Effects of male pre-ejaculatory vocalizations on female receptive behavior in the rat (Rattus norvegicus)

Both male and female rats (Rattus norvegicus) emit ultrasounds during copulation. Bursts of ultrasounds from males that occur as ejaculation approaches are lower in frequency, longer in duration, and louder than those that appear earlier in the ejaculatory series. We attempted to determine if these pre-ejaculatory calls affect female sexual behavior. The behavior of females paired with devocalized males was compared with that of females paired with intact males, and in a second experiment tape-recorded vocalizations were played to females paired with devocalized males. Females were less receptive when paired with devocalized males; they were less likely to remain stationary when males attempted to mount. Playback of both types of calls restored female immobility toward control levels, although pre-ejaculatory calls were more effective than mating ones. Pre-ejaculatory calls restored running and training levels somewhat toward control level but to a lesser extent than female immobility.     

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.     

Unconditioned sexual incentive motivation in the male Norway rat (Rattus norvegicus)

Sexual incentive motivation was evaluated in a procedure consisting of a large open field where incentive animals were confined behind wire mesh openings. When sexually inexperienced male rats (Rattus norvegicus) were exposed to the receptive female-male incentives, they spent more time close to the female. If the incentives were receptive female-nonreceptive female, the receptive female was preferred. However, when the alternatives were nonreceptive female-male, no preference was obtained. Castration abolished preference for the receptive female, and treatment with testosterone propionate restored it. Estradiol plus oil is as efficient as estradiol plus progesterone for giving the ovariectomized female incentive properties. The living female can be replaced with female odor. Sexual experience did not have any long-term effects on the female's incentive value, but immediately preceding limited sexual activity enhanced it     

Stimulus control of maternal responsiveness to Norway rat (Rattus norvegicus) pup ultrasonic vocalizations

Mother rats (Rattus norvegicus; 6 to 8 days postpartum) approach and maintain proximal orientation to a pup that is emitting ultrasonic vocalizations (USVs) far more than do virgin females (W. J. Farrell & J. R. Alberts, 2002). We used a playback regimen to examine the roles of acoustic and nonacoustic cues in regulating maternal proximal orientation toward vocalizing pups. When presented with recorded USVs, mothers of 6- to 8-day-old pups and nulliparous virgin females exhibited equivalent levels of proximal orientation toward the playback speaker. Mothers did show enhanced proximal orientation toward recorded USVs, however, if a silent pup was positioned below the speaker. Pup odors appear to be crucial for the maternal response to vocalizing pups, as peripherally induced anosmia attenuated maternal proximal orientation toward a vocalizing pup. Furthermore, spatial contiguity between olfactory and auditory stimuli was required for a maximal maternal response.     

Reexamination of spectral mechanisms in the rat (Rattus norvegicus)

The spectral mechanisms of the rat were studied in electrophysiological and behavioral experiments. The photopic electroretinogram (ERG) was recorded (a) to rapidly flickering lights, (b) during the cone phase of dark adaptation, and (c) with a flicker photometric procedure. Each procedure indicated the presence of a single cone mechanism having a lambda max of about 510 nm. Increment-threshold spectral sensitivity functions measured in a behavioral test situation yielded the same conclusion. A behavioral experiment failed to produce any evidence for the presence of color vision in this rodent. The consistent conclusion from both behavioral and electrophysiological experiments is that the rat retina contains only a single photopic spectral mechanism.     

Receptivity of the female rat (Rattus norvegicus) after male devocalization: a ventral perspective

Male rats (Rattus norvegicus) emit at least two patterns of vocalization during copulation, the mating call and the pre-ejaculatory call. Both calls promote immobility of the female during lordosis, but the pre-ejaculatory calls are more effective. We undertook, through ventral observations of the mating pair, to determine if the female failed to assume or maintain the lordosis posture when mounted by a devocalized male and also to determine if the devocalized male was providing adequate stimulation to induce receptive behavior. Females were more likely to move away from the devocalized males before assuming the full lordosis posture. Furthermore, they were more likely to move away before the males had a chance to engage in intromissive behavior. However, when the females remained immobile long enough for the males to achieve a mount or intromission, there was little difference in the behavior of either animal that resulted from the devocalization of the male.     

Potentiation of acoustic startle behavior in the rat (Rattus norvegicus) at the onset of darkness

In Experiment 1 (N = 8), a 20-ms light pulse, given at various times before a noise burst, inhibited reflex expression with a single trough at a lead time of 70 ms, whereas a dark pulse facilitated the reflex with two peaks at 40 and 160 ms. In Experiment 2 (N = 18) facilitation by dark onset had a single peak, and inhibition by light onset a single trough; thus, the double peak of the dark pulse may result because inhibition from light onset at the end of the dark pulse was briefly impressed on the facilitatory effect of dark onset. In Experiment 3 (N = 12), diazepam (2.5 mg/kg, but not 1 mg/kg) eliminated dark facilitation but not light inhibition. These diazepam data reveal a basic similarity, perhaps identity, of the mechanisms responsible for the effect of dark onset and those producing reflex facilitation by Pavlovian fear conditioning and prolonged background noise, because all are moderated by a GABAergic system.     

Acoustic startle threshold of the albino rat (Rattus norvegicus)

The startle threshold of the albino Sprague-Dawley rat runs parallel to the curve of the hearing threshold. The difference between the startle and hearing threshold is 87 dB (SPL) at a background noise level of 75 dB (SPL). At 110 dB (SPL), the threshold has a range from 2 kHz to 50 kHz with a minimum at 10 kHz and a second minimum at 40 kHz. Amplitude and latency of the startle response are not only dependent on the sensation level of the acoustic stimulus but also on the frequency. At threshold, only the head movement component of the startle response is elicited.     

Maternal age and male behavior in relation to successful reproduction by female rats (Rattus norvegicus)

This research demonstrated that multiple ejaculatory series may be critically important for successful reproduction by multiparous old females near the end of their active reproductive lives. Multiple series were essential for successful pregnancy, apparently because more than one series was necessary to induce functional luteal activity. Thus, (a) females receiving just one series failed to show the cessation of regular estrous cycles which characteristically follows induction of a functional luteal phase; (b) direct measurement of plasma progestins by radioimmunoassay revealed that only those females receiving more than one series had circulating levels significantly higher than unmated, normally cycling controls; and (c) administration of exogenous progesterone increased significantly the incidence of successful pregnancy among females allowed to mate through only one series. In addition to facilitating the occurrence of pregnancy, multiple series also increased both the number of eggs released at ovulation and the number of embryos present and developing 12 days later. It is concluded that multiple series may function as a behavioral mechanism for prolonging the female's active reproductive life.     

Life-span changes in the copulatory behavior of male rats (Rattus norvegicus)

The copulatory behavior of 5-, 10-, 15-, and 20-month-old sexually naive male rats (Long-Evans) was studied during 2-hr tests with receptive females. There was no apparent change in sexual arousal as measured by latency to initiate copulation across age, with subjects from all groups exhibiting comparable latencies to first mount and first mount with intromission. The numbers of ejaculations achieved were also similar across ages. Significant age differences were found for frequency of mounts, with 20-month-olds having the highest mean frequency. The persistent mounting by older males appeared to account for significant group differences in interintromission interval and ejaculation latency. It is suggested that motor deficits may impair the ability of older males to achieve intromission, increasing the number of mount bouts as well as the number of mounts per bout, thus extending the length of each copulatory series.     

Milk availability modulates weaning in the Norway rat (Rattus norvegicus)

Two experiments were conducted to investigate the role of milk availability in the development of independent feeding and drinking in the Norway rat pup. Beginning on Day 14 postnatally, pups were exposed to different levels of milk supply by changing litter size from 8 to 4 versus 12 pups (Experiment 1) or limiting temporally the pups' access to a lactating dam to 8, 14, or 24 hr daily (Experiment 2). Both manipulations accelerated weaning in milk-poor pups in comparison with same-age pups with relatively greater milk supplies. By adding solid food and water to their diet, early weaning pups compensated for the negative energy consequences of milk reduction and achieved premanipulation growth rates. Milk availability thus appears to affect weaning, and it is suggested that the developmental changes in the nutritive energy balance between mother and offspring contribute to the emergence of independent ingestion.     

Cooperative behavior of laboratory rats (Rattus norvegicus) in an instrumental task

Cooperation is a cognitively demanding, complex social behavior, found primarily in primates. Here we investigated mutualism in rats (Rattus Norvegicus), a simple form of cooperation in which two subjects work on operant task, receiving immediate and simultaneous sucrose reward for a joint action. To receive the sucrose reward, familiar pairs of rats were required to nose poke simultaneously. Following 44 training days, we examined the relation of social contact and ultrasonic vocalizations to the rat's cooperative behavior by testing the effects of inserting opaque, wire-mesh, or no partition--between subjects. Cooperative behavior (simultaneous nose-poking): (a) increased gradually during initial training; (b) decreased with the opaque partition (restricting visual, acoustic, and physical communication); (c) increased with a wire mesh partition restricting only physical contact); and (d) increased with the number of 50 kHz USV "happy" calls and the intensity of social interaction. The possibility of studying the development of cooperative behavior in laboratory rats using a simple procedure based on commercially available equipment may prove useful in modeling determinants of social behavior.     

Maternal responsiveness to infant Norway rat (Rattus norvegicus) ultrasonic vocalizations during the maternal behavior cycle and after steroid and experiential induction regimens

When removed from the nest and placed in a cool environment, Norway rat (Rattus norvegicus) pups emit ultrasonic vocalizations that can elicit maternal search behavior. The authors examined the behavior of pregnant dams, mothers, and virgin females during exposure to a pup that was either warm and silent or cool and vocalizing. Results indicate potentiated maternal reactions to a vocalizing pup: Mothers approached and maintained proximal orientation to a vocalizing pup far more than did virgin females. Elevated levels of proximal orientation appeared within hours of birth, increased during the 1st week postpartum, and declined by the time of weaning. Estrogen plus progesterone administration facilitated virgin females' proximal orientation toward vocalizing pups, whereas prolonged exposure to pups in the absence of hormones was without effect, suggesting that the ontogeny of the maternal response is regulated, at least in part, by maternal hormones.     

Perioral somatosensory determinants of nursing behavior in Norway rats (Rattus norvegicus)

Prior to assuming the upright crouching posture over their pups during nursing bouts, lactating rats typically engage in several oral behaviors, including nuzzling, licking and rearranging pups. By acutely depriving dams of various aspects of perioral stimulation from pups (with anesthesia of the mystacial pads or of the tongue, with mouth suturing, or with muzzling), we found the following: (1) Distal stimulation from pups maintains proximity-seeking behavior, but is insufficient to stimulate nursing behavior. (2) Lack of tongue feedback decreases pup licking and hastens the onset of crouching. (3) Snout, but not tongue, contact with pups is required for hovering over them. (4) The position of the dam while hovering over her litter enables the pups to gain access to her ventrum, thereby provoking her upright, crouching posture. (5) Older pups are capable of bypassing the dam's perioral attentions and stimulating crouching directly by burrowing under the dam's ventrum.     

Development of the auditory orientation response in the albino rat (Rattus norvegicus)

The development of head orientation to auditory stimulation was examined in rat pups at Postnatal Days 8, 11, 14, 17, and 20. The animals were tested in a quiet environment with single bursts of 65 dB (SPL) broad-band noise. A reflexive head turn toward the sound was first seen on Postnatal Day 14 and subsequently on Days 17 and 20. This result demonstrates that the onset of directional auditory responses occurred between Day 11 and Day 14. The role of binaural cues in early sound orientation was examined in 17-day-old pups with monaural ligation of the external meatus. These animals were unable to localize a sound source and consistently turned toward the side of the unligated ear regardless of the position of the stimulus. Thus binaural cues were shown to be important for head orientation to sound in early development. In a separate study, head orientation to high and low frequency tone pips was examined. Directional responses were first seen on Day 12 for a 16-kHz tone and Day 14 for a 2-kHz tone. These results indicate an earlier onset for orientation to high frequency sounds in the rat.     

Sexual free behavior in male rats (Rattus norvegicus).

Male rats (Rattus norvegicus) were given continuous access to estrous female rats for 24 hr each day for 10 days. During the first 12 hr, the rats achieved an average of 10 ejaculations, followed by a 1- to 2-day recovery period with little sexual activity. During the last 7 days, the rats maintained a reasonably stable equilibrium level of 3 ejaculations per day. These occurred predominantly during the dark phase of the diurnal cycle, they frequently occurred in a cluster, and they usually occurred shortly after the introduction of a novel estrous female. Except for quantitative differences, these results are generally consistent with conventional research but systematically extend the generality of the results to the context of the free behavior situation. The availability of sexual activity had no appreciable effect on food and water intake, but it did decrease the amount of running activity.     

Prior exposure to stress reduces the diversity of exploratory behavior of novel objects in the rat (Rattus norvegicus)

Although a large literature has documented the varied effects of stress on an organism, relatively little attention has been devoted to investigating stress effects in ecologically relevant situations. Our experiment was conducted to assess the effects of stress on rats' (Rattus norvegicus) exploration of novel objects in a seminaturalistic and familiar environment that is relatively free of the constraints that have been placed on rats in prior investigations of stress. The results show that prior exposure to stress decreased the rats' diversity of exploration but did not affect general activity in comparison with animals not exposed to stress. We propose that the effect of stress on the qualitative aspects of exploratory behavior may be due to effects on organisms' sensitivity to predation.     

Prenatal expression of species-typical action patterns in the rat fetus (Rattus norvegicus)

We investigated sensory and behavioral responsiveness of the rat fetus. On Days 19, 20, or 21 of gestation, rat fetuses received intraoral infusions of a biologically important stimulus, milk, or a novel chemical stimulus, lemon. Using a technique to directly observe behavior in utero, we found that rat fetuses discriminate between intraoral infusions of milk and lemon, exhibiting different levels and patterns of overall activity after infusion. Milk was found to evoke a low magnitude, delayed increase in overall fetal activity from Day 19 through Day 21, whereas lemon evoked a high-magnitude, spiked pattern of activity that diminished from Day 19 to Day 21. Late in gestation these two stimuli elicited species-typical action patterns. Milk infusions elicited a stretch response much like the one shown by pups at the nipple; lemon infusions elicited face wiping typical of older pups and adults exposed to aversive gustatory stimuli.