Effects of isolation-rearing on intracranial self-stimulation reward of the lateral hypothalamus: baseline assessment and drug challenges

There is evidence that isolation rearing produces down-regulation of the dopamine D2 receptor. Therefore, isolation rearing should also modify the effects of D2 antagonists on intracranial self-stimulation (ICSS) reward. This study investigated the effect of isolation rearing on ICSS reward, and modulation of that reward by SCH23390, Raclopride and MK-801. Sprague-Dawley rats were reared alone (isolates) or in pairs from day 21 postnatal to day 75 postnatal. At this time, all rats were implanted with monopolar stimulating electrodes in the lateral hypothalamus. The ICSS rate-frequency curve-shift method was used to assess reward and operant motor function at baseline and after administration of SCH-23390 (D1 antagonist: 0.02, 0.06, 0.2 mg/kg), Raclopride (D2 antagonist: 0.01, 0.025, 0.06 mg/kg), and MK-801 (non-competitive NMDA receptor antagonist: 0.1, 0.2 mk/kg). Isolation-reared rats displayed similar measures of both basal reward and motor function when compared to socially reared controls. Isolation-reared rats were subsensitive to the reward decreasing effects of Raclopride. Socially reared rats were observed to have more variant baseline reward measures, and could be divided into distinctly different groups with different basal reward function. Isolation-rearing down-regulates D2 function but does not affect basal reward function, but some unknown factor in the social rearing environment did have a substantial effect on basal reward function.     

Increased Drinking following Social Isolation Rearing: Implications for Polydipsia Associated with Schizophrenia

Primary polydipsia, excessive drinking without known medical cause, is especially associated with a diagnosis of schizophrenia. We used animal models of schizophrenia-like symptoms to examine the effects on schedule-induced polydipsia: post-weaning social isolation rearing, subchronic MK-801 treatment (an NMDA-receptor antagonist) or the two combined. Male, Sprague-Dawley rats reared in groups or in isolation beginning at postnatal day 21 were further divided to receive subchronic MK-801 (0.5 mg/kg twice daily) or saline for 7 days beginning on postnatal day 62. Following a 4-day withdrawal period, all groups were trained on a schedule-induced polydipsia paradigm. Under food-restriction, animals reared in isolation and receiving food pellets at 1-min intervals developed significantly more drinking behavior than those reared with others. The addition of subchronic MK-801 treatment did not significantly augment the amount of water consumed. These findings suggest a predisposition to polydipsia is a schizophrenia-like behavioral effect of post-weaning social isolation.     

Behavioral reactions to stimulation of the emotiogenic zones of the brain in the rats with different individual experience

Emotional reactions evoked by electric stimulation of the hypothalamus and amygdala were studied in white outbred rats, grown either in conditions of isolation or in community. The method of self-stimulation in shuttle box was used. The values of self-stimulation reaction were significantly lower and those of avoidance reactions were higher in animals bred in isolation. Their food-procuring behaviour disappeared faster at stimulation of the negative emotiogenic zones. The observed differences are due to plastic reorganization of the brain reinforcing systems.     

Le comportement du Chevrotain africain,Hyemoschus aquaticus Ogilby (Artiodactyla,Ruminantia)

The purpose of this study was to test the influence of social isolation during infancy on learning behavior in rats. Rats, reared isolated (n = 55) or paired (n = 60) for a period of 50 days following weaning, were tested on different learning tasks in an operant conditioning chamber. Isolation rearing produced significantly better learning when the training period directly followed the different housing conditions. After a 28 day period of social isolation for the subjects raised paired during infancy no differences in learning between isolated and paired reared animals could be observed. The differences are discussed in relation to the effects of stimulus deprivation.     

Effect of Time of Separation from Mother on Behavior in Open Field and on State of the Sympathoadrenal System in Rats Reared under Conditions of Social Isolation

The level of catecholamine excretion with urine and behavior in the open field test were studied in male Wistar rat pups separated from female at the age of 17, 21, 30, and 36 days and reared subsequently for 40 days under conditions either of social isolation or in the group with their kin. It was shown that in rats weaned at the age of 17 and 21 days, the subsequent rearing in isolation led to an increase of the urinary excretion of adrenaline and a decrease, of noradrenaline. After isolation at the 30th day, there was revealed an increase of the excretion level both of adrenaline and of noradrenaline. The social isolation also leads to an increase of horizontal motor activity in rats weaned at the age of 17 days and to a decrease of this activity in rats weaned at the 30-day age. After isolation at the age of 21 days, an earlier extinguishing of the activity it was observed at repeated testing. Isolation at the age of 36 days, on the contrary, led to a decrease of the extinguishing process. Thus, it is shown that the social isolation produces changes in rat behavior and has a stress effect on the animals; the effect of isolation depends on duration of the rearing of rats in the nest with the female.     

Postweaning environmental and social factors influencing the onset and expression of agonistic behavior in Norway rats

Two studies were conducted to determine the importance of the postweaning environment and social milieu in regulating the expression of intraspecific aggression in Norway rats. In Experiment 1, male rats were housed either individually or in pairs at 21 days of age. In addition, one-half of the singly housed and paired animals were given experiences with intruders during maturation. At 85 days of age, all animals were given a brief intruder test and then removed from their postweaning environment and provided individually with homecages for a two week period until tested for aggression toward intruders. Results of intruder tests given during maturation indicated agonistic exchanges appeared earlier and more frequently in cages housing a single resident than cages with cohabiting males. However, agonistic exchanges between singly reared residents and intruders had detrimental consequences in adulthood especially under long-term combat situations. That is, although individually reared animals, with early fighting experiences, were capable of initiating intraspecific aggression, such individuals were unable to defend their homecage over a long period of time as evidenced by the high number of wounds and tendency to lose body weight during adult fighting.In Experiment 2, male Norway rats were reared in pairs from 21 days of age and identified as dominant or subordinate on the basis of intracolony social interactions shown during maturation. At 80 days of age, animals were paired with individually reared males in an unfamiliar cage for a 20 day period and examined for agonistic behavior toward intruders at 100 days of age. Group-reared subordinate males exhibited defensive behavior during confrontations with individually reared animals and incurred more wounds and lost more body weight than their cohabiting partner. In addition, subordinate males showed significantly fewer offensive postures toward intruders than individually reared cohabitants. In contrast, group-reared dominant animals did not differ from individually reared males in display of agonistic patterns, in number of wounds, and body weight changes during the period of cohabitation. These findings demonstrate that early rearing factors have pronounced effects on agonistic behavior. Animals experiencing defeat during development are more likely to lose agonistic confrontations in unfamiliar territory than either animals dominant in their early social interactions or animals without the experience of winning or losing agonistic encounters. These results have implications for the understanding of agonistic behavior and predicting outcomes of animal contests, and reveal important differences in agonistic experiences among animals reared in groups.     

Social isolation improves working memory at reversal but not primary radial-arm maze learning in rats

Social isolation starting from the 21st day of birth affected neither a short-term nor a long-term memory in male rats at primary acquisition learning in an 8-arm radial maze. A number of the short-term and long-term memory errors were substantially decreased during primary learning but the difference between groups was not significant. Isolates were faster to start a search in an individual trial and took less time to finish offa trial. During the reversal learning, when baited and non-baited arms were reversed, the isolates outperformed of socially reared rats on working but not reference memory task. In overall they made twice less working memory errors than socially reared animals. During the reversal learning the isolates were also faster than non-isolates in initiation and completion of a trial. Maternal separation of rat's pups on the postnatal days 1-21 for 4 hr per day did not affect either working or reference memory on both primary and reversal learning. The data obtained are discussed on basis of influence of stress in early postnatal life on hypothalamo-pituitary axis and its effects on behavior of adult animals.     

Adult rats exposed to early-life social isolation exhibit increased anxiety and conditioned fear behavior, and altered hormonal stress responses

Social isolation of rodents during development is thought to be a relevant model of early-life chronic stress. We investigated the effects of early-life social isolation on later adult fear and anxiety behavior, and on corticosterone stress responses, in male rats. On postnatal day 21, male rats were either housed in isolation or in groups of 3 for a 3 week period, after which, all rats were group-reared for an additional 2 weeks. After the 5-week treatment, adult rats were examined for conditioned fear, open field anxiety-like behavior, social interaction behavior and corticosterone responses to restraint stress. Isolates exhibited increased anxiety-like behaviors in a brightly-lit open field during the first 10 min of the test period compared to group-reared rats. Isolation-reared rats also showed increased fear behavior and reduced social contact in a social interaction test, and a transient increase in fear behavior to a conditioned stimulus that predicted foot-shock. Isolation-reared rats showed similar restraint-induced increases in plasma corticosterone as group-reared controls, but plasma corticosterone levels 2 h after restraint were significantly lower than pre-stress levels in isolates. Overall, this study shows that isolation restricted to an early part of development increases anxiety-like and fear behaviors in adulthood, and also results in depressed levels of plasma corticosterone following restraint stress.     

Nest Making and Oxytocin Comparably Promote Wound Healing in Isolation Reared Rats

Background

Environmental enrichment (EE) fosters attachment behavior through its effect on brain oxytocin levels in the hippocampus and other brain regions, which in turn modulate the hypothalamic-pituitary axis (HPA). Social isolation and other stressors negatively impact physical healing through their effect on the HPA. Therefore, we reasoned that: 1) provision of a rat EE (nest building with Nestlets®) would improve wound healing in rats undergoing stress due to isolation rearing and 2) that oxytocin would have a similar beneficial effect on wound healing.

Methodology/Principal Findings

In the first two experiments, we provided isolation reared rats with either EE or oxytocin and compared their wound healing to group reared rats and isolation reared rats that did not receive Nestlets or oxytocin. In the third experiment, we examined the effect of Nestlets on open field locomotion and immediate early gene (IEG) expression. We found that isolation reared rats treated with Nestlets a) healed significantly better than without Nestlets, 2) healed at a similar rate to rats treated with oxytocin, 3) had decreased hyperactivity in the open field test, and 4) had normalized IEG expression in brain hippocampus.

Conclusions/Significance

This study shows that when an EE strategy or oxytocin is given to isolation reared rats, the peripheral stress response, as measured by burn injury healing, is decreased. The findings indicate an association between the effect of nest making on wound healing and administration of the pro-bonding hormone oxytocin. Further elucidation of this animal model should lead to improved understanding of how EE strategies can ameliorate poor wound healing and other symptoms that result from isolation stress.     

Temporal shift between daily sperm movement and mating (sperm reflux) in the Asian comma butterfly,Polygonia c-aureum

Temporal changes in mating behavior and daily sperm movement were examined in adult males of the butterfly Polygonia c-aureum L. (Lepidoptera: Nymphalidae), reared under L15:D9 photoperiod at 21 °C. In this butterfly, when extra sperm is present in the duplex (storage organ) at mating, sperm reflux occurs: extra sperm are moved from the duplex to the lower and middle portions of the vasa deferentia by peristaltic movement of the lower portion of the vasa deferentia. It is known in several moth species that daily sperm movement from the testis to the upper portion of the vasa deferentia and from the upper vasa deferentia to the duplex occurs in a circadian manner. If sperm reflux and daily sperm movement occur at the same time, it could create confusion, because these sperm movements are directed oppositely. In this study, we examined the temporal changes of sperm movement and mating behavior in P. c-aureum in order to clarify whether the two sperm movements are separated in time. Our results showed that most matings occurred during the second half of the photophase, whereas daily eupyrene (nucleate) sperm movement from the upper to the middle portion of the vasa deferentia occurred immediately after light on (early photophase), indicating the presence of a time lag between mating and daily sperm movement. As sperm reflux occurs during mating, these two oppositely directed sperm movements could be performed without conflict.     

Early social environment interferes with the development of NADPH-diaphorase-reactive neurons in the rodent orbital prefrontal cortex

The influence of early parental deprivation on the development of NADPH-diaphorase-(NO-synthase) reactive neuron numbers in subregions of the orbital prefrontal cortex (ventrolateral orbital, lateral orbital, and agranular insular cortex) was quantitatively investigated in the precocious lagomorph Octodon degus. Forty-five-day-old degus from three groups were compared: (1) repeated parental separation: degus that were repeatedly separated from their parents during the first three postnatal weeks and thereafter raised in undisturbed social conditions; (2) chronic isolation: degus that were raised under undisturbed social conditions until postnatal day 21, and then were reared in chronic social isolation; and (3) control: degus that were reared undisturbed in their families. Compared to the control animals the ventrolateral orbital prefrontal cortex and agranular insular cortex of the two deprived groups displayed significantly decreased density of NADPH-diaphorase-reactive neurons (down to 62% in the ventrolateral orbital prefrontal cortex of males, 70% in the agranular insular cortex, and in the lateral orbital prefrontal cortex 80% in both genders). These results confirm that early changes of social environment interferes with the development of limbic circuits, which might determine normal or pathological behaviors in later life.     

Early predictors of self-biting in socially-housed rhesus macaques (Macaca mulatta)

The development of self-biting behavior in captive monkeys is little understood and poses a serious risk to their well-being. Although early rearing conditions may influence the expression of this behavior, not all animals reared under similar conditions self-bite. The purpose of this study was to examine the effects of three rearing conditions on biting behavior and to determine whether early infant behavior can predict later self-biting. The subjects were 370 rhesus macaques born at the National Institutes of Health (NIH) Animal Center between 1994 and 2004. They were reared under three conditions: mother-reared in social groups (n=183), peer-reared in groups of four (n=84), and surrogate-peer-reared (n=103). Significantly more surrogate-peer-reared animals self-bit compared to peer-only or mother-reared animals. There was no sex difference in self-biting, but this result may have been affected by a sex bias in the number of observations. The durations of behaviors exhibited by the surrogate-peer-reared subjects were recorded in 5-min sessions twice a week from 2 to 6 months of age while the animals were in their home cages and play groups. In the play-group situation, surrogate-peer-reared subjects who later self-bit were found to be less social and exhibited less social clinging than those that did not self-bite. Home-cage behavior did not predict later self-biting, but it did change with increasing age: surrogate clinging and self-mouthing decreased, while environmental exploration increased. Our findings suggest that surrogate rearing in combination with lower levels of social contact during play may be risk factors for the later development of self-biting behavior.     

The weight of the clan: Even in insects, social isolation can induce a behavioural syndrome

Social isolation has dramatic consequences on the development of individuals of many vertebrate species, and it induces a set of behavioural disturbances rending them unable to process environmental as well as social stimuli appropriately. We hypothesized that isolation syndrome is a ubiquitous trait of social life that can be observed in a wide array of species, including invertebrates. Here we report that gregarious cockroaches (Blattella germanica) reared in isolation showed (i) stronger exploration-avoidance, (ii) reduced foraging activity, (iii) reduced willingness to interact socially, and (iv) reduced ability to assess mating partner quality than conspecifics reared in groups. We demonstrate the occurrence of a behavioural syndrome induced by social isolation, similar to syndromes described in vertebrates, revealing the importance of social interactions and group-living in this non-eusocial insect species. We suggest that investigating social isolation effects on individual development should provide interesting results to assess social cohesion of species and thus constitute an additional tool for comparative studies focusing on the evolution of social life.     

The effects of rearing method on social behaviors of mentored,captive‐reared juvenile California condors

Puppet‐reared and parent‐reared captive‐bred California condor (Gymnogyps californianus) juveniles were studied before their release into the wild. Behavioral data were collected during social interactions within two cohorts of juveniles (N = 11) and their adult mentors (N = 5). The purposes of this study were to (1) document the social behaviors of mentored juvenile California condors, and (2) compare social behaviors for two different rearing methods (puppet‐reared versus parent‐reared) during two phases of the mentoring process (San Diego Wild Animal Park versus release sites). Of the 17 behaviors examined by 2 × 2 analyses of variance (ANOVAs), two significant interactions between the rearing method and mentoring phase were found: pulls feathers and feeds alone. For both behaviors, parent‐reared condors engaged in these activities more often at the zoo and less often at the release pens than did the puppet‐reared condors. The main effect of rearing was also significant for two behaviors: near others, and receives contact aggression from other. Parent‐reared birds were more likely to be near another bird and receive contact aggression, regardless of mentoring phase, than puppet‐reared birds. The effect size for 16 of the 17 behaviors was greater for the rearing method than for mentoring phase. Rearing method differences may persist long‐term, as parent‐reared adult mentors were significantly more aggressive than puppet‐reared adult mentors. Dominance relations were examined for both cohorts, with the first cohort exhibiting a strong linear relationship (h′ = 0.86, P = 0.018), whereas the second cohort exhibited a moderate but non‐significant linear hierarchy (h′ = 0.63, P = 0.21). The rearing method had no effect on dominance among the juveniles, but adults were probably dominant to juveniles (P = 0.052; the difference was nearly significant). Although social behaviors between the two rearing groups were similar in most respects, this study is the first to document measurable differences between puppet‐ and parent‐reared captive‐bred California condor juveniles. Zoo Biol 27:1–18, 2008. © 2007 Wiley‐Liss, Inc.     

Early social environment interferes with the development of NADPH–diaphorase‐reactive neurons in the rodent orbital prefrontal cortex

The influence of early parental deprivation on the development of NADPH–diaphorase–(NO‐synthase) reactive neuron numbers in subregions of the orbital prefrontal cortex (ventrolateral orbital, lateral orbital, and agranular insular cortex) was quantitatively investigated in the precocious lagomorph Octodon degus. Forty‐five‐day‐old degus from three groups were compared: (1) repeated parental separation: degus that were repeatedly separated from their parents during the first three postnatal weeks and thereafter raised in undisturbed social conditions; (2) chronic isolation: degus that were raised under undisturbed social conditions until postnatal day 21, and then were reared in chronic social isolation; and (3) control: degus that were reared undisturbed in their families. Compared to the control animals the ventrolateral orbital prefrontal cortex and agranular insular cortex of the two deprived groups displayed significantly decreased density of NADPH–diaphorase‐reactive neurons (down to 62% in the ventrolateral orbital prefrontal cortex of males, 70% in the agranular insular cortex, and in the lateral orbital prefrontal cortex 80% in both genders). These results confirm that early changes of social environment interferes with the development of limbic circuits, which might determine normal or pathological behaviors in later life. © 2004 Wiley Periodicals, Inc. J Neurobiol, 2005     

Context-dependent aggregation in Common Frog Rana temporaria tadpoles: influence of developmental stage, predation risk and social environment

1. This study examines the aggregation behaviour and activity of larvae of the Common Frog Rana temporaria in relation to the early social environment, ontogeny and the presence of chemical cues from a predatory fish to address three main questions: (i) Does previous social interaction influence aggregation behaviour in later developmental stages? (ii) To what extent does aggregation behaviour depend upon the risk level perceived by the individual? (iii) Does aggregation behaviour change through ontogeny?
2. Tadpoles were reared from eggs either in small groups or in complete isolation. In relatively early stages of development, tadpoles showed no preference for the side of the test container containing siblings over the side containing no larvae regardless of the social context experienced (isolation or contact with siblings).
3. The presence of chemical cues from a potential predator did not trigger the aggregation behaviour of tadpoles, but it had a remarkable effect on their activity and spatial distribution. Tadpoles exposed to water preconditioned by a predator spent significantly less time swimming and avoided the central area of the test container more frequently than did controls exposed to unconditioned water.
4. Tadpoles were more active, avoided the central section and associated preferentially with conspecifics (siblings) at later stages of development regardless of the social conditioning they had previously experienced.
5. Individuals reared in groups were twice as active as individuals reared in isolation. This suggests that the early social environment experienced by larvae can influence future behaviour and thereby growth and development rates.
6. The expression of conspecific attraction is probably linked to an ontogenetic shift in larval behaviour. However, reduced activity, rather than aggregation, appears to be the basic antipredator mechanism in larval Common Frog.     

Behavioral interactions in cocaine-treated rats

Effects of a single dose of cocaine (10 mg/kg,i.p.) were studied on spontaneous motor activity (SMA), stereotypy. self-stimulation of the posterior hypothalamus or area ventralis tegmentum (A10 area) and nonreinforced bar-pressing in rats. Interactions among these behaviors were investigated under different experimental designs. Cocaine caused an initial increase of SMA, which was later decreased significantly, as stereotypy became more prominent. Cocaine also caused facilitation of self-stimulation responding that further increased SMA. In both groups of self-stimulating rats, stereotypy disappeared completely during self-stimulation. Following cessation of self-stimulation, SMA and bar-pressing (that became nonreinforced) decreased, and stereotypy reappeared in both groups of rats.     

Acceleration of sexual maturation in female rats by male stimulation.

In comparison to controls reared in isolation, female rats reared in the presence of an adult male from weaning attained puberty 5 days earlier when living in groups or almost 9 days earlier when living singly with a male. Accelerated sexual development occurred in the absence of accelerated physical growth.     

Effect of Isolation-Rearing on Conditioned Dopamine Release In Vivo in the Nucleus Accumbens of the Rat

Abstract: Intracerebral microdialysis in conjunction with HPLC coupled to electrochemical detection was used to investigate the effect of isolation-rearing in the rat on extracellular dopamine (DA) and its metabolites in vivo, in the shell region of the nucleus accumbens, in response to footshock and in relation to a conditioned emotional response. Male Lister hooded rats were reared from weaning for 6–8 weeks in either social isolation or groups of five. In the training phase, rats were exposed to a novel environment for 10 min where they experienced mild footshock. Footshock caused an immediate increase in basal extracellular DA levels in both rearing groups relative to control rats. However, the increase in extracellular DA was prolonged in the case of the isolation-reared rats and significantly greater than in group-reared rats. Exposure to the novel environment without shock (control groups) did not significantly alter basal extracellular DA in the nucleus accumbens shell; 140 min later rats were returned to the testing box (contextual stimulus) without receiving footshock. The contextual stimulus increased basal extracellular DA in the nucleus accumbens of both groups of rats with respect to controls; however, this increase was significantly greater and more prolonged in isolates. Extracellular levels of the metabolites 3,4-dihydroxyphenylacetic acid and homovanillic acid did not differ between isolation- and group-reared rats, and they were not significantly affected by either footshock or the contextual stimulus. These results suggest that exposure to footshock and a contextual stimulus are associated with increases in basal extracellular DA levels in the nucleus accumbens shell. The results also support evidence in favour of an isolation-induced enhancement in dopaminergic activity in the nucleus accumbens, which probably underlies aspects of the behavioural syndrome associated with isolation.     

Epidemiology of Ostertagia ostertagi in dairy cow from different breeding systems

The epidemiology of Trichostrongylidae infections in three dairy cow groups reared under different conditions and in different habitats was studied with particular emphasis on Ostertagia ostertagi. In cows bred intensively in swampy lowlands (Po River Valley) the prevalence of infection was 43% and the mean intensity 567. In cows bred traditionally (Friuli, Northern-Eastern Italy), the prevalence in permanently housed cows was 24% (mean intensity: 13). In grazing cows, it was 68% (mean intensity: 120). The distribution of parasite numbers (O. ostertagi) per host was studied with the negative binomial model. The observed data were also compared by the method of cumulative relative frequencies. The results suggest that parasitism is also important in cows reared intensively.