The choose-short effect in rat memory for event duration: the subjective-shortening model.

Two predictions derived from the subjective-shortening model were tested in rats. The predictions concerned the temporary occurrence of the choose-short effect with extended training at a given retention interval (RI) and the occurrence of a temporary choose-long effect, when RIs shorter than those used during training were applied. In a first experiment, using a stepwise delay procedure with training 0-s RI sessions interpolated between each series of increasing RIs, results showed: (1) a choose-short effect during the stepwise increase in the delay procedure, (2) a temporary occurrence of the choose-short effect during testing at a given RI and (3) a choose-long effect in half of the animals, when a RI shorter than that used previously was applied. These contrasting results suggest that the disappearance of the choose-short effect could be, as proposed by the model, either the consequence of the foreshortening of the reference memory (for rats choosing-long) or the consequence of an adaptation of the working memory (for rats which did not choose long). Results were discussed in relation with the procedure which could have contributed, by the interposition of 0-s RI sessions, to maintain a stable reference memory. In order to test this interpretation, a second experiment, using the classical stepwise delay procedure without training sessions interpolated, was carried out. In these conditions, rats did never present a significant choose-long effect when the RI was shortened. These results suggest that rats maintained a stable reference memory and could improve their performances during retention testing sessions either by an adaptation of their working memory or by the adoption of an alternative strategy which consisted in learning to maintain an orientation towards the location of the correct lever.

Effects of age on short-term memory for time in rats.

Working memory (WM) for event duration was investigated in four different age groups of rats (6, 12, 18, and 24 months), using a delayed, symbolic-matching-to-sample procedure. In an initial experiment, all age groups responded as though a long sample duration were a short one when retention intervals (RI) were applied. This effect is called the choose-short effect. In a second experiment, in order to test whether, in compliance with the subjective-shortening model, there would be a "shortening" of the sample duration according to the length of the RI, the psychophysical function relating the probability of choosing the long comparison stimulus to sample duration and the point of subjective equality (PSE) were determined across successive RIs in two age groups (6 and 18 months). A significant shift of the psychophysical function towards a longer duration and a significant increase of the PSE appeared as a function of the RI in 6-month-old, but not in 18-month-old rats. These results support the idea that the choose-short effect is due to a shortening of the event duration in the WM in young animals, whereas there is no evidence that the sample forgetting occurred on a time dimension in older rats.

Behavioural and glial changes in old rats following environmental enrichment.

The effects of enriched environment on short-term memory for event durations and on astrocytes (cell density, cell area and % of GFAP immunoreactivity) in hippocampus (Hi), frontal cortex (FC) and corpus callosum (CC) were analysed in old rats housed from weaning to the end of behavioural testing (23 months) either in standard (SC) or in enriched (EC) conditions and in young adults (5 months) all housed in SC. Old SC and EC and young SC rats trained (for 2 months) or not, in a Symbolic Delayed Matching to Sample Task, had to discriminate and remember two (2- and 10-s) signals after short retention intervals. Results confirm the aging-related acquisition and memory deficit. EC reduced the slowness of acquisition, reversed the short-term memory deficit and promoted the retention of the short signal (choose short effect). Old SC naive rats had many hypertrophied astrocytes with long processes in Hi and CC while old EC rats had decreased astrocytes number and size. The behavioural testing resulted in young adult SC rats in Hi and CC, in increased astrocytes number, size and GFAP% and in their decrease in old SC rats. EC and testing have additive effects (very low astrocytes number, size and GFAP%) to compensate for the aging-induced gliosis, mostly in Hi.

Peak procedure performance in young adult and aged rats: acquisition and adaptation to a changing temporal criterion.

Twenty-four-month-old and 4-month-old rats were trained on a peak-interval procedure, where the time of reinforcement was varied twice between 20 and 40 sec. Peak times from the old rats were consistently longer than the reinforcement time, whereas those from younger animals tracked the 20- and 40-sec durations more closely. Different measures of performance suggested that the old rats were either (1) systematically misremembering the time of reinforcement or (2) using an internal clock with a substantially greater latency to start and stop timing than the younger animals. Old rats also adjusted more slowly to the first transition from 20 to 40 sec than did the younger ones, but not to later transitions. Correlations between measures derived from within-trial patterns of responding conformed in general to detailed predictions derived from scalar expectancy theory. However, some correlation values more closely resembled those derived from a study of peak-interval performance in humans and a theoretical model developed by Cheng and Westwood (1993), than those obtained in previous work with animals, for reasons that are at present unclear.

Effects of aluminum exposure on behavioral parameters in the rat.

Adult rats were treated by intraperitoneal injection of aluminum gluconate for 3 months. Rats were submitted to the radial maze test to determine the influence of chronic aluminum intoxication on cognitive and noncognitive behavioral processes. Both learning abilities (working memory and reference memory) and rapidity (time spent to respond and to master a trial) were analyzed. Aluminum concentration was evaluated in the brain, serum, and liver to assess aluminum body burden. While hippocampus and neocortex showed a significant increase in aluminum concentration, aluminum treatment did never affect the animal's performance during cue learning or when the insert cues were removed. The only behavioral difference observed was a decrease in rapidity: both the total time to finish a trial and the latency to make the first choice were lengthened in aluminum-intoxicated rats.

Age-related differences for duration discrimination in rats.

The effects of age on duration discrimination were analysed with a symbolic matching-to-sample task, where a compound signal (light and sound) was presented at each trial for a duration of either 2 or 10 s. Four groups of rats (6, 12, 18, and 24 months old) were trained to press one lever if the signal was short and the other if it was long. Results show that, in comparison with younger rats (6 and 12 months), presenescent and senescent rats (18 and 24 months) were slower to reach the acquisition criterion. However, when the performance criterion was met, no age-related difference was found: the percentages of correct responses were equivalent, whatever the duration of the stimulus. These results are in accordance with other data, which have often shown that the cognitive impairment reported in old animals results more from a slowness to learn than from an incapacity to discriminate between different durations.

Differential environmental modulations on locomotor activity, exploration and spatial behaviour in young and old rats.

The effects of environmental enrichment on motor activity, exploration and spatial performances were studied in young and old rats. Both young (4 mo old) and old (22 mo old) rats were housed from weaning to testing either in standard or in enriched conditions. All rats were submitted successively to spontaneous alternation test and to object exploration test. Results show that locomotion is decreased by age and enrichment but that the quality of exploration expressed by corrected alternation scores or by the response to spatial change is improved by enrichment sometimes in old, sometimes in young rats. Enrichment tends to accelerate the acquisition of spatial informations in young rats, but it does not succeed to restore the reactivity to spatial change of old rats in the object exploration test. These results, although they do not rule out a persistance of a continued behavioural plasticity during aging, also support the idea that the beneficial effects of environmental stimulations do not succeed to restore high cognitive function, such as the capacity to have a spatial representation, in old animals.

Cognitive and noncognitive processes involved in selective object exploration: comparison between young adult and old rats.

The age effects on locomotor activity, object-oriented exploration, habituation, and response to a spatial change were studied in young adult and old rats using an object exploration test. In this test the spatial response was evaluated by the renewal of exploration of a familiar object after its repositioning. The specificity of the spatial response was determined by comparison with control animals not submitted to a spatial change. Male Wistar rats 6 and 24 months old were used. Results showed a significant decrement in locomotor activity, object exploration, and spatial reactivity in old rats. The habituation curve and the reactivity to a new object were preserved. Detail analyses suggest that the spatial deficit of old rats is due to an incapacity to detect the spatial change and not to their poor locomotor or exploratory activity. These results corroborate those obtained in spatial orientation tasks and support the idea that the lack of spatial response observed in old animals is more related to cognitive impairments than to other factors such as sensory, motor, or motivational differences.

Cholinergic blockade and response timing in rats.

The effects of central cholinergic blockade on the temporal regulation of behaviour were studied with a two-level DRL schedule. Five-month-old male Wistar rats had to press lever A and then wait for a minimum of 5 s before pressing lever B to obtain the reinforcer (sweetened milk). After a stable baseline performance, subjects were injected in random order with the general cholinergic blocker, scopolamine, 0.15 and 0.5 mg/kg, the peripheral cholinergic blocker, methylscopolamine, 0.15 and 0.5 mg/kg, and a combination of the cholinesterase inhibitor, physostigmine, 0.2 mg/kg, and scopolamine, 0.5 mg/kg. Each drug treatment was separated by 2 days of saline treatment. Results showed that scopolamine at 0.5 mg/kg significantly impaired the temporal regulation of the A-B response sequence: the median A-B inter-response time (IRT) was shortened and the coefficient of variation of the A-B IRT distribution was increased, thus revealing a loss in the sensitivity to time. This disruption of accurate timing behaviour lowered efficiency. The drug changed neither the duration of the B-A interval nor the A-B response rate, but significantly increased the rate of the superfluous B-B sequences. Methylscopolamine was without effects and physostigmine totally or partially reversed all the scopolamine effects. These results suggest that scopolamine at 0.5 mg/kg specifically affected the mechanism(s) underlying response timing, and that the effects were not secondary to changes in activity or motivation. They partly corroborate data obtained in other procedures and support the idea that the central cholinergic system is involved in the temporal regulation of behaviour.

Acquisition and long-term retention of a two-lever DRL schedule: comparison between mature and aged rats.

The effects of ageing on temporal regulation, general activity and memory were analysed in a two-lever DRL schedule. The task consisted in pressing on lever A and then waiting a minimum of time before pressing lever B to get the reinforcer. Adult and senescent rats were submitted to preliminary training followed by 5 DRL 5-second training sessions and 3 retention testing sessions after a 21-day break. Results showed that, relative to adults, senescent rats were slower to reach the 5-second DRL criterion, emitted fewer temporally regulated A-B response sequences and an equivalent amount of repetitive superfluous A-A and B-B response sequences. The quality of temporal regulation was evaluated by the coefficient of variation (CV) and the median of the A-B interresponse-time distribution. In training, aged rats exhibited a higher CV only during the first 10-minute periods of the sessions, and emitted median IRTs similar to those of adults. The B-A intertrial-intervals were longer in aged than in adult rats. No age-related differences appeared for efficiency. Finally, long-term retention was not affected in either age group. The results favour an interpretation in terms of temporary recall memory deficit with a preservation of temporal regulation capacity, rather than age-related motor and motivational differences.

Difference in scopolamine sensitivity during ageing in rats: dichotomy between behavioural repertoire and learning performances.

The effects of different doses of scopolamine in adult and aged rats were compared in two behavioural experiments. The first experiment involved visual reversal learning, while the second experiment investigated individual and social behavioural patterns. Results showed that the same dose of scopolamine was more effective in impairing the reverse discrimination in aged than in adult rats. In contrast, low doses which were effective in modifying behavioural patterns in adults, were inactive in aged rats. This difference in scopolamine sensitivity (supersensitivity versus hyposensitivity), observed between the two types of behaviour, is discussed in relation to age-related functional receptor adaptation, selective changes in muscarinic receptors in the different brain areas and dopaminergic- cholinergic interactions.

Age-related scopolamine effects on social and individual behaviour in rats.

The modulation of spontaneous (social and individual) behaviour as a function of the age of the rat (1, 3, 6, 12, 18 and 24 months) and of scopolamine dose (0.1, 0.2, 0.3 and 0.5 mg/kg) was studied. Observations were conducted during the dark phase of the reverse light/dark schedule using a reintroduction procedure. Results showed a marked effect of scopolamine on most of the behavioural patterns considered. Environmental interaction was enhanced whilst agonistic and social active interactions (social grooming) and play fighting were reduced by the drug. A slight hyposensitivity in the youngest rats and a marked hyposensitivity to the drug in the oldest ones were observed. The relationship to biochemical data and human sensitivity on the one hand and to learning and memory tasks and cholinergic specificity on the other hand, are discussed.

Scopolamine effects on juvenile conspecific recognition in rats: Possible interaction with olfactory sensitivity.

The social recognition of a juvenile conspecific by an adult male rat was evaluated as the decrease in investigation time when the same juvenile individual was reintroduced 30min after the first exposure period. The results showed that scopolamine impaired this transient individual recognition. A drop in investigation time was also observed in both tests (first and second exposure), with the same and with different juvenile individuals, in scopolamine treated animals. A second experiment showed that scopolamine disturbed the chemosensory preference for familiar odour observed in the control group. In the light of these two experiments, and according to the chemosensory mediation of social recognition in the rat, it is impossible to rule out a lack of odour discrimination in the absence of social recognition after scopolamine injection.

Scopolamine disrupts visual reversal without affecting the first discrimination.

The effect of scopolamine (0.5 mg/kg) was determined in a brightness discrimination test (Y maze) motivated by electrical shocks (escape avoidance). Male adult Sprague Dawley rats were used. Results show that scopolamine impairs significantly the visual reversal without affecting the first brightness discrimination. The qualitative analysis reveals that the anticholinergic drug-induced deficit involves both perseveration, i.e. failure to suppress inappropriate response, and a tendency to adopt a position habit. A parallelism with hippocampal and frontal lobe damage symptoms is discussed and an interpretation in terms of disinhibition and incapacity to solve a more difficult problem is proposed.

Scopolamine-induced deficits in acquisition of a complex spatial learning.

The role of scopolamine was studied in a complex spatial orientation task. The procedure involved an increasing difficulty of the task: at the pretraining stage a cue (box) was placed at the reinforcement spot and the animal could give a correct response by adopting either a cue-strategy or an orientation response (i.e., go to the arm on the right of a visual landmark). In the subsequent spatial training, the box was removed, so that the orientation response was the only correct one. Results show that scopolamine-injected animals are able to use a cue-strategy but are unable to acquire a spatial orientation strategy: this more complex task asks for more time and for more sustained attention. When the response is already partly acquired, scopolamine has less effect. The cholinergic system would thus be involved in the quality or even the complexity of the response rather than in the retention itself. Though a state-dependent effect may not be excluded, it by itself cannot explain the observed differences. Finally, an impairment of the maintenance of attention could be responsible for the deficits observed in the acquisition of the complex task.

Responsiveness to "social" releasers in cattle. II. Relation between social status and responsiveness, and possible effect of previous familiarization with the test conditions.

The data of two previous experiments, investigating the effects of social releasers in Friesian-Dutch heifers were analysed in order to verify whether the frequency of approach responses to models and calls of conspecifics was modified by pre-experimental factors. First, responsiveness was correlated to the social status of individuals; a social hierarchy was assessed on the basis of agonistic interactions recorded for a 2-month period prior to the experiments with models. A significant positive correlation was found between the total number of responses and the total number of escape-avoidance reactions to aggressive acts. The same result was found when individual ranks were assessed on a hierarchy re-presenting a dominance order established on the basis of the number of aggressive acts causing escape-avoidance reactions. In this case, also, animals with highest ranks presented a significantly higher number of responses to models than animals with lowest ranks. In our second experiment, a significant positive correlation was found between all criteria used to assess individual ranks in a hierarchy, while this result was not found necessarily in the first experiment. The other pre-experimental factor under consideration was a possible familiarization with the testing conditions (models/calls of conspecifics, handling, and other procedures); some of the group-members tested during the second experiment had already been tested in the first one, carried out a year before. Many results demonstrated that the frequency of responses to social releasers was significantly lower for the previously tested subjects than for the animals tested for the first time.

Responsiveness to "social" releasers in cattle. I. A study of the differential and additive effects of visual and sound stimuli, with special reference to the law of heterogeneous summation.

Two experiments (Exp.1, Exp.2) were carried out to investigate the effects of social releasers, i.e. models and calls of conspecifics, on the approach responses of Friesian-Dutch heifers. In Exp.1, models consisted of either a vague representation of a conspecific's shape, or of a life-sized photograph of a conspecific. Sound (S) stimulations (calls) and movement (M) were combined to these visual (V) models. In Exp.2, V models and S stimulations were presented either separately or in combination. Models consisted of a ♂ and of a ♀ life-sized photograph and S stimulations consisted of recorded calls of ♂♂ or of ♀♀ conspecifics. In Exp.1, a significant increase in responsiveness was found after addition of sound S to the V models, especially to the photographs. Adding S to V models resulted in a much higher increase in responsiveness than adding M to V models. An additive effect was also found in Exp.2: combined presentations of V and S increased significantly the number of responses and of responding subjects as compared to either V or S separate presentations. In both Exp.1 and Exp.2, the total number of responses was a little more than twice as high for combined presentation of V and S as it was for V separate presentations, or for S separate presentations (Exp.2). In Exp.2, a complete verification of Seitz's law of heterogeneous summation of the stimuli was attempted by a further comparison, demonstrating that the releasing effect of V and S presented in combination (simultaneously) was equivalent to the number of responses found after addition of the partial effects of V and S presented separately (successively).