The ultrasonic postejaculatory vocalization and the postejaculatory refractory period of the male rat.

After ejaculation, the male rat emits an ultrasonic (22-kHz.) vocalization. This sound is produced repeatedly until about three fourths of the postejaculatory interval has elapsed. In this study, the occurrence of the vocalization was described, and physiological and behavioral evidence was presented that the postejaculatory vocalization reflects an inhibitory state that underlies the postejaculatory refractory period. The vocalization period was characterized by a predominance of slow-wave, spindling, sleep-like electroencephalographic activity. Electrical shock was able to stimulate mating responses only after the cessation of the vocalization period. It was concluded that an absolute refractory period of the postejaculatory interval lasts until the end of the vocalization period and that the time from the termination of the vocalization until the resumption of mating is a relative refractory period.     

Testing social acoustic memory in rats: Effects of stimulus configuration and long-term memory on the induction of social approach behavior by appetitive 50-kHz ultrasonic vocalizations

Rats emit distinct types of ultrasonic vocalizations (USVs), which serve as situation-dependent affective signals. In appetitive situations, such as rough-and-tumble-play, high-frequency 50-kHz USVs occur, whereas low-frequency 22-kHz USVs can be observed in aversive situations, such as social defeat. USVs serve distinct communicative functions and induce call-specific behavioral responses in the receiver. While aversive 22-kHz USVs serve as alarm calls and induce behavioral inhibition, appetitive 50-kHz USVs have a pro-social communicative function and elicit social approach behavior, supporting the notion that they serve as social contact calls to (re)establish or maintain contact among conspecifics. The aim of the present study was to use the rat's ability to communicate in the ultrasonic range via high-frequency 50-kHz USVs in order to develop a test for social acoustic memory in rats with relevance for human verbal memory. Verbal learning and memory is among the seven cognitive domains identified as commonly deficient in human schizophrenia patients, but particularly difficult to model. We therefore tested whether the induction of social approach behavior by playback of appetitive 50-kHz USVs is dependent on (1) acoustic stimulus configuration and (2) social long-term memory, and whether (3) social long-term memory effects can be blocked by the administration of scopolamine, a muscarinic acetylcholine antagonist producing amnesia. Results show that social approach behavior in response to playback of natural 50-kHz USVs depends on acoustic stimulus configuration and occurs only when sound energy is concentrated to a critical frequency band in the ultrasonic range. Social approach behavior was detected during the first exposure to playback of 50-kHz USVs, whereas no such response was observed during the second exposure 1week later, indicating a stable memory trace. In contrast, when memory formation was blocked by i.p. administration of scopolamine (0.5mg/kg or 1.5mg/kg) immediately after the first exposure, rats displayed social approach behavior during the second exposure as well. Induction of social approach behavior in response to repeated playback of natural 50-kHz USVs may therefore provide a new and rather unique approach for testing social acoustic memory in rats with relevance to human verbal memory.     

Analysis of ultrasonic vocalizations emitted by intruders during aggressive encounters among rats (Rattus norvegicus)

This investigation was concerned with the identification of the ultrasonic vocalizations produced by intruders during aggressive interactions and the role of these signals in agonistic behavior of rats. In the first experiment, experienced resident males were paired with both devocalized and intact vocalizing naive intruder males. Devocalization of the intruder males resulted in a drastic decrease in 50-kHz vocalizations and the elimination of all 22-kHz vocalizations. This almost total absence of ultrasonic vocalizations was not accompanied by any change in resident aggressive behavior or intruder defensive and submissive behavior. In a second experiment, naive intruders were tested with either deafened or intact resident males. Similarly, preventing residents from hearing intruder ultrasounds had no detectable effect on any aggressive behavior. These experiments are not consistent with the correlative evidence that intruder-produced 22-kHz vocalizations inhibit the aggressive behavior of the resident. The results also show that most of the ultrasonic vocalizations emitted during aggressive encounters are probably produced by the intruder.     

Ultrasonic vocalizations as indices of affective states in rats

Adult rats spontaneously vocalize in ultrasonic frequencies. Although these ultrasonic vocalizations (USVs) have been described as by-products of locomotor activity or social signals, accumulating evidence suggests that they may also index anticipatory affective states. Converging ethological, pharmacological, and brain stimulation research indicates that whereas long low-frequency (> 0.3-s, approximately 22-kHz) USVs occur during anticipation of punishment or avoidance behavior, short, high-frequency (< 0.3-s, approximately 50-kHz) USVs typically occur during anticipation of reward or approach behavior. Thus, long 22-kHz USVs may index a state of negative activation, whereas short, 50-kHz USVs may instead index a state of positive activation. This hypothesis has theoretical implications for understanding the brain circuitry underlying mammalian affective states and clinical applicability for modeling hedonic properties of different psychotropic compounds.     

Juvenile stress potentiates aversive 22-kHz ultrasonic vocalizations and freezing during auditory fear conditioning in adult male rats

Traumatic experiences that occur during adolescence can render individuals vulnerable to mood and anxiety disorders. A model in juvenile rats (age: 27-29 days) was developed previously to study the long-term effects of adolescent stress exposure on behaviour and physiology. This paradigm, termed juvenile stress, involves subjecting juvenile rats to different stressors on consecutive days over a 3-day period. Here, we investigated the effects of the juvenile stress paradigm on freezing behaviour and aversive 22-kHz ultrasonic vocalizations (USVs) during auditory fear conditioning in adult male rats (age: 68-90 days). We found that rats previously subjected to juvenile stress increased aversive 22-kHz USVs (total calls and time spent calling) compared with controls during fear-conditioning training. The acoustic USV parameters between control and juvenile stress rats were largely equivalent, including duration, peak frequency and amplitude. While rats did not differ in freezing behaviour during fear conditioning, juvenile stress rats exhibited greater cue-conditioned freezing upon testing 24 h later. Our results show that juvenile stress elicited different long-term changes in freezing and aversive USVs during fear conditioning. Furthermore, they highlight the importance of assessing USVs to detect experience-dependent differences between control and stress-exposed animals which are not detectable by measuring visible behaviour.     

Overt behavior and ultrasonic vocalization in a fear conditioning paradigm: a dose-response study in the rat

The behavioral analysis of laboratory rats is usually confined to the level of overt behavior, like locomotion, behavioral inhibition, instrumental responses, and others. Apart from such visible outcome, however, behaviorally relevant information can also be obtained when analyzing the animals' ultrasonic vocalization, which is typically emitted in highly motivational situations, like 22-kHz calls in response to acute or conditioned threat. To further investigate such vocalizations and their relationship with overt behavior, we tested male Wistar rats in a paradigm of Pavlovian fear conditioning, where a tone stimulus (CS) was preceding an aversive foot-shock (US) in a distinct environment. Importantly, the shock dose was varied between groups (0-1.1 mA), and its acute and conditioned outcome were determined. The analysis of visible behavior confirms the usefulness of immobility as a measure of fear conditioning, especially when higher shock doses were used. Rearing and grooming, on the other hand, were more useful to detect conditioned effects with lower shock levels. Ultrasonic vocalization occurred less consistently than changes in overt behavior; however, dose-response relationships were also observed during the phase of conditioning, for example, in latency, call rate and lengths, intervals between calls, and sound amplitude. Furthermore, total calling time (and rate) were highly correlated with overt behavior, namely behavioral inhibition as measured through immobility. These correlations were observed during the phase of fear conditioning, and the subsequent tests. Importantly, conditioned effects in overt behavior were observed, both, to the context and to the CS presented in this context, whereas conditioned vocalization to the context was not observed (except for one rat). In support and extent of previous results, the present data show that a detailed analysis of ultrasonic vocalization can substantially broaden and refine the spectrum of analysis in behavioral work with rats, since it can provide information about situational-, state-, and subject-dependent factors which are partly distinct from what is visible to the experimenter.     

Is sexual-aggressive vocal communication related to asymmetric mechanisms in the brain?

In Mongolian gerbil males, a reliable asymmetric relationship has been discovered between a stereotyped ultrasonic vocalizattion and the volume of a discrete nucleus within the preoptic-anterior hypothalamic area, the sexually dimorphic area, pars compacta (SDApc). The steroid-sensitive ultrasonic, emitted at high levels during courtship interactions, is associated with the absence of aggressive encounters and appears to be necessary for the formation and maintenance of pair-bonds with females. Interactions between unfamilliar, non-pair-bonded adult male and estrous female gerbils result in male aggression, reduction of male ultrasonic calling, and disruption of normal female dartmale follow sociosexual sequences. Castration further reduces ultrasonic calling, inhibits normal sociosexual sequences, and increses aggression in interaction between unfamiliar compared to pair-bonded individuals. Bothg brain structure and lateralization of vocal function in males depend on the action of sex steroids during sexual differentiation and in adulthood. Thus, the gerbil provides a new model to study steroid-dependent lateralization of brain mechanisms of vocal behavior mediating aggressive-sexual relationships. © 1994 Wiley-Liss, Inc.     

Analysis of ultrasonic vocalizations emitted by residents during aggressive encounters among rats (Rattus norvegicus)

This investigation was concerned with the extent to which aggressive resident rats emit 40-70-kHz vocalizations and the effect of these signals on intruders. In Experiment 1, deafened and intact intruder males were given two encounters with resident animals. Deafened intruders engaged in a higher duration of immobile or freezing postures than intact animals. Experiment 2 indicated that the augmentation of freezing found among deafened intruders was not due to an inability to detect ultrasounds made by residents since intruders encountering devocalized resident males showed no reliable differences in specific motor patterns from intruders paired with intact residents. The results further demonstrated that 40-70-kHz vocalizations are produced almost entirely by intruding animals since there were no significant changes in occurrence of these calls when resident males were devocalized. Under the constraints of the testing procedures employed, the role of ultrasonic communication during the initial formation of agonistic relations could not be determined experimentally.     

Sensory cues that elicit ultrasonic vocalizations in female rats (Rattus norvegicus)

Female rats emit ultrasonic vocalizations during copulation. Female vocalizations are associated with darting and other proceptive behavior. In addition, females frequently call while approaching the male. A series of experiments was undertaken to determine whether female rats emit ultrasonic vocalizations in response to other rats or to their odors. Females were exposed either to bedding soiled by males or females or to devocalized conspecifics separated from the subject by a barrier. Females vocalized more to male cues than to female cues, regardless of whether the odor cues were from soiled bedding or from a devocalized conspecific. In addition, subjects vocalized more when presented with a devocalized female rather than a neutral stimulus. Furthermore, devocalized adult males, separated by a barrier, were more effective than either devocalized castrated or juvenile males in eliciting vocalizations; in turn, castrates and juveniles were more effective than neutral stimuli. Female calling was likely induced by odors from the male. In one experiment, the female subject was positioned in such a way that she could neither see or touch the stimulus male. Presentation of tape-recorded male vocalizations did not affect calling by the female. Vocalizations emitted by females in response to male odors may attract males or may facilitate subsequent copulatory behavior by the male.     

Differential rates of exhaustion and recovery of several parameters of male rat sexual behavior.

Sexual exhaustion of the male rat was studied with respect to several copulatory measures. The ejaculatory latency, intercopulatory interval, and intromission frequency demonstrate a U-shaped curve, exhibiting high values for the initial ejaculatory series, falling to a minimum at an intermediate ejaculatory series, and again increasing at exhaustion. The absolute refractory period of the postejaculatory interval (measured at vocalization termination) increases linerly, whereas the relative refractory period (the remaining portion of the postejaculatory interval) is a postively accelerating function. Partial recovery tests demonstrated that the preejaculatory measures and absolute refractory period have substantially returned to baseline values by Day 6, while the relative refractory period is still extended. The significance of these data to the theoretical modeling of sexual behavior is discussed.     

Facilitation of the effect testosterone on male sexual behavior in rats deprived of REM sleep

The possibility of rapid eye movement sleep deprivation (REMd) altering the effect of testosterone on masculine sexual behavior was investigated. Adult castrated male Wistar rats with no sexual experience were randomly assigned to the following three groups: REMd (using the water tank technique) for 7 days, large platform control for 7 days, and undisturbed sleep control. All subjects were treated with 1 mg testosterone propionate daily for 14 days. Masculine sexual behavior was assessed 3 consecutive days prior to steroid administration and was evaluated daily during the treatment. Frequencies and latencies of mounts, intromissions, and ejaculations, as well as the postejaculatory refractory period were recorded. One hundred percent of the REMd subjects presented mounts, intromissions, and ejaculations sooner than the control group. In almost all parameters, a clear facilitation of sexual activity was observed in the REMd group.     

Ejaculate disruption in two species of voles (Microtus): on the PEI matching law.

We permitted male prairie and montane voles (Microtus ochrogaster and M. montanus) five thrusts, without ejaculation, with a female at variable times after a 1st male ejaculated. In both prairie and montane voles, there were fewer sperm, in relation to control conditions, in the female's tract 1 hr after ejaculation if the female received thrusts immediately or 15 min after the ejaculate. There was no such effect after a 50-min delay. There was no significant decrease in litter production in prairie voles caused by thrusts delivered either immediately or after a 15-min delay. Sperm transport in these species is susceptible to disruption for a longer period than in deer mice or rats. The proposal that the postejaculatory interval protects a male from disrupting its own sperm transport (the PEI matching law) appears not to hold for these species.     

Social contacts and production of 50-kHz short ultrasonic calls in adult rats

The goal of the study was to provide evidence that the production of 50-kHz calls by adult rats is driven by potential or direct social contacts. The calls have been studied during daily visits to a cage by single or paired rats. Repeated exposure of rats to the cage frequently visited by other rats or direct contact between rats significantly increased the number of 50-kHz calls. The increase in production of 50-kHz calls was reduced by 78% after intrapreoptic-anterior hypothalamic injection of MK-801, an N-methyl-D-aspartate-type glutamate receptor antagonist. Calls emitted in all situations had a similar acoustic profile. It was found that 50-kHz calls were produced in anticipation of, and/or during, direct social contacts among adult rats and were predominantly initiated by olfactory stimuli. The calls seem to express an appetitive behavioral state in which the central glutamatergic mechanism is implicated.     

Ultrasonic vocalizations by male house mice (Mus musculus) to novel odors: roles of infant and adult experience

Two experiments were conducted to examine 70-kHz ultrasonic courtship vocalizations by adult male mice (Mus musculus) to novel odors following exposure to these odors in infancy and/or adulthood. The research was performed in hopes of better understanding the experiences giving rise to vocalizations to naturally occurring chemosignals. Experiment 1 demonstrated that adult males normally do not vocalize to the urine of female rats but would come to do so if adult female mice odorized with female rat urine were repeatedly encountered postpubertally. On the other hand, encountering their own mother odorized with female rat urine from birth until weaning did not promote vocalizations to the urine of female rats. Experiment 2 was designed to examine vocalizations to the urine of female mice whose urinary odor was altered by the ingestion of fenugreek, a spice. Although the magnitude of the effect was smaller in this experiment, greater amounts of vocalization again were seen by males that as adults encountered females that had ingested fenugreek. Males with such experience also showed a small but significant elevation in vocalizations to the fenugreek odor itself. Again, experience with the novel odor during infancy was not associated with elevated vocalizations during adulthood either to fenugreek-altered urine or to the fenugreek odor itself. Thus vocalizations to two different novel odors occurred only after an adult male had encountered an adult female odorized with the novel odor. On the other hand, none of the novel odor experiences eliminated vocalizations to the naturally occurring chemosignal in female mouse urine.(ABSTRACT TRUNCATED AT 250 WORDS)     

不同性别比和童年经济状况对未婚男性性态度的影响

基于进化心理学的生命史理论,通过两个实验探讨性别比和童年经济状况同男性性态度间的关系。实验1和实验2分别采用图片和文字材料启动被试的性别比感知,结果发现感知性别比和童年经济状况对未婚男性的"随意性行为"态度表现出显著的交互效应：童年家庭经济状况差的未婚男性在感知到周围女性数量相对充裕的情境下,其性开放态度会较感知到周围男性数量充裕时明显上升;童年家庭富裕的男性的性开放态度在不同感知性别比下保持相对稳定。     

Effect of masker level on infants' detection of tones in noise

In adult listeners, the signal-to-noise ratio at masked threshold remains constant with increases in masker level over a wide range of stimulus conditions. This relationship was examined in 7-month-old infants by obtaining masked thresholds for .5- and 4-kHz tones presented in four levels of continuous masking noise. Adults were also tested for comparison. Masker spectrum levels ranged from 5 to 35 dB/Hz for .5-kHz tones, and from -5 to 25 dB/Hz for 4-kHz stimuli. Thresholds were determined for stimuli of both 10 and 100 msec in duration. The results indicated that infants' performance was more adult-like for 4-kHz stimuli. Although mean thresholds for both 10- and 100-msec, 4-kHz tones were approximately 7 dB higher in infants than in adults, E/N0 at threshold remained essentially constant over the 30-dB range of maskers employed. By contrast, infants' thresholds for .5-kHz tones were exceptionally high at lower levels of the masker. Threshold E/N0 decreased significantly as masker level increased from 5 to 35 dB/Hz, and this decrease was significantly greater for 10- than for 100-msec stimuli. Temporal summation of .5-kHz tones, measured as the difference between thresholds obtained at the two signal durations, was greater for infants than for adults at low levels of the masker. However, because infants' thresholds improved more rapidly with level for 10- than for 100-msec tones, age differences in temporal summation were no longer significant when masker spectrum level was 35 dB/Hz. These results suggest that the relationship between signal-to-noise ratio at masked threshold and level of the masker is dependent on both signal frequency and duration during infancy.     

Ultrasonic vocalizing by adult female mice (Mus musculus)

Though an extensive body of literature exists concerning the emission of 70-kHz ultrasonic vocalizations by adult male mice (Mus musculus), almost nothing is known about the ultrasonic vocalizing of adult females. A series of five experiments was conducted to examine adult female mouse ultrasound emission. Results indicated female ultrasonic vocalizing to be typically displayed among female mouse dyads and comparable to the ultrasonic vocalizing levels obtained among male-female pairs. Genotypically based variations in the production of ultrasounds by females were also noted. Like male mice, socially naive females readily ultrasonically vocalized to anesthetized female conspecifics and rarely vocalized to anesthetized males. Unlike males, socially experienced females emitted few ultrasounds to either female urine or female-soiled cage shavings. Although social experience served to increase the ultrasound emission of male mice to female sex cues, the production of ultrasounds by females to these cues was decreased by social experience. Implications with regard to the existence of a behavioral/functional sexual dimorphism in adult mouse ultrasonic vocalizing are discussed.     

Effect of masker level on infants’ detection of tones in noise

In adult listeners, the signal-to-noise ratio at masked threshold remains constant with increases in masker level over a wide range of stimulus conditions. This relationship was examined in 7-month-old infants by obtaining masked thresholds for .5- and 4-kHz tones presented in four levels of continuous masking noise. Adults were also tested for comparison. Masker spectrum levels ranged from 5 to 35 dB/Hz for .5-kHz tones, and from ?5 to 25 dB/Hz for 4-kHz stimuli. Thresholds were determined for stimuli of both 10 and 100 msec in duration. The results indicated that infants’ performance was more adultlike for 4-kHz stimuli. Although mean thresholds for both 10- and 100-msec, 4-kHz tones were approximately 7 dB higher in infants than in adults, E/N₀ at threshold remained essentially constant over the 30-dB range of maskers employed. By contrast, infants’ thresholds for .5-kHz tones were exceptionally high at lower levels of the masker. Threshold E/N0 decreased significantly as masker level increased from 5 to 35 dB/Hz, and this decrease was significantly greater for 10- than for 100-msec stimuli. Temporal summation of .5-kHz tones, measured as the difference between thresholds obtained at the two signal durations, was greater for infants than for adults at low levels of the masker. However, because infants’ thresholds improved more rapidly with level for 10- than for 100-msec tones, age differences in temporal summation were no longer significant when masker spectrum level was 35 dB/Hz. These results suggest that the relationship between signal-to-noise ratio at masked threshold and level of the masker is dependent on both signal frequency and duration during infancy.     

不同性别比和童年经济状况对未婚男性性态度的影响

基于进化心理学的生命史理论,通过两个实验探讨性别比和童年经济状况同男性性态度间的关系。实验1和实验2分别采用图片和文字材料启动被试的性别比感知,结果发现感知性别比和童年经济状况对未婚男性的"随意性行为"态度表现出显著的交互效应：童年家庭经济状况差的未婚男性在感知到周围女性数量相对充裕的情境下,其性开放态度会较感知到周围男性数量充裕时明显上升;童年家庭富裕的男性的性开放态度在不同感知性别比下保持相对稳定。     

Copulation and intermale aggression in rats

Attack behavior of reliably aggressive male Long-Evans rats against unfamiliar male intruders was observed immediately following copulation to one or more ejaculations. Compulatory series to five ejaculations did not differ from copulation to a single ejaculation or from a noncopulatory control in affecting aggressive behavior. Repetitive biting attacks occurred in all conditions, with comparable wounding. Evidently, the male postejaculatory state of insensitivity to sexual stimuli does not extend to stimuli eliciting intermale aggression. A second experiment determined the attack-eliciting capacity of foreign males placed in the home cage of an actively copulating male. As intromissions increased and the interval to ejaculation decreased, the probability of intermale aggression and interruption of copulation diminished. The results are discussed in reference to sexual and aggressive strategies of the copulating male.