Pair-housed Rats Research Articles

Age differences in conditioned place preferences and taste aversions to nicotine.

Adolescents and adults differ in their behavioral sensitivities to drugs of abuse, including nicotine. Studies have shown that both rewarding and aversive properties of drugs of abuse can exist concomitantly. The present study investigated the ontogeny of these opposing qualities across a range of doses using a combined conditioned taste aversion and place preference paradigm in pair-housed rats that were not deprived of food or water. Results indicated that adolescents were more sensitive to the rewarding properties of nicotine than adults. In contrast, although all doses produced a taste aversion at both ages in the same rats, the aversion was weaker at lower than high doses in adolescents whereas adults showed strong aversion at all doses, suggesting modest attenuation in nicotine's aversive properties among adolescents relative to adults. Thus, attenuated aversive and accented appetitive sensitivities of adolescents to nicotine can be experienced simultaneously in the same animals. © 2016 Wiley Periodicals, Inc. Dev Psychobiol 58: 660-666, 2016.

The effects of social contact on cocaine intake under extended-access conditions in male rats.

Social learning theories of drug use propose that drug use is influenced by the behavior of peers. We previously reported that cocaine self-administration under limited-access conditions can be either facilitated or inhibited by social contact, depending on the behavior of a peer. The purpose of this study was to determine whether social contact influences cocaine self-administration under conditions that are more representative of problematic patterns of drug use. Male rats were assigned to either isolated or pair-housed conditions in which a social partner either had access to cocaine or did not have access to cocaine. Pair-housed rats were tested in custom-built operant conditioning chambers that allowed both rats to be tested simultaneously in the same chamber. In Experiment 1, rats were tested for 14 consecutive days during daily 6-hr test sessions. In Experiment 2, different doses of cocaine were tested in 23-hr test sessions conducted every 3 days. All groups of rats escalated their cocaine intake in Experiment 1; however, pair-housed rats with a partner without access to cocaine had lower levels of intake throughout the 14 days of testing. In Experiment 2, pair-housed rats with a partner without access to cocaine had lower levels of cocaine intake than did rats with a partner with access to cocaine, and this effect was observed at all doses of cocaine tested. These data indicate that the behavior of a social partner (i.e., whether or not that partner is also self-administering cocaine) influences cocaine self-administration under conditions that model problematic patterns of drug use. (PsycINFO Database Record

Versatile 3D-printed headstage implant for group housing of rodents

BackgroundAn unfavourable yet necessary side-effect of stereotaxic surgery involves the social isolation of post-surgery rats, in order to protect their wound site or skull-mounted implant from damage. Social isolation can cause a myriad of behavioural and physiological changes that are detrimental to the well-being of rats, with potential negative implications for a range of experimental paradigms.New method. Female Sprague Dawley rats (n=40) were implanted onto the skull with a novel 3D-printed headstage socket that surrounded an electrode connector. The socket accommodated a removable stainless-steel headcap for the purposes of protecting the implant. Rats were pair-housed following surgery, and their behaviour was monitored for up to several weeks under two experimental conditions that involved EEG recording and deep-brain stimulation, as well as behavioural test sessions inside an open-field maze. Rat weights were compared between individually- and pair-housed rats at up to 3 weeks post-surgery. ResultsThese experiments were successfully carried out using pair-housed rats, with no damage or complications observed regarding the implant and its headcap. Rats were able to carry out a range of normal behaviours including running, grooming, foraging and sleeping. Compared to individually-housed rats, pair-housed rats gained less weight over the 3 weeks post-implantation period.Comparison with existing method(s). This method offers additional protection compared to group-housed post-surgical rats that lack the protective headcap. It is also potentially more practical and versatile than a fully-implantable device for the safe post-surgery group housing of rodents. ConclusionsThis implant design can reduce the cost of rodent upkeep, whilst potentially avoiding a myriad of behavioural and physiological changes that are known to result from social isolation.

Alcohol-induced changes in opioid peptide levels in adolescent rats are dependent on housing conditions.

BackgroundEndogenous opioids are implicated in the mechanism of action of alcohol and alcohol affects opioids in a number of brain areas, although little is known about alcohol's effects on opioids in the adolescent brain. One concern, in particular when studying young animals, is that alcohol intake models often are based on single housing that may result in alcohol effects confounded by the lack of social interactions. The aim of this study was to investigate short- and long-term alcohol effects on opioids and the influence of housing conditions on these effects.MethodsIn the first part, opioid peptide levels were measured after one 24-hour session of single housing and 2-hour voluntary alcohol intake in adolescent and adult rats. In the second part, a model with a cage divider inserted during 2-hour drinking sessions was tested and the effects on opioids were examined after 6 weeks of adolescent voluntary intake in single-and pair-housed rats, respectively.ResultsThe effects of single housing were age specific and affected Met-enkephalin-Arg6Phe7 (MEAP) in particular. In adolescent rats, it was difficult to distinguish between effects induced by alcohol and single housing, whereas alcohol-specific effects were seen in dynorphin B (DYNB), beta-endorphin (BEND), and MEAP levels in adults. Voluntary drinking affected several brain areas and the majority of alcohol-induced effects were not dependent on housing. However, alcohol effects on DYNB and BEND in the amygdala were dependent on housing. Housing alone affected MEAP in the cingulate cortex.ConclusionsAge-specific housing- and alcohol-induced effects on opioids were found. In addition, prolonged voluntary alcohol intake under different housing conditions produced several alcohol-induced effects independent of housing. However, housing-dependent effects were found in areas implicated in stress, emotionality, and alcohol use disorder. Housing condition and age may therefore affect the reasons and underlying mechanisms for drinking and could potentially affect the outcome of a number of end points in research on alcohol intake.

Overexpression of DeltaFosB in nucleus accumbens mimics the protective addiction phenotype, but not the protective depression phenotype of environmental enrichment.

Environmental enrichment produces protective addiction and depression phenotypes in rats. ΔFosB is a transcription factor that regulates reward in the brain and is induced by psychological stress as well as drugs of abuse. However, the role played by ΔFosB in the protective phenotypes of environmental enrichment has not been well studied. Here, we demonstrate that ΔFosB is differentially regulated in rats reared in an isolated condition (IC) compared to those in an enriched condition (EC) in response to restraint stress or cocaine. Chronic stress or chronic cocaine treatment each elevates ΔFosB protein levels in the nucleus accumbens (NAc) of IC rats, but not of EC rats due to an already elevated basal accumulation of ΔFosB seen under EC conditions. Viral-mediated overexpression of ΔFosB in the NAc shell of pair-housed rats (i.e., independent of environmental enrichment/isolation) increases operant responding for sucrose when motivated by hunger, but decreases responding in satiated animals. Moreover, ΔFosB overexpression decreases cocaine self-administration, enhances extinction of cocaine seeking, and decreases cocaine-induced reinstatement of intravenous cocaine self-administration; all behavioral findings consistent with the enrichment phenotype. In contrast, however, ΔFosB overexpression did not alter responses of pair-housed rats in several tests of anxiety- and depression-related behavior. Thus, ΔFosB in the NAc the shell mimics the protective addiction phenotype, but not the protective depression phenotype of environmental enrichment.

The effects of social learning on the acquisition of cocaine self-administration

BackgroundSocial learning models of substance use propose that drug-use behaviors are learned by observing and mimicking the behavior of others. The aim of this study was to examine the acquisition of cocaine self-administration in three groups of experimentally naïve rats: rats that were tested in isolation, rats that were tested in the presence of another rat that had access to cocaine and had previously been trained to self-administer cocaine, and rats that were tested in the presence of another rat that did not have access to cocaine. MethodsMale rats were reared in isolated or pair-housed conditions and implanted with intravenous catheters. Pair-housed rats were then assigned to drug-experienced or drug-naïve conditions. In the drug-experienced condition, one rat of each pair was trained to self-administer cocaine in isolation before the reintroduction of its partner. In the drug-naïve condition, one rat of each pair did not have access to cocaine for the duration of the study. For each group, the acquisition of cocaine self-administration was measured over 15 days in rats with access to cocaine but no prior operant training. ResultsRats tested with a drug-experienced partner were faster to acquire cocaine self-administration and emitted more active lever presses than rats tested with a cocaine-naïve partner. Data for the isolated control group fell between the other two groups on these measures. ConclusionThese data indicate that the acquisition of cocaine self-administration can either be facilitated or inhibited by social contact. Collectively, these results support a social learning model of substance use.

Post-weaning social isolation impairs observational fear conditioning

Many mammals can utilize social information to learn by observation of conspecifics (social learning). Social learning of fear is expected to be especially advantageous for survival. However, disruption of social development in early life can impair social cognition and might also be expected to disrupt social learning. Social isolation during a critical period of adolescence disrupts social development. The purpose of this study was to determine whether disruption of social development through post-weaning social isolation leads to impairments of social fear learning. Rats were reared in isolation or pair-housed from immediately post-weaning, for 3 weeks. Social fear learning in rats was acquired by observation of tone-footshock pairings administered to a conspecific. Isolation-reared rats displayed less conditioned freezing than pair-housed rats when tested the next day. This reduction of conditioned freezing was correlated with conspecific-oriented behaviors during conditioning, was measured despite similarities in demonstrator behaviors, and occurred despite a manipulation that equalized freezing during conditioning between the pair-housed and isolation-reared rats. The results could not be explained by abnormal sensitization to a repeated tone or deficits in freezing or direct fear conditioning. These results demonstrate that observational fear conditioning is impaired by social isolation, and provide a model to study impaired social affective learning. Impaired social cognition, manifested as inability to recognize or appropriately interpret social cues, is a symptom of several psychiatric disorders. Better understanding of the mechanisms of impaired social fear learning can lead to novel treatments for social cognition symptoms of psychiatric disorders.

Differential effects of environmental enrichment, social-housing, and isolation-rearing on a rat gambling task: Dissociations between impulsive action and risky decision-making

Decision-making deficits, measured using the Iowa Gambling Task (IGT), are observed in many psychiatric populations. Additionally, evidence suggests that the environment also influences the development of these same disorders. To determine the direct influence of the environment on decision-making by utilizing the rat gambling task (rGT), a risky decision-making test modeled after the IGT. Male rats reared in isolation, in pairs, or in an enriched environment were trained on the rGT as adults. During the rGT, animals chose from four different options. The optimal strategy on the rGT and IGT is the same: to favor options associated with smaller immediate rewards and less punishment/loss. Impulsive action is also measured during rGT performance by recording the number of premature responses made, similar to the five-choice serial reaction time task. Compared to pair-housed rats, isolated and environmentally enriched rats were slower at learning the optimal strategy. However, following training, only isolation-reared rats chose the disadvantageous options more often. Amphetamine altered decision-making on the rGT in socially housed animals, yet isolates were unaffected. Conversely, amphetamine increased premature responding similarly in all groups. This increase was attenuated by prior administration of a dopamine D(1) or D(2) antagonist; however, the ability of amphetamine to alter decision-making was not blocked by either drug. Housing environment affects animals' ability to learn and perform a decision-making task. Additionally, amphetamine's effect on impulsive action appears to be mediated by the dopaminergic system, whereas its effect on risky decision-making may be mediated by other neurotransmitters.

Interaction of environment and chronic methylphenidate on anxiety‐like behavior and dopamine receptors in adolescent rodents

Environmental rearing conditions are reported to alter behavior and dopamine (DA) transmission in rodents. Administration of DA‐ergic drugs, such as methylphenidate (MPH), may have varying effects depending on environmental exposure. This study explored the interactive effects of environment and chronic MPH on DA receptors and anxiety‐like behavior in adolescent rats. Sprague‐ Dawley rats (PND 21; n=8/group) were housed in isolated/impoverished (II), pair‐housed (PH), or socially/environmentally enriched (EE) conditions for 4 weeks. At 1 week, rats were implanted with ALZET minipumps that delivered saline or MPH (2 or 8 mg/kg/day) for 3 weeks. Post treatment, rats were tested for locomotor behavior and anxiety‐like behavior in an elevated plus maze (EPM). Rats were sacrificed at PND 50; brains processed for in vitro autoradiography to quantify DA D1 and D2 receptors in striatal regions. II rats had greater locomotor activity and spent less time in the open arms of the EPM than EE and PH rats. Environment did not affect the density of D1 or D2 receptors in striatal regions. MPH did not impact any measure of behavior. However, there was a dose‐dependent decrease in DA D2 receptor density in caudate putamen and a similar trend in accumbens across housing groups. While D1 receptor density was not affected by MPH across housing groups, preliminary results suggest that D1 receptors were increased by 8 mg/kg/day MPH in II rats, but decreased in PH and EE animals. Differences in locomotor and anxiety‐like behavior between II, PH, and EE animals highlight the importance of considering housing conditions in rodent studies. MPH did not impact behavior, but did decrease D2 receptor density across housing conditions. Despite no main effect of MPH on D1 receptors, there was a trend towards an interaction between MPH and housing. Thus, chronic MPH causes alterations in DA systems, which may appear more pronounced in animals with a history of isolation.

Environmental enrichment protects against the effects of chronic stress on cognitive and morphological measures of hippocampal integrity

Chronic stress has detrimental effects on hippocampal integrity, while environmental enrichment (EE) has beneficial effects when initiated early in development. In this study, we investigated whether EE initiated in adulthood would mitigate chronic stress effects on cognitive function and hippocampal neuronal architecture, when EE started one week before chronic stress began, or two weeks after chronic stress onset. Adult male Sprague Dawley rats were chronically restrained (6h/d) or assigned as non-stressed controls and subdivided into EE or non-EE housing. After restraint ended, rats were tested on a radial arm water maze (RAWM) for 2-d to assess spatial learning and memory. The first study showed that when EE began prior to 3-weeks of chronic stress, EE attenuated chronic stress-induced impairments in acquisition, which corresponded with the prevention of chronic stress-induced reductions in CA3 apical dendritic length. A second study showed that when EE began 2-weeks after the onset of a 5-week stress regimen, EE blocked chronic stress-induced impairments in acquisition and retention at 1-h and 24-h delays. RAWM performance corresponded with CA3 apical dendritic complexity. Moreover, rats in EE housing (control or stress) exhibited similar corticosterone profiles across weeks, which differed from the muted corticosterone response to restraint by the chronically stressed pair-housed rats. These data support the interpretation that chronic stress and EE may act on similar mechanisms within the hippocampus, and that manipulation of these factors may yield new directions for optimizing brain integrity and resilience under chronic stress or stress related neuropsychological disorders in the adult.

Environmental enrichment alters gene expression of steroidogenic enzymes in the rat hippocampus

Neuroactive steroids are synthesized in the central and peripheral nervous systems. The purpose of this study was to analyze the effects of environmental enrichment on neuroactive steroidogenesis in the rat hippocampus. Environmental enrichment rats were housed in a group of nine in a large cage and three groups of pair-housed rats were housed in a standard cage for 8 weeks. The levels of mRNAs for steroidogenic enzymes and proteins in hippocampus were quantified by real-time RT-PCR. Environmental enrichment increased the mRNA expression levels of 5α-reductase-1 and 3α-hydroxysteroid dehydrogenase, which catalyze synthesis of allopregnanolone from progesterone. Hence, environmental enrichment appears to affect allopregnanolone synthesis.

Sex differences, or not, in spatial cognition in albino rats: acute stress is the key

Male rats, Rattus norvegicus, typically outperform females in tests of spatial cognition. However, as stress affects cognition differently in the two sexes, performance differences may be an artefact of stress. Rats face at least two sources of stress during an experiment: the test situation (acute) and housing conditions (chronic, e.g. isolation). We used a task (the Morris water maze, MWM) that allowed testing of both spatial working and reference memory to investigate whether chronic stress (isolation housing) and/or acute stress (the task) has a differential impact on spatial cognition in male and female albino rats. Irrespective of age at the onset of isolation housing, isolated rats were not spatially impaired relative to pair-housed rats. However, the acute stress of the task led to adult males apparently outperforming adult females: adult females took longer to reach the platform than did males because they spent more time in thigmotaxis (swimming close to the wall) during testing. In juvenile rats, the stress caused by swimming in the MWM resulted in both males and females being highly thigmotactic and no sex difference in performance. We conclude that stress can lead to apparent differences between the sexes in performance on a spatial cognition task.

Sex differences in spatial cognition are not caused by isolation housing

Summary In mammals, males typically have better spatial ability than do females. However, most of the data come from laboratory tests and it is possible that factors impacting on the captive animal cause the observed sex differences in spatial cognition. A common influence on cognitive ability is stress, which may have its effect acutely, in the testing situation, or chronically, due to the housing conditions. We used a spatial working and reference memory task (the Morris water maze) to investigate if isolation housing had a differential impact on spatial cognition in male and female rats. Either as juveniles or as adults, rats were housed in pairs or in isolation. We also manipulated the duration of isolation housing. Regardless of housing condition, we found a sex difference in spatial ability only in the youngest rats. However, we found no evidence that isolated rats were spatially impaired relative to pair-housed rats. We also found no difference in body weight, food intake or bar biting behaviour (indicators of welfare in rodents) between pair and isolated rats. We conclude that isolation housing causes insufficient stress to cause sex differences in spatial cognition.

Social status modulates basal IL-1 concentrations in the hypothalamus of pair-housed rats and influences certain features of stress reactivity

Recent findings from our laboratory and others indicate that exposure to stress can increase expression of the pro-inflammatory cytokine interleukin-1 (IL-1). In a series of studies examining this response, we observed pronounced differences in baseline levels of hypothalamic IL-1 of pair-housed rats. We hypothesized that these pair-wise differences might be a result of prolonged social stress associated with dominance/submissiveness, and that the submissive animal would show heightened baseline levels of IL-1. In order to test this hypothesis, we utilized a food competition paradigm (access to cheerios) to assess dominance within a dyad prior to the assessment of hypothalamic IL-1 levels. Based on the results of this test, clear dominance hierarchies were observed in ∼50% of the dyads, a ratio comparable to what has been reported previously. More importantly, this dominant/submissive categorization could be used to predict pair-wise differences in hypothalamic IL-1 with greater than 90% accuracy. Specifically, the submissive rat in each dyad (determined a priori) consistently evinced hypothalamic IL-1 levels that were nearly double that of its dominant cage mate. Further studies demonstrated that submissive rats showed a more rapid and pronounced hyperthermic response to novel environment stress relative to dominant rats. Interestingly, social status had no effect on corticosterone reactivity, even when the nature and intensity of the stressor was varied. Finally, maintenance of a clear dominance hierarchy obfuscated hypothalamic IL-1 responses to footshock exposure, with the most robust increases in hypothalamic IL-1 provoked by footshock being observed in pairs where there was no clear dominance hierarchy. Together, these findings suggest that social status can have a significant impact on stress reactivity and neuroimmune consequences of stressor exposure even in the unperturbed home cage environment.

Anabolic androgenic steroid affects social aggression and fear-related behaviors in male pair-housed rats

This study examines the effect of chronic administration of the anabolic androgenic steroid nandrolone decanoate (ND) on dominant and subordinate male rats in a pair-housed condition. Pair-housed rats were assessed for dominance status based on their behavior and alterations in body weights. Throughout the study the rats were allowed limited social interactions on a daily basis. At all other times, a Plexiglas divider kept the rats separated, allowing olfactory and visual contact between the cage mates while preventing significant physical contact. One week into the study all subjects were subcutaneously implanted with a pellet that continuously infused either ND (15 mg/kg/day) or placebo for 21 days. Following the pellet implant, behavioral tests including reassessment for dominance status, and a conditioned fear test were conducted over a period of approximately 2 months to investigate possible long-term changes. The main finding is that during the allowed social interactions, the dominant ND-pretreated rats spent more time on highly aggressive behaviors than the dominant placebo-treated rats. In addition, the probability for highly aggressive behaviors was maintained for the ND-treated rats throughout the study, whereas it was decreased for the placebo-treated rats. The ND-treated subordinate rats showed less fear in a potential threatening situation compared to placebo-treated controls. These findings support the relatively long-term behavioral changes that have been seen in humans after abuse of ND and other anabolic androgenic steroid compounds.

The effects of changes in housing on feeding and wheel running

The experiments explored the effects on feeding when rats were moved between individual and paired housing. In Experiment 1, rats moved to paired housing showed a 3-day suppression in feeding (initially 23%) compared to chronically individual- or pair-housed rats. In Experiment 2, half of the rats from the two control groups of Experiment 1 were moved between individual and paired housing on alternate days. Only the rats moved to paired housing showed a feeding suppression (initially 40%), but the nature of the suppression differed from Experiment 1; it appeared that only one rat of each pair showed a feeding suppression. Experiment 3 examined simultaneous introduction of running wheels and moves to paired housing. The feeding suppression induced by the move to paired housing was more immediate and shorter lived than the wheel-induced suppression. Unlike wheel access, paired housing produced only a temporary suppression of body weight. These experiments suggest that the relatively simple manipulation of moving rats from individual to paired housing results in a temporary stress-induced decrease in feeding.

Voluntary ethanol intake of individually- or pair-housed rats: Effect of ACTH or dexamethasone treatment

The effect of ACTH or dexamethasone treatment on ingestion of 10% ethanol, 0.5 M NaCl and water was studied in individually- and pair-housed rats. Crowding or decreasing the amount of space per rat by increasing the number of rats per cage from 1 to 2, together with the associated increase in social interactions caused a large increase in ethanol intake. In pair-housed rats and in rats housed alone, ACTH treatment caused a large increase in Na intake but no change in ethanol intake. In pair-housed rats and in rats housed alone, dexamethasone treatment caused no change in either ethanol or Na intake. Thus, it would appear that the induction or maintenance of a high ethanol intake of rats during crowding, a presumed social stressor, can not be attributed entirely to either an increase in blood ACTH levels with the subsequent increase in glucocorticoid hormones or to a decrease in blood ACTH and natural glucocorticoid hormone levels. However, the possibility that ACTH and/or adrenocorticoid hormones, combined with other physiological or environmental factors, causes stressor-induced ethanol intake cannot be excluded.

The effect of the beta-carboline FG 7142 on the behaviour of male rats in a living cage: an ethological analysis of social and nonsocial behaviour.

The effects of FG 7142, a beta-carboline benzodiazepine receptor partial inverse agonist, on the social behavior of pair-housed rats were investigated. Four 6-min dyadic social encounters in a living cage were observed in a paradigm in which one member of a pair of rats was injected. The four injection groups (n = 8) were vehicle control, and FG 7142 at 2.5, 5.0 and 10.0 mg/kg, respectively. All injections were administered 2 min before the start of the first observation trial. Compared to the effects of vehicle alone, FG 7142 decreased aggressive behaviour but did not change the level of total social interaction. Thus there were compensating increases in approaching and avoiding behaviours following the administration of FG 7142. Locomotion declined marginally and immobility increased in FG 7142-injected rats. FG 7142 decreased the incidence of self-grooming. The evidence is consistent with a relatively selective reduction in intraspecies aggression in male rats after the injection of the beta-carboline inverse agonist.

Behavioural, physiological and immunological consequences of social status and aggression in chronically coexisting resident-intruder dyads of male rats

The behavioural and physiological consequences of social status and reciprocal fighting in resident-intruder dyads of Long Evans male rats were evaluated. Before a chronic cohabitation of 10 days, residents and intruders were individually housed for one month to increase their aggressiveness. Control animals included isolates, i.e., animals kept individually housed throughout the experiment and pair-housed rats, i.e., pairs of rats housed together from their rats in the laboratory. In 19 out of 20 dyads, a clear dominance relationship developed with an advantage to the resident in 68% of the cases. Dominants showed more exploratory activity than subordinates in an open-field test at the end of the cohabitation period; subordinates groomed longer than animals from other experimental groups. Dominants had lower pain thresholds than individually and pair-housed animals. Both dominants and subordinates had higher tyrosine hydroxylase enzymatic activities in the left adrenal than isolated and pair-housed rats. Subordinates lost body weight and had higher plasma corticosteroid concentrations than animals from the other experimental groups. In addition, they had smaller thymus glands and reduced spleen lymphocyte responses to mitogenic stimulation in vitro, in comparison to dominant animals. These results show that subordination in the dyadic resident-intruder paradigm leads to a complex syndrome of behavioural and physiological changes, some of which may be modulated by the intensity of aggressive interactions.

Spatial interaction between cage and test environments: Position preferences of young isolated and pair-housed rats

This study further investigates the relationship between cage size and activity in the open field. Male hooded rats were housed at weaning either individually or in pairs in small, medium, or large cages and observed after 15 days in an apparatus providing a choice between three environments of identical construction and dimensions to the different home cages. Three experimental conditions were employed. Both isolated and pair-housed rats were tested alone in the unfamiliar apparatus and some isolated rats were first habituated to the apparatus. Time spent in each size apparatus component was recorded. The results failed to explain earlier findings of a positive relationship between open-field behavior and cage size in isolated rats, in terms of spatial preference. Isolated component preferences were inversely related to cage size, while pair-housed preferences were positively related to cage size. Component preference was not related to component area or perimeter distance. The implications of this spatial behavior for exploration and emotionality theories are discussed.