Spatial Memory and Response Strategies in Rats: Age, Sex and Rearing Differences in Performance

Rats reared in social isolation made more errors on a spatial memory task and made errors earlier in each trial than socially reared rats. The difference in performance only occurred when rats were isolated prior to 50 days of age, and it survived IOO days of subsequent social housing. IOO days of isolation after 50 days of age did not influence performance on the spatial memory task. Subsequent experiments suggest that spatial abilities may not differ between groups but that isolates are slower to learn to make a particular response and to locate a particular arm when spatial and response cues are irrelevant. In contrast to previous experiments, clear response strategies were seen in the present experiments. These were prevalent in the young (54-days-old) rats, were less common at 90 days and had completely disappeared by 180 days. Response strategies were more common in male rats and in socially reared rats.     

Preoperative enrichment and behavioral recovery in rats with septal lesions

To assess the behavioral effects of preoperative differential housing male rats were placed in either enriched or isolated environments at weaning prior to receiving either sham operations or septal lesions when 57 days of age. Rats with septal lesions showed reduced habituation of ambulation and initially made fewer rears in an empty open field but made more object-contacts coupled with a lack of habituation in the object-filled field. Septal rats also showed severe impairments when tested in a 12-arm radial maze with 7 arms baited and 5 arms unbaited. Preoperative enrichment did not significantly affect these lesion-induced changes. Nevertheless, enrichment significantly lowered ambulation (but did not affect habituation) in the open field and increased the number of manipulatory relative to nonmanipulatory contacts. However, preoperatively enriched septal rats showed a deficit in spontaneous alternation (45%) in contrast to the high levels (83%) shown by intact enriched rats, whereas both intact and septal isolated rats showed similar levels of spontaneous alternation (68%). These results conflict with earlier reports that preoperative enrichment "protects" rats against the deficits produced by septal lesions.     

Response timing and development: Fixed-interval performance in precociously weaned rats

Young rats weaned at 16 days were taught to press a lever by shaping at 18 days and trained for 11 days (from 20 to 30 days of age) on a fixed-interval 60-sec schedule, at a rate of 5 half-hour sessions per day. Comparison with rats weaned at 16 or 20 days, trained at 21 days, and subjected for 8 days (from 23 to 30 days of age) to the same procedure showed that the temporal regulation of behaviour, as evaluated by the Curvature Index, was mostly training-dependent. Post-reinforcement pause durations depended upon the mixed influence of age and training. The evolution of running rates mostly depended upon training, whereas trends in overall rates seemed to depend essentially upon the age of the subjects. Precocious weaning per se or raising conditions prior to weaning did not significantly influence the temporal regulation of behaviour under FI. The results support previous ideas that overall response rate and response timing were dissociated and indicate high sensitivity to periodic stimuli in weanling rats. Finally, they show that, within an information processing model of timing, FI data of weanling rats might be compatible with their having an “internal clock” that runs at a higher rate than in older subjects.     

Effects of age at differential housing and the duration of individual housing/grouping on lntermale fighting behavior and adrenocortical activity in TO strain mice

A study of the effects of the duration of individual and group housing on intermale fighting and adrenocortical activity was conducted in TO strain mice. It was found that fighting and threat increased with progressive isolation up to an asymptote at 56–58 days. ‘Basal’ adrenocortical function differed little under the 2 housing conditions, but after ‘stress,’ mice isolated for short periods, which had not fought when tested for aggression, had lower titers than group-housed counterparts. However, mice that had been isolated for longer periods, and had fought in aggression tests, had higher corticosterone titers than comparable group-housed animals. The effects of a short duration (28–30 days) of differential housing, commencing at different ages, were also studied. The shorter duration adrenocortical changes were largely confirmed. In general, the earlier the age at which the differential housing was imposed, the greater the behavioral differences between animals under each housing condition. It is suggested that this is largely a consequence of a loss of behavioral plasticity in older mice. The data provides little support for the concept of the “isolation stress syndrome,” or for the view that the characteristic fighting exhibited by individually housed mice is a consequence of “social deprivation”.     

Weight Loss in Rats Produced by Running: Effects of Prior Experience and Individual Housing

To investigate factors affecting activity-based anorexia (ABA) or activity-stress (AS), rats were given 2-hr access to a running wheel immediately prior to their daily 1.5-hr food access during the light cycle. This produced a reduction in food intake, a steady increase in running, and a large drop in body weight with a prolonged delay before weight recovery began. Experiment 1 found that these effects were reduced in rats with prior experience of eating at this time of day. In contrast, prior experience of running in the wheel when on ad lib food enhanced these effects in Experiment 2, where a subsequent change for half the subjects to individual housing produced a further decrease in body weight. The latter factor was investigated from the outset of Experiment 3 and again individually housed rats showed greater weight loss than did group-housed rats. This experiment also found that in rats of the same age a low initial body weight predicts greater vulnerability to ABA. It was concluded that ABA results from activity-induced reduction of feeding, which prolongs adaptation to a new feeding schedule and is accentuated by social isolation.     

Cytotoxic lesions of the hippocampus do not impair social recognition memory in socially housed rats

Socially housed mice with cytotoxic lesions of the hippocampus do not exhibit social recognition memory 30 min following exposure to a juvenile mouse, however the social recognition memory of singly housed rats is unimpaired. The present study tests the hypothesis that social housing of rats could render social recognition memory hippocampally dependent as seen for mice. Rats were housed with juveniles or with adults. Two social recognition one-animal tests paralleling those used with mice were carried out. Seven social discrimination two-animal tests were also given. Sham operated and hippocampally lesioned rats had normal social memory at 30 min whether socially housed for 24, 48 h, 7 or 8 days prior to testing. These findings support other results indicating the hippocampus proper is not required for normal social memory in rats. In a final experiment, rats socially housed in groups of three since weaning, were tested for 30 min and 24 h social memory. Unlike mice, rats socially housed throughout life exhibited social memory only at 30 min, but not at 24 h. Manipulations that extend social memory in rats may be required to render social memory hippocampally dependent or rats and mice may differ in the neural mediation of social memory.     

Experience-dependent changes in dendritic arbor and spine density in neocortex vary qualitatively with age and sex

Male and female Long-Evans hooded rats were placed in the complex environments for 3 months either at weaning (22 days), in young adulthood (120 days), or in senescence (24 months). The dendritic morphology of both the apical and basilar fields of layer III pyramidal cells was analyzed in both parietal and visual cortex. There were two novel results. First, although spine density was increased significantly with complex-housing in adulthood, it was decreased significantly by the same housing during development. Second, dendritic length was increased in both parietal and occipital cortex at all ages in males and was increased in adult females as well, but juvenile females showed no change in dendritic length in the occipital cortex and only a small effect on the apical field in parietal cortex. Thus, there are qualitative differences in the changes in spine density at different ages and the dendritic changes in response to complex versus isolated housing vary with sex, and in females, the changes vary with age as well. These results may explain some of the apparent inconsistencies in reports of spine and dendrite changes in the literature.     

Ontogeny of sex differences in defensive burying behavior in rats: effect of social isolation

The aim of the present experiments was to clarify sex differences in socio-developmental factors that affected defense behavior in rats. Sex differences in the defensive burying behavior of rats, and related social factors, were explored in three developmental stages: juvenile, puberty, and adult; 30, 50, and 80 days of age, respectively. The duration of burying, digging into bedding material, stretch-attend postures, and crouch/freezing were measured in a shock-prod test. For males, the duration of burying was longer in the juvenile and pubertal stages than in adulthood. For females, no age differences in the duration of burying were found. Males showed longer burying durations than females in both the juvenile and pubertal stages. For both sexes, the highest duration of digging was found in the juvenile stage, and females showed longer durations of digging than males. Both male and female rats isolated during the juvenile stage, from 26 to 40 days of age, showed smaller durations of burying behavior compared to pair-reared rats. This effect of juvenile isolation was maintained among both adult males and females even when they were returned to pair rearing after isolation. Isolation during adulthood, from 66 to 80 days of age, increased burying behavior in males, but decreased it in females. The durations of digging, stretch-attend postures, and crouch/freezing were not affected by isolation. The decrease in defensive burying and its increase resulting from isolation in adult male rats, suggest that the emergence of adult-like social relationships in males suppressed the duration of burying. Male and female rats isolated during the juvenile stage maintained lower levels of burying, suggesting that social experience as juveniles is important for the emergence of defensive burying behavior.     

Fighting,nonagonistic social behavior,and exploration in isolation‐reared rats

Sixty hooded Long‐Evans rats were assigned to one of three conditions: isolation‐rearing (from 14–30 days of age), brief isolation (from 28–30 days of age), and normal (no isolation). From day 14 to day 21, the isolation‐reared animals were maintained in an incubator set at 35°C to prevent hypothermia and were separated from each other by Plexiglas. They learned to nourish themselves by lapping milk that continuously flowed down one side of the enclosure. Thereafter, they were housed in standard hanging wire cages. Unlike previous studies of isolation at this age, there were no significant weight differences between the isolation‐reared group and the two control groups. At 30 days of age, the three groups were compared on fighting (serious and play), nonagonistic social behaviors (following and crawling under), and exploratory behavior (open‐field ambulation). The isolation‐reared group was significantly different from the normal group on all measures (isolates were higher on all measures except exploration). The isolation‐reared animals showed significantly more serious fighting (aggression) than the brief isolation animals, whereas these two groups did not differ on other behaviors. This methodology allows for the study of isolation‐rearing without the nutritional confounds found in previous research and shows an effect on aggression that is not accounted for by recent isolation. Aggr. Behav. 25:211–223, 1999. © 1999 Wiley‐Liss, Inc.     

Establishment of a schizophrenic animal model through chronic administration of MK-801 in infancy and social isolation in childhood

BackgroundAlthough an increasing amount of evidence supports a “two-hit” hypothesis for the neurodevelopmental model of schizophrenia, there has been no development in animal models to test this hypothesis.MethodsAn animal model was established by chronic administration of 0.1, 0.3, and 0.5 mg/kg MK-801 in P7-P21 rats followed by four weeks of social isolation in childhood and then five days of social housing. Animal behaviors were measured by the open field (OF) test, the novel object recognition (NOR) test, the prepulse inhibition (PPI) test, and the elevated plus maze (EPM) test.ResultsWe found a significant decrease in the NOR index in adolescent rats compared to saline control rats when administering 0.5 mg/kg of MK-801 (P = 0.02). We found that social isolation had no significant effect on NOR index, though social isolation significantly increased the total distance traveled and significantly decreased the resting time in adolescent rats in the OF test (P < 0.001 and P = 0.003, respectively). In contrast, we observed that MK-801 administration showed no significant effects on either total distance traveled or resting time. Both MK-801 administration and social isolation had no significant effect on the percent of PPI and startle amplitudes in adolescent rats. Social isolation significantly reduced the open arm entries in adolescent rats in the EPM test (P = 0.023), but it did not reduce the ratio to enter the open arms and the stay time in open arm. Administration of MK-801 showed no significant effect on the indexes of entering the open arms in the EPM test on adolescent rats.ConclusionMK-801 intervention in infancy is associated with the damage of long-term visual memory, whereas social isolation in childhood is associated with the increased spontaneous activity and anxiety levels. Administration of MK-801 in infancy and social isolation in childhood are two independent factors on the neurodevelopmental defects.     

Sodium amylobarbitone and responses to nonreward

Three experiments are reported testing two alternative hypotheses concerning the behavioural effects of sodium amylobarbitone (SA): (1) that it blocks the after-effect of nonreward; (2) that it blocks conditioned frustration, elicited by stimuli associated with nonreward. In support of (2) Experiment I showed that SA given in acquisition abolished the partial reinforcement extinction effect (PREE) when rats were run at one trial a day in an alley for food reward on a continuous (CRF) or partial (PRF) reinforcement schedule. Experiment II showed that, in the goal section, the effect of the drug on the PREE was due to its presence during acquisition and was not due to state dependency; but the effect of the drug in the start section was consistent with state dependency of the PREE. In Experiment III, in opposition to (1) and again in support of (2), SA given to rats trained to show patterned running for water reward on a single alternation schedule blocked patterning by increasing running speeds on nonreward trials, not by decreasing running speeds on rewarded trials.     

The effects of isolation rearing on behavioural inhibition in the rat

The possibility that isolation-rearing in the rat affects the development of inhibitory mechanisms was studied in a series of experiments. It was found that socially-isolated rats were (1) slower to learn both a lever-panel alternation, and a two-lever alternation schedule of reinforcement, (2) more persistent than controls in pressing a lever for food when a supply of identical “free food” was introduced into the operant chamber, but (3) similar to control rats in their response to preloading with food, a procedure which inhibited lever pressing to the same extent in the two groups. Finally, it was shown in a separate experiment that the effects of increased food deprivation on lever pressing in the presence of free food were qualitatively different from the effects of social isolation, and therefore the social/isolate difference cannot be interpreted as motivational. The possible contributions of neophobia to the difference are discussed. It is concluded that isolates may well suffer from a disinhibitory defect, but that there are probably other effects of isolation in addition.     

Effects of pre- and postpubertal social experience on sexual and social behavior in male and female brown lemmings (Lemmus sibiricus)

The effects of prepubertal social contact and postpubertal mating experience on sexual behavior were examined in male and female brown lemmings (Lemmus sibiricus). Subjects of both sexes were either isolated at weaning (18 days) or then housed with siblings for another 10 days before being isolated. Before testing for sexual behavior at the age of 75 days, half of each group of males received a series of exposures to estrous females and half of each group of females was housed with a stud male. The results of mating tests indicated that whereas prior sexual experience facilitated subsequent copulatory behavior in both sexes, prepubertal interactions subsequently facilitated contact social and sexual behavior in males but did not increase the behavioral scores of females, which engaged in high levels of sexual behavior regardless of prepubertal experience. The effect of prior copulatory activity on male sexual performance could not be attributed to increasing age (i.e., maturational processes) because the behavior of males tested once but at different ages did not differ appreciably. Prepubertal social interaction appeared to predispose males to benefit from sexual experience later in life.     

Social enrichment improves social recognition memory in male rats

The social environment is thought to have a strong impact on cognitive functions. In the present study, we investigated whether social enrichment could affect rats’ memory ability using the “Different Objects Task (DOT),” in which the levels of memory load could be modulated by changing the number of objects to be remembered. In addition, we applied the DOT to a social discrimination task using unfamiliar conspecific juveniles instead of objects. Animals were housed in one of the three different housing conditions after weaning [postnatal day (PND) 21]: social-separated (1 per cage), standard (3 per cage), or social-enriched (10 per cage) conditions. The object and social recognition tasks were conducted on PND 60. In the sample phase, the rats were allowed to explore a field in which 3, 4, or 5 different, unfamiliar stimuli (conspecific juveniles through a mesh or objects) were presented. In the test phase conducted after a 5-min delay, social-separated rats were able to discriminate the novel conspecific from the familiar ones only under the condition in which three different conspecifics were presented; social-enriched rats managed to recognize the novel conspecific even under the condition of five different conspecifics. On the other hand, in the object recognition task, both social-separated and social-enriched rats were able to discriminate the novel object from the familiar ones under the condition of five different objects. These results suggest that social enrichment can enhance social, but not object, memory span.     

Post-weaning social and cognitive performance of piglets raised pre-weaning either in a complex multi-suckling group housing system or in a conventional system with a crated sow

We studied the social and cognitive performance of piglets raised pre-weaning either in a conventional system with a sow in a farrowing crate (FC) or in a multi-suckling (MS) system in which 5 sows and their piglets could interact in a more physically enriched and spacious environment. After weaning at 4 weeks of age, 8 groups of 4 litter-mates per pre-weaning housing treatment were studied under equal and enriched post-weaning housing conditions. From each pen, one pair consisting of a dominant and a submissive pig was selected, based on a feed competition test (FCT) 2 weeks post-weaning. This pair was used in an informed forager test (IFT) which measured aspects of spatial learning and foraging strategies in a competitive context. During individual training, submissive (informed) pigs learned to remember a bait location in a testing arena with 8 buckets (the same bucket was baited in a search visit and a subsequent relocation visit), whereas dominant (non-informed) pigs always found the bait in a random bucket (search visits only). After learning their task, the informed pigs’ individual search visit was followed by a pairwise relocation visit in which they were accompanied by the non-informed pig. Effects of pre-weaning housing treatment were not distinctly present regarding the occurrence of aggression in the FCT and the learning performance during individual training in the IFT. During paired visits, informed and non-informed pigs changed their behaviour in response to being tested pairwise instead of individually, but MS and FC pigs showed few distinct behavioural differences.     

Spatial learning in pigs: effects of environmental enrichment and individual characteristics on behaviour and performance

This study investigated the effects of both environmental enrichment and individual behavioural characteristics on spatial cognitive capabilities of pigs, using a novel latent spatial learning paradigm based on Tolman’s detour experiments (1948). Pigs were housed either in ‘barren’ pens or in pens enriched with straw bedding from birth. Pigs were restrained in a Backtest at 10 and 17 days postpartum. Based on their escape behaviour in this test, which has been shown to reflect their behavioural style, six ‘high-resisting’(HR) and six ‘low-resisting’ (LR) pigs were selected from each housing environment (n = 24 in total). At 12 weeks of age, pairs of pen mates (LR and HR) were exposed to a maze three times (exploration trials). Pigs were then placed individually in the maze, and social reinstatement proved to be a strong incentive to find the exit leading to the home pen. We subsequently blocked the direct route to the exit, forcing animals to find a detour (memory test 1, MT1). This test was repeated once to investigate the relative improvement, i.e. detour learning (memory test 2, MT2). Housing condition and Backtest response strongly affected exploration patterns. In spite of this, no effects on performance during the subsequent memory tests were found. Performance was substantially improved in MT2, indicating that once a goal is apparent, pigs are able to solve a complex spatial memory task easily. In conclusion, social reinstatement provided a good incentive to complete a spatial task, and the substantial improvement in performance between MT1 and MT2 stresses the need for task complexity when testing spatial memory in pigs. Housing conditions or individual behavioural style did not affect spatial memory during MT1 or MT2. However, housing environment and behavioural style strongly affected explorative behaviour of pigs in an unfamiliar maze during both exploration trials and memory tests. This implicates that apparent effects of environmental enrichment on spatial learning and memory in pigs might reflect differences in explorative patterns rather than in cognitive processes.     

Effect of auditory isolation on activity-based anorexia

The objective was to study male and female rats exposed to Activity-Based Anorexia (ABA; simultaneous exposition to food restriction and free access to an activity wheel) under two conditions of sound attenuation, by means of different arrangements of wheels (standard versus isolation) in the laboratory room. Regardless of the sound attenuation condition, all but one male and one female with access to wheels had to be removed from the experiment, but extended ABA endurance was found in rats in the sound attenuation condition. Furthermore, significantly lower levels of running were observed in both males and females under the sound attenuation arrangement in the isolation condition. The results suggest that external stimulation plays a role in the activity displayed by rats exposed to ABA, and that the reduction of external stimulation diminishes running but does not protect rats from developing ABA.     

Effects of previous housing conditions on shock-induced aggression.

This study investigated shock-induced aggression as a function of housing rats in single or communal cages for varying periods of time before testing. Rates of fighting were unaffected by housing conditions when rats spent seven or 14 days in either of these settings before testing. Placing rats in these settings for 21 or 28 days before testing, however, did affect rates of fighting. Communal caging of subjects for 28 days before testing was particularly deleterious to shock-induced aggression. How rats are housed outside the experimental chamber is a variable affecting shock-induced aggression and should be considered both in designing future studies and as a topic for future investigation.     

Influence of timing of post-weaning isolation on play fighting and serious aggression in the male golden hamster (Mesocricetus auratus)

Effects of timing of social isolation on play fighting and serious fighting were studied at different ages in male golden hamsters. Litters were isolated at 21, 35, and 65 days of age, and tested in a resident-intruder paradigm. Behaviors were compared within grous and with a fourth group of socially reared conspecifics. The earlier the pups were isolated, the more they engaged in play activities. Later, in adulthood, the aggression level of the same animals was retested using the same paradigm. The three isolated groups showed a high level of aggression, with significant differences among them. When compared with socially reared subjects, a reliable difference in the level of aggression was also found. These results support the view that early social experience is important, suggesting that isolation during early critical periods of socialization has a significant impact on play fighting, whereas short periods of isolation may be enough to trigger adult agonistic behavior. © 1994 Wiley-Liss, Inc.     

The effects of pre- and postweaning rearing conditions on latent inhibition and partial reinforcement extinction effect in male rats

Male rats were either handled daily or left undisturbed (nonhandled) during the first 21 days of life. At weaning (Day 22) animals in each early treatment condition were assigned to two housing conditions, isolated or grouped, creating four early/late treatment conditions: Handled-Grouped (HG), Handled-Isolated (HI), Nonhandled-Grouped (NHG), and Nonhandled-Isolated (NHI). At maturity, all animals were tested in the latent inhibition (LI, Experiment 1) and the partial reinforcement extinction effect (PREE, Experiment 2) paradigms. In LI, animals receiving prior nonreinforced preexposure to a stimulus (PE) show subsequently retarded conditioning to that stimulus as compared to nonpreexposed (NPE) controls. In the PREE, partially reinforced (PRF) animals exhibit higher resistance to extinction as compared to continuously reinforced (CRF) controls. LI and PREE were obtained in the HG condition, but were impaired in NHG and HI animals. The pattern of impairment was different in the two conditions. The absence of LI in NHG males was due to increased suppression in the PE group, whereas in HI males, was due to decreased suppression in the NPE group. Likewise, the attenuation of the PREE in NHG males stemmed primarily from decreased resistance to extinction in the PRF group, whereas in HI males, it resulted from increased resistance to extinction in the CRF group. The combination of nonhandling and isolation led to the reinstatement of both LI and PREE: normal LI and PREE were obtained in the NHI animals. These results demonstrate that early rearing experience interacts with later experience differentially depending upon the later experience.