Development of Exploration and Investigation in the Norway Rat (Rattus norvegicus)

A cross-sectional study of the development of spontaneous exploratory behavior in the laboratory rat was conducted. Long-Evans hooded rat pups aged 11 to 28 days (n = 158, from 25 litters) were placed into a dimly illuminated arena for 10 min on 2 consecutive days. Analysis of videotaped records from the second session for each rat pup showed a relatively sudden emergence of locomotor exploration between 12 and 16 days of age. The behavioral topography of interactions with objects developed less uniformly, with some features (e.g., bout length) relatively unchanging and other features (e.g., number of bouts, number of different behaviors displayed) increasing with age. The data for locomotor exploration are consistent with an explanation that depends on maturation-driven emergence of species-typical behavior, but that explanation cannot account for the pattern of development for object investigation. These results provide further evidence that some important aspects of the character of exploratory behavior depend, at least partially, on experience, and that exploration is not in fact a single behavior system. In addition, they provide important baseline information for further studies of the structure and function of exploratory behavior.     

Behavior of Maudsley reactive and nonreactive rats (Rattus norvegicus) in three consummatory contrast paradigms

Maudsley reactive (MR/Har) and nonreactive (MNRA/Har) rats (Rattus norvegicus) were tested in successive, simultaneous, and anticipatory contrast procedures. The MR/Har rats showed smaller successive negative contrast effects than the MNRA/Har rats when shifted from 32% to 4% sucrose, and the degree of contrast was smaller in animals of both strains than that typically obtained with unselected Sprague-Dawley derived rats. Chlordiazepoxide (4 and 8 mg/kg), which typically reduces contrast, did not influence degree of contrast in rats of either strain. Animals of both strains showed positive and negative contrast in the simultaneous contrast procedure, but degree of contrast in both cases was smaller in rats of the MR/Har strain. Animals of both strains also showed anticipatory contrast when a 0.15% saccharin solution preceded 32% sucrose in once-per-day pairings. In terms of latency to initiate licking, the MNRA/Har rats showed a contrast effect, but the MR/Har rats showed a "reinforcement" effect--shorter latency when saccharin preceded sucrose than when saccharin preceded saccharin. Open-field tests showed typical strain differences: The MNRA/Har rats ambulated more, reared more, defecated less, and showed less thigmotaxis than the MR/Har rats.     

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.     

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.     

Olfactory discrimination of individual littermates in rats (Rattus norvegicus)

Adult male rats (Rattus norvegicus) were trained in a Y-maze to discriminate the presence of a littermate from its absence. Transfer of training in subsequent tests indicated that (a) the animals were capable of distinguishing among individuals when relatedness and familiarity were held constant, (b) this ability was not due to training, and (c) their performance was based solely on odor cues. The results are discussed in relation to social behavior in rats.     

Environmental influences on flank marking and urine marking by female and male rats (Rattus norvegicus)

Sprague-Dawley rats (Rattus norvegicus) were observed in a familiar environment. In Experiment 1 a leader entered a clean chamber, and an opposite-sex follower entered the chamber next. Both sexes began to flank mark after several sessions. Males flank marked more, and females locomoted more, but both sexes urine marked and investigated objects equally often. Leaders and followers did not differ on any measure. In Experiment 2 we measured floor marks more precisely and manipulated the number of objects and the presence of scent marks. Flank marking was more frequent in the presence of conspecific urine but did not vary with the number of objects or the sex of the rats. More objects elicited more investigating and urine marking and produced fewer floor marks but increased the number of marks in the central area in relation to the periphery. The results indicate that rats' flank marking is behaviorally distinct from urine marking and differentially affected by environmental variables.     

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.     

The easy-to-hard effect in human (Homo sapiens) and rat (Rattus norvegicus) auditory identification

The authors examined whether progressively training humans and rats to perform a difficult auditory identification task led to larger improvements than extensive training with highly similar sounds (the easy-to-hard effect). Practice improved humans' ability to distinguish sounds regardless of the training regimen. However, progressively trained subjects were more accurate and showed more generalization, despite significantly less training with the stimuli that were the most difficult to distinguish. Rats showed less capacity to improve with practice but still benefited from progressive training. These findings indicate that transitioning from an easier to a more difficult task during training can facilitate, and in some cases may be essential for, auditory perceptual learning. The results are not predicted by an explanation that assumes interaction of generalized excitation and inhibition but are consistent with a hierarchical account of perceptual learning in which the representational precision required to distinguish stimuli determines the mechanisms engaged during learning.     

Recognition of food by individual, food-naive, weaning rats (Rattus norvegicus)

To control and observe first feeding experiences, pups were reared without access to food. At weaning age, individual pups received simultaneous access to a palatable, noncaloric diet and a similar diet containing starch. Each diet contained a flavor cue. Within 20 min of sampling both diets pups preferentially ingested the caloric alternative. Further tests indicated that pups learned a preference for flavor cues paired specifically with ingestion of starch. In caloric discrimination tests in which the diets were not distinguished by artificial flavors, pups did not show the rapid preference for the caloric diet seen in earlier experiments. We suggest that recognition and preference for the caloric diet is based on its rapid postingestive effects rather than readily preferred sensory features of the starch. Our findings demonstrate the abilities of individual, food-naive pups to rapidly recognize food and indicate a role for associative learning in the onset of independent feeding.     

The influence of preexperimental experience on social discrimination in rats (Rattus norvegicus)

The authors used laboratory rats (Rattus norvegicus) of known relatedness and contrasting familiarity to assess the potential effect of preexperimental social experience on subsequent social recognition. The authors used the habituation-discrimination technique, which assumes that multiple exposures to a social stimulus (e.g., soiled bedding) ensure a subject discriminates between the habituation stimulus and a novel stimulus when both are introduced simultaneously. The authors observed a strong discrimination if the subjects had different amounts of preexperimental experience with the donors of the 2 stimuli but a weak discrimination if the subjects had either equal amounts of preexperimental experience or no experience with the stimuli. Preexperimental social experience does, therefore, appear to influence decision making in subsequent social discriminations. Implications for recognition and memory research are discussed.     

Discriminability, loudness, and masking in the rat (Rattus norvegicus): a confirmation and extension

In Experiment 1, rats discriminated between two sound pressure levels (SPL) of a pure tone: standard (STD) SPLs of 84 and 74 dB and comparison (CO) SPLs 4, 14, and 24 dB below STD were tested in quiet and 60 dB noise at 4 and 12.5 kHz (24 conditions). The decibel difference between STD and CO accounted for only 43.52% of the variance in the signal detection measure of sensitivity, d', across conditions, whereas the loudness difference (LD = STD0.35 - CO0.35) accounted for 89.82% of the variance in d'. These results confirm and extend previous observations that: (a) equal decibel differences are not equally discriminable; (b) loudness for the rat increases as a power function of SPL with an exponent of 0.35: and (c) masked loudness is a linear function of loudness in quiet. In Experiment 2, the assumptions of normal distribution and equal variance implicit in the use of the d' measure were examined. Receiver operating characteristic curves that were well approximated by straight lines of unit slope in normal-normal coordinates were obtained and thereby validated the use of d' in Experiment 1.     

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.     

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     

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.     

Loudness bisection and masking in the rat (Rattus norvegicus).

The bisection method of animal psychophysical scaling was examined as a measurement procedure. The critical assumptions of bisection scaling, as described by Pfanzagl (1968), were tested to determine if a valid equal-interval scale could be derived. A valid scale was derived in which loudness for the rat (Rattus norvegicus; n = 13) was a power function of sound pressure for 4-kHz tones. Masking noise reduced the discriminability of tonal stimuli but did not affect the bisection point. This result is consistent with an interval scale representation of loudness and demonstrates scale meaningfulness. Loudness bisection data that have been reported in the literature for 3 species (humans, rats, and pigeons) are in substantial agreement with our results.     

Habituation of the hiding response to cat odor in rats (Rattus norvegicus)

Cat odor-induced hiding was examined in rats (Rattus norvegicus) using an apparatus with a "hide box" at one end and a piece of a worn cat collar at the other end. Rats spent most of their time hiding on exposure to the cat collar, but this response gradually habituated over repeated daily exposures. Hiding was reversed by administering the anxiolytic drug midazolam (0.375 mg/kg). Rats showed increased anxiety on the elevated plus-maze after exposure to the collar. This response was absent in habituated rats, suggesting that habituation of hiding reflects decreased odor-induced anxiety. It was established that rats located in the hide box of the apparatus might not have detected the odor of the cat collar placed at the other end of the apparatus. This implies that habituation occurs after relatively modest levels of odor exposure. Overall, the results cast some doubt on claims that predatory odors in rats are akin to phobic stimuli in humans.     

Numerical competence in rats (Rattus norvegicus): Davis and Bradford (1986) extended

The task of rats (Rattus norvegicus) was to enter 1 box of a defined ordinal number among an array of boxes. In Experiments 1 and 2, the rats correctly chose the 4th box from arrays of 6 and 12 boxes, respectively. In Experiments 3 and 4, in which the ordinal number of the correct box was increased in a graduated fashion, they were able to select the correct box even when its position was higher than 10th among 12 and 18 boxes, respectively. In Experiment 5, the possibility that the rats had used cues to the openability of the box doors was ruled out. In Experiments 6 and 7, the rats succeeded in the task even when the total number of boxes was varied from trial to trial. In Experiment 8, both small- and large-sized boxes were used to control for the possibility of using the cumulative length of the boxes as a cue. Overall, the results suggest that performance was based on numerical cues. Intentional acts were occasionally observed but appeared not to be essential for solving the task.     

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.     

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.