Vasotocin stimulates appetitive responses to the visual and pheromonal stimuli used by male roughskin newts during courtship

It is now well established that vasotocin (AVT) and its mammalian homologue vasopressin influence various social behaviors in vertebrates, but less is known about the mechanisms through which these peptides modulate behavior. In male roughskin newts, Taricha granulosa, AVT stimulates a courtship behavior, amplectic clasping. Three general explanations for how AVT affects male courtship behavior have been considered: by enhancing a central state of sexual motivation, by affecting sensorimotor integration mechanisms in individual sensory modalities, or by influencing a nonspecific state of attention, arousal, or anxiety. AVT administration enhanced appetitive responses to visual and olfactory sexual stimuli, as would be expected if AVT affects a state of sexual motivation that affects behavioral responses to sexual stimuli regardless of the sensory modality in which they are processed. However, AVT selectively enhanced responses to female olfactory stimuli (sex pheromones), but similarly enhanced responses to female and food-related visual stimuli (worms), thus questioning the utility of such a motivational mechanism, as responses to female stimuli were not selectively enhanced in all sensory modalities. We therefore propose that exogenous AVT independently influences olfactory processes associated with orientation/attraction toward a female sex pheromone and visual processes associated with orientation/attraction toward a visual feature common to females and worms. In further experiments AVT administration failed to stimulate feeding behavior but did decrease locomotor activity. Thus, AVT does not stimulate courtship behavior in this species by enhancing the animals' general state of attention or by decreasing general anxiety, as responses to nonsexual, attractive stimuli were not uniformly enhanced, nor by stimulating general arousal, as activity levels did not increase. Rather, the data support the conclusion that AVT affects courtship by influencing specific sensorimotor processes associated with behavioral responses to individual releasing stimuli, which suggests a mechanistic framework for understanding socially motivated behavior is this species.     

Sex steroids and vasotocin interact in a female amphibian (Taricha granulosa) to elicit female-like egg-laying behavior or male-like courtship.

Female egg-laying behaviors and male amplectic clasping behaviors in the rough-skinned newt (Taricha granulosa) are similar in that animals clasp an object. In the case of egg-laying, females clasp submerged inanimate objects, whereas in amplexus, males clasp conspecific females. Considering these behavioral similarities and differences, we investigated the possibility that gonadal steroids and vasotocin (AVT) interact to control egg-laying behaviors, as has been shown for the control of amplexus in Taricha males. Intact, gravid T. granulosa females injected ip with AVT, unlike those injected with saline, exhibited egg-laying behaviors and oviposition. In ovariectomy-steroid-implant studies, no saline-injected female exhibited egg-laying behaviors, whereas AVT-injected ovariectomized females exhibited egg-laying behaviors if implanted with estradiol (E2), testosterone, or dihydrotestosterone (DHT), and not if implanted with empty capsules. When given a choice between clasping aquatic vegetation or other females (amplectic clasping), following an AVT injection, unoperated and sham-operated control females and ovariectomized females with E2 implants did not preferentially clasp aquatic vegetation over other females. In contrast, AVT-injected ovariectomized females with DHT implants preferentially clasped other females. Thus, exposure of Taricha females to estrogens or androgens appears to determine whether the AVT-induced clasping is egg-laying or amplectic clasping.     

Androgens and male mating behavior in rough-skinned newts, Taricha granulosa

Cyproterone acetate was administered either orally or intraperitoneally to intact, adult male newts, Taricha granulosa. The number of males that exhibited the courtship behavior of clasping when tested with nuptial females was not altered by the antiandrogen treatments. In males which were unresponsive to nuptial females, the occurrence of clasping was not evoked by injections for 4 days of testosterone, dihydrotestosterone, or 11-ketotestosterone. Further, the incidence of clasping was not significantly elevated by injections of prolactin and/or testosterone for 30 days. The effect of sexual activity on testosterone and dihydrotestosterone levels in male newts was determined by radioimmunoassay of plasma collected from males which were: (1) isolated from females; (2) allowed to clasp a female for 2 min; or (3) allowed to clasp a female for 1 hr. The testosterone and dihydrotestosterone levels were unchanged during this period of clasping. In February and again in June, plasma androgen concentrations were measured in males which differed in their propensity to initiate courtship when paired with females. Androgen levels were similar for males that clasped a female and males that never attempted to clasp a female. Plasma androgen levels in the male newt are apparently not correlated with sexual responsiveness.     

Duration of androgen treatment modifies behavioral response to arginine vasotocin in Taricha granulosa

Five and eighteen days after surgery, injection of arginine 8-vasotocin (AVT) stimulated clasping behavior in castrated controls (unimplanted or cholesterol-implanted) but not in androgen-implanted, castrated newts (Taricha granulosa). Conversely, AVT injected 33 days after castration increased the incidence of courtship behavior of androgen-implanted males, but not of unimplanted castrates. Plasma androgen concentration of androgen-implanted castrates (as determined by radioimmunoassay) was maintained at a level typical of intact males at the peak of the breeding season. Because the AVT-androgen interaction could occur at the level of the pituitary gland, castrated, androgen-implanted newts were hypophysectomized and injected with AVT. Hypophysectomy did not abolish the behavioral response to AVT.     

Sexual differentiation of pheromone processing: Links to male-typical mating behavior and partner preference

Phoenix et al. (Phoenix, C., Goy, R., Gerall, A., Young, W., 1959. Organizing actions of prenatally administered testosterone propionate on the tissues mediating mating behavior in the female guinea pig. Endocrinology 65, 369–382.) were the first to propose an essential role of fetal testosterone exposure in the sexual differentiation of the capacity of mammals to display male-typical mating behavior. In one experiment control male and female guinea pigs as well as females given fetal testosterone actually showed equivalent levels of mounting behavior when gonadectomized and given ovarian steroids prior to adult tests with a stimulus female. This finding is discussed in the context of a recent, high-profile paper by Kimchi et al. (Kimchi, T., Xu, J., Dulac, C., 2007. A functional circuit underlying male sexual behaviour in the female mouse brain. Nature 448, 1009–1014.) arguing that female rodents possess the circuits that control the expression of male-typical mating behavior and that their function is normally suppressed in this sex by pheromonal inputs that are processed via the vomeronasal organ (VNO)-accessory olfactory nervous system. In another Phoenix et al. experiment, significantly more mounting behavior was observed in male guinea pigs and in females given fetal testosterone than in control females following adult gonadectomy and treatment with testosterone. Literature is reviewed that attempts to link sex differences in the anatomy and function of the accessory versus the main olfactory projections to the amygdala and hypothalamus to parallel sex differences in courtship behaviors, including sex partner preference, as well as the capacity to display mounting behavior.     

Visual sex discrimination in goldfish: seasonal, sexual, and androgenic influences

The olfactory signals used by goldfish for sexual and aggressive communication have been studied extensively, but little work has addressed the role of other sensory modalities in social communication in this species. We therefore investigated the role that visual stimuli play in sex discrimination and the ability of androgens, which masculinize courtship behavior, to affect behavioral responses toward female visual stimuli. We found that males selectively orient toward female visual stimuli during the breeding season but not outside it, whereas prostaglandin F2-alpha (PGF2alpha)-injected females do not differentially approach male and female visual stimuli, even during the breeding season. Implanting adult females with testosterone (T) and 11-ketotestosterone (KT), however, induced orientation responses toward female visual stimuli similar to those observed in males. These results indicate that visual sexual stimuli are likely important for reproductive signaling in goldfish, potentially helping males identify ovulating females from a distance in a shoal of fish, and that androgens can influence mechanisms associated with orientation responses toward such stimuli.     

Historical perspective: Hormonal regulation of behaviors in amphibians

This review focuses on research into the hormonal control of behaviors in amphibians that was conducted prior to the 21st century. Most advances in this field come from studies of a limited number of species and investigations into the hormonal mechanisms that regulate reproductive behaviors in male frogs and salamanders. From this earlier research, we highlight five main generalizations or conclusions. (1) Based on studies of vocalization behaviors in anurans, testicular androgens induce developmental changes in cartilage and muscles fibers in the larynx and thereby masculinize peripheral structures that influence the properties of advertisement calls by males. (2) Gonadal steroid hormones act to enhance reproductive behaviors in adult amphibians, but causal relationships are not as well established in amphibians as in birds and mammals. Research into the relationships between testicular androgens and male behaviors, mainly using castration/steroid treatment studies, generally supports the conclusion that androgens are necessary but not sufficient to enhance male behaviors. (3) Prolactin acts synergistically with androgens and induces reproductive development, sexual behaviors, and pheromone production. This interaction between prolactin and gonadal steroids helps to explain why androgens alone sometimes fail to stimulate amphibian behaviors. (4) Vasotocin also plays an important role and enhances specific types of behaviors in amphibians (frog calling, receptivity in female frogs, amplectic clasping in newts, and non-clasping courtship behaviors). Gonadal steroids typically act to maintain behavioral responses to vasotocin. Vasotocin modulates behavioral responses, at least in part, by acting within the brain on sensory pathways that detect sexual stimuli and on motor pathways that control behavioral responses. (5) Corticosterone acts as a potent and rapid suppressor of reproductive behaviors during periods of acute stress. These rapid stress-induced changes in behaviors use non-genomic mechanisms and membrane-associated corticosterone receptors.     

Hormone effects on male sex behavior in adult South African clawed frogs, Xenopus laevis

Intact, adult male Xenopus laevis were injected with human chorionic gonadotropin (HCG) and tested with intact HCG-primed females. Under these conditions, males displayed high levels of sex behavior (clasping of females). By 2 weeks after castration, these males had ceased clasping. Testosterone and dihydrotestosterone reinstated clasping in male castrates. Following removal of testosterone or dihydrotestosterone pellets, castrated males ceased to clasp. No male was ever observed to clasp following estradiol implanted in pellets or in silastic capsules. In experiments on castrated, adult, female Xenopus laevis, both testosterone and testosterone propionate pellets reliably produced male sex behavior in the form of clasping. The clasping of testosterone-implanted female and male castrates was very similar in form and duration. The behavioral effectiveness of testosterone in both sexes and the ineffectiveness of estradiol in eliciting clasping is paralleled by autoradiographic localization of sex steroids in brain where the distribution of testosterone-concentrating, cells is the same for males and females, but different from the distribution of estradiol-concentrating cells.     

Multiple mechanisms of phenotype development in the bluehead wrasse

Despite having detailed information on mechanisms mediating sex-typical behavior in many species, we have little understanding of whether the same mechanisms regulate these behaviors when they are performed in the same species under different social contexts. In the five field experiments of this study of bluehead wrasses (Thalassoma bifasciatum), a sex-changing fish, we examined the roles of arginine vasotocin (AVT) and the potent teleost androgen 11-ketotestosterone (11KT) in mediating sexual and aggressive behaviors typical of dominant males. We demonstrated that AVT appears necessary for the assumption of dominant territorial status in males and females, but is sufficient only in the socially dominant terminal phase (TP) male phenotype. Specifically, an AVP V(1) receptor antagonist prevented both TP males and females from gaining dominance over recently vacated territories. However, unlike TP males in a previous study, neither females nor initial phase males responded to AVT treatment with increases in display of TP male typical behaviors when under social conditions that inhibit sex change. Treating females with 11KT did not alter responsiveness to AVT, but did induce male coloration and courtship behavior that was not observed in oil-treated females. Combined with the results of a previous study, these results indicate that the ability of AVT to induce male-typical behavior differs among sexual phenotypes and that this differential responsiveness appears to be dependent on social context and not directly on exposure to 11KT. Furthermore, since 11KT can induce courtship behavior in females that is not affected by AVT, there may be different hormonal mechanisms mediating courtship behavior under different social contexts.     

Brain arginine vasotocin concentrations related to sexual behaviors and hydromineral balance in an amphibian

Arginine vasotocin (AVT), a potent stimulator of sexual behaviors and regulator of hydromineral balance in male rough-skinned newts (Taricha granulosa), was measured in 11 brain areas using microdissection and radioimmunoassay procedures. A 60-min test for sexual behaviors was used to segregate males into two groups: sexually responsive (initiated amplectic clasping behaviors) and sexually unresponsive (exhibited no sexual behaviors). Compared to sexually unresponsive males, sexually responsive males had significantly higher concentrations of immunoreactive (ir) AVT in the dorsal preoptic area, optic tectum, ventral infundibular nucleus, and cerebrospinal fluid. These results provide evidence for a behavioral action of endogenous AVT in T. granulosa. In another study, irAVT was measured in normal males (control newts maintained in water) and males that were dehydrated for 6 hr. Compared to normal males, dehydrated males had significantly lower concentrations of irAVT in the ventral preoptic area, but not in the other 10 areas of the brain. That different brain areas are associated with sexual behaviors and hydromineral balance suggests that there are some neuroanatomical separations between the behavioral and hydromineral aspects of the vasotocinergic system in this amphibian.     

Arginine vasotocin induces sexual behavior of newts by acting on cells in the brain

To investigate whether arginine vasotocin (AVT) acts on target cells in the brain of Taricha granulosa (a urodele amphibian), the behavioral effects of intracerebroventricular (ICV) and intraperitoneal (IP) injections of AVT were compared. Male newts exhibited the greatest sexual activity (amplectic clasping) following an ICV injection of 0.1 μg AVT. Another study showed that nanogram quantities of AVT, administered ICV, stimulated the behavior. An ICV injection of an antagonist to arginine vasopressin, d(CH₂)₅Tyr(Me)AVP, or an anti-AVT immune serum significantly inhibited the sexual behavior. Intracranial implants of 17β-estradiol (E₂) or 5α-dihydrotestosterone (DHT) in castrated males maintained the behavioral response to an injection of AVT. Another study found that an IP injection of DHT or E₂ did not increase the incidence of newt sexual behavior during the 8 hours following the injection.     

Methyl-testosterone induces male-typical ventilatory behavior in response to putative steroidal pheromones in female round gobies (Neogobius melanostomus)

Male round gobies (Neogobius melanostomus) transiently increase their rate of ventilation when exposed to nanomolar concentrations of three putative steroidal pheromones (etiocholanolone, ETIO; estrone, E1; estradiol-glucuronide, E2-3g), whereas females exhibit this response only to ETIO. In this study we implanted females with Silastic capsules that were empty (Blank) or filled with methyl-testosterone (MT) to determine whether androgen induces a male-typical pattern of ventilatory response. As with untreated females in our previous studies, Blank females increased ventilation only in response to ETIO. In contrast, MT females tested 2-3 weeks postimplant responded not only to ETIO, but also to E1 and E2-3g. These results provide the first demonstration of an androgen inducing a male-typical behavioral response to a steroidal pheromone in adult female fish. Together with our previous findings of sexually isomorphic sensory responses to steroid odorants, the present results also suggest that a central action of androgen is responsible for sexually dimorphic behavioral responses to putative pheromonal steroids in Neogobius.     

Androgen Induction of Male Sexual Behaviors in Female Goldfish

The effectiveness of testosterone (T) and 11-ketotestosterone (K) in inducing male-typical sex behaviors in goldfish was examined by implanting intact adult females with one empty (blank) Silastic implant (B females), one implant containing T or K, or one T and one K implant (T + K females). Behavior of the four female groups was compared to that of untreated males and males containing a blank implant. Male-typical behaviors (coutship, spawning) and associated behavioral changes (increased activity, reduced spontaneous feeding) were assessed 3.5 and 4.5 months after implant in 30-min tests in which the test female or male was allowed to interact with a stimulus female in which sexual receptivity and attractivity had been induced by acute prostaglandin F_2αinjection. Prostaglandin-induced female-typical spawning behavior in the test females and males was also assessed 4.5 months after implant in a 60-min test for female-typical behavior in which the test fish was injected with prostaglandin and placed immediately with a sexually active male. Blood samples 5 months postimplant showed that implants generated physiological levels of T and K. In both tests for male-typical behaviors, K and T + K females exhibited the full suite of behaviors shown by spawning males, e.g., male-typical courtship and spawning, increased swimming activity, and reduced spontaneous feeding. Although behaviors of K and T + K females did not differ, those of T + K females were more often equivalent to those of males and significantly different from those of B females. T females exhibited marginal male-typical behaviors which never differed significantly from those of B females. Androgen-treated females exhibited female-typical; spawning behaviors equivalent to that of males and B females. The results show that adult female goldfish can be behaviorally masculinzed without behavioral defeminization, and suggest that male-typical sex behaviors in goldfish are dependent on K, although other steroids also may be required. The inducible behavioral bisexuality of goldfish, a gonochoristic species, is discussed in terms of the prevalence of hermaphroditism in teleosts.     

Courtship behaviour in Japanese macaques during heterosexual and homosexual consortships

Female Japanese macaques (Macaca fuscata) in the Arashiyama population near Kyoto, Japan, are unusual, in that they exhibit what many would consider to be male-typical sexual characteristics. Specifically, they mount other females within the context of temporary, but exclusive, sexual relationships (i.e., homosexual consortships) and they sometimes exhibit a preference for female sexual partners, even when given the choice of a sexually motivated male alternative. In this study, we examined whether female Japanese macaques also exhibited male-typical patterns of courtship behaviour during homosexual consortships. Data were collected on courtship behaviour during heterosexual and homosexual consortships in free-ranging Japanese macaques from the Arashiyama (Japan) population. We analyzed the occurrence of 12 different courtship behaviours during 3374 heterosexual inter-mount intervals and 1412 homosexual inter-mount intervals. Sex differences between heterosexually consorting males and females existed for only two of the 12 courtship behaviours we investigated: inclined-back presentations and sexual vocalizations. Dominant and subordinate homosexually consorting females were sex-typical in their expression of inclined-back presentations and sexual vocalizations. Consequently, facultative same-sex sexual partner preference, mounting and consortships do not co-occur with male-typical patterns of courtship behaviour in female Japanese macaques.     

Love Is Blind: Indiscriminate Female Mating Responses to Male Courtship Pheromones in Newts (Salamandridae)

Internal fertilization without copulation or prolonged physical contact is a rare reproductive mode among vertebrates. In many newts (Salamandridae), the male deposits a spermatophore on the substrate in the water, which the female subsequently takes up with her cloaca. Because such an insemination requires intense coordination of both sexes, male newts have evolved a courtship display, essentially consisting of sending pheromones under water by tail-fanning towards their potential partner. Behavioral experiments until now mostly focused on an attractant function, i.e. showing that olfactory cues are able to bring both sexes together. However, since males start their display only after an initial contact phase, courtship pheromones are expected to have an alternative function. Here we developed a series of intraspecific and interspecific two-female experiments with alpine newt (Ichthyosaura alpestris) and palmate newt (Lissotriton helveticus) females, comparing behavior in male courtship water and control water. We show that male olfactory cues emitted during tail-fanning are pheromones that can induce all typical features of natural female mating behavior. Interestingly, females exposed to male pheromones of their own species show indiscriminate mating responses to conspecific and heterospecific females, indicating that visual cues are subordinate to olfactory cues during courtship.     

Discrimination of conspecific sex and reproductive condition using chemical cues in axolotls (<Emphasis Type="Italic">Ambystoma mexicanum</Emphasis>)

Chemosensory cues play an important role in the daily lives of salamanders, mediating foraging, conspecific recognition, and territorial advertising. We investigated the behavioral effects of conspecific whole-body odorants in axolotls, Ambystoma mexicanum, a salamander species that is fully aquatic. We found that males increased general activity when exposed to female odorants, but that activity levels in females were not affected by conspecific odorants. Although males showed no difference in courtship displays across testing conditions, females performed courtship displays only in response to male odorants. We also found that electro-olfactogram responses from the olfactory and vomeronasal epithelia were larger in response to whole-body odorants from the opposite sex than from the same sex. In males, odorants from gravid and recently spawned females evoked different electro-olfactogram responses at some locations in the olfactory and vomeronasal epithelia; in general, however, few consistent differences between the olfactory and vomeronasal epithelia were observed. Finally, post hoc analyses indicate that experience with opposite-sex conspecifics affects some behavioral and electrophysiological responses. Overall, our data indicate that chemical cues from conspecifics affect general activity and courtship behavior in axolotls, and that both the olfactory and vomeronasal systems may be involved in discriminating the sex and reproductive condition of conspecifics.Abbreviations EOG electro-olfactogram - VNO vomeronasal organ     

Activation of sexual behavior by implantation of testosterone propionate and estradiol benzoate into the preoptic area of the male Japanese quail (Coturnix japonica)

Intracranial implantation of minute pellets of gonadal steroids was performed to determine neuroanatomical loci at which steroids activate sexual behavior in the Japanese quail (Coturnix japonica). In this species, systemic treatment of castrated males with either testosterone propionate (TP) or estradiol benzoate (EB) restores male-typical copulatory behavior (head grabbing, mounting, and cloacal contact movements). In addition, EB activates female-typical receptive behavior (crouching). Adult male castrated quail were implanted intracranially with 300-micrograms pellets containing TP, EB, or cholesterol (CHOL) and behavior was tested with intact males and females. Either TP or EB pellets in the preoptic area (POA) activated male-typical copulatory behavior. Mounting was specifically activated without concomitant activation of other steroid-sensitive sexual and courtship behaviors. TP and EB implants in adjacent nuclei containing receptors for these steroids and CHOL implants in POA had no effect on male-typical copulatory behavior. Eighteen percent of all males tested for female-typical receptivity crouched, but no specific effect of EB was seen at any site. The similarity of the POA sites for activation of mounting by TP and EB is consistent with the hypothesis that cells within the POA aromatize testosterone to estrogens, which directly stimulate the cellular processes leading to behavioral activation.     

Rapid, Corticosterone-Induced Disruption of Medullary Sensorimotor Integration Related to Suppression of Amplectic Clasping in Behaving Roughskin Newts (Taricha granulosa)

Endogenously secreted or injected corticosterone (CORT) rapidly suppresses courtship clasping in male roughskin newts (Taricha granulosa) by an action on a specific neuronal membrane receptor. Previous studies, using immobilized newts, showed that CORT administration rapidly depresses excitability of reticulospinal neurons and attenuates medullary neuronal responsiveness to clasp-triggering sensory stimuli. The present study used freely moving newts to examine clasping responses and concurrently record sensorimotor properties of 67 antidromically identified reticulospinal and other medullary reticular neurons before and after CORT injection. Before CORT, reticulospinal neurons fired in close association with onset and offset of clasps elicited by cloacal pressure. Reticulospinal neurons also showed firing correlates of nonclasping motor events, especially locomotion. Neuronal activity was typically reduced during clasping and elevated during locomotion. Medullary neurons that were not antidromically invaded (unidentified neurons) usually showed sensorimotor properties that resembled those of reticulospinal neurons. Intraperitoneal CORT (but not vehicle) reduced the probability and quality of hindlimb clasping in response to cloacal pressure, especially within 5–25 min of injection. Simultaneously, responses of reticulospinal and unidentified neurons to cloacal pressure and occurrence of clasping-related activity were attenuated or eliminated. CORT effects were relatively selective, altering clasping-related neuronal activity more strongly than activity associated with nonclasping motor events. The properties of CORT effects indicate that the hormone impairs clasping by depressing processing of clasp-triggering afferent activity and by disrupting the medullary control of clasping normally mediated by reticulospinal neurons. The rapid onset of these CORT effects implicates a neuronal membrane receptor rather than genomic action of the steroid.     

Intersexual cooperation during male clasping of external female genitalia in the spider Physocyclus dugesi (Araneae, Pholcidae)

Sexual conflict may influence the shape and evolution of body structures that males use to grasp females during mating. Not only sexual coercion but also intersexual cooperation may be involved during clasping behavior. Among pholcid spiders, secondary sexual modifications of the male chelicerae, such as apophyses with spines or tooth-like processes, function to grasp the female by specific parts of her external genitalia such as grooves or apophyses of the epigynum. We analyzed how the female and the male respond when their structures for clasping are experimentally modified in the pholcid Physocyclus dugesi. We used three treatment groups for virgin females that differed in the manipulation of the epigynum apophyses (uncovered, partially covered, and fully covered by a plaster) and two groups of males (uncovered and fully covered cheliceral apophyses). We found that females are mainly cooperative to courting males not only when the female genital apophyses were experimentally covered but also when the male cheliceral apophyses were covered. The current data also indicate behavioral flexibility in males during courtship, especially when they had difficulty in genital intromission. Our experimental results, together with previous observational studies, support a modulated-cooperative scenario between the sexes for cheliceral clasping and genital intromission in pholcid spiders.     

A neurobehavioral model for rapid actions of corticosterone on sensorimotor integration.

Stress-induced corticosterone (CORT) secretion that causes a rapid blockade of courtship clasping by male roughskin newts (Taricha granulosa) is mediated by a specific neuronal membrane receptor for CORT. Amplectic clasping, which can be triggered by pressure on the ventral body surface and cloaca, is controlled by the influence of medullary neurons on the spinal cord. Using clasping as a simple neurobehavioral model, we have focused our analysis of CORT effects on clasping by examining the steroid's effects on neurophysiological properties of medullary neurons, especially medullary reticulospinal neurons, the principal output cells from the brain to the spinal cord. Systemic CORT caused, within 3 min of injection, diverse reductions in reticulospinal neuron excitability. Another rapid CORT effect on medullary neurons was to depress responsiveness to pressure on the cloaca. Experiments with chronically implanted, freely moving newts revealed that the rapid CORT effects are quite specific to neural processes related to clasping. CORT injections rapidly blocked clasping in response to cloacal stimuli and concurrently depressed neuronal responses to cloacal pressure and firing associated with clasping. Activity of reticulospinal neurons was often associated with nonclasping movements and this activity was rarely altered by CORT. Thus, CORT mainly affected aspects of neuronal function related to clasping. In other neurophysiological experiments, we found that the neuropeptides vasotocin and corticotropin-releasing hormone modified the neural effects of CORT. Prior exposure of medullary neurons to either of these neuropeptides caused systemic CORT administration to rapidly potentiate neuronal responses to cloacal stimuli, indicating that the direction and potency of CORT effects depend critically on the prevailing neuroendocrine state of the brain.