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PubMed2004

Maintenance in working memory or response selection? Functions of NMDA receptors in the pigeon "prefrontal cortex".

The prefrontal cortex is involved in various aspects of working memory like stimulus maintenance and response selection functions. Neurobehavioral studies and neurocomputational models assume a role for NMDA receptors in prefrontal cortex for maintenance processes, while our previous studies on NMDA receptors in the avian prefrontal cortex-analogue, the nidopallium caudolaterale (NCL), showed them to be involved in response selection functions. Various tasks used in PFC-related research address in fact both functions, so they cannot disambiguate their separate contributions to performance. In order to investigate the role of NMDA receptors in avian NCL for stimulus maintenance and response selection, we trained pigeons in a delayed matching-to sample (DMTS) task, requiring both functions, and a simultaneous matching to sample (SMTS) task, requiring only response selection. After reaching criterion, pigeons had to perform the tasks alternately under local NMDA receptor blockade in NCL (DL-AP5) and after infusion of vehicle (saline solution). Blockade of NCL-based NMDA receptors led to significant increases in error rates in both DMTS and SMTS--compared with the same subjects' performance during training and in the control condition. However, there was no additional increase in errors due to the additional maintenance component, so the impairment appears to be due to deficits in adequate selection of responses, the function necessary for both tasks. We conclude that NMDA receptors in the pigeon NCL participate in response selection rather than stimulus maintenance in tasks requiring the processing of context information.

Animals↗
PubMed2005

The effect of orbital prefrontal cortex lesions on performance on a progressive ratio schedule: implications for models of inter-temporal choice.

In a previous experiment [Kheramin S, Body S, Mobini S, Ho M-Y, Velazquez-Martinez DN, Bradshaw CM, et al. Effects of quinolinic acid-induced lesions of the orbital prefrontal cortex on inter-temporal choice: a quantitative analysis. Psychopharmacology 2002;165: 9-17], destruction of the orbital prefrontal cortex (OPFC) in rats altered choice between two delayed food reinforcers, enhancing preference for the larger reinforcer. Theoretical analysis based on a quantitative model of inter-temporal choice [Ho M-Y, Mobini S, Chiang T-J, Bradshaw CM, Szabadi E. Theory and method in the quantitative analysis of 'impulsive choice' behaviour: implications for psychopharmacology. Psychopharmacology 1999;146:362-72] indicated that the lesion had increased the relative value of the larger of the two reinforcers due to a general reduction of absolute reinforcer value. The present experiment tested this hypothesis using a reinforcement schedule that did not entail either explicit choice or delayed reinforcement. Ten rats received quinolinic acid-induced lesions of the OPFC, and ten rats received sham lesions. The rats were trained under a progressive-ratio schedule of food reinforcement for 60 daily sessions. Response rates in successive ratios were a bitonic (inverted-U) function of ratio size. Analysis of the data using a three-parameter equation derived from a quantitative model of ratio schedule performance [Killeen PR. Mathematical principles of reinforcement. Behav. Brain Sci. 1994;17:105-72] revealed that the parameter specifying hypothetical reinforcer value was significantly lower in the OPFC-lesioned group than in the sham-lesioned group, consistent with the hypothesis that destruction of the OPFC resulted in devaluation of the food reinforcer.

Animals↗
PubMed2005

Sex-selective effects of neonatal isolation on fear conditioning and foot shock sensitivity.

Our previous work demonstrates enduring effects of the early life stress of neonatal isolation (ISO). ISO facilitates appetitive response learning in adult female, but not male rats, and enhances corticosterone levels and stress responsivity in infant and juvenile rats of both sexes. Corticosterone acts at brain areas such as hippocampus that are rich in glucocorticoid receptors, differentiate postnatally, are sexually dimorphic, and involved in learning. Thus, ISO is hypothesized to alter aversive learning in adult rats in a sex-specific manner. This study tests this hypothesis using context and cue fear conditioning. Pups were isolated for 1h a day on postnatal (PN) days 2-9 or were non-handled and were then tested in adulthood (PN70-90). In Experiment 1, context- and cue-elicited freezing and ultrasonic vocalizations (USVs; 22 kHz range) were measured. Experiments 2-4, respectively, examined three unconditioned foot shock responses (flinch, jump, vocalization), unconditioned fear (time in center of an open, novel arena), and appetitive (stroking-induced; 50 kHz range) USVs. ISO had a sex-selective effect on context-induced USVs that may reflect changes in foot shock sensitivity. ISO increases foot shock sensitivity and tends to enhance context-induced fear in female rats, whereas ISO tends to impair context-induced fear in male rats. Overall, male rats show greater conditioned fear, and female rats show greater unconditioned fear as well as enhanced responses to the aversive and appetitive stimuli. The sex-specific effect of ISO on context fear may reflect neuronal reorganization in stress responsive areas and/or sex differences in some unconditioned responses.

Animals↗
PubMed2004

Early environmental experience alters baseline and ethanol-induced cognitive impulsivity: relationship to forebrain 5-HT1A receptor binding.

The relationship between impulsivity and drug abuse is poorly understood despite evidence that impulsive behaviour both predicts, and is a consequence of, drug use. Moreover, although there are clear individual differences in the propensity to addiction, this relationship has not been investigated with respect to impulsive behaviour. We tested whether early environmental experience would influence behavioural measures of impulsivity, and further, whether this experience would alter impulsive choice following ethanol intoxication. Thirty-six male, Long-Evans rats were reared in either isolated (1 rat/cage), standard (2 rats/cage), or enriched (group housed with toys) conditions. After a 3-month rearing period, animals were tested in two operant tasks measuring either motor (go/no-go) or cognitive (delay-to-reinforcement) impulsivity. Rats were then re-tested following 0, 0.3, 0.6, 0.9, and 1.2 g/kg ethanol. Forebrain 5-HT1A binding was assessed post-mortem using in vitro receptor autoradiography with the agonist [3H]8-OH-DPAT (3H-8-hydroxy-2-[di-n-propylamino]tetralin). Rearing condition did not influence baseline motor impulsivity, but isolation rearing led to decreased baseline cognitive impulsivity. Ethanol did not affect motor impulsivity, but dose-dependently increased impulsive choice in the delay-to-reinforcement task. Enriched rats were more impulsive overall, and isolation-reared rats only showed a shift in impulsive behaviour after 1.2 g/kg. Isolation rearing decreased, and enrichment rearing increased 5-HT1A binding in the frontal pole of the cortex following experience in the delay-to-reinforcement task. Isolation-reared rats also showed a significant decrease in 5-HT1A binding in the dentate gyrus of the ventral hippocampus following experience in the delay-to-reinforcement relative to the go/no-go task. These data indicate that differential rearing has a significant influence on cognitive impulsivity, and that altered serotonergic function may underlie these differences.

Age Factors↗
PubMed2004

Satiety enhancement by selective orexin-1 receptor antagonist SB-334867: influence of test context and profile comparison with CCK-8S.

Acute systemic treatment with the selective orexin-1 (OX1R) antagonist SB-334867 reduces food intake in rats, an effect associated with an acceleration in behavioural satiety and unrelated to gross behavioural disruption, alterations in palatability, or toxicity. However, as enhanced satiety is behaviourally indexed by an earlier-than-normal transition from eating to resting, and since orexin-A has been implicated in mechanisms of arousal, it remains possible that sedation contributes to the anorectic effect of acute OX1R blockade. Previous work has shown that, when treated with SB-334867 (30 mg/kg, i.p.) 30 min before a 1h test with palatable food, rats begin to show appreciable levels of resting 10-15 min earlier than under control conditions (i.e. around 20 min versus 30-35 min into the session). The present results demonstrate that a 20 min increase in the injection-test interval (i.e. 50 min) had no significant impact on the anorectic, behavioural or weight gain effects of SB-334867 in non-deprived male rats. Most importantly, this altered treatment regimen led to a temporal profile of resting virtually identical to that previously observed with the more conventional 30 min injection-test interval. Although parallel studies indicated that the OX1R antagonist accelerated the onset of resting (and suppressed most active behaviours) even in the absence of food, an equianorectic dose of the natural satiety-related signal cholescystokinin octapeptide (CCK-8S; 5 microg/kg, i.p.) also produced very similar behavioural effects regardless of the presence of food. Together with evidence that SB-334867 preserves the structural integrity of natural feeding behaviour, does not induce nausea/illness or alter taste/palatability and fails to influence EEG measures of arousal/sleep, the present findings are consistent with the view that acute OX1R antagonism selectively enhances satiety. However, unlike the immediate short-circuiting of the satiety sequence induced by CCK-8S, the slower response to SB-334867 implies a more indirect mechanism of action.

Analysis of Variance↗
PubMed2005

Electrolytic lesions to nucleus accumbens core and shell have dissociable effects on conditioning to discrete and contextual cues in aversive and appetitive procedures respectively.

The nucleus accumbens (n. acc.) has been implicated in conditioning to both discrete and contextual cues but its precise role is as yet controversial because conflicting patterns of effect have been reported. These inconsistencies may relate to the extent to which the lesions used encroach on different subfields of n. acc. and the use of different task variants. The present study compared the effects of selective lesions of shell and core subfields of nucleus accumbens (n. acc.) across aversive and appetitive trace conditioning variants. In both experiments, an auditory stimulus was contiguous with footshock or food, or presented at a trace interval. A continuous flashing light in each case provided an experimental background stimulus. Conditioning to the cues provided by the experimental chambers was also assessed. Rats with electrolytic lesions to the n. acc. shell and core showed different patterns of effect in aversive (Experiment 1) and appetitive (Experiment 2) variants of this procedure. In Experiment 1, the core lesion reduced the difference between trace and contiguously conditioned groups, in responding to the discrete noise stimulus. However, neither lesion had any detectable effect on contextual conditioning. In Experiment 2, the shell lesion clearly increased contextual conditioning, selectively in the trace conditioned group, but neither lesion had any detectable effect on discrete cue conditioning. Thus, whilst the shell and core lesions produced dissociable effects on discrete cue and contextual conditioning, the conclusions to be drawn depend on the procedural variant in use.

Animals↗
PubMed2005

Announced rewards counteract the impairment of anticipatory behaviour in socially stressed rats.

It is known that stress can influence the sensitivity to rewarding stimuli. Previous observations revealed that socially stressed rats do not display an appetitive behavioural response in anticipation of a reward. A previous study showed that this insensitivity to rewards (anhedonia) could be restored by chronic administration of an antidepressant. Several lines of evidence exist for the role of dopamine in the mechanism of action of antidepressant treatments concerning their therapeutic effect on anhedonia. Therefore, it was hypothesized that regular activation of the reward system, that involves mesolimbic dopaminergic systems, could counteract the effect of social stress on reward-sensitivity. For this, it was investigated whether a treatment of regular reward announcements could prevent the development of anhedonia. This was confirmed by the result that socially stressed rats that received this treatment were able to display anticipatory behaviour which is characterized by increased activity after presentation of a stimulus that was previously associated with a sucrose reward. Surprisingly, a non-treated socially stressed group, that did not show an anticipatory response for sucrose, did display anticipatory behaviour for another type of reward (enriched cage). It seems that, although one might assume the existence of an anhedonic state based upon the absence of anticipatory activity towards a sucrose reward, this assumption cannot be generalised to other types of reward. It will be discussed whether this might be caused by the highly rewarding properties of the enriched cage which probably has a therapeutical efficacy of its own.

Analysis of Variance↗
PubMed2005

Hemispheric specialization and dual processing in strongly versus weakly lateralized chicks.

Lateralization of various functions is well established in the domestic chick and other vertebrates. The function of lateralization is a question under investigation now. In a previous study, it was shown that chicks incubated in the dark (Da), which prevents the development of lateralization of visual foraging and predator detection, did not perform well when two tasks requiring simultaneous use of the expertise of the right and left eye systems were presented. Performance of a task requiring the chick to find grains against a background of pebbles deteriorated in the Da chicks in the presence of a predator. In the study reported here, the two simultaneous tasks were tested in both binocular and monocular conditions to establish the role of each eye system. Learning of the pebble-floor task was poor in Da chicks tested binocularly or monocularly and in the light-exposed (Li) chicks using the left eye (LE). The time taken to complete the pebble-floor task in the presence of a predator was significantly longer for the Da chicks. This was because pecking was interrupted for longer times when the predator was presented, and the Da chicks made more distress calls than the Li. The latency to detect the predator was longest in the Li chicks using the right eye (RE) (i.e., the Li were lateralized). In the Li chicks, sustained initial viewing of the predator was by the LE. There was no LE/RE difference in the Da chicks. The intensity of responses decreased with continued presentation of the predator but forced use of the RE slowed this habituation. Exposure to light before hatching affects lateralization of both learning of the pebble-floor task and the detection of an overhead predator. Attending to the foraging and predator vigilance tasks simultaneously is impaired in the Da chicks and the superior ability of Li (lateralized) chicks is due to a specific effect of light stimulation of the RE prior to hatching. We have shown that superior performance on the dual task results from the ability to allocate food searching to one hemisphere (left) and predator vigilance to the other (right) hemisphere, achieved only by Li chicks.

Animals↗
PubMed2005

Alteration and recovery of appetitive behaviour following nerve section in the starfish Asterias rubens.

The starfish Asterias rubens is an invertebrate deuterostome whose nervous system shows remarkable regenerative properties. To understand when full functionality of a damaged part of the nervous system recovers, and to follow nerve regeneration in detail, we carried out behavioural experiments with 29 starfishes that had the nerve in one of the arms sectioned in a mid-arm position. Loss and recovery of normal behaviour was followed by video analysis of animal performance in an appetitive behavioural test. When compared to 13 control (unoperated) animals, the appetitive response of freshly sectioned animals is normal initially, progressively deteriorates up to 40 days after the lesion, and then gradually improves until 60 days, when recovery is complete. This is true only when one of the leading arms in the appetitive test is a sectioned arm; turning the starfish so that both the leading arms facing the prey are unlesioned, results in normal behaviour even at 40 days after the cut. Thus, regeneration is a multi-step process whose time course coincides with anatomical regeneration. At intermediate times the animals have coordination problems in an appetitive behaviour test and these give some insights into how arms may inter-communicate to organize concerted movements.

Animals↗
PubMed2005

Habituation and dishabituation of exploratory and appetitive responses in the honey bee (Apis mellifera L.).

Sucrose-induced antennal and proboscis motor responses of the honey bee were comparatively studied using muscle responses. The activity of the proboscis muscle responses was correlated with sucrose stimulus concentration, but that of antennal muscle responses was not. Both responses habituated following repetitive stimulation with a low sucrose concentration and dishabituated upon stimulation with a high sucrose concentration. Despite the different response properties of the responses, their habituation kinetics were indistinguishable. The absence of sensory adaptation was confirmed by taste hair recordings. The potential for habituation was related to the functions of the responses studied. The exploratory antennal response used for continuous sampling of the environment habituated to a smaller extent than the appetitive proboscis response.

Adaptation, Physiological↗
PubMed2005

Amphetamine-induced 50 kHz calls from rat nucleus accumbens: a quantitative mapping study and acoustic analysis.

Emission of 50 kHz ultrasonic calls in rats is known to be associated with appetitive behavioural situations and positive social interactions. The purpose of the study was to pharmacologically characterize amphetamine-induced 50 kHz calls and to perform quantitative mapping of this response in the nucleus accumbens. Injections of amphetamine into the nucleus accumbens induced species-typical 50 kHz calls in adult rats. The acoustic parameters of the calls were not affected by different amphetamine doses or combination of agents. The increase in the number of calls occurred predominantly from the accumbens shell and to a lesser degree from the core region. This effect was dose-dependent within the range of 1-20 microg of amphetamine and was reversed by pretreatment with D1 or D2 dopamine antagonists (SKF-83566 or raclopride) administered to the same brain site. However, another D2 dopamine receptor antagonist, haloperidol, which is known to increase the accumbens dopamine level, was ineffective in reversing the increase in call number at the dose studied. On the contrary, intraacumbens haloperidol, when injected alone, caused an increase in 50 kHz calls. It is concluded that the release of dopamine, predominantly in the accumbens shell region, is responsible for production of 50 kHz calls and the calls may indicate an appetitive state compatible with anticipation of reward and positive affect. Both D1 and D2 subtypes of dopamine receptors may be necessary to induce 50 kHz calls and signal the appetitive state.

Amphetamine↗
PubMed2005

Competition for sucrose pellets in tetrads of male Wistar, Fischer or Sprague-Dawley rats: is intra-group ranking reflected in the level of anxiety?

Competition for palatable food or fluids within groups of rats has been previously used to mirror intra-group ranking. The paradigm of competition for sucrose pellets in non-food-deprived male Wistar rats was here extended from triads to tetrads aiming at evaluating whether the number of poor-performing rats, those animals being likely to model aspects of human psychopathologies (anxiety/depression/social withdrawal), could be increased. To evaluate potential superiority over the previously used Wistar strain, establishment and stability of the ranking was also assessed in tetrads of male Fischer and Sprague-Dawley rats. Clear and stable rank orders were seen in around 60-70% of both triads and tetrads of Wistar rats: a high-performing, a medium-performing and one (in triads) or two (in tetrads) poor-performing rats were identifiable, indicating that the number of poor-performing rats had increased in tetrads. Comparable rank orders were also seen in tetrads of Fischer and Sprague-Dawley rats. At the end of an extended period of repeated testing, tetrads of these two strains, as well as some selected Wistar tetrads, were tested in the elevated zero-maze and plasma corticosterone levels were determined. The differentiation in competition-performance among cage mates was not paralleled by a difference in performance in the elevated zero-maze or in plasma corticosterone levels in any of the three strains. These data indicate that the level of anxiety in a non-social paradigm, the elevated zero-maze, does not reflect the competition-performance within the home cage and thus, the dominant/subordinate status in this food-competition paradigm may not reflect/being caused by different levels of anxiety.

Analysis of Variance↗
PubMed2006

Excitotoxic lesions of the gustatory thalamus eliminate consummatory but not instrumental successive negative contrast in rats.

Lesions of the gustatory thalamus (GT) prevent the occurrence of between-session contrast effects (i.e., anticipatory negative contrast and consummatory successive negative contrast [cSNC]) involving liquid rewards. These deficits are attributed to a disruption of the reward comparison mechanism that computes the value of the current reward relative to the expected reward. Instrumental successive negative contrast (iSNC), which occurs following a surprising reduction in the magnitude of a solid food reward, is also keyed off the detection of a disparity between the value of the expected and actual rewards. The present study examined whether neurotoxic lesions of the GT prevent the occurrence of iSNC just as they abolish cSNC. In Experiment 1, both GT-lesioned and neurologically intact rats trained with 10 food pellets and shifted to one food pellet showed a significant iSNC effect. In Experiment 2, the same GT-lesioned rats failed to show cSNC when shifted from 1.0M sucrose to 0.15% sodium saccharin. The obtained pattern of results suggests that separate reward comparison mechanisms with different neural substrates underlie expression of cSNC and iSNC.

Animals↗
PubMed2006

Defensive responses of Wistar and Sprague-Dawley rats to cat odour and TMT.

Cat odour and trimethylthiazoline (TMT) are two predator odours commonly used to study defensive behaviour in rats. However their reported efficacy varies markedly across laboratories. We assessed whether rat strain differences might explain such variation. Wistar and Sprague-Dawley rats were tested for unconditioned and conditioned responses to both odours. Cat odour produced robust unconditioned and conditioned defensive behaviour, with notably stronger effects in Wistar rats. TMT produced limited unconditioned avoidance, but failed to elicit conditioned responses in either strain. Results support suggestions that faeces-derived odours such as TMT are less predictive of a predator threat than those derived from fur or skin, and identify the possibility that strain differences affect the defensive response seen to predator odours.

Analysis of Variance↗
PubMed2006

The mGluR2/3 agonist LY379268 attenuates context- and discrete cue-induced reinstatement of sucrose seeking but not sucrose self-administration in rats.

We recently reported that systemic injections of the mGluR(2/3) agonist LY379268, which decreases evoked glutamate release, attenuate context- and discrete cue-induced reinstatement of heroin seeking, but not heroin self-administration. Here, we report that systemic injections of LY379268 attenuate context- and discrete cue-induced reinstatement of sucrose seeking, but not sucrose self-administration. These results suggest that similar neuronal mechanisms mediate cue-induced relapse to opiate- and palatable food seeking.

Amino Acids↗
PubMed2006

Appetitive sensitization by amphetamine does not reduce its ability to produce conditioned taste aversion to saccharin.

Previous exposure to amphetamine attenuates its ability to induce conditioned taste aversion (CTA). Because amphetamine, unlike emetic agents like LiCl, possesses appetitive properties that sensitize when it is administered repeatedly, the present study assessed the contribution of sensitization to this US-pre-exposure effect (US-PEE). It was found that not all sensitizing regimens of systemic amphetamine injections produce a US-PEE. In addition, previous exposure to amphetamine in the VTA, where it acts to induce sensitization but not CTA, did not produce a US-PEE. It is concluded that amphetamine sensitization alone does not modulate this drug's ability to produce CTA. Implications of these findings for anatomically based associative and non-associative models of CTA and the US-PEE are discussed.

Amphetamine↗
PubMed2006

Light experience and the development of behavioural lateralization in chicks III. Learning to distinguish pebbles from grains.

Chicks searching for food grains against a background of unfamiliar pebbles usually peck pebbles less when using the right eye (RE), or both eyes, than when using the left eye (LE), provided that the embryo's RE has been exposed to light (Li), as is normal. When pecking is fast this right/left difference is mainly due to a heightened ability of RE chicks to inhibit premature pecks (and inappropriate responses in general). Dark incubation (Da) abolishes this ability in RE chicks, and RE and LE chicks show similar frequent pebble pecks. We show now that, under conditions that cause cautious pecking, both Li and Da chicks show a new effect: in both cases LE chicks peck pebbles more than RE chicks, probably because of the novelty of pebbles. Interest in novelty in LE chicks is known to be unaffected by light in incubation. Age-dependent effects are also important. RE and LE chicks, which had either the LE or RE exposed to light before hatching, were tested on days 3, 5, 8 or 12 post-hatching, under conditions giving normal fast pecking. Artificial exposure of the embryo's LE to light reversed the lateralization: in general, chicks using the light-exposed eye performed well at all ages. Irrespective of which eye system had heightened ability to inhibit pebble pecks, RE performance differed from usual on 2 days, whereas LE chicks showed no age-dependent changes. Changes confined to the RE system, therefore, affect behaviour independently of lateralization of the ability to inhibit inappropriate response.

Age Factors↗
PubMed2004

Excitatory backward conditioning in an appetitive conditioned reinforcement preparation with rats.

Four experiments were conducted to examine appetitive backward conditioning in a conditioned reinforcement preparation. In all experiments, off-line classical conditioning was conducted following lever-press training on two levers. Presentations of a sucrose solution by a liquid dipper served as an unconditioned stimulus (US) and two auditory stimuli served as conditioned stimuli (CSs); one was paired with the US in either a forward (Experiment 1a) or a backward (Experiments 1b, 2, and 3) relationship, and the other served as a control CS, which was not paired with the US. In testing, each lever-press response produced a presentation of one of the CSs instead of appetitive reinforcers. The response to a lever was facilitated, compared to the response to another lever, when the response produced the backward CS presentation as well as when it produced the forward CS presentation; that is, the backward CS served as an excitatory conditioned reinforcer.

Animals↗