characteristics except their chromosome formula." Kallman (1952a, b, 1953) expressed himself as substantially in agreement with this, basing his agreement on his findings in twin studies that 60 per cent of the co-twins of homosexuals displayed no evidence of homosexual behavior, whereas 100 per cent of 44 pairs of one egg twins were concordant "as to overt practice and quantitative rating of homosexual behavior after adolescence." Kallman thus concluded that homosexuality is "a gene controlled disarrangement between male and female maturation (hormonal) tendencies," and that "overt homosexual behavior in the adult male may be viewed as an alternative minus variant in the integrative process of psychosexual maturation, comparable in the sexually reproductive human species to the developmental aspects of left handedness in a predominantly right-handed human world."

Pare (1956), using Barr's technicjue for the identification of the sex chromatin pattern of cells, adduced strong evidence against Lang's theory that male homosexuals are genotypically female. In a series of 50 markedly homosexual men (average rating on the Kinsey 6-point rating scale was 4.5), the sex chromatin pattern was male in all cases and the incidence of sex chromatin spots did not differ from that of normal male controls. Barr and Hobbs (1954) reported similar findings in a series of transvestites ; the sex chromatin pattern was always in accord with the external genital morphology. The studies of Bleuler and Wiedemann (1956), Hal)Och (1957), and Raboch and Nedoma <1958) are in agreement with these other investigations. Thus the case is clear that, so far as homosexuality and transvestism are concerned, there is no correlation with the sex chromatin pattern of the body cells. Admittedly, it can still be argued that sex chromatin determinations give no direct information regarding genes and further t'lucidation of


this aspect of the problem must await further developments in cytologic techniques.

In the past several years there is increasing evidence from psychiatric and psychologic research to support a view that homosexuality and certain other disorders of psychologic sex have their origins in social learning. The phenomenon of gender role inversion mentioned earlier provides some insight into some of the factors influencing such atypical and disordered learning. Brown (1958a), for example, postulated that males disj^laying gender role inversion the early years of life characteristically "involve a father who is psychologically ineffective and socially distant so far as the boy is concerned; or a father who is chronically abusive and cruel to the boy. In addition such a family constellation will involve a mother or a mother substitute who is 'idolized' by the boy and to whom he is excessively close and attached." Brown further postulated two additional parent-child relationships which could provide the basis for inversion: (1) "a family in which the same-sex parent himself or herself is i)redominantly or considerably inverted in sexrole structure, thus exposing the child to a distorted model with which to identify," and (2) "a family in which the parent or parents actually encourage and rear a child of one biologic sex to feel and think and behave like the opposite sex."

The clinical findings of Kolb and Johnson (1955) and of Litin, Giffin and Johnson (1956), although couched in the loosely defined language of psychoanalysis, are in essential agreement with Brown's view. These authors explain deviant sex behavior such as homosexuality in terms of "subtle attitudes within the family" developing from "unconscious or, less frequently, conscious fostering of deviant sexual behavior early in life within the family setting."

It is the opinion of the authors of this chapter that neither the purely genetic explanation nor the purely environmental explanation supplies all the answers to the questions loosed by the disorders of psychologic sex. Certainly the evidence of human hermaphroditism points strongly to the tremendous influence of rearing and social learning in the establishment of normal gender role (i)sychologic sexj and, by anal


1428


HORMONAL REGULATION OF BEHAVIOR


ogy, disordered psychologic sex. As stated earlier, however, to make such a statement is not to endorse an oversimplified theory of environmental determinism. The experiences entering into social learning require the active presence of a person — a person whose body morphology, physiologic functioning and cerebral and cognitional equipment are the heritage of genetic factors.^" It can also be allowed that some as yet unidentified inherited perceptual or cognitional factor may reinforce or make it easier for disorders of psychologic sex to evolve during the course of a person's growing up. Nonetheless, life experience and social learning must be accorded an important place in the establishment of psychologic sex, either ordered or disordered, for the evidence militates too strongly against a theory of innate, preformed, and inherited behavioral imperatives, hormonal or otherwise.

^"An important difficulty facing an investigator of genetic factors in behavior is the one of knowing precisely what to look for. Such gross items as "intelligence" or "over-all adaptability," although much studied in the past, have proven inadequate as measures of genie influence on behavior. The work of such students of animal behavior as Lorenz, Beach, W. C. Young, J. P. Scott, Guhl, and others, shows clearly that the problem is not as simple as earlier believed. There seems to be general agreement among such investigators that the inherited aspects of a given type of behavior are simple behavioral units often related to basic physiologic and neurophysiologic processes; for the human, at least, it is evident that such anatomic considerations as body morphology must be added. Lower animals displaj' different capacities to learn and to respond emotionally and physiologically according to the nature of the stimulus; such capacities are clearly related to genetic endowment. An example is the finding by Scott and Charles (1953, 1954) that the ease with which beagles could be taught to track a rabbit, terriers to fight, and spaniels to retrieve and socialize could be broken down into a step-wise sequence of learning processes. Scott (1953) suggested that the study of genie influences on behavior is likely to be most fruitful if approached in terms of these smaller units or processes. In Scott's words: "A gene can act only by modifying some physiologic process, whether it be growth in the embryo or pigment formation in the adult, and clear-cut genetic results can be expected in the study of behavior only where a trait is measured which is either a physiologic process or largely based on one. If behavior is measured in terms of over-all adjustment, dozens and even hundreds of physiologic processes are likely to be involved and these may be organized so that a deficiency in one is cancelled by an excess in another."


B. HERMAPHRODITISM AND PSYCHOPATHOLOGY

No one acquainted with Freudian libido theory^" can fail to be aware that sexuality in the broad sense conceived of by Freud has been accorded a position of great importance in theories concerning the origins of psychiatric disorders. For some, the libido theory constitutes an accepted psychiatric doctrine despite the shaky assumptions (see Bieber, 1958) on which it is based. ' ■

It was in these respects that the evidence of human hermaphroditism was examined by Money, Hampson and Hampson (1956). Representing the several varieties of hermaphroditism, 94 hermaphroditic patients were studied psychologically and psychiatrically, and their psychologic healthiness was judged on a 4-point scale. Of the 94 patients, 63 were evaluated as healthy, 16 mildly nonhealthy, 14 moderately nonhealthy, and one, a mental defective, severely nonhealthy. Functional psychosis was conspicuously absent in this group of hermaphroditic patients. In general their psychologic nonhealthiness was in the direction of excessive inhiI)ition and rcs(M-ve. Commonly these pa '■ Briefly, the libido theory (.see Freud, 1915) assumes the existence of instinctive sexual forces somehow related to "psychic energy" (a dubious concept in any case) and ultimately responsible for all behavior. Freud postulated an "ego" and a "superego" as the psychic apparatus by which libidinal energy is controlled and made socially acceptable and useful. In elaborating his libido theory, Freud envisioned a 3-stage process of psychosexual development consisting of (1) an oral stage related to later dependency relationships, (2) an anal stage related to authority relationships, and (3) a genital stage related to sexuality in the genital erotic sense. P.sychopathologic states, he postulated, were simply regressions to earlier stages of psychosexual, or libidinal organization. Thus Freud considered disordered sexuality, or more precisely, disordered libido, as the primary or basic cause for the neuroses and possibly other psychologic conditions as well. Contrary to views held by other scientists, psychoanalysis has classically held the position that all human emotion, whether affection, warmth, aggression, or anger, has its origins in sexuality as elastically encompassed by the libido theory. Bieber (1958) takes to task those who anachronistically continue to espouse such facile explanatory constructs: "Current psychoanalytic thinking is infiltrated with teleology. We are assigning purpose to behavior facilely. ... A teleologic theory cannot account for all psychologic mental functioning. It is inaccurate and misleading."


tients tended to lag in psychologic growth and personality development relative to their contemporaries of comparable IQ. Spells of depressed moodiness, sullen and anxious bash fulness and shyness, and extreme diffidence and guardedness in matters pertaining to sexual and romantic situations were noted. With respect to the gender role these individuals had established, in 95 per cent there was an equivocal correspondence between gender role and of the sex of assignment and rearing whether or not a contradiction existed between this pair of variables and 1 or more of the other 5 sexual variables. Only 5 of the 94 gave any evidence of psychologic nonhealthiness on grounds of a demonstrably ambiguous gender role and orientation. None of the patients studied was psychotic and none required psychiatric hospitalization. This low incidence of seriously incapacitating mental disorder is corroborated by Money's unpublished survey of the literature on hermaphroditism in which he found a 2 per cent incidence of psychosis among 248 postadolescent hermaphrodites.

In the 94 hermaphroditic patients referred to above, several factors stood out as having important bearing on psychologic healthiness, including psychosexual orientation. The first important factor was the nature and degree of the person's visible genital or secondary sexual ambiguity; the more publicly conspicuous the individual's ambiguous appearance, the more difficult it proved for him to transcend his handicap and come to terms with it psychologically. In view of the importance of body appearance, as discussed earlier, this finding is not surprising. The surprise is that so many ambiguous-looking patients were able, appearance notwithstanding, to grow up and achieve a rating of psychologically healthy, or perhaps only mildly nonhealthy. The healthy rating was certainly more common in the patients whose body morphology, irrespective of gonads and sex chromatin pattern, was unambiguous looking than it was in the l)atients whose sexual appearance was equivocal. Thus, patients whose sex of rearing was contradicted by their gonadal and chromosomal sex were not necessarily destined to be rated nonhealthy.

A second factor of importance in later psychologic adjustment was the consistence


of gender role training and experience which the individual had received during the growing-up years. After the early months of life, reassignment of sex, as commented on in an earlier section, turns out to be extremely conducive to later psychologic nonhealthiness. Corrective surgical, hormonal, or psychologic procedures were of greater benefit if they were instituted during infancy or childhood rather than at a later age. This point is well illustrated by the psychologic difficulties which hyperadrenocortical women in their thirties or forties have experienced when treatment with cortisone permitted, for the first time, some degree of iDelated body feminization. Having already attained some measure of acceptance of their lot as virilized women physically unattractive to men, these women have found themselves gauche, inexpert, and lacking in the self-assurance required for success in the role of girl-friend or wife. As a result episodes of anxiety and depression were common in such patients. By contrast the girls treated in their pre-teen years to permit feminizing body changes, found it considerably easier, psychologically, to negotiate the transition to adolescence and adulthood.

The observation that a multiplicity of factors is involved in the etiology of psychopathology, where it occurs, in hermaphroditic individuals makes clear the danger of embracing pat explanatory concepts, such as those based on libido theory, for the neuroses or any other psychopathy.

VII. Concluding Remarks

On the foregoing pages the authors have considered only a handful of the issues and problems germane to human sexual behavior. Without doubt many other considerations, and many other research contributions could and perhaps should have been included. The reader who is aware of the vast psychiatric and psychologic literature pertaining to sexual functioning in humans will understand the difficulties of a thoroughly comprehensive discussion of the topic in a limited allotment of pages. On the other hand, it is hoped that a central theme will, nonetheless, have been spelled out, a theme first envisioned in the first edition of this book bv Frank Lillie who wrote:


1430


"....There is no such biologic entity as sex. What exists in nature is a dimorphism within species into male and female individuals, which differ with respect to contrasting characters, for each of which in any given species we recognize a male form and a female form, whether these characters be classed as of the biologic, or psychologic, or social orders. Sex is not a force that produces these contrasts; it is merely a name for our total impression of the differences. It is difficult to divest ourselves of the prescientific anthropomorphism which assigned phenomena to the control of personal agencies, and we have been particularly slow in the field of the scientific study of sex-characteristics in divesting ourselves not only of the terminology, but also of the influence of such ideas."

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Young WC. Sex and internal secretions. (1961) 3rd Eda. Williams and Wilkins. Baltimore.
Section A Biologic Basis of Sex Cytologic and Genetic Basis of Sex | Role of Hormones in the Differentiation of Sex
Section B The Hypophysis and the Gonadotrophic Hormones in Relation to Reproduction Morphology of the Hypophysis Related to Its Function | Physiology of the Anterior Hypophysis in Relation to Reproduction
The Mammalian Testis | The Accessory Reproductive Glands of Mammals | The Mammalian Ovary | The Mammalian Female Reproductive Cycle and Its Controlling Mechanisms | Action of Estrogen and Progesterone on the Reproductive Tract of Lower Primates | The Mammary Gland and Lactation | Some Problems of the Metabolism and Mechanism of Action of Steroid Sex Hormones | Nutritional Effects on Endocrine Secretions
Section D Biology of Sperm and Ova, Fertilization, Implantation, the Placenta, and Pregnancy Biology of Spermatozoa | Biology of Eggs and Implantation | Histochemistry and Electron Microscopy of the Placenta | Gestation
Section E Physiology of Reproduction in Submammalian Vertebrates Endocrinology of Reproduction in Cold-blooded Vertebrates | Endocrinology of Reproduction in Birds
Section F Hormonal Regulation of Reproductive Behavior The Hormones and Mating Behavior | Gonadal Hormones and Social Behavior in Infrahuman Vertebrates | Gonadal Hormones and Parental Behavior in Birds and Infrahuman Mammals | Sex Hormones and Other Variables in Human Eroticism | The Ontogenesis of Sexual Behavior in Man | Cultural Determinants of Sexual Behavior


Reference: Young WC. Sex and internal secretions. (1961) 3rd Eda. Williams and Wilkins. Baltimore.


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