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T
his article provides a review of the past and current literature on the neurobiology of
sexual function. The influence of endocrine, neurotransmitter, and central nervous sys-
tem influences on male and female sexual function are discussed for sexual desire, arousal,
and orgasm or ejaculation stages of sexual responding. Endocrine factors reviewed in-
clude the following: androgens, estrogens, progesterone, prolactin, oxytocin, cortisol, and phero-
mones. Neurotransmitters and neuropeptides discussed include nitric oxide, serotonin, dopa-
mine, epinephrine, norepinephrine, opioids, acetylcholine, histamine, and g-aminobutyric acid.
Central nervous system influences on sexual function are discussed briefly with reference to brain-
stem regions, the hypothalamus, and the forebrain. Arch Gen Psychiatry. 2000;57:1012-1030
Within the past decade, increasing re- tending animal findings to the complex ex-
search attention has been paid to the neu- perience of human sexual function.
robiology of sexual function. This has been Wherever possible in the present re-
fostered, in part, by a growing awareness view, the effects of endocrine, neurotrans-
of the deleterious effects of pharmacologi- mitter, and CNS influences are discussed
cal agents on sexual behavior, by an in- separately as they pertain to the sexual re-
creased recognition of the high incidence sponse phases of desire, arousal, and or-
of sexual difficulties present in men and gasm (including ejaculation in males). This
women and, most recently, by the enor- classification of sexual disorders draws
mous success of using sildenifil citrate heavily on Masters and Johnson’s Human
(Viagra) for the treatment of male erec- Sexual Response2 model and Kaplan’s3 tri-
tile dysfunction. In this article, we provide phasic model of sexual response in which
a concise review of the past and current desire, arousal, and orgasm are conceptu-
literature on the endocrine, neurotrans- alized as distinct and sequential phases. In
mitter, and central nervous system (CNS) actual clinical practice, however, sexual de-
influences on male and female sexual func- sire, arousal, and orgasm difficulties more
tion. We would like to acknowledge the often than not coexist—suggesting an in-
enormity of the field at the outset, and em- tegration of phases, and desire does not
phasize that this article is meant as a broad necessarily precede arousal—arousal re-
overview of the field. Wherever appli- sponses may also ignite desire (for a re-
cable, within each section, we refer the view of problems associated with this clas-
readers to more in-depth, specialized re- sification system in women, see Leiblum4).
views. While the focus of this article is on
human research, in areas such as the brain STAGES OF HUMAN SEXUAL
localization of sexual function where little RESPONSE
human data exist, we also briefly summa-
rize the findings from the animal litera- Sexual desire is commonly defined as the
ture (for a more detailed review of the ani- broad interest in sexual objects or expe-
mal literature in this field, see Pfaus1). One riences. Because there is no objective physi-
must, of course, exercise caution when ex- ological criterion for desire, it is gener-
ally inferred by self-reported frequency of
sexual thoughts, fantasies, dreams, wishes,
From the Department of Psychology, University of Texas, Austin. and interest in initiating and/or engaging
Males Females
*↑ indicates increase; ↓, decrease; 0, no change; DHEA, dehydroepiandrosterone; ellipsis, not applicable; HRT, hormone replacement therapy; and ICI, intracavernous
injection.
Males Females
(Continued)
Males Females
*↑ indicates increase; ↓, decrease; ellipsis, not applicable; MAOI, monoamine oxidase inhibitor; SSRI, selective serotonin reuptake inhibitor; 0, no change;
HSDD, hypoactive sexual desire disorder; and ED, erectile dysfunction.
vasocongestion and lubrication in fe- arousal disorder is under way. Find- a1-receptors produces erection. In
males and this may adversely influ- ings from animal studies suggest se- women, by contrast, adrenergic ac-
ence sexual arousal and desire. Find- rotonin may facilitate, inhibit, or tivation facilitates vasocongestion
ings from uncontrolled studies and have no effect on sexual behavior de- and suppression of adrenergic ac-
animal studies tentatively suggest pending on which serotonin recep- tivity impairs sexual arousal and or-
prolactin may have an inhibitory in- tor subtype is involved. Studies on gasm. Norepinephrine levels in-
fluence on drive in males and fe- the effects of antidepressants on hu- crease during sexual arousal in men
males. Controlled human studies man sexual function suggest activa- and women. Minimal research sug-
suggest the levels of prolactin and tion of the serotonin2 receptor im- gests increasing the level of NE may
oxytocin increase during sexual pairs all stages of the sexual response facilitate erectile responding in men;
arousal in men and women. Abnor- in males and females. Case reports comparable studies have not been
mally high levels of cortisol de- in males showing a facilitatory in- conducted in women. Long-term
crease sexual drive in men and fluence of antiparkinsonian medi- opioid use impairs erection in men
women possibly owing to in- cations (which enhance dopamine possibly via suppression of circulat-
creased corticotropin-releasing hor- activity) and an inhibitory influ- ing hormones such as testosterone.
mone. Minimal research on phero- ence of antipsychotic medications Case reports indicate opioid antago-
mones and sexual response suggest (which suppress dopamine activ- nists may restore erectile function-
a facilitatory influence on sexual at- ity) on desire and erection argue for ing in dysfunctional men. Limited
tractiveness in men. a facilitatory influence of dopamine studies suggest opioids have an
Nitric oxide (via the conver- on male sexual behavior. Research analogous effect in women. Acetyl-
sion of guanosine triphosphate to in male rats indicating dopamine fa- choline facilitates penile erection via
cGMP) is essential for penile and cilitates sexual drive, erection, and the relaxation of smooth muscles of
possibly clitoral vasocongestion. ejaculation corroborates these hu- the corpus cavernosum. The role of
Sildenafil prolongs the action of man findings. Limited research con- acetylcholine in female vasoconges-
cGMP and is effective in treating ducted in females suggests a facili- tion is unknown. Case studies sug-
erectile dysfunction of organic, psy- tatory role of dopamine on sexual gest histamine facilitates erection in
chogenic, and mixed causes. Re- desire and orgasm. Adrenergic ac- men with erectile failure. One com-
search on the effectiveness of silde- tivity (ephedrine) inhibits erectile re- parable case study has been re-
nafil treatment for female sexual sponding in men and blockade of ported in women. The H2 and pos-